U.Seeliger C.Odebrecht J.P.

Castello Subtropical Convergence Environments

Springer
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U. Seeliger C.Odebrecht J. P. Castello (Eds.)

Subtropical Convergence
Environments
The Coast and Sea in the Southwestern Atlantic

With 66 Figures

Springer
Dr. ULRICH SEEUGER
Dr. CLARISSEODEBRECHT
Lie. JORGE PABLOCASTELW
Department of Oceanography
Universidade do Rio Grande
96201 Rio Grande, RS
Brazil

ISBN·13: 978·3_642-64418-4 Springer-Verlag Berlin Heidelberg New York

Library of Congreu Cataloging-in-Publication Data

Subtropical convergence environments: the coast and sea in the

southwestern Atlanticl Ulrich Sedig. r, Clari~ Odebrecht, Jorge
P~blo Castello (eds.).
p. cm_
Includes bibliographical references and indeL
IS " I\(_ I3: 971i-.l-64!·6H 18-4 .... IS " I\(· I3: 97Ji...J..6.1!-'O-I67-11
nO I: 10.1007/978-.\-641-'0-167-6

I. Coastal ei:ology--South Atlantic Ocean. 2. Marine ecology-

- South Atlantic Ocean. 3. Marine biology--South Atlantic Ocean.
4. Biological diversity-·South Atlantic Ocean. I. Seeliger, U.
(Ulrich),1944- . II. Odebrecht, Clarisse, 1952·
III. Castello, Jorge Pablo.
QH93.6.S83 1997
574.5'2636'091636·-dc20 96-32225

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Preface

Our understanding of coastal and marine ecosystems, like coasts and seas
themselves, is continuously evolving. Earlier work on these environments
and their biota in the warm-temperate southwestern Atlantic has amassed
useful data, which now need to be analysed as the evaluation, estimation,
and management of these resources depend on comprehending the system
as a whole. The considerable amount of information now dispersed in
scientific papers, reports, and dissertations requires new ways of organiza-
tion. To encompass the pertinent knowledge, to deal with conflicting inter-
ests, to identify research needs, and to meet future demands, an attempt
has been made to synthesize the rapidly expanding database, although this
effort can only be viewed as one stage in the process of understanding these
systems. It is hoped that the book will be useful to many different users,
including other scientists, resource managers, students, and interested
laypersons.
First the book describes the environmental setting and the biota of the
Patos Lagoon estuary, the coastal region, and the continental shelf. Following
sections relate these components in space and time to illustrate functional
aspects within and the integrated character between systems. The last
sections of the book emphasize the impact of fisheries and other human
activities on the coast and sea and suggest socio-economically feasible, multi-
faceted management strategies. An inventory of all marine taxa in the warm-
temperate southwestern Atlantic is added as an appendix.
The supportive funding for the book was provided by the Brazilian
Government (Program for the Support of Scientific and Technological
Development in Environmental Science-PADCT/CIAMB/CNPq/CAPES) and
the World Bank. Weare grateful to all of our colleagues who authored
individual contributions, for extracting the pertinent information from the
enormous body of original literature upon which this book is based, and for
generously sharing their unpublished data. Special thanks go to Drs. Jose
Tundisi and Wolfgang Pfeiffer for their encouragement to edit this book.

Rio Grande, Brazil Ulrich Seeliger

October 1996
Contents

Introduction and Overview

U. SEELIGER and C. ODEBRECHT ....................................... 1

2 Regional Climate
A.H.F. KLEIN .......................................................... 5

3 Coastal Plain and Patos Lagoon

M.L. ASMUS ........................................................... 9

4 Environment and Biota of the Patos Lagoon Estuary............. " 13

4.1 Geological Setting
L.J. CALLIARI ......................................................... 13
4.2 Hydrographic Characteristics
C.A.E. GARCIA ....................................................... 18
4.3 Environmental Chemistry
L.F.NIENCHESKI and M.G. BAUMGARTEN ............................. 20
4.4 Tidal Marsh and Wetland Plants
C.S.B. COSTA ......................................................... 24
4.5 Submersed Spermatophytes
U. SEELIGER .......................................................... 27
4.6 Benthic Macroalgae
U. SEELIGER .......................................................... 30
4.7 Microalgae
e. ODEBRECHT and P.e. ABREU ...................................... 34
4.8 Bacteria and Proto zooplankton
P.C. ABREU and e. ODEBRECHT ...................................... 37
4.9 Zooplankton
M. MaNTO, A.K. DUARTE, and I.M. GLOEDEN ........................ 40
4.10 Benthic Invertebrates
C.E. BEMVENUTI ..................................................... 43
4.11 Physiological Adaptations of Invertebrates and Fish
E.A. SANTOS and A. BIANCHINI. ...................................... 47
X Contents

4.12 Ichthyoplankton
e. SINQUE and J.H. MUELBERT ....................................... 51
4.13 Fish Fauna
J.P. VIEIRA and J.P. CASTELLO ........................................ 56
4.14 Bird Fauna
C.M. VOOREN ........................................................ 62
4.15 Marine Mammals
M.C. PINEDO ......................................................... 63

5 Energy Flow and Habitats in the Patos Lagoon Estuary ............ 65

5.1 Primary Production Cycles
U. SEELIGER, e.S.B. COSTA and P.e. ABREU .......................... 65
5.2 Trophic Structure
C.E. BEMVENUTI ..................................................... 70
5.3 Irregularly Flooded Marginal Marshes
C.S.B. COSTA ......................................................... 73'
5.4 Unvegetated Intertidal Flats and Subtidal Bottoms
C.E. BEMVENUTI ..................................................... 78
5.5 Seagrass Meadows
U. SEELIGER .......................................................... 82
5.6 Rubble Structures and Hard Substrates
R.R. CAPfTOLI ........................................................ 86

6 Coastal and Marine Environments and Their Biota.................. 91

6.1 Geomorphological Setting
L.J. CALLIARI ......................................................... 91
6.2 Physical Oceanography
C.A.E. GARCIA ....................................................... 94
6.3 Chemical Characteristics
L.F. NIENCHESKI and G. FILLMANN ................................... 96
6.4 Coastal Foredune Flora
U. SEELIGER .......................................................... 98
6.5 Coastal Foredune Fauna
N.M. GIANUCA ...................................................... 102
6.6 Bacterioplankton
P.C. ABREU .......................................................... 104
6.7 Phytoplankton
C. ODEBRECHT and V.M.T. GARCIA ................................. 105
6.8 Protozooplankton
C. ODEBRECHT ...................................................... 109
6.9 Zooplankton
M. MONTO, I.M. GLOEDEN, A.K. DUARTE, and e. RES GALLA Jr ...... 110
Contents XI

6.10 Benthic Beach Invertebrates

N.M. GIANUCA ...................................................... 114
6.11 Continental Shelf Benthos
R.R. CAPITOLI ...................................... ; ................ 117
6.12 Ichthyoplankton
C. SINQUE and J.H. MUELBERT ...................................... 120
6.13 Pelagic Teleosts
J.P. CASTELLO ....................................................... 123
6.14 Demersal and Benthic Teleosts
M. HAIMOVICI. ...................................................... 129
6.15 Early Life History of Silverside Fishes
G. PHONLOR and J.C. COUSIN ....................................... 136
6.16 Demersal Elasmobranchs
C.M. VOOREN ....................................................... 141
6.17 Cephalopods
M. HAIMOVICI. ...................................................... 146
6.18 Marine Mammals and Turtles
M.C. PINEDO ........................................................ 150
6.19 Sea and Shore Birds
C.M. VOOREN ....................................................... 154
6.20 Genetic-Biochemical Differentiation among Marine Organisms
J.A. LEVy •.••....•••......•••••••..••••..••••.•.••••••••••••••.••••.• 159

7 Relationships and Function of Coastal and Marine Environments 161

7.1 The Foredune System
U. SEELIGER ......................................................... 161
7.2 The Beach and Surf Zone
V.M.T. GARCIA and N.M. GIANUCA................................. 166
7.3 The Continental Shelf and Slope
J.P. CASTELLO, M. HAIMOVICI, e. ODEBRECHT, and e.M. VOOREN .. 171

8 Estuarine-Marine Interactions
P.e. ABREU and J.P. CASTELLO ...................................... 179

9 Fisheries
M. HAIMOVICI, J.P. CASTELLO, and C.M. VOOREN ................... 183

10 Natural and Human Impact

U. SEELIGER and e.S.B. COSTA ...................................... 197

11 Management Considerations
M.L.AsMUS and P.R.A. TAGLIANI ................................... 205
XII Contents

Appendix .................................................................. 209

References................................................................. 259

Subject Index .............................................................. 295

List of Contributors

Abreu, P.C., Dept. of Oceanography, Universidade do Rio Grande,

96201-900 Rio Grande, RS, Brazil
Asmus, M.L., Dept. of Oceanography, Universidade do Rio Grande
Baumgarten, M.G., Dept. of Chemistry, Universidade do Rio Grande
Bemvenuti, C.E., Dept. of Oceanography, Universidade do Rio Grande
Bianchini, A., Dept. of Physiological Sciences, Universidacle do Rio Grande
Calliari, 1.J., Dept. of Geological Sciences, Universidade do Rio Grande
Capitoli, R.R., Dept. of Oceanography, Universidade do Rio Grande
Castello, J.P., Dept. of Oceanography, Universidade do Rio Grande
Costa, C.S.B., Dept. of Oceanography, Universidade do Rio Grande
Cousin, J.C.B., Dept. of Morpho-Biological Sciences, Universidade do Rio
Grande
Duarte, A.K., Dept. of Oceanography, Universidade do Rio Grande
Fillmann, G., Dept. of Oceanography, Universidade do Rio Grande
Garcia, C.A.E., Dept. of Physics, Universidade do Rio Grande
Garcia, V.M.T., Dept. of Oceanography, Universidade do Rio Grande
Gianuca, N.M., Dept. of Oceanography, Universidade do Rio Grande
Gloeden, I.M., Dept. of Oceanography, Universidade do Rio Grande
Haimovici, M., Dept. of Oceanography, Universidade do Rio Grande
Levy, J.A., Dept. of Chemistry, Universidade do Rio Grande
Montu, M., Dept. of Oceanography, Universidade do Rio Grande
Muelbert, J.H., Dept. of Oceanography, Universidade do Rio Grande
Niencheski, L.F., Dept. of Chemistry, Universidade do Rio Grande
Odebrecht, C., Dept. of Oceanography, Universidade do Rio Grande
Phonlor, G., Dept. of Oceanography, Universidade do Rio Grande
XIV List of Contributors

Pinedo, M.C., Dept. of Oceanography, Universidade do Rio Grande

Santos, E.A., Dept. of Physiological Sciences, Universidade do Rio Grande
Seeliger, U., Dept. of Oceanography, Universidade do Rio Grande
Sinque, C., Dept. of Oceanography, Universidade do Rio Grande
Tagliani, P.R.A., Dept. of Oceanography, Universidade do Rio Grande
Vieira, J.P., Dept. of Oceanography, Universidade do Rio Grande
Vooren, C.M., Dept. of Oceanography, Universidade do Rio Grande

Klein, A.H.F., Faculty of Marine Sciences, Universidade do Vale do Itajaf,

88302-202 Itajaf, SC, Brazil
Resgalla, Jr.C., Faculty of Marine Sciences, Universidade do Vale do Itajaf
1 Introduction and Overview
U. Seeliger and C. Odebrecht

The South Atlantic between Africa and South America is under the domain
of the high pressure centre of the Atlantic anticyclone which controls the
climate and determines the large-scale oceanographic circulation. The
northward flowing eastern boundary Benguela Current, the South Equato-
rial Current, and the western boundary currents, represented by the
southward and northward flowing Brazil and Malvinas Currents, respec-
tively, are the main currents of the anticyclonic subtropjcal gyre between 15
and 40° S, with the centre at surface between 25 and 30° S(Reid 1989). The
western boundary currents converge between approximately 32 and 40° S
(Castello and Moller 1977) and are forced offshore, giving rise to the Sub-
tropical Convergence in the southwestern Atlantic (Fig. l.1). Western
boundary currents also receive seasonally large continental freshwater
runoff (Castello and Moller 1977; Hubold 1980a; Ciotti et al. 1995) from the
La Plata River drainage basin (3 170000 km2; Urien 1972) and, through the
Patos Lagoon estuary, from the Patos-Mirim Lagoon watershed (201626 km2;
Herz 1977).
The northern reaches of the Subtropical Convergence directly influence
about 700 km of southwestern Atlantic coast, comprising approximately
100000 km2 of southern Brazilian inshore, nearshore, continental shelf, and
slope regions between Sta. Marta Grande Cape, Brazil (28° 40' S) and Uruguay
(34° 40' S) (Fig. 1.1). The variability of the western boundary current circu-
lation, the position of the Subtropical Convergence, and the volume of conti-
nental runoff are the principal forcing factors in this area. Together, they
provide the foundation for physical and chemical characteristics which
determine the biological structure and ecological processes. In consequence,
shelf and slope environments are not dissociated from tropical and sub-
antarctic waters nor do they function in isolation from the La Plata River, the
huge Patos Lagoon and its estuary, or the fringing shorelines; thus, the coast
and sea are integrated and interdependent. Interactions between the
oligotrophic Brazil Current, the nutrient-rich waters of the Malvinas Current,
and continental runoff make the region an important nursery and feeding area as
well as r~production grounds for fishery stocks of subtropical and antarctic origin
which utilize the Brazil and Malvinas Currents for long distance transport.
These interactions are also responsible for the high biological production of the
coastal zone between the coastal plain and the continental shelf break, which is
clearly demonstrated by satellite images (Longhurst et al. 1995).

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
 2 U. Seeliger and C. Odebrecht

29' 100 km

30·

URUGUAY

36'

ARGENTINA

Fig. 1.1. Location of warm-temperate in- and offshore regions under the influence of
the Subtropical Convergence in the southwestern Atlantic. Brazil (BC), Malvinas (MC),
Benguela (BGC), and South Equatorial (SEC) Current

On a global scale, coastal zones are affected by climate and sea-level

changes, whereas exploitation by man leads to a gradual decline of
resources at a regional level. Initially, the plentiful resources of the south-
western Atlantic coast and sea were exploited by indigenous human popu-
lations and early European settlers without harm to the environment. Since
it was first colonized the coastal zone has served as a natural channel of
export, promoting the establishment of human settlements and industrial
development, and the considerable resource potential has invited exploita-
tion by national and foreign fishing fleets. During the last few decades,
Introduction and Overview 3

accelerated demographic growth and increasing economic demands have

caused a decline of the seemingly limitless natural resources, and human
activities might be on the verge of irreversibly changing the nature of the
coast and sea in this region. Unfortunately, regional impacts tend to accu-
mulate and eventually might become significant on a global scale. Studies
of large marine ecosystems point to a global depletion of regional seas
(Sherman et al. 1993).
The basis required for predicting the role of natural and man-mediated
processes contributing to changes in the coastal zone is a detailed knowl-
edge of biodiversity. Biodiversity is expressed by the sum of genes, indi-
viduals, species, communities, and ecosystems within a geographically
defined region and is the biological response of an ecosystem to past envi-
ronmental conditions. Tampering with biodiversity might thus severely
curtail the capacity of ecosystems to reply with viable solutions to natural
changes and man's impact. Owing to the great size and relative inaccessi-
bility of the marine ecosystems, answers to the causes of loss and mecha-
nisms of preservation of biodiversity are severely hampered by a lack of
comprehensive surveys of marine taxa and overviews of the processes that
control their diversity. New interdisciplinary approaches are needed,
existing regional inventories of biodiversity should be capitalized on, and
on-site monitoring of marine biodiversity is essential. The first testimony
of marine biodiversity (see Appendix) of the Subtropical Convergence
marine ecosystem in the warm-temperate southwestern Atlantic not only
provides the necessary inventory to help trace changes in biodiversity with
time but also furnishes essential background information for urgent man-
agement actions to support sustainability.
Despite an increasing commitment to solve current coastal management
problems and to reduce the threat of diminishing fishery stocks, a compre-
hensive appraisal of the biological resources, an assessment of their sus-
tainable exploration, and an evaluation of oceanographic processes in the
warm-temperate southwestern Atlantic are lacking in the international
literature. Contrary to the historic situation in other parts of the world,
which suffered consequences of uncontrolled resource exploitation decades
ago, the information now available may still prevent further deterioration
of these marine ecosystems and their resources.
2 Regional Climate
A.H.F. Klein

The climate of coastal regions between latitudes 29 and 34° S is under the
control of the high pressure centre of the Atlantic anticyclone (Monteiro
1968; Nobre et al. 1986). The latitudinal migration of the centre and the
passage of polar frontal systems (Fig. 2.1) in 6-10-day intervals (Hartmann
et al. 1986; Stech 1990) have a seasonally modifying influence on the cli-
mate (Kousky and Ferreira 1981; Fortune and KOusky 1983; Paz 1985;
Vieira and Rangel 1988; Gan 1992). The proximity of the Subtropical
Convergence and the stabUizing influence of the extensive inshore Patos-
Mirim Lagoon system have imprinted warm-temperate characteristics on
the coast and sea (Semenov and Berman 1977), while the distribution
patterns of flora (Coutinho and Seeliger 1986; Cordazzo and Seeliger 1987)
and fauna (Semenov 1978) classify the area as a warm-temperate biogeo-
graphical transition zone.
The marked influence of the Atlantic anticyclone leads to dominant NE
winds (5 m S-I mean velocity) throughout the year, followed by SW winds
(8 m S-I mean velocity) during a passage of a cold front (Stech and
Lorenzetti 1992) which are more common in winter than in summer
(Delaney 1965; Godolphin 1976; Zeltzer 1976; Tommazelli 1993). The
region is subject to an average of 223 days of sunshine and 404 cal cm-2
mean daily solar radiation, with mean monthly radiation of 603 cal cm-2
dati in December and 213 cal cm-2 dati in June. The regional temperature
regime is a function of season and number and intensity of cold front pas-
sa~es (Nobre et al. 1986). Mean annual temperatures vary between 19°C and
17 C in the north and south of the region, while monthly mean low and
high temperatures vary between 13°C and 24°C in July and January, respec-
tively (IBGE 1986). During the winter, some cold front passages may cause
frost in the early morning hours of clear and dry days (Fortune and Kousky
1983; Nobre et al. 1986). Total mean anl'lual precipitation (1200-1500 mm)
may strongly vary from year to year and is principally related to the path
and frequency of cold front passages (Paz 1985; Hartmann et al. 1986;
Nobre et al. 1986; Gan 1992). Mean monthly rainfall is highest during the
winter and spring (June to October), but a second peak may occur in sum-
mer (Castello and Moller 1978), when daily precipitation occasionally sur-
passes 100 mm (Gomes et al. 1987). The summer months are associated with
a seasonal water deficit (Fig. 2.2), although precipitation and evaporation
result in an average annual water surplus of 200-300 mm (IBGE 1986).

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
6 A.H.F. Klein

Fig. 2.1. Barometric pressure chart with approaching polar frontal system

In the southwestern Atlantic, distinct interannual variations in precipitation,

with either a high amount of rainfall or dry periods, seem to be a
consequence of the effect of the El Nino-Southern Oscillation cycle on the
glbbal climate (Nobre et al. 1986; Gan 1992), but the processes involved are
still not well understood. This phenomena directly influences the amount of
continental freshwater runoff and thus the biogeochemical processes in
coastal and marine ecosystems of the Southwest Atlantic (Ciotti et al. 1995).
Regional Climate 7

mm
200

J F M A M J J A s 0 N D

Fig. 2.2. Annual water balance of coastal regions at 32' S

3 Coastal Plain and Patos Lagoon
M.L. Asmus

The Patos Lagoon as a Natural Unit

Wetland and deep-water habitats (sensu Cowardin et al. 1979) of different

types characterize the coastal plain of Brazil's extreme south, though the
Patos-Mirim Lagoon complex is the dominant feature. With a surface area
of 10 227 km2, the Patos Lagoon classifies as the world's largest choked
lagoon (Kjerfve 1986). It stretches in a NE-SW direction from 30° 30'S to 3t
12'S near the city of Rio Grande where the lagoon conneCts to the Atlantic
Ocean. The lagoon receives waters from a 201 626 km2 drainage basin,
either directly from tributaries or through the Sao Gonc;:alo Channel from
the Mirim Lagoon watershed (Fig. 3.1). Elevated precipitation and complex
river flow patterns in the drainage basin result in highly dynamic and
rapidly changing hydrographic processes.
The Patos Lagoon can be divided into five biological units (Fig. 3.2), rep-
resented by Guafba Bay, Tapes Bay, Casamento Lagoon, the central lagoon
body, and the estuary. Each unit is composed of deeper open and shallow
protected waters. The Guaiba River is the major freshwater tributary. River
flow dynamics and water quality fluctuate significantly. A delta is formed at
the lower reaches of the river as a result of sediment deposition when the
river flow energy dissipates upon entering upper Guafba Bay. Both Tapes
Bay and Casamento Lagoon are semi-open water bodies with distinctive
hydrodynamic characteristics. The central open water body of the Patos
Lagoon itself is composed of four elliptic cells which represent approxi-
mately 80% of the lagoon area. The shorelines are dominated by freshwater
marshes and sandy beaches. There are 971 km2 of estuarine area (approxi-
mately 10% of the lagoon) in the south which exchanges waters with the
Atlantic Ocean through a 20 km long and 0.5-3 km wide inlet. Shallow,
protected, marginal bays and a deeper, central, open water body constitute
two ecologically distinct estuarine environments (Bonilha and Asmus
1994).

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
10 M.L. Asmus

Fig. 3.1. The Patos and Mirim Lagoon watersheds

Settlement and Land Use History

Prior to the arrival of the first European colonists, the indigenous Chana and
Tupi-Guarani Indians occupied the coastal plain. The Indians settled along
the lagoon and ocean shores, where they exploited abundant fish, shellfish,
and shrimp resources (Vieira 1983). The modern occupation history of the
coastal plain began with the arrival of politically opposed Portuguese and
Spanish Jesuits in 1605 and 1626, respectively. Following the Treaty of Madrid
in 1750, which ratified the agreement of a status quo after the foundation of
the town of Rio Grande in 1737, the coastal region became the centre of
intense Portuguese colonization efforts. In order to reinforce its diplomatic
achievements and to warrant sovereignty, Portugal occupied the territory in
1747 with settlers from the Azores, who entered through the port of Rio Grande.
Coastal Plain and Patos Lagoon 11

t
50 km

CAMAQUA RIVER '

ATLANTIC OCEAN

Fig. 3.2. The Patos Lagoon with major biological units

Colonization quickly radiated along the littoral towards Porto Alegre in the north
and Chui in the south (Fig. 1.1). The new settlers were endowed with land for
cultivation and cattle breeding, which became the principal economic activity of
the region (Viera and Rangel 1988). Agglomerations of settlements along the
Patos Lagoon margins eventually gave rise to the present 16 counties, which
together represent 90% of the population of the State of Rio Grande do Sul.
12 M.L. Asmus: Coastal Plain and Patos Lagoon

From the very beginning, these settlements not only exploited the abun-
dant fishery resources of the Patos Lagoon as a food source, but also used
the lagoon as a cultural, social, and commercial link. Advances in agricul-
tural techniques, principally irrigated rice cultivation, as well as the
implantation of modern means of transport, caused profound modifica-
tions in occupation and development of urban centres around the lagoon
(Habiaga et aI., unpubI.). Counties which in the past depended on local
lacustrine ports for their economic growth were forced to exploit tourism
as a new means of development. Today, virtually all port activities are
assumed by the ports of Porto Alegre, Pelot as, and Rio Grande, which move
approximately 22 500 000 tons of goods yearly. Over the last few decades,
rapid and uncontrolled demographic and industrial growth around the
lagoon has altered its natural processes and magnified environmental con-
flicts. Elevated runoff from the Guafba River tributaries is responsible for
adding ever increasing sediment loads, nutrients, hazardous heavy metals
(Baisch et aI. 1989), and agrotoxins, whilst na'.jgation and port activitie~
and the establishment of fertilizer and fish processing plants and petroleum
refining have led to deterioration of the waters in the lower estuary
(Almeida et aI. 1993). Escalating conflicts between resource exploration and
human impact call for urgent management of the Patos Lagoon and the
coastal ecosystem (Asmus et aI. 1984).
4 Environment and Biota
of the Patos Lagoon Estuary

4.1
Geological Setting

1.J. Calliari

Regional Geological History

The Patos Lagoon is part of the coastal province which is geologically

defined by the Pelotas Basin and the basement (Villwock 1978). The base-
ment is characterized by the SuI-Rio Grandense shield, composed of
crystalline Precambrian rock formations, and by Paleozoic and Mesozoic
sedimentary and volcanic sequences of the Parana Basin, both of which
provided the sediments for the marginal Pelotas Basin. Owing to successive
transgressions and regressions during the Cenozoic, clastic continental,
transitional, and marine sediments of about 8000 m thickness accumulated
in the Pelotas Basin. The emersed portion comprises the present flat coastal
plain between 29 and 35° S which can geomorphologically be divided into
an inner alluvial plain, the Lombas Barrier, the Guaiba-Gravatai Lagoon
system, a multiple barrier complex, the Patos-Mirim Lagoon system, and
the continental shelf (Fig. 4.1.1; Delaney 1965; Villwock 1984). Geo-
morphologically, the estuary of the Patos Lagoon is surrounded by tertiary
and quaternary deposits of the low lands of the Patos-Mirim Lagoon
system and the multiple barrier complex, which are characterized by eolian
deposits, beach ridges, sand plains, stable and active dune fields, and
lagoonal terraces (Fig. 4.1.2; Godolphin 1976; Zeltzer 1976; Paim et al.
1987).

Estuarine Evolution

The ev()lutionary history of the Patos-Mirim Lagoon system was decisively

influenced by the development of a multiple sand barrier complex as a
result of eustatic sea-level changes during the Quaternary (Villwock 1978).
At least four successive transgression-regression cycles deposited pre-
sumably discontinuous barrier islands parallel to the coast. The isolation

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
14 L. J. Calliari

\
100 km

m:m Lombas Barrier

=
t;;;;;;;;;;;j
Guaiba·Gravatai Lagoon
System

D Multiple Barrier Complex

m I\\\Wl Patos· Mirim Lagoon
~ System

R§i!~ Continental Shelf

Fig. 4.1.1. Geology and geomorphology of the coastal province. (Modified after Villwock
1984)

of the Patos-Mirim Lagoon system was initiated by the formation of barrier III
during the late Pleistocene (120 000 B.P.; Villwock et al. 1986). Coastal waves
and wave-generated longshore currents progressively closed the barrier, except
for a channel through which waters of the extensive drainage basin forced their
way to the ocean. Thus, the entire system can be classified as a partially closed
lagoon (Nichols and Allen 1981). The geomorphological characteristics support
the model of lagoon evolution postulated by Zenkovitch (1967) and Rosen
(1975). However, although geomorphological events during the late Pleistocene
created the original estuarine conditions, erosion and deposition processes of
Holocene morpho dynamics were responsible for the evolution of the modern
Paws Lagoon estuary. The last Holocene transgression (5500 B.P.) moulded
barrier III deposits into a continuous scarp and deposited an external barrier
(IV), composed of recent beach ridges and adjacent dunes (Villwock et al.
1986). The progradation of barrier IV, through deposition of seven regressive
Geological Setting 15

...

t;,::;{;~ EOLIAN DEPOSITS

o BEACH RIDGES
D LAGOONAL TERRACES
~ PLEISTOCENE
'"::., TERTIARY
Q PRECAMBR.lAN

ATLANTIC OCEAN :N

t
20 km

Fig.4.1.2. Geology and geomorphology of the Patos Lagoon estuary. (Modified after
Paim et al. 1987)

beach ridges (Godolphin 1976) over a 13 km wide and 63 km long area

immediately south of the estuary, has given rise to two different theories of
local holocenic estuarine evolution. Godolphin (1976) postulated that
interactions between lagoonal outflow -and coastal hydrodynamics, like
waves and longshore currents, formed the main process influencing the
evolution of the Patos Lagoon estuary, whilst Long and Paim (1987)
suggested that cyclic migrations of the main channel in relation to coastal
orientation were responsible.
16 1.J. Calliari

Estuarine Morphodynamics

The Patos Lagoon has a maximum width of 59.8 km, and the main axis
extends 180 km. Average depth in the lagoon body and the estuary is 5 m,
but depth distributions are highly variable. The bottom topography of the
main lagoon body is characterized by natural and artificial channels (8-9 m
depth), large adjacent areas (>5 m depth), and shallow marginal bays. In
the estuary, large shoal areas between 1 and 5 m deep predominate, and the
maximum depth (18 m) is found in the inlet channel which connects to the
Atlantic Ocean (Calliari 1980; Toldo 1991). A large number of shoals and
spits, which define embayments, are typical of shallow «1 m) lagoon and
estuarine margins. They are evidence of recent morphodynamic estuarine
adjustments. Their development is largely a function of bidirectional winds
blowing parallel to the main lagoon axis, wave approach (Zenkovitch 1959),
microtidal oscillations, and the initial shape of the lagoon basin (Rosen
1975). Spits in the lower estuary are evidence of similar morpho dynamic
adjustments, but tidal inflow and flushing patterns are likely to have
influenced their development. Geomorphological and sedimentological
data indicate that lagoonal spits were deposited over the substrate of
submerged banks, owing to a gradual water level rise (1 m) during the last
3 centuries (Toldo 1991).

Sediment Sources

The region's high level of precipitation causes considerable fluvial silt and clay
transport from the large drainage basin, and thus the estuary receives sedi-
ments from a variety of sources. Owing to a gradual decrease (1 m/120 km) in
elevation towards the estuarine inlet, the reduced flushing current veloci-
ties of the Patos-Mirim Lagoon complex lead to large sediment deposition
from the water column during transport (Martins 1963; Hartmann et al.
1980). Apart from fluvial sediment input, the hydrodynamic erosion of
estuarine margins, especially those comprised of Pleistocene and eolic
Holocene formations, lagoonal terraces, and marshes, also contribute
significant amounts of sediments. Additionally, sandy sediments of
marginal dunes along the eastern shore are added during periods of strong
NE winds to adjacent estuarine bays. Finally, high current velocities of
inflowing seawater promote the deposition of sandy marine sediments in
the lower estuary, and the benthic estuarine fauna and flora represent
important sources ofbiodetritus.
Geological Setting 17

5km

Fig. 4.1.3. Distribution of bottom types in the Patos Lagoon estuary

18 C.A.E. Garcia

Depositional Environments and Sediment Characteristics

Six different bottom types can be related to depth and to hydrodynamic

energy levels in the estuary (Calliari 1980; Fig. 4.1.3). Silty clay sediments
are typical of deep channel and shallow protected bay environments
(Martins 1963, 1966, 1971). Transitional environments change with
decreasing depth from silty clay, clayey silt, mixed sand-silt-clay, clayey
sand to silty sand bottoms. Sandy bottoms are predominantly associated
with the shallow marginal portions of large shoals but occasionally occur in
high current channels. Also, marginal marshes, sandy shores, and islands
are common depositional environments.
Estuarine sediments are predominantly composed of quartz (90%), feld-
spar (5%), heavy minerals (0.5-1.50/0), and calcium carbonate derived from
molluscs and cirripeds (Martins 1963; Calliari 1980). Owing to the sedimen-
tary diversity of the source area and the complex estuarine circulation,
significant heavy mineral distribution patterns are lacking, though epidote
and hornblend prevail in areas under the influence of the Sao Gon~alo
Channel. Furthermore, anthropogenic activities have modified the sand,
silt, and clay fraction of superficial sediments in the lower estuary through
the introduction of organic carbon compounds (1.50/0 in fine fraction) and
heavy metals (Cu, Zn, Cd, Pb, Cr, Mn, Fe, Ni, Co; Baisch et al. 1988).

4.2
Hydrographic Characteristics

C.A.E. Garcia

Freshwater Inflow

Geomorphological evolution, implemented by high coastal wave energy in

combination with littoral sediment drift, formed the barrier that separates
the Patos Lagoon and the Atlantic Ocean. The lagoon and the ocean are
connected by a single inlet with a cross-section (0.5-3 km) which is small
compared with the lagoon surface area. Both regional geomorphological
events and the nature of the inlet characterize, more than any other
parameter, the system as a choked coastal lagoon (Kjerfve 1986).
Hydrographic estuarine and lagoon characteristics are closely linked to
and dominated by hydrological/riverine cycles. About 850/0 of the water
from the immense Patos-Mirim Lagoon drainage basin originates from the
Guafba river, the Camaqua river, and from Sao Gon~alo Channel falling
into the upper, central, and lower lagoon, respectively (Fig. 3.2). Freshwater
discharge varies considerably between the summer/fall and winter/spring
(6 to 5300 m3 S-1 from the Camaqua basin and 41 to 25 000 m3 S-1 from the
Guafba basin; Herz 1977). Especially during years under the influence of
Hydrographic Characteristics 19

El Nino, runoff greatly exceeds average values. Water flow through the Sao
Gonyalo Channel is controlled by artificial locks to impede saltwater pene-
tration from the estuary into the Mirim Lagoon. Only during periods of
elevated water accumulation in the Mirim Lagoon are these locks opened.
As a consequence, freshwater discharge into the upper estuary is variable
(700-3000 m 3 S"l), usually being more intense during the winter and spring.

Tides and Currents

The southern Brazilian coast is located in a region of minimal tidal influ-

ence (Defant 1961). The mean tidal amplitude is 0.47 m, and tides are pre-
dominantly mixed-diurnal with an F-parameter of 2.42. The tidal wave
(0 1 = 25.8h), with an amplitude of 10.8 cm, represents the principal compo-
nent (Herz 1977). Under typical summer conditions, the energy spectrum
of the water level time series in the estuarine inlet refle.l;;ts significant signals
of diurnal (T=25.6 h), semi~diurnal (T=12.5 h), and quarter-diurnal (T=6.2 h)
tidal components (Moller et al. 1996). As is typical for choked lagoons, the
inlet channel of the Patos Lagoon acts as a set-down filter and therefore
largely confines tidal influence to the channel and strongly attenuates
amplitudes as the tidal wave advances into the estuary and lagoon. Owing
to a large surface to volume ratio and to a gradual decrease in elevation of
seawater, maximum current velocities in the main lagoon body are about
0.3 m s"\ with a frequent inversion of direction (Moller, unpubl.). However,
in the inlet flushing current velocities may reach 1.7-1.9 m S"l after pro-
longed periods of heavy rainfall (DNPVN 1941), whilst peak inflowing
seawater current velocities approach 1.3 m S"l.

Forcing Factors of Lagoon and Estuarine Circulation

Most choked coastal lagoon systems are wind forced, rather than tidal driven,
and experience limited short-term marine variability (Kjerfve 1986). The
Patos Lagoon region is influenced by a predominantly NE-SW wind regime.
Winds from the NE quarter (mean velocity 3.6-5.1 m S"l) occur during 22% of
the year and winds from the SW quarter (mean velocity 5.7-8.2 m S"l ), during
12%. Seasonal wind patterns are represented by frequent and strong NE
quarter winds between September and April and by SW quarter winds from
May to October (Delaney 1965; DHN 1974). These winds blow along the NE-
SW main axis of the lagoon body and were identified early on as the principal
forcingJactor in the Patos Lagoon system (Bicalho 1883; von Ihering 1885;
Malaval 1922). They decisively control circulation, salinity distribution, and
water levels, though fluvial discharge may also generate seasonal pressure
gradients with elevated water levels after the onset of the rainy period.
20 L.F. Niencheski and M. G. Baumgarten

Wind direction influences both local and large-scale circulation patterns.

Under NE winds a pressure gradient rises along the main lagoon axis. A pres-
sure gradient between the inlet channel and retreating adjacent coastal waters
favours flushing of the lagoon water (Motta 1969). In contrast, SE and SW
winds cause inversion of flow in the main lagoon body and raise the water
level at its northern limits (Ferreira, unpubl.). Preliminary data indicate a
time lag of 4-6 h for wind-induced seiching and set-up/set-down cycles
(Moller, unpubl.). However, since systematic wind-driven circulation pat-
terns are poorly developed and highly variable, the time-lag responses are
likely to be unstable (Rosauro and Endres 1991). Also, modelling attempts
(Bonilha 1974; Rosauro 1986) indicate that in the major part of the lagdon,
water levels and vortex formations are strongly influenced by wind direction
and velocity, respectively (Almeida et al. 1990). During the summer, SW
winds tend to generate a stationary wave with a nodal line along the main
axis of the lagoon and several cells with inverse current flow proximate to the
margins (Moller et al. 1996). In the inlet channel, outflowing currents are
forced along the western margin under the influence of NE winds, whilst SE
and SW winds direct inflowing currents towards the shallower eastern bank
(Gafn!e 1927).
As a consequence of reduced tidal influence in the inlet and in the estuary,
the salinity distribution lacks tidal variability but does correlate with wind
forcing and variations in freshwater input on scales of hours to weeks. During
periods of low fluvial discharge (i.e. summer/fall), onshore SE and SW winds
force seawater through the inlet into the lower estuary and occasionally as far
as 150 krn into the lagoon. In contrast, NE quarter winds together with high
fluvial discharge significantly decrease estuarine salinities (Calliari 1980;
Costa et al. 1988a). Fluvial discharge in excess of 3000 m3 S-1 causes pronounced
salinity stratification in the inlet, and higher values extend the estuarine mixing
zone into coastal waters (Moller et al. 1991). Although our understanding of
the system's hydrodynamics is still limited, it is evident that both the regional
climate and the hydrological cycles are the principal forcing factors control-
ling lagoon and estuarine circulation patterns and salinity variations.

4.3
Environmental Chemistry

L.F. Niencheski and M. G. Baumgarten

Temperature and Salinity

The time-space variability of physico-chemical estuarine parameters and

their component concentrations are a result of interactions between forcing
factors, regional sediment characteristics, and anthropogenic activities. In a
general manner, estuarine water temperature and salinity, with higher
Environmental Chemistry 21

values during the summer, are related to the region's seasonal temperature
cycles and wind and precipitation patterns, respectively (Vilas Boas 1990).
Temperature and salinity profiles characterize different estuarine condi-
tions. Homogeneous conditions, ranging from freshwater to full-strength
seawater, are associated with high fluvial discharge and NE quarter winds
or minimal freshwater discharge and SW quarter winds, respectively. In
contrast, periods of strong freshwater runoff and cQncomitant SW wind-
induced seawater penetration cause vertical stratification because fresh-
water advances in the deeper channels over the saltwedge. Also, lateral
temperature and salinity stratification may occur in the inlet during
reduced freshwater discharge (Kantin 1983). Therefore, rather than being
under tidal influence, the extent of the saltwedge or unpredictable short-
term changes from stratified to homogeneous conditions are principally
controlled by the combined effects of highly variable runoff and wind pat-
terns (Baptista 1984; Moller et al. 1991). Since the saltwedge strongly influ-
ences mixing processes of estuarine waters and the formation of salinity
gradients, which directly act upon the liberation andlor removal of ele-
ments in the water column and the interfaces (Niencheski and Windom
1994), the estuary of the Patos Lagoon is chemically speaking highly unsta-
ble (Niencheski et al. 1986). A general absence of conservative mixing
between lagoon and ocean waters increases this instability even further
(Baptista 1984) because water masses are frequently a result of prior mixing
(Vilas Boas 1990). Furthermore, owing to the physiographic heterogeneity
of the estuary with deep channels, open shallow waters, and different types
of marginal bays, each environment displays site-specific physico-chemical
characteristics which strongly vary with the season and from year to year.

Suspended Matter and Dissolved Oxygen

The principal sources of suspended matter are the rivers in the northern
(Jacui, Sinos, Gravatai, Cai, TaquarO and central (Camaqua) portion of the
Patos Lagoon, the Sao Gon<;:alo Channel, and erosion processes in the southern
part. Furthermore, interactions between geomorphological features of the
narrow inlet channel and inflowing seawater during strong SW winds favour
the resuspension of deposited sediments in the lower estuary (Niencheski
and Windom 1994). As a result, suspended matter loads in the estuary tend
to increase towards the ocean and depend strongly on precipitation patterns
in the Patos-Mirim watershed. Despite high mean suspended matter loads in
the channel (50 mg I -1) and shallow water environments (30 mg I -1), dissolved
oxygen in the water tends to be close to saturation or even at supersaturation
levels, especially in the inlet channel with pronounced hydrodynamic activity.
The shallow depth «1 m) of the larger part of the estuary favours water
oxygen levels in equilibrium with the atmosphere.
22 L.F. Niencheski and M. G. Baumgarten

Nutrients

Calculations of the annual estuarine nutrient budget demonstrate high dis-

solved nitrogen, phosphate, and silicate concentrations (Niencheski and
Windom 1994) without any clear seasonal pattern. Since fluvial nutrient lev-
els, added to the upper Patos Lagoon, suffer substantial reduction during
transport before reaching the estuary, high dissolved and particulate nutrient
concentrations are likely to be a result of industrial effluents and remobiliza-
tion of bottom sediments in the lower estuary (Niencheski and Windom
1994).
The average phosphate concentration in estuarine channels and marginal
bays north of the city of Rio Grande (Fig. 3.2) is approximately 1 ~, but may
reach 3 ~ during the summer (Kantin 1983). The higher phosphate con-
centrations in semi-enclosed bays south of Rio Grande (Fig. 4.3.1) are
probably due to emission of industrial effluents, especially those from the local

6
A
5
o UPPER ESTUARY
i' ~ LOWER ESTUARY
=.4
O INLET
w
~
0:( 3
J:
Q.
(I)
0 2
J:
Q.
-- ~ ------

0
6
B
5
i' : SOUTH
=. 4
w
~
8 NORTH
0:( 3
J:
D.
(I)
0 2
J:
Do.

M J J A SON 0 J F M A M J J
1990 1991

Fig. 4.3.1. Phosphate distribution in the inlet and the main channel of the upper and
lower estuary (A) and in shallow estuarine waters north and south of Rio Grande city
(B). Dotted line represents maximum levels at which phosphate appears to be buffered
by sediments according to Liss (1976)
Environmental Chemistry 23

fertilizer plants (Baumgarten et al. 1995). Phosphate and suspended matter

concentrations in the estuary are low during freshwater influence but tend to
increase with salt water intrusion (Niencheski and Windom 1994), either
because phosphate adsorption onto particulate matter decreases, owing to
increased competition with Crand sot seawater ions for similar exchange
sites (Liss 1976), or because desorption processes liberate phosphates from
suspended matter. Several sources add nitrate and ammonia to the estuary.
Elevated nitrate and ammonia concentrations in the upper estuary are partially
a consequence of aquatic macrophyte and algal decomposition and minerali-
zation processes in the lagoon (Kantin 1983). The concentrations in waters of
the Sao GonlYalo Channel derive from agricultural activities in the Mirim
Lagoon watershed and from raw domestic sewage discharge by the city of
Pelotas. High surface water concentrations in the estuarine inlet (Vilas Boas
1990) and in waters around the port of Rio Grande (Baumgarten et al. 1995)
seem to be a consequence of atmospheric input by local fertilizer industries
and urban sewage. The Patos Lagoon water is the dominant source of
dissolved silicate (approx. 175 J.!.M) to the estuary. Although silicate concen-
trations oscillate along salinity gradients, they tend to decrease in a conserva-
tive manner towards the ocean, except when freshwater dominates the
estuary.

Metals

Particulate metal concentrations in estuarine waters strongly depend on

resuspension processes of bottom sediments. Sediment resuspension in the
inlet channel by inflowing seawater leads to salinity, suspended matter, and
particulate metal concentration gradients which decrease towards the upper
estuary, whilst wind and wave action may locally contribute to high sus-
pended matter and consequently to high particulate metal concentrations;
thus, a dear seasonal pattern of metal distribution is lacking. Particulate met-
als are either a result of natural weathering or anthropogenic contribution.
Their origin can be traced by using aluminium as a geochemical normalizer
to establish metaVAl ratios and to determine enrichment factors. The
enrichment factor ofNi, V, Ba, and Fe in estuarine waters with high suspen-
sion loads (>50 mg n is relatively constant, which suggests that these metals
occur at natural levels. However, the enrichment factors of other metals (Cu,
Zn, Pb, Li, Cr, Mn, Cd, As, Ag, P) tend to be higher in water with low suspended
solids, indicating an anthropogenic contribution (Niencheski et al. 1994).
Specifically elevated estuarine copper concentrations might be introduced by
the Patos Lagoon which receives drainage waters from mining activities on
the Camaqua River. Sporadic increases of suspended copper and lead levels
may also reflect metal input by industrial activities in the Patos Lagoon
watershed (Baumgarten 1987; Vilas Boas 1990; Niencheski and Windom 1994).
24 C.S.B. Costa

4.4
Tidal Marsh and Wetland Plants

C.S.B. Costa

Species Composition

As in Uruguay and Argentina (Costa and Davy 1992), the intertidal

marshes of the islands and margins of the Patos Lagoon estuary are essen-
tially flooded by brackish water and are occupied by annual and perennial
tidal marsh and wetland plants. There are approximately 70 species in the
lower estuarine marsh flora (Appendix). The tropical (Paspalum vaginatum,
Acrosticum aureum) and cold temperate (A trip lex patula, Limonium
brasiliensis) elements of the flora characterize southern Brazilian marshes
as part of a warm-temperate biogeographical transition zone. Although
species distribution occurs over a wide vertical range with a high degree of
overlap, the distribution of dominant species is related to topography
(Fig. 4.4.1). Sites with more than 50% cover of the marsh grasses Spartina
alterniflora, Scirpus americanus, Scirpus maritimus, Spartina densiflora,
and Juncus effusus occur more frequently in areas flooded during 64.0, 37.4,
25.5, 20.1, and 3.1% of the year, respectively. The local occurrence of
monospecific patches additionally suggests interspecific competion as a
determinant of plant distribution. Even though dominant species cover
more than 50% of the marsh surface, secondary species (Typha domingensis,
Bacopa monnieri, Alternanthera phyloxeroides, Ischaemum minus, Acrosticum
aureum, Polypogon monspeliensis) may locally reach up to 90% cover.

Plant Community Structure

The vertical inundation gradient, the horizontal salinity gradient, and the
presence of indicator species characterize six marsh plant communities
(Costa and Copertino, unpubl.). Despite their proximity to the water table,
upper marshes between 0.8 to 1.2 m above mean low water (MLW) only
inundate during exceptionally high water levels in the estuary. Two com-
munities, represented by the indicator species Ischaemum minus, Polypogon
monspeliensis, Cyperus polystachyus, Eleocharis spp., Alternanthera
phyloxeroides (Schafer 1985; Cordazzo and Seeliger 1988a), and by Cyperus
obtusatus, Panicum repens, Bacopa monierii, Paspalum vagina tum, Hydro-
cotyle bonariensis, Juncus acutus (Reitz 1961; Cordazzo and Seeliger 1988a;
Rocha and Costa 1988), characterize transitions to wetlands and dune
slacks, respectively. In upper marsh areas (0.3-0.8 m above MLW), which
are rarely inundated by either fresh or brackish water, a third community
of dense stands of Juncus effusus becomes dominant (Capftoli et al. 1978).
Tidal Marsh and Wetland Plants 25
100
SPARTINA ALTERNIFLORA
~
80
u>- SCIRPUS AMERICANUS
z
w
;j
aw 60 SCIRPUS MARITI MUS
Ik:
LL SPARTINA DENSIFLORA
z 40
~

a
0
JUNCUS EFFUSUS
0 20
...J
LL

o )50% COVER
.(l.70 -0.40 .(l .10 +0.20 +0.50 +0.80 • HIGHEST
MEAN WATER LEVEL (m) FREQUENCY

Fig. 4.4.1. Vertical range of distribution of dominant salt marsh species in the Patos Lagoon
estuary. FFOM Frequently flooded oligohaline marsh; FFMM frequently flooded
mesohaline marsh; OFM occasionally flooded marsh; RFM rarely flooded marsh

The majority of small islands and erosive banks in the estuary are com-
prised of occasionally flooded mid-marshes (0.1-0.3 m MLW) dominated by a
dense Spartina densiflora community, which is also typical of nutrient-rich
coastal marshes in Uruguay and northeastern Argentina (Parodi 1940;
Mobberley 1956; Carnavale et al. 1987; Cordazzo and Seeliger 1988a). A com-
bination of structural, physiological, and life-cycle characteristics permits
Spartina densiflora to thrive in the mid-marshes despite high ion concentra-
tions, low water and redox potential, impeded gas exchange during submer-
sion, and perturbation by currents (Davy and Costa 1992). During drought
periods the lignified leaves partially obtain support from a large number of
siliceous bodies. A thick foliar epidermis, sunken stomates, vascular bundles
protected by a lignified cell layer, and a C4 metabolism effectively control the
plant's water budget, thus avoiding the toxic and osmotic effects of excessive
NaCI accumulation. Apart from active excretion by foliar salt glands, NaCI
uptake is partially impeded by several. suberized root cell layers. Finally,
aerenchymatic leaf, stem, rhizome, and root tissues oxidate the rhizosphere,
thereby preventing accumulation of CO 2 and toxic substances during anaero-
bic respiration in flooded sediments (Perazzolo and Pinheiro 1991; Bastos et
al. 1993a,b). The perennial Salicomia gaudichaudiana co-occurs with Spar-
tina densiflora but becomes particularly dominant in adjacent mud flats
where prolonged inundation and large salinity fluctuations impede growth of
the latter. Other accompanying species include Vigna luteola and Senecio
tweediei which, owing to their vine-like growth, avoid light attenuation by the
Spartina densiflora canopy (> 1.5 m).
26 C.S.B. Costa: Tidal Marsh and Wetland Plants

As elsewhere along the southwestern Atlantic coast (Chebataroff 1952;

Oliveira and Nhunch 1986; Soriano-Sierra 1990), the frequently inundated
lower marshes (between 0.10 and 0.50 m MLW) can be divided into oligo-
«6) and mesohaline (>6) salinity zones (Coutinho and Seeliger 1984) and
are dominated by mono-specific Scirpus american us and Spartina altern i-
flora communities, respectively (Fig. 4.4.2). Bluegreen and macro algae are
common components of both communities (Coutinho and Seeliger 1986;
Yunes et al. 1990). Highly productive stands of Scirpus maritimus may
opportunistically dominate transitional areas between mid-marshes and
lower mesohaline marshes (Costa, in press). Owing to wave and wind
deposited debris, these sites also favour the growth of the annual nitro-
philous species Chenopodium album, A trip lex patula, and Atriplex hastata,
the biannual Apium graveolens, and the perennial grass Eriochloa punctata.

+1.20 ... MEAN WATER LEVEL (m)

-0.50
V> T\IVF TDS RFM OFM FFOM FFMM

I
30

e:l
PAlOS
LAGOON

e4D 4D

I:
I ~
G) Z
::J
«
0
(J)
z
~ ~ ~ ~ ~
~ ~
G G ::!i

~ ~
G) G)
~
~ II 3
Fig. 4.4.2. Spatial distribution of different salt marsh communities in the Patos Lagoon
estuary and their ordination according to mean water level and average salinity.
TWF Transition to wet fields; TDS transition to dune slacks; RFM rarely flooded marsh;
OFM occasionally flooded marsh; FFOM frequently flooded oligohaline marsh; and
FFMM frequently flooded mesohaline marsh
Submersed Spermatophytes 27

4.5
Submersed Spermatophytes

U. Seeliger

Ecology of Ruppia maritima L.

The larger part of the shoals and shallow «1.5 m) nearshore areas, with their
reduced circulation and sandy sediments, is colonized by submersed higher
vegetation. The principal colonizer is Ruppia maritima (Fig. 4.5.1), though
Zannichellia palustris, Potamogeton striatus, Myriophyllum brasiliense, and
Ceratophyllum demersum may become locally important during prolonged
periods of low salinity (Cafruni et al. 1978; Cafruni 1983; Moreno 1994;
Appendix).
The dispersal mechanisms of Ruppia maritima involve seeds and dis-
lodged plants. Germination of seeds is regulated by environmental condi-
tions acting upon the seeds after shedding. Since germination responses
differ significantly between sites, populations appear to have adapted their
life history to site-specific environmental conditions. In ephemeral marsh
pools, for example, exposure to desiccation stress breaks the seed coat and
stimulates the action of the seed's enzyme systems, thus preparing seeds for
immediate germination during subsequent flooding. In permanently
flooded estuarine shoals, low salinity and increasing temperatures may
have a synergistic effect on seed germination. Since lower salinities occur in
the winter and spring and higher temperatures in the spring and summer, the
seeds will tend to germinate in spring when temperature (15°C) and salinity
conditions are optimal (Seeliger et al. 1984; Koch and Seeliger 1988), though
low germination rates persists during the entire growth cycle.
Incident light, water temperature, and salinity support perennial growth
of Ruppia maritima in the estuary, but annual growth cycles are more
common. Growth may commence any time between the early spring and
early summer and cease between the late summer and late fall. The highest
growth, leaf density, and biomass are attained during the summer (Cafruni
1983; Moreno 1994). The formation of reproductive shoots with flowers
and fruits may continue in the perennial population, but generally initiates
in annual populations during peak growth in the summer (Fig. 4.5.2).
Resource allocation into reproductive structures is highest in intertidal
populations under exposure stress during summer. In lower subtidal
populations flower and fruit formation is absent, suggesting the existence
of depth-related sexual and vegetative reproductive strategies (Costa and
Seeliger 1989).
28 U. Seeliger

Fig. 4.5.1. Habit of Ruppia maritima plant (A), leaf tip (B), inflorescence (C), cluster of
fruits on long peduncle (D), and seed (E)

Several factors, acting either alone or in concert, cause die-off of annual

Ruppia populations in the late summer/fall. Experimental culture studies
suggest that underwater light regimes, rather than water temperature and
salinity, may limit growth and production of Ruppia populations in the
estuary (Knak 1986). Additionany, desiccation may interfere with continued
growth since the depth distribution of populations is restricted to a narrow
range. Upper growth limits are controlled by air exposure (>20% year -1)
and lower limits, at about 70 cm below mean low water (ML W), by low
underwater irradiance. After dry summers and during NE winds in the fan,
estuarine water levels tend to remain for prolonged periods 50-60 cm below
MLW (Costa et al. 1988a), which causes exposure of large areas and
subsequent mortality of Ruppia beds (Costa and Seeliger 1989). Further-
more, shading by algal mats and epiphytes may induce the death of popu-
lations. Owing to the basal growth of Ruppia leaves, epiphytic colonization
Submersed Spermatophytes 29

120
L5'
• VEGETATIVE
100
• UNDERGROUND

Fig. 4.5.2. Partitioning of mean monthly Ruppia maritima biomass in beds with annual
(Arraial Bay) and perennial (Mendanha Bay) growth

is densest on the older leaf apex during the summer and fall (Ferreira and
Seeliger 1985), resulting in sub-optimal light for continued growth. Also,
drift algae tend to become entangled in Ruppia leaves and flowering stems.
As a consequence, the drag force on the plants increases, making them
susceptible to dislodging by waves and currents. These stressful conditions
are likely to promote thinning of beds and may contribute to population
die-off (Teixeira da Silva 1995).

Biomass and Production

Average total biomass production of Ruppia beds is about 25 g dry weight m-\
though summer peaks can exceed 120 g dry weight m-\ owing largely to the
contribution of reproductive shoot biomass (Cafruni 1983; Moreno 1994).
The average belowground biomass represents approximately 30% of the
total biomass, but below/aboveground biomass ratios decrease during
meadow development and especially after the formation of reproductive
shoots (Fig. 4.5.2). The occurrence, permanence, and size of Ruppia beds as
30 U. Seeliger

well as their biomass and production depend on whether populations

follow annual or perennial growth cycles, probably as a function of under-
water light conditions and hydrodynamic patterns. Large and unpredict-
able natural variations of these factors may cause the total extension of
Ruppia beds in the estuary to differ between years by more than 40%, with
total interannual standing plant stock estimates oscillating between 3200
and 5200 metric tons (Moreno 1994).

4.6
Benthic Macroalgae

U. Seeliger

Species Composition and Distribution

The Patos Lagoon estuary is the site of growth for 94 benthic algal species,
represented by filamentous and colony-forming Cyanophyta (40), Chloro-
phyta (26), Phaeophyta (3), Xanthophyta 0), and Rhodophyta (24)
(Coutinho 1982; Appendix). The spatially and temporally unpredictable,
and thus stressful environmental conditions in the estuary are likely to
account for Cyanophyta becoming the largest and most diverse component
of the flora. Red and brown algae of marine origin progressively disappear
towards the upper estuary, but species diversity remains approximately
constant because the contribution of euryhaline green and bluegreen alga
elements increases (Coutinho and Seeliger 1984). The horizontal distribu-
tion patterns define three algal groups (Fig. 4.6.1) corresponding to annual
salinities of 1-12 in the upper estuary, 4-24 in the lower estuary, and l3-34
in the inlet. Although these salinity ranges may not express the salinity
tolerances of species, they do indicate salinity preferences of different
species groups. Superimposed on the salinity-dependent distribution are
algal affinities to substrate stability. Algal colonization and growth in the
lower estuarine shoals depend largely on the temporary presence of more
or less stable substrates, though some species also thrive as floating mats in
bays with reduced circulation. In contrast, the boulders of the jetties at
both sides of the estuarine inlet provide a permanent substrate for growth
and development. Substrate permanence clearly separates the local flora
into two species groups (Coutinno and Seeliger 1984). The permanent
rocks of the jetties and the influence of less turbid coastal waters also
support short vertical distribution gradients of resident algal populations.
At estuarine sites, algae lack vertical distribution patterns because high
suspension loads cause strong attenuation of the incident light (Coutinho
1982; Moreno 1994).

Fig. 4.6.1. Similarity based on frequency of occurrence of 64 algae in different annual

salinity regimes in the Patos Lagoon estuary.
Benthic Macroalgae 31

CORRELATION COEFFICIENT
-0.2 0 0.2 0.4 0.6 0.8 1.0

MERISMOPEDIA PUNCTATA
UPPER ESTUARY POLYSIPHONIA SP.
GLEOCAPSA CREPIDINUM
SPIROGYRA SP.

OSCILLATORIA OKENI
CLADOPHORAFLEXUOSA
ACROCHAETIUM FLEXUOSUM

~
OSCILLATORIA OBSCURA
ENTEROMORPHA L1NZA
SPIRULINA LABYRINTHIFORMIS.
LYNGBYA CONFERVOIDES
RHIZOCLONIUM KERNERI
CHROOCOCCUS TURGIDUS
POLYSIPHONIA SUBTILISSIMA
·C ECTOCARPUS SILICULOSUS
MICROCOLEUS TENERRIMUS
OSCILLATORIA LIMOSA
VAUCHERIA LONGICAULIS
CLADOPHORA CRISPATA
MICROCOLEUS CHTHONOPLASTES
LOWER ESTUARY PHORMIDIUM FRAGILE
ULVARIA OXYSPERMA
L.- ENTEROMORPHA INTESTINALIS
CHROOCOCCUS MINOR
ENTEROMORPHA BULBOSA

l:: ENTEROMORPHA CLATHRATA

ULOTHRIX FLACCA
RHIZOCLONIUM RIPARIUM
CHROOCOCCUS PALLIDUS
ENTEROMORPHA FLEXUOSA
PHORMIDIUM USTERII
CALOTHRIX PARIETINA
L YNGBYA AESTUARII
OSCILLATORIA ANIMALIS
XENOCOCCUS CLADOPHORAE
OSCILLATORIA SUBBREVIS
NODULARIA HARVEYANA

PETALONIA FASCIA
GIGARTINA TEEDII
POLYSIPHONIA TEPIDA
GRATELOUPIA FILICINA

t: HYPNEA MUSCIFORMIS
BRYOPSIS PLUMOSA
CERAMIUM BREVIZONATUM
GIFFORDIA MITCHELLAE
CLADOPHORA FASCICULARIS
CLADOPHORA SCITULA
ACROCHAETIUM GLOBOSUM
ULVA FASCIATA

~
GELIDIUM CRINALE
PORPHYRA PUJALSII
INLET CARRADORIA VIRGATA
GELIDIUM FLORIDIANUM
BRYOCLADIA THYSIGERA
CHAETOMORPHA AEREA

1
PHAEOPHILA VIRIDIS
GYMNOGONGRUS GRIFFITHSIAE
GRATELOUPIA CUNEI FOLIA
KYLINIA CRASSIPES
ERYTHROCLADIA SUBINTEGRA
ULVA LACTUCA
ENTEROMORPHA COMPRESSA
BANGIA ATROPURPUREA

Fig. 4.6.1.
32 U. Seeliger

Seasonal Patterns

The seasonal distribution of the 94 algal species shows three patterns of

algal periodicity. A group of 46 species is present throughout the year, and
two groups of 24 species each occur either during the summer/fall or dur-
ing the winter/spring. A seasonal alternation of eurythermal cool-water
algae, as part of the winter flora, with eurythermal tropical elements,
comprising the summer flora, is characteristic of floras in warm-temperate
biogeographic transition zones (Coutinho and Seeliger 1986). Seasonal
growth cycles occur in 55 green, red, and brown benthic algae and can be
related to light, temperature, photoperiod, and salinity. During
summer/fall, the growth peaks of 15 species significantly correlate to high
salinities, either alone or in combination with other factors. The favourable
influence of high salinity on the growth of the majority of these species
suggests stressful salinity conditions during the remaining seasons. Peak
growth of 11 species during the winter/spring largely depends on either low
water temperatures or their combined effect with low light. The growth of
the remaining 27 species either varies little or presents several peaks during
he year, probably owing to site-specific conditions rather than to seasonal
variations in environmental factors (Coutinho and Seeliger 1986). None of
the green, brown, and red algae reproduces throughout the entire year;
reproductive periodicity is restricted to one or two seasons. The lowest and
highest reproductive activity occurs in January and May, respectively. As
for growth, photoperiod, salinity, and temperature (either alone or in com-
bination) might explain the formation of different reproductive structures
in 45 species. Lack of reproduction in some Chlorophyta (23%) and
Rhodophyta (12%), or the absence of one life-cycle phase, might be a
response to increased estuarine stress (Coutinho 1982).

Biomass and Production

Benthic algal production is influenced by abiotic conditions and the hetero-

geneity of the environment. The first selects for the principal primary
producer species, the second for the size and site of production. Although
the general absence of consolidated substrate in the estuary probably limits
the growth of most species, it does not impede substantial biomass forma-
tion of others. Chlorophyta (Enteromorpha sp., Rhizoclonium riparium,
Ulothrix flacca) and Xantophyta (Vaucheria longicaulis) are the most
prominent producer species, but some Cyanophyta (Lyngbya confervoides,
lvlicrocoleus chthomoplastes) are also conspicuous and important estuarine
primary producers. Optimal biomass production of each species is linked
to physico-chemical conditions and the permanence of stable substrate.
Although Enteromorpha sp. (10-35 g dry weight m-2 ) and Rhizoclonium
Benthic Macroalgae 33

riparium (47-817 g dry weight m-2 ) form biomass throughout the year, des-
iccation during exposure tends to reduce biomass formation during some
months. Biomass formation of Lyngbya confervoides (221 g dry weight m-2 ),
Microcoleus chthomoplastes (10 g dry weight m-\ and Vaucheria longicaulis
(21-161 g dry weight m-2) is largely restricted to the summer, fall, and
winter, respectively (Fig. 4.6.2). Large quantities of macroalgae which grow
on sediments, shell debris, pebbles, and Ruppia plants are often dislodged
by waves and currents and occasionally aggregate into extensive floating
mats. As long as drift algae remain in the photic zone, continuous growth
results in further biomass formation (Coutinho 1982).

50%
220
0% Rhizoclonlum rlparlum

200
50%
~
180 0
0% ~
Jo
r
> 50% 160 tJ:J
U
2 Enteromorpha sp. is
:;:
w 140
aw 0%
:::I Jo
II)

..
II)

120 iii"
..
0:
u. 50%
Co

~
I- 0% C'O,..,. <\"~" 100
'<
~
5w
to
liS·
0: 50% 80 :t
3
Mlcrocoleus chthonoplastes N

\
0% 60
~

50% 40
Vaucherla longicaulls

0% 20

50%
A S 0 N 0 J F M A M J J

Fig. 4.6.2. Total average monthly biomass and relative monthly frequency of occurrence
of principal algal producer species
34 C. Odebrecht and P. C. Abreu

4.7
Microalgae

C. Odebrecht and P. C. Abreu

Taxonomic Composition and Cell Abundance

Quantitative studies of the Patos Lagoon estuary phytoplankton have been

done only recently (Abreu 1987; Odebrecht et al. 1987; Bergesch 1990;
Torgan and Garcia 1990; Abreu 1992; Persich 1993; Abreu et al. 1994a;
Yunes et al. 1994; Bergesch et al. 1995). Throughout the year, diatoms, mo-
nads, and a taxonomically heterogeneous group of small «20 Jim) flagel-
lates (Prasinophyceae, Cryptophyceae, Haptophyceae, Chlorophyceae) are
the principal organisms (Appendix). Cyanobacteria (Cyanophytes) and
dinoflagellates are more abundant in periods of lower and higher salinity,
respectively. Owing to their functional role as primary producers
(Lindholm 1985), the. number and biomass of the' autotrophic ciliate Meso-
dinium rubrum indicate that it also makes up an important part of the
phytoplankton community. Except during periods with high chlorophyll a
concentrations owing to diatoms or cyanobacteria, unidentified small flag-
ellates and monads are the most abundant organisms (10 6 _109 cells r\
representing the major phytoplankton fraction in the estuary (Fig. 4.7.1).
Because their identification poses special problems, more detailed studies
are required to understand their dynamics and their contribution to the
estuarine phytoplankton community.
Diatoms, dinoflagellates, and cyanobacteria follow distinct seasonal
patterns as a result of meteorological conditions which govern nutrient
availability, salinity, and light regimes. High phytoplankton abundance in
n
the spring (10 6 _10 9 cells is related to blooms of euryhaline diatoms, like
Skeletonema costatum and Cerataulina daemon (Fig. 4.7.1), which are
introduced into the estuary from coastal nearshore waters. Higher salinity
(>20) and temperature in the summer and/or fall favour maximum abun-
n
dance (104 _10 5 cells of Mesodinium rubrum, dinoflagellates (Prorocentrum
minimum, Peridinium quinquecorne), and the diatoms Chaetoceros,
Rhizosolenia, Coscinodiscus, and Odontella. High numbers of the limnic
cyanobacteria Microcystis aeruginosa (109 cells n and the oligohaline
diatom Skeletonema subsalsum (106 cells rl) occur during flushing and low
salinity conditions «5). Also, interannual differences in species
composition are largely due to haline conditions which vary with precipi-
tation and runoff.
Large-scale meteorological factors control the phytoplankton composi-
tion in the entire estuary in addition to local hydrographic factors and
winds, which strongly interact with benthic and planktonic micro algal
communities in the extensive shoals (Bergesch et al. 1995). The brackish
pennate diatoms Cylindrotheca (=Nitzschia) closterium, Bacillaria paradoxa,
 Microalgae 35

Terpsinoe americana, Melosira spp., Surirella spp., and the centric Pleurosira
laevis are frequently resuspended from the sediments into the water column.
Phytoplankton cell concentrations per volume tend to be higher in shoals
than in the deeper open estuarine waters and are about three orders of
magnitude lower than those of benthic micro algae (Bergesch et alI995).
Epiphytic micro algae are important colonizers on submersed plants of
Ruppia maritima and on other available substrates. Colonization on Ruppia
starts in the spring with the diatom Cocconeis placentula, which adheres to
the leaves through mucus strands liberated by the raphe. Subsequently,
Synedra fasciculata becomes fixed on live or empty frustules of Cocconeis,
forming a three-dimensional mosaic which permits colonization of 4mphora,
Nitzschia, and naviculoid diatoms. During the final stage of colonization at
the end of the summer, diatoms like Pleurosira laevis, Melosira, Navicula,
Rhopalodia, and Mastogloia completely cover the leaf apex (Ferreira and
Seeliger 1985).

1.5
• Temperature B
o Salinity
;--
...- 10
S·
'-"

......
~
U 0.5

Ilill Centric diatoms BCerataulina daemon 12l Rhizoso/enia spp .

III Pennate diatoms rJiJMesodinium rubrum .Chaecoccro$ "PP·
l2'l Chaetoceros "PP· ~Ske/etonema cos/alUm
1.0 Q Gymnodinium sp . 2.0 • CylindrorhecR c/osterium

c • Prorocentrum minimum
D

-
...
<=>
' ;;; 0.5
'"
S
';;; 1.0
......
......
~ ~
U U

o 0.s..-,1Ilil4........
MAMJJASONDJFM MAMJJASONDJFM

Fig. 4.7.1. A Water temperature and salinity and B cell concentration of autotrophic
nanoflagellates (3-4 pm); C diatoms and dinoflagellates «20 pm); and D Mesodinium
rubrum and diatoms (>20 pm) in the Patos lagoon estuary (1989-1990)
36 c. Odebrecht and P. C. Abreu
Chlorophyll Q

Pronounced daily and weekly variability and annual and interannual

oscillations of chlorophyll a are common in the estuary. Distinct seasonal
pulses with high concentrations (30-75 mg m-3 ) occur in the spring and
summer, but occasionally high concentrations are also observed in winter
(Bergesch 1990; Proens:a et al. 1994; Abreu et al. 1994a). While annual and
interannual variability is principally related to seasonal light regimes and
meteorological conditions, like precipitation and wind direction and speed,
small-scale variability is largely a function of direct wind impact. Similarly,
wind-induced resuspension together with high productivity in shallow
areas «2 m) generates concentrations of phytoplankton biomass twice as
high as those in the deeper channels (>3 m) (Bergesch 1990; Proens:a 1990),
resulting in marked horizontal variability of chlorophyll a. In both deep
water and shoal environments, pico- «2 J.lffi) and nanoplankton (2-20 J.lffi)
represent an annual mean of more than 70% of the total chlorophyll a con-
centration (Bergesch and Odebrecht 1987; Odebrecht et al. 1988; Proens:a·
1990; Persich 1993; Abreu et al. 1994a). During spring/summer, the contri-
bution of microphytoplankton (>20 J.lffi) rises and the mean contribution
may approximate 30% of the total phytoplankton chlorophyll a.

Phytoplankton Primary Production

Lower (2-5 mg C m-3 h- I ) and higher (160-350 mg C m-3 h- I ) values of phyto-

plankton primary production in the estuary are associated with winter
(June-August) and spring/summer, respectively (Abreu et al. 1994a). Dur-
ing winter, net production may be negative, suggesting a prevalence of
heterotrophic processes in the water column. Productivity per volume in
channel environments tends to be a third of that in the shoals. The largest
portion of the assimilated carbon is taken up by the <20 J.lffi fraction
(annual mean = 70.3% of total particulate production). Dissolved produc-
tion represents 5-60% (annual mean = 22.6%) of total primary production
(Abreu et al. 1994a).
Seasonal differences in primary production are linked to irradiance and
temperature regimes, and to the availability of nitrogen compounds. Owing
to resuspension of seston loads, highly turbid waters (coefficient of light
reduction >10 m- I ) inhibit photosynthetic activity, especially during the
winter when the mean irradiance in the water column is frequently below
the critical value of 7.8 J.lffiol m-2 S-I (40 gcal dai\ Riley 1967) for phyto-
plankton growth. Coastal diatoms (Skeletonema costatum, Chaetoceros
spp.), dinoflagellates (Prorocentrum minimum), and Mesodinium rubrum
seem to be adapted to high light conditions (>400 J.lffiol m-2 S-I), while
nanoflagellate communities appear to photosynthesize more efficiently at
low light «50 JlIllol m-2 S-I; Abreu 1992). Nitrogen compounds for
Bacteria and Proto zooplankton 37

primary production are introduced into the estuary either during

freshwater outflow or during intrusion of nutrient-rich seawater (Abreu et
al. 1995a).

4.8
Bacteria and Protozooplankton

P. C. Abreu and C. Odebrecht

Bacteria

During the last decade increased research efforts and modern analytical
techniques have drastically changed our understanding of microbial ecology
in aquatic environments. New facts have led to the "microbial loop" concept
which postulates that the dissolved fraction of organic matter becomes avail-
able to higher trophic levels through bacterial uptake of dissolved organic
carbon, which in turn is grazed upon by flagellates. Energy and material are
channelled through ciliates and microzooplankton to higher consumers
(Azam et al. 1983). Recent investigations into the ecology of microbial organ-
isms of the Patos Lagoon estuary and adjacent coastal waters have shown that
bacteria contribute significantly to the carbon budget of the estuarine food-
web (Abreu et al. 1992). Bacterial biomass exceeds phytoplankton biomass
during several months of the year, but together they represent approximately
58% of the particulate organic carbon. Microbes are likely to play an impor-
tant role in the lagoon, estuarine, and coastal foodwebs (Abreu 1992), and
preliminary data suggest that the phytoplanktonic, bacterial, heterotrophic
flagellate, and ciliate biomasses are closely related (Fig. 4.8.1).

2500 D PHYTOPLANKTON
i;
o ATT. BACTERIA
2000 !il l1 FREE BACTERIA
()
1:11
:::s. [Ill FLAGELLATES
;-1500 ~ CILIATES
o .POC
m
~ 1000
()

500

M A M J ASONDJ FM
MONTHS

Fig. 4.8.1. Seasonal variation of particulate organic carbon (POC) and carbon-based
biomass of phytoplankton, free and attached bacteria, heterotrophic flagellates, and
ciliates in the Patos Lagoon estuary
 lJ>
00

:-0
o
~
(3
c
~
::l
0-
- o
o0-
a 10pm c ~
nln
::r
,..

Fig. 4.8.2. Feeding of Strombidium sp. (75 x 25 11m) by engulfing.a chain of Skeletonema (a,b) and details of damaged Skeletonema cells (c)
Bacteria and Proto zooplankton 39

A first evaluation of bacterial production in the coastal plume of turbid

estuarine waters and in immediately adjacent clear coastal waters demonstrated
that although in turbid waters bacterial production (2.09-6.55 mg C m-3 h-1) is
much lower than phytoplankton production (79.02 mg ~ m-3 h- 1), bacteria
may consume as much as 60% of the phytoplankton-produced organic
matter (Abreu et al. 1995b). In order to evaluate the importance of the
detrital chain in the estuarine foodweb, future studies will have to focus on
the role of bacterial succession and activity during degradation processes of
the large estuarine macrophyte and macro algal biomass and on grazing
and transfer rates of microbial organic matter to higher trophic levels.

Protozooplankton

Protozooplanktonic organisms in the estuary are represented by taxonomi-

cally diverse groups of heterotrophic flagellates (2-3 J..lm), dinoflagellates
and ciliates, like loricate (tintinnids) and aloricate oligotrichids (Strombidiidae)
(Appendix). The abundance of flagellates (l95-3800x10 3 cells n and ciliates
(1-62 cells n oscillates during the year (Abreu et al. 1992), but large
numbers (104 -5 cells n of loricate ciliates (principally Leprotintinnus and
Tintinnopsis) and aloricates of various sizes (Strombidium spp.) also occur.
Heterotrophic flagellate (21 I..lg C n n
and ciliate (34 I..lg C peak carbon
biomass estimates are highest during the spring, while values between
3 and 41..lg C r1 occur during the winter (Abreu 1992; Abreu et al. 1992). The
availibility of bacterial biomass directly controls the biomass of hetero-
trophic flagellates. However, microcosm experiments indicate that flagel-
late biomass is insufficient to sustain continuous growth of ciliate predators
because ciliate production (168.9 mg C m-3 dail) exceeds flagellate produc-
tion (39.3 mg C m- 3 dai\ It is likely that ciliates, rather than exclusively
grazing on flagellates, occasionally ingest bacteria, small phytoplankton
cells, and organic detritus to satisfy their nutritional requirements (Abreu
et al. 1992). Ciliates (i.e. Strombidium sp.) may also feed on large phyto-
plankters by engulfing and creeping along chains of Skeletonema, sucking
soft substances from the cells, and leaving semi-digested and damaged
chains behind (Fig. 4.8.2).
40 M. Montu, A.K. Duarte, and I.M. Gloeden

4.9
Zooplankton

M. .Montti, A.K. Duarte, and I.M. Gloeden

Species Composition and Distribution

Zooplankton in the estuary is composed ofholoplanktonic organisms (with

their entire lives as plankton) and meroplankton (temporarily planktonic).
Copepods are typical holoplankters, whilst barnacles and molluscs are
typical meroplankters, spending their early lives floating in the estuary
until they find a suitable substrate to metamorphose into attached forms.
Other meroplanktonic organisms, like the pink shrimp, the blue crab, and
some snails metamorphose into mobile forms.
The species composition of zooplankton (Appendix) in the estuary is
directly related to local hydrographic conditions. Saltwater penetration and
freshwater flushing introduce marine and freshwater species, respectively.
In general, circulation patterns created by bottom characteristics and the
influence of winds tend to concentrate species of marine origin along the
eastern and those of freshwater origin along the western margin of the
estuary. The estuarine salinity regime not only influences the distribution
of species with different origins but also species diversity which augments
with salinity increases. In order to describe patterns of species introduction
and distribution, the hydrographic estuarine conditions can be divided into
periods of inflowing seawater with high salinity, periods of salt and fresh-
water mixing, and periods of freshwater dominance during pronounced
flushing (Fig. 4.9.1; Montti 1980; Castello 1985).
After saltwater inflow, species of marine origin, principally Acartia
tonsa, Euterpina acutifrons, Oncaea conifera, Sagitta tenuis, larvae of
cirripeds (Balanus improvisus) and of echinoderms dominate the zoo-
plankton community. The larvae of most meroplanktonic species
(polychaetes, molluscs, crustaceans) of adjacent coastal waters are intro-
duced during this period and may reach the upper limits of the estuary
(Montti 1980), though juveniles of the mysids are generally restricted to the
lower estuary or to the inlet channel. During flushing periods, larval densi-
ties are drastically reduced and freshwater species like the copepods Noto-
diaptomus carteri, Notodiaptomus incompositus, Mesocyclops annulatus,
and a large number of planktonic (Diaphano-soma sarsi, Eubosmina tubicen)
and of small pleustonic cladocerans (Simosa, Alona, Chydorus, Pleuroxus,
Biapertura, Macrothrix, Eurycercus, Camp to-cercus, Kurzia, Pseudosida)
b~~come dominant. These cladocerans are regularly found in the shoals and
often associate with sub- and intertidal vegetation (Montti 1980; Montti and
Gloeden 1986). Periods of fresh-saltwater mixing promote the introduction
of species of both marine (Le. larvae of cirripeds, Paracalanus parvus,
Zooplankton 41

SPRING SUMMER

FALL WINTER

r----l PLANKTONIC PLEUSTONIC

L--' FRESHWATER SPECIES FRESHWATER SPECIES

Fig. 4.9.1. Distribution of zooplankton in surface waters of the Patos Lagoon estuary.
Distribution of pleustonic freshwater species does not imply an exclusion of other
species
42 M. Montu, A.K. Duarte, and I.M. Gloeden: Zooplankton

Euterpina acutifrons) and freshwater (Notodiaptomus incompositus,

Diaphanosoma brachyurum, Moina micrura) origin as well as a dominance
of Acartia tonsa. Typical estuarine species, like Acartia tonsa and Euterpina
acutifrons, only depend to some extent on tidal transport mechanisms for
their introduction. They persist in the estuary due to their euryhaline
nature and by maintaining reproduction and migration processes that
coordinate with the water circulation patterns. Continuous reproduction of
these species, the permanent presence of naupliil copepodites stages, the
abundance of females in excess of males, and migrations to deeper, high
salinity strata and to protected shoals where Acartia tonsa may reach
densities of up to 40 016 org. m-3 indicate successful strategies to warrant
the permanence of these species in the estuary (Duarte 1986).

Seasonal Patterns

Seasonal variations ill species composition, density, diversity, and growth

cycles are likely to be influenced by pronounced differences between sur-
face water temperatures in the winter (9-1S°C) and summer (2S-28°C).
Owing to more frequent saline conditions and higher water temperatures,
species of marine origin are common components of the zooplankton
community during the summer and fall. The cladocerans Pleopys
polyphemoides, Podon intermedius, and Penilia avirostris occur in the
lower estuary where also Ctenophores, Siphonophores, Medusae, and
Chaetognats (Sagitta tenuis) occasionally reach high densities. Some
species of subtropical waters, like Sagitta enflata and Centropages velifica-
tus, only enter the estuary during these warmer months. Also, mysids of
coastal origin are typically seasonal, with lower densities during winter.
These species follow a successional sequence. Mysidopsis tortonesi,
Metamysidopsis elongata, and sporadically Promysis atlantica dominate
from the spring to fall and are replaced by Neomysis americana in winter.
Some species of limnic origin, like the cladoceran Moina micrura, Diapha-
nosoma sarsi, and Ceriodaphnia cornuta coexist throughout the year.
Spring water temperatures tend to promote cyclomorphosis in Ceriodaph-
nia, resulting in cornuta and rigaudi forms (Montu 1980). Estuarine water
temperatures also seem to influence the reproduction and the size structure
of adult Acartia tonsa. Although reproduction occurs throughout the year,
reduced reproduction and lack of resting eggs during colder month result
in the dominance of larger individuals in winter (Duarte 1986). Also, the
diversity, density, and biomass of zooplankton follow seasonal patterns.
Larval densities tend to increase in the spring and summer when elevated
nUmbers (12 473 org. m-3 ) of Balanus improvisus larvae may competitively
exclude other zooplankton species. The highest total zooplankton densities
(40016 org. m-3 ) and biomass (0.2-11 rnl m-3 ) occur in the summer, whilst
Benthic Invertebrates 43

intermediate densities are typical of the spring (15 526 org. m-3 ) and fall
(14190 org. m-3 ) and the lowest densities occur during the winter (6251 org. m-3;
Montu 1980).
At sites with prolonged water residence, reduced circulation, and ele-
vated levels of domestic and industrial effluents, estuarine pollutants
appear to cause changes in the zooplankton community structure and
anatomical anomalies. Under these conditions, species diversity and den-
sity are reduced and 14.29% of individuals of Acartia tonsa display mal-
formations, expressed by adult females with legs of copepoditos stages IV
and V, inters exes (females with legs of male copepodite stages IV and V),
and ruptures of chitin between segments with extrusion of tissue.Anoma-
lies are likely to be a consequence of exposure to pollutants during several
life-cycles, suggesting that Acartia tonsa might be a suitable indicator for
deteriorating environmental conditions in estuaries (Montu and Gloeden
1982).

4.10
Benthic Invertebrates

C. E. Bemvenuti

Species Composition and Diversity

The benthic community consists of animals that live in the sediments as

infauna by burrowing or forming permanent or semi-permanent tubes,
extending just above the sediment surface; animals that live on the sedi-
ment surface as either mobile or sedentary epifauna; and animals that form
specialized live-bottom communities. As common to estuarine-lagoonal
environments, euryhaline marine and estuarine, rather than limnic, species
are the dominant components of the benthic macrofauna in the Patos
Lagoon estuary (Appendix). Typical estuarine invertebrates are repre-
sented by about 15 species, whilst only 3 limnic species (Tanais stanfordi,
Palaemonetes argentinus, Heleobia parchapei) occur in the estuary during
periods of pronounced freshwater influence (Capftoli et al. 1978). The
polychaetes Laeonereis acuta, Nephtys jluviatilis, Heteromastus similis, the
tan aid Kalliapseudes schubartii, and the pelecipod Erodona mactroides, all
components of the infauna, show the highest frequency of occurrence and
dominance in mudflats (Bemvenuti 1983, 1987a, 1992), seagrass beds
(Asmus 1984), and open water environments.
Epifaunal organisms, like the gastropod Heleobia australis and the crabs
Chasmagnathus granulata and Metasesarma rubripes occur at high densi-
ties in surface layers of bottom sediments and in marginal salt marshes
(Capitoli et al. 1978; D'Incao et al. 1992). The euryhaline and highly mobile
decapods Callinectes sapidus and Cyrtograpsus angulatus use the marginal
44 C. E. Bemvenuti

bays during the summer as growing sites, but migrate to deeper waters
during the lower fall temperatures. The pink-shrimp Penaeus paulensis is
commercially the most important decapod in the estuary. From the spring
to early summer, larvae of Penaeus paulensis enter the estuary from the
ocean. The shallow and protected habitats of estuarine shoals provide ideal
conditions for their development into juveniles and sub-adults. By the fall,
most of the females have matured and migrate back to the ocean to repro-
duce (D'Incao et al. 1990; D'Incao 1991). Polychaetes (i.e. Sigambra grubii,
Onuphis setosa, Magelona riojai, Hemipodus olivieri) and peracarid crusta-
ceans, represented by isopods (i.e Synidotea marplatensis), cumaceans
(i.e. Dyastilis sympterigiae), and amphipods (i.e. Bathyporeiapus bisetosus),
are major marine macrofaunal components in the estuary inlet.
The benthic macrofaunal community in the estuary is composed of only
38 species (Capitoli et al. 1978; Bemvenuti et al. 1992), most of which are
r-strategists with pronounced seasonal and interannual variations in abun-
dance. Low species diversity and high species abundance tend to be indica-
tive of stressful estuarine conditions (McLusky 1981). Since choked estu-
arine characteristics and unpredictable wind and precipitation patterns
promote unstable bottom substrates, modify hydrodynamics, and cause a
general absence of conservative salinity gradients in the Patos Lagoon estu-
ary, they may account for the low diversity of the macrobenthic fauna
(Bemvenuti et al. 1992).

Vertical Distribution Patterns and Abundance

The upper zone of marshes, which fringe the estuary, are dominated by
insects, but isopods and the amphipod Orchestia platensis also occur. The
omnivorous crab Chasmagnathus granulata occurs at high densities in
burrows in the lower marshes (Capitoli et al. 1978), whilst the preferred
habitat of the crab Metasesarma rubripes is a small fissure at the base of
Spartina alterniflora along eroded edges of the lower marsh. The density of
infaunal marsh species, like Orchestia platensis (3023 indo m-2 ) and juvenile
Laeonereis acuta, is low in comparison with subtidal environments
(Capitoli et al. 1977). Extensive unvegetated mudflats are characteristically
inhabited by the deposit feeder Laeonereis acuta, whose range extends
from the intertidal to about 1 m depth and reaches densities of 5127 indo m-2
and a biomass of 28.26 g m-2 (Bemvenuti 1987a, 1992). In spite of its wide
depth distribution and pronounced seasonality, the gastropod Heleobia
australis is most abundant (45 616 indo m-2 ) and achieves the highest bio-
mass (246 g m-2) in shallow waters (Capitoli et al. 1978). During periods oflow
density, Heleobia australis is replaced by the tube-dweller Kalliapseudes
schubartii (l2 808 indo m-2; Bemvenuti 1983), whilst Erodona mactroides
becomes abundant (20 300 indo m-2 , 216 g m-2 ) after intense recruitment
Benthic Invertebrates 45

during the summer (Bemvenuti et al. 1978). Although shallow vegetated areas
of the estuary, dominated by Ruppia maritima, may not be essential for the
establishment of the benthic macrofauna, they nevertheless provide
favourable conditions by either offering protection to young individuals of
Kalliapseudes and Erodona or leafy substrate for juveniles of Penaeus
paulensis and Heleobia. Since the biomass of these species is positively related
to the Ruppia biomass, shallow vegetated areas of the estuary may have an
important habitat function (Asmus 1984). In addition, deeper open waters
(2-6 m) are dominated by the deposit feeder Heleobia australis, the
suspension feeders Erodona mactroides and Kalliapseudes schubartii, and the
carnivore or detritus feeder Nephtys jluviatilis. The highest mean densities of
Heleobia (l2 927 indo m-2 ) and mean biomass of Erodona (281 g m-2 ) are
found in the mid and upper reaches of the estuary, respectively (Bemvenuti et
al. 1978). Benthic populations in deep channel environments are poor and
typically composed of deposit feeders, though only Heleobia australis occurs
in higher densities (Capitoli et al. 1978; Bemvenuti et al. 1992).

Recolonization Processes and Predation

A complex combination of abiotic and biotic factors controls recolonization

processes and influences the resilience of soft-bottom substrates in the
estuary. Furthermore, resilience processes to macrofaunal colonization are
not limited to the species level, but also affect the size categories of invading
individuals as a result of their changing niche requirements during
development (Giller 1984). After defaunation, the soft bottom substrates in
the Patos Lagoon estuary appear to be initially recolonized through
recruitment of Laeonereis acuta larvae and invasion by adults of Nephtys
jluviatilis. During intense predation by macro predators, these species give
way to lateral migration by large individuals of Heteromastus similis. Owing
to their limited dispersion, Kalliapseudes schubartii, large individuals of
Laeonereis, and epifaunal peracarids will rarely participate in the first stages
of colonization. Further colonization depends on habitat conditions, the
predation pressure on initial invaders, and their reproduction increment. The
available spaces might subsequently be occupied through recruitment by
Heteromastus and Nephtys larvae (Bemvenuti 1992).
Exposure makes epifaunal peracarids, like isopods, amphipods, and
tanaidaceans, susceptible to predation (Nelson 1979a; Virnstein et al. 1984),
and they may therefore serve as indicators of predation pressure (Bemvenuti
1987a,b). The highest densities of peracarids occur in estuarine shoals where
dense algal mats offer natural shelter; the abundance of crustaceans is
associated with a reduced number of predators and protection and food
offered by macroalgae during the fall and winter (Bemvenuti 1987b).
Experimental field studies suggest that macropredators like fish and
46 C. E. Bemvenuti: Benthic Invertebrates

decapods keep the density of sedentary epi-and infauna, an important

component of their diet (Araujo 1984; Asmus 1984; Bemvenuti 1990), below
the carrying capacity of the estuary. However, both macropredators and
infauna reach their highest abundance during the summer and early fall on
vegetated (Asmus 1984) and unvegetated (Bemvenuti 1983, 1987a,b, 1992)
subtidal bottoms. High infaunal densities are partially maintained through
escape mechanisms (i.e. burrowing) from predation, where success of escape
is a function of the size and mobility of the organisms (Virnstein 1979), but
they also depend on species-specific utilization of shallow estuarine habitats
and on efficient reproductive strategies, which permit the rapid replacement
of depleted populations (Bemvenuti 1992). Trophic interactions and
modifications of the soft-bottom community in the estuary are best
represented by an intermediary trophic level (Fig. 4.10.1; sensu Ambrose
1984a). At this level an infaunal polychaete (Nephtys fluviatilis) and small
crabs regulate the abundance of non-predatory infauna of specific sediment
strata through predation and bioturbation (Bemvenuti 1992, 1994).

SMALL PREDATORY
DECAPODS INFAUNA

1 JUVENILE
SUPERFICIAL
INFAUNA
JUVENILE
1
SUBSUPERFICIAL
INFAUNA

SEDENTARY EPIFAUNA ADULT INFAUNA

Fig. 4.10.1. Conceptual diagram of predation on the soft-bottom community in shallow

estuarine shoals
Physiological Adaptations ofInvertebrates and Fish 47

4.11
Physiological Adaptations of Invertebrates and Fish

E.A. Santos and A. Bianchini

Among benthic estuarine invertebrates decapods play an important eco-

nomic and ecological role. The shrimp Penaeus paulensis represents one of
the principal estuarine fishery resources, and the crab Chasmagnathus
granulata decisively interacts with the energy transfer in submersed and
emersed estuarine habitats (D'Incao et al. 1992). Although the respiratory
system of Chasmagnathus granulata is still highly dependent on the aquatic
environment, the species is able to exploit the upper intertidal region
effectively (Santos and Colares 1986; Santos et al. 1987). Aquatic decapods
have acquired ecological and evolutionary advantages to withstand
prolonged exposure to the atmosphere through morphological and
physiological adaptations. Metabolic adaptations, which may involve the
crustacean hyperglycaemic hormone (CRR), enable Chasmagnathus
granulata to survive in the atmosphere (Santos et al. 1988; Santos and
Stefanello 1991; Nery et al. 1993; Schmitt and Santos 1993b), and ion regu-
lation processes during emersion appear to be strongly water-dependent
(Schmitt and Santos 1993a). In general, the metabolism of Chasmagnathus
granulata differs from that of typical aquatic decapods, like Callinectes
sapidus (Santos and Colares 1990), being characterized by metabolic
depression and dependence on lipids as the energy source under stressful
conditions. In order to fulfill their vital functions, each life stage of Penaeus
paulensis and Chasmagnathus granulata has physiologically adapted to the
steep and highly unpredictable estuarine salinity gradients and to occa-
sional high, localized concentrations of nitrogen compounds and possibly
heavy metals (Seeliger and Knak 1982a,b; Baumgarten and Niencheski
1990a; Almeida et al. 1993) and pesticides. Also, some economically
important molluscs, like Mesodesma mactroides and the abundant Donax
hanleyanus, as well as the isopod Excirolana armata must tolerate salinity
variations in shallow coastal nearshore waters.

Salinity and Temperature

The ability to cope with osmotic stress in media with highly unpredictable
and fluctuating salinity regimes is clearly an essential prerequisit for the
successful settlement of Penaeus paulensis and Chasmagnathus granulata
populations in the Patos Lagoon estuary. Chasmagnathus granulata is an
euryhaline species with wide (0-40) salinity tolerance, owing to a combina-
tion of efficient hyper-and hypo-osmoregulation of its extracellular fluid
(Fig. 4.11.1), ionic (Na\ K\ ci\ Cn haemolymphatic regulation, intracel-
48 E.A. Santos and A. Bianchini

lular fluid regulation, and cell volume control (Bromberg 1992). Although
energy cost studies have emphasized the relative importance of carbo-
hydrate and lipids for osmoregulatory processes (Fig. 4.11.2; Santos and
Nery 1987; Nery and Santos 1993), the redistribution of energy otherwise
available for somatic growth or reproduction to physiological mechanisms
employed during salinity adaptation is likely to have a significant ecologi-
cal consequence. Salinity also controls the responses of Chasmagnathus
granulata to pH gradients. Under high salinity and low pH conditions the
osmotic equilibrium is disturbed, but pH values above 4.0 are well tolerated
(Miranda 1994), and the survival chances of Penaeus paulensis larvae
increase with pH.
In addition, Mesodesma mactroides (15-35), Donax hanleyanus (10-35)
(Souza-Santos 1991), and Excirolana armata (7-70) display considerable
salinity tolerance. Mesodesma mactroides is aNa \ K\ and Ca2+ conformer,
and salinity tolerance depends entirely on intracellular fluid regulation.
However, the volume adjustment of the branchial and muscle cells appears

-;-:-1200
<II
~

.><
~1000
E
<II
0
.§. 800
>-
t-
:::::i
c:( 600
..J
0
:E
en - ISOSMOTIC
0 400
c -( '- SUMMER
0 ... WINTER
0
..J
III
200

200 400 600 800 1000 1200

MEDIUM OSMOLALITY (mOsm kg water·1)

Fig. 4.11.1. Blood osmotic concentration in Chasrnagnathus granulata in winter and

summer, after 15 days of exposure to different salinities following a 30-day acclimation
period at a salinity of 20
Physiological Adaptations ofInvertebrates and Fish 49

Fig. 4.11.2. Blood glucose levels of Fig. 4.11.3. Mortality of Chasmagnathus

Chasmagnathus granulata exposed to granulata (96 h) after exposure to different
different salinities as a function oftime, salinity and temperature combinations
following a IS-day acclimation period
at a salinity of 30

to be regulated by intracellular variations of both inorganic ions and

organic effectors, which are independent of carbohydrate energy sources.
The metabolic responses of Donax hanleyanus and Excirolana armata vary
with salinity (Souza-Santos 1991). At low salinities «10), oxygen con-
sumption is reduced in both species. Under these conditions, metabolic
depression may decrease the energy expenditure of Donax hanleyanus. At
higher salinities (approx. 20), protein and carbohydrate metabolism is
stimulated without a concomitant increase in food uptake, which causes a
pronounced decline in production. An increase of oxygen consumption at
higher salinities (30) is likely to be associated with osmoregulatory proc-
esses and a concomitant reduction of protein metabolism.
Both field (D'Incao et al. 1992) and laboratory (Miranda 1994) data indi-
cate the preference of Chasmagnathus granulata for temperatures between
20 and 25°C. At lower temperatures mortality increases with lower salinity
«20; Fig. 4.11.3), possibly as a result of changes in ion (i.e. cr, Ca2+) and
osmoregulatory mechanisms (Miranda 1994). Similar to other land and
semi-terrestrial crabs, Chasmagnathus granulata remains in burrows and
becomes inactive at low temperatures. A relation between high tempera-
tures and reproductive activities is indicated by high recruitment and the
presence of ovigerous females throughout the summer (D'Incao et al.
1992). Local populations of Penaeus paulensis are eurythermal, with larvae
tolerating temperatures between 8 and 38°C, though the best development
occurs at intermediate temperatures (Boff and Marchiori 1984).
50 E.A. Santos and A. Bianchini: Physiological Adaptations ofInvertebrates and Fish

Pollutants

Since crustaceans are ammoniotelic, they tend to tolerate high concentrations

of nitrogen compounds. Ammonia tolerance, however, appears to be salinity
dependent. At salinities of 20, the LC50 (96 h) for Chasmagnathus granulata is
525 mg N-NH3 rl. A significant increase of osmotic concentration and Croccurs
after 96 h of hyperosmotic shock with rising ammonia concentrations. In
contrast, at low salinities sublethal ammonia levels cause a significant increase
of haemolymphatic K+ concentrations, possibly owing to inter-ference with
ionic transport processes. A potassium imbalance might interfere with the
oxidative branchial metabolism and consequently alter the energy balance of
the organism. In contrast, Penaeus paulensis displays low ammonia tolerance,
especially during larval development. Healthy larval development seems to be
inhibited above ammonia concentrations of 2.8 mg r l (Ostrensky Neto 1991),
and long-term exposure to low concentrations {0.14 mg N-NH3 n may reduce
growth up to 43%. The ammonia tolerance of Excirolana armata is similar to
that of the other crustaceans, toxicity being partially related to the species water
balance and osmoregulation.
Both Penaeus paulensis and Chasmagnathus granulata show significant
responses to potential estuarine pollutants like copper (CuSO 4) and pesticides
(methyl parathion), respectively. An LC 50 (96 h) for Penaeus paulensis post-
larvae at 1.33 ppm Cu and a decline of wet weight increase (42.69%) at 0.22 ppm
are likely to be either a function of enzyme inhibition for protein synthesis and
disturbance of osmoregulatory processes or an indirect effect of copper leading
to reduced feeding. In Chasmagnathus granulata pesticide toxicity (LC50 -96 h)
increases with temperature, possibly owing to hypoxia and higher oxidative
enzymatic de-sulphurization of the compound (Monserrat and Bianchini 1995).

Adaptations in Fishes

Some teleost fish also display physiological adaptation to environmental gradi-

ents associated with estuaries and coastal waters. In general, juveniles of the
flatfish Paralichthys orbignyanus are euryhaline (0-45) over a wide range of
temperatures. Lower oxygen consumption (0.1 mg g-l h- l) is associated with low
salinity and temperature (ltC; Wasielesky 1994). Juveniles of Paralichthys
orbignyanus and MugU platanus are also resistant (LC50 96 h) to elevated
ammonia (0.67 and 0.90 mg r\ respectively) and nitrite (30.57 and 80.97 mg r\
respectively) concentrations. Higher salinity increases the ammonium and
nitrite tolerance in Paralichthys orbignyanus and ammonium tolerance in
MugU platanus (Wasielesky 1994; Bianchini et al. 1996). Paralichthys orbign-
yanus has also low tolerance to pH stress. An increase of the ventilatory rate at
pH 5.2 to 4.7 and 100% mortality at pH 4.0 is probably related to an acid-base
disequilibrium, owing to plasmatic HC03-depletion (Wasielesky 1994).
Ichthyoplankton 51

4.12
Ichthyoplankton

C. Sinque and J. H. Muelbert

Species Composition and Abundance

The Patos Lagoon estuary plays an important role in the early life history of
many local fish species. The estuary serves as a nursery ground for coastal
and estuarine species. A variety of habitats, which provide an abundant
food supply and protection against predators, make this estuary a suitable
environment for the development of eggs and larvae (Weiss and Krug 1977;
Weiss and Souza 1977a,b; Castello and Krug 1978; Weiss 1981; Muelbert
1986; Pereira 1986; Muelbert and Weiss 1991). The dynamic nature of the
estuary also contributes to the sporadic presence of many oceanic species
in the ichthyoplankton.
Eggs and larvae of at least 29 fish species are found in the estuarine and
adjacent coastal waters (Appendix). These species belong to different eco-
logical categories (Chao et al. 1982), and their presence and abundance in
early stages of development reflect the degree to which they utilize the
estuary. Estuarine resident fish (those who complete their entire life-cycle
in the estuary) are represented by the eggs and larvae of the flatfish Achirus
garmani and by larvae of Atherinidae, Syngnathus folletti, Blenniidae,
Gobiidae, and Gobiesocidae. These species are not that abundant (0.4-
3.3%) and are usually found in shallow inland estuarine regions.
A large number of marine species depend on the estuary for their devel-
opment, such as those with an abundant number oflarvae, e.g. the Clupeidae
Brevoortia pectinata, the Engraulididae Lycengraulis grossidens (=L. olidus),
and the Sciaenidae Micropogonias furnieri. They are present in egg and larval
stages and together account for 88% of the eggs and 66% of the larvae found
in the estuary. A less abundant group, which only opportunistically uses the
estuary as a nursery ground, is present only in the larval stage. It includes
many commercially important species, like the Sciaenidae Macrodon
ancylodon, Menticirrhus spp., and Paralonchurus brasiliensis, and the
flatfishes Paralichthys sp. and Symphurus jenynsi. In addition, Trichiurus
lepturus, present in both egg and larval stages, can be included in this group.
Freshwater species, mainly the Siluriforme Parapimelodus nigribarbis, use the
estuary opportunistically during periods of intense freshwater runoff.
A last group of species, composed of larvae of marine origin, is occasionally
caught in more saline waters. These species only occur in the estuary under
conditipns of salt water intrusion. Included in this group are Anchoa marinii,
Ophyctus gomesi, Urophycis brasiliensis, Porichthys porosissimus, Epinephelus
sp., Prionotus punctatus, Cynoscion spp., Umbrina canosai, and Peprilus paru.
52 C. Sinque and J. H. Muelbert

Spatial Distribution

The degree of transport and the location of spawning are the key factors
controlling the spatial distribution of ichthyoplankton. With the exception
of the estuarine species Achirus garmanii, the eggs of all other fish species
in the estuary are of marine origin (Weiss 1981; Chao et al. 1982), and thus
eggs are generally more abundant closer to the mouth of the estuary. The
distribution within the estuary reflects spawning location and utilization of
the estuary as a nursery ground. Species such as Engraulis anchoita and
Anchoa marinii that spawn further offshore and do not depend on the
estuary for their development, are confined in their distribution to the
estuary inlet. Species like Micropogonias furnieri and Achirus garmani
which use the estuary during their development lay eggs distributed
throughout the estuary.
This pattern of horizontal distribution is also observed for larvae. The
larvae of oceanic species (Anchoa marinii, Porichthys porosissimus,
Prionotus punctatus, Synagrops sp., Parona signata, Cynoscion striatus,
Umbrina canosai, Mugil spp., Trichiurus lepturus, Peprilus paru, Symphurus
jenynsi) occur in the estuary during periods of strong saltwater intrusion
(Muelbert and Weiss 1991), whilst larvae of coastal water species (Brevoortia
pectinata, Lycengraulis grossidens (=1. olidus), Macrodon ancylodon,
Menticirrhus spp., Micropogonias furnieri, Paralonchurus brasiliensis) use the
entire estuary as a nursery area. Atherinidae, Syngnathus folletti, Blenniidae,
Gobiidae, and Achirus garmani reside and are uniformly distributed in the
Patos Lagoon estuary. The freshwater species Parapimelodus nigribarbis
exhibits a decrease in abundance towards the ocean.
The vertical distribution of fish eggs is influenced by their buoyancy and
by the salinity structure of the water column. In the Patos Lagoon estuary
the maximum number of eggs is related to salinities between 25 and 30
(Weiss 1981). Since these values generally correspond to surface waters
near the estuarine inlet and upper estuarine bottom waters, they induce a
vertical distribution pattern of the eggs and may reflect an ideal density for
egg buoyancy (Muelbert 1986). Although most fish larvae in the estuary
show a uniform vertical distribution, some groups of larvae seem to present
different patterns. Less saline surface waters favour Brevoortia pectinata,
Lycengraulis grossidens, Parapimelodus nigribarbis, and Atherinidae, while
Micropogonias furnieri and Trichiurus lepturus are frequently found in
more saline bottom waters. During anomalous periods of freshwater run-
off, this pattern may change, and Parapimelodus nigribarbis may become
more abundant at the bottom than near the surface (Muelbert and Weiss
1991). Based on the available information on the spatial distribution of
iththyoplankton, Weiss (1981) has proposed a general model for the
dynamics of fish eggs and larvae in the estuary of the Patos Lagoon
(Fig. 4.12.1).
 .. OCEAN ESTUARY LAGOON- &
~
o
1
::l

30 SALINITY
Fig. 4.12.1. General model summarlzmg the different early life-history stages of fish in the Patos Lagoon estuary.
The width of the arrows represents the relative abundance of different stages of development (egg stages: shaded;
VI
larvae: clear. Modified after Weiss 1981) V.l
 U1
>1>0
BREVOORTIA PECTINATA I_
LYCENGRAULIS SP. _ • •-
ENGRAULIS ANCHOITA

ANCHOA MARINI!
..
MICROPOGONIAS FURNIERI
J -
TRICHIURUS LEPTURUS

ACHIRUS GARMANI •

OTHERS I.- •
TOTAL I_
EGGS
I m-3J WINTER
• ..
SUMMER

Fig. 4.12.2. Seasonal distribution of fish egg abundance (eggs mO') during a 2-year cycle (1981-1983) o
U>
S·
.g
n>
§
p..
';-<

~
~
J:
6:
n>
:4
Ichthyoplankton 55

BREVOORTIA PECTINATA
lYCENGRAULIS SP.
ANCHOA MARINII
PARAPIMElODUS VAlENCIENSIS
PORICHTHYS POROS1SSIMUS
HYPORHAMPHUS KRONEI
SYNGNA THUS FOllETTI
MUGILSP.
ATHERINIDAE
PRIONOTUS PUNCTATUS
SYNAGROPS SP.
PARON A SIGNATA
MICROPOGONIAS FURNIERI
MACRODON ANCYLODON
PARALOCHURUS BRASILIENSIS
UMBRINA CANOSAI
CYNOSCION STRIA TUS
MENTIC1RRHUS SP.
BLENNIIDAE
TRICHIURUS LEPTURUS
PEPRILUS PARU
GOBIONELLUS SP.
GOBIOSOMA PARRI
PARALICHTHYS SP.
ACHIRUS GARMANI
SYMPHURUS JENYNSI
GOB1ESOX STRUMOSUS
OTHERS
TOTAL

Fig. 4.12.3. Seasonal distribution of larval abundance (larvae m-') during a 2-year cycle
(1981-1983 )

Temporal Distribution

Temperature rather than salinity seems to control the seasonal distribution

of spawning and of ichthyoplankton in the estuary. In general, the highest
abundance of eggs (Fig. 4.12.2) and larvae (Fig. 4.12.3) occurs during the
summer. With the exception of Engraulis anchoita, the eggs of all identified
56 J.P. Vieira and J.P. Castello

species are present in the spring (18%) and summer (80%). During the fall
and winter egg abundance and diversity are low. Eggs and larvae follow
similar seasonal distribution patterns (Fig. 4.12.3). The highest larval
abundance and diversity occur in the summer, and the lowest larval abun-
dance in the fall, with Brevoortia pectinata, Lycengraulis grossidens, and
Atherinidae as the dominant taxa, and Achirus garmani, Parapimelodus
nigribarbis, Hyporhamphus kronei, Syngnathus folletti, Mugil spp., Micro-
pogonias jurnieri, Paralonchurus brasiliensis, and Gobiosoma parri also
present. Although in winter and spring the ichthyoplankton concentration is
generally higher than in the fall, fewer species are present in the estuary.
Fish larvae in the Patos Lagoon estuary exhibit two different patterns of
seasonal distribution (Muelbert and Weiss 1991). One group of larvae is
present throughout the year, while another exhibits a discontinuous sea-
sonal cycle (Fig. 4.12.3). Brevoortia pectinata, Lycengraulis grossidens, and
Atherinidae compose the first group, suggesting that these species repro-
duce throughout the entire year. The second group predominates from
early spring to late summer and includes Micropogoilias furnieri, Macrodon
ancylodon, Blenniidae, Gobionellus spp., Peprilus paru, Trichiurus lepturus,
Paralichthys sp., and Achirus garmani.

4.13
Fish Fauna

J.P. Vieira and J.P. Castello

Estuarine Use and Reproductive Strategies

The abundant food resources and protection against predation provided by

estuarine shoals make estuaries ideal nursery grounds (Potter et al. 1983;
Loneragan et al. 1989; Vieira and Musick 1993, 1994) and may have deci-
sively influenced the evolution of migratory recruitment patterns of juve-
nile fish (Miller and Dunn 1980; Castello 1986; Vieira and Musick 1993,
1994). About two-thirds of the commercial fishery in the USA is based on
estuarine-related species (McHugh 1966, 1985), and similar estimates apply
in southern Brazil (Chao et al. 1986). Several commercially important
species utilize the Patos Lagoon estuary as their principal nursery during
part of their life-history, while others may use shelf areas under the influ-
ence of estuary runoff. Therefore, only a limited number of species is
entirely estuarine-dependent, but most species are facultative or oppor-
tunistic migrators, as has been observed elsewhere (Day et al. 1989).
:The ichthyofauna of the Patos Lagoon estuary is composed of approxi-
mately 110 species (Appendix; Chao et al. 1982, 1985; Vieira 1991a), but
only a few of these are abundant or frequent. Depending on whether
species complete their entire life-cycle in the estuary (estuarine residents
Fish Fauna 57

sensu McHugh 1967) or whether during specific life-cycle phases marine

and freshwater species migrate into the estuary (estuarine-dependent sensu
Blaber et al. 1989; estuarine-opportunists sensu Lenanton and Potter 1987),
seven categories oflocal estuarine use by fish can be distinguished.
Estuarine residents are represented by Odontesthes argentinensis, Atherinella
(=Xenomelaniris) brasiliensis, Jenynsia lineata, Ramnogaster arcuata,
and several species of the Blenniidae, Gobiidae, and Poecilidae which are
largely restricted to estuarine shoals, whilst Achirus garmanii and Genidens
genidens are common to deeper waters. The marine species Mugil platanus,
Paralichthys orbygnianus, and the sciaenids Pogonias cromis and Micropogonias
furnieri spawn in the sea or in waters proximate to the estuary, but their
post-larval, juvenile, and sub-adult life-cycle stages seem to be obligatory
users of estuarine shoals and deeper waters as nursery grounds. Early life-
cycle stages of another group of marine species (Macrodon ancylodon,
Menticirrhus americanus, Paralonchurus brasiliensis, Symphurus jenynsi,
Porichthys porosissimus, Urophycis brasiliensis, Lye.engraulis grossidens,
Anchoa marin ii, Trichiurus lepturus, Pomatomus saltatrix) use deeper
estuarine waters only facultatively or opportunistically as a nursery.
However, under favourable conditions sub-adults may persist in the estuary
throughout the year. Yet another considerable number of marine species
(like Cynoscion guatucupa, Umbrina canosai, Ctenosciaena gracilicirrhus,
Prionotus punctatus, Peprilus paru, Lagocephalus laevigatus, and Trachinotus
marginatus) occasionally penetrate the estuary under favourable conditions
as juveniles or sub-adults. Netuma barba and Netuma planifrons are
anadromic species which spend the major part of their life-cycle in the sea
but migrate through the estuary to reproduce in limnic and pre-limnic zones.
The juvenile stages of these species also use the estuary as a nursery and
feeding ground. Additionally, juveniles and sub-adults of the freshwater
species Parapimelodus nigribarbis and Pimelodus maculatus use the estuary
opportunistically. Especially during periods of intense freshwater flushing,
their eggs, larvae, and juveniles spread throughout the entire estuarine area.
Finally, a large group of freshwater (chiclids and charachiforms) and tropical
marine species (Carangidae, Serranidae, Pomacentridae, Gerreidae,
Balistidae), coastal elasmobranchs, and anguiliforms are occasional visitors
to the estuary.

Specific Diversity and Space-Time Segregation

Most of the species which compose the fish assemblage in the Patos Lagoon
estuary are of marine origin. In general, the type of fish encountered and
migration patterns of their life-cycle phases are similar to those observed in
estuaries elsewhere (Vieira and Musick 1994). Only a few species dominate
the assemblage and are commonly caught by one or more fishing methods
58 J.P. Vieira and J.P. Castello

throughout the year (Chao et al. 1982, 1985). Of the approximately

110 species (Chao et al. 1985),22 are caught by all fishing methods employed
(i.e. bottom trawl, mid-water trawl, beach seine). The bottom trawl method
nets 81 species. Micropogonias jurnieri, Netuma barba, and Genidens
genidens alone total more than 50% of the catch, whilst seven sciaenids, two
ariids, and one batrachoid represent 90% in number and weight. The mid-
water trawl method catches 43 species, all of which are also found in bottom
trawls. Although the proportion of engraulidids and Trichiurus lepturus
increases in mid-water trawls, Micropogonias jurnieri continues to be an
important member. The species (n = 61) composition of beach seine catches
differs from the other two methods. Atherinidae, Jenynsiidae ;-::: Anablepidae,
and juveniles of Mugilidae represent more than 70% of the catch in numbers,
though Micropogonias jurnieri is still an important contributor.
Based on the fishing methods employed and on the spatial and temporal
distribution of species, the estuarine ichthyofauna can be divided into
deeper water, shallow water, and pelagic associ'!tions (Vieira and Musick
1994). Depth gradients further separate the deeper water association into a
group below 4 m depth and another between 2 and 4 m. The major compo-
nents of the deeper water estuarine association are bottom-oriented
epibenthic and demersal fish, like Micropogonias jurnieri, Netuma barba,

" CJ BOITOM TRAWL (> 4 m)

BOITOM TRAWL « 4 m)
BEACH SEINE « 2 m)

Fig. 4.13.1. Size

frequency distribution
(lO-mm size class) of
all species caught by
o 40 80 120 160 200 240 bottom trawls and
beach seines at
20 60 100 140 180 220 >250 different depths in the
TOTAllENGHT CLASS (mm) Patos Lagoon estuary
Fish Fauna 59

Genidens genidens, and several species of Sciaenidae (Macrodon ancylodon,

Menticirrhus american us, Paralonchurus brasiliensis, Cynoscion guatucupa,
Umbrina canosai), which comprise an important portion of the southern
Brazilian fishing industry. Species collected by bottom trawls in channels
and adjacent areas (>4 m depth) are largely represented by post-larvae,
young-of-the-year, and sub-adults between 10 and 250 mm total length
(TL) (Fig. 4.13.1). Several young-of-the-year sciaenid species and Porichthys
porosissimus, which uses the estuary as a nursery ground, may reach high
abundance in the summer. However, throughout the year juveniles and
sub-adults of Micropogonias furnieri, Netuma spp. and sub-adults of
Paralonchurus brasiliensis, Macrodon ancylodon, and Menticirrhus americanus,
which use the Patos Lagoon as a feeding ground, represent the dominant
component in channels and adjacent areas (Fig. 4.13.2a). With the
exception of Netuma barba, all dominant species are represented by
individuals of less than 70 mm TL, and several sciaenids (e.g. Ctenosciaena
gracilicirrhus, Umbrina canosai, Cynoscion guatucupa} are restricted to this
size class. Netuma barba and Anchoa marinii dominate the size class
between 70-90 mm TL, whilst larger (>90 mm TL) individuals are mostly
represented by the sciaenids (Micropogonias furnieri, Paralonchurus
brasiliensis, Macrodon ancylodon, Menticirrhus americanus), and Ariidae
(Netuma barba, Genidens genidens). Species collected by bottom trawls in
open estuarine waters (2-4 m depth) range from 10 and 250 mm TL (Fig. 4.13.1;
4.13.2b), and Netuma barba and Anchoa marinii dominate the size class
between 70 and 90 mm TL. However, individuals below 70 mm TL are less
abundant and are principally represented by Micropogonias furnieri. The
larger size class (> 100 mm TL) is mainly composed of Micropogonias
furnieri, Netuma barba, and Genidens genidens.
Fish species of the shallow water association «2 m) caught by beach
seines (littoral fishes sensu Day et al. 1989) are typically dominated by small
estuarine resident fish (Atherinidae, Jenynsiidae = Anablepidae) and juve-
niles of estuarine-dependent marine species of the Mugilidae and Micro-
pogonias furnieri (5 to <200 mm TL; Figs. 4.13.1; 4.13.2c). Together, these
species make up 90% in number of beach seine catches and display three
clearly defined annual abundance peaks (Chao et al. 1985). Young indi-
viduals of three species of Mugilidae «50 mm TL) compose 42% (M.
platanus 34.6%) of the catch. Mugil curema and Mugil gaimardianus occur
during summer/fall, whilst Mugil platanus is present throughout the year,
with a peak abundance from late winter to early summer. However,
although absent from bottom trawl or beach seine samples, large mono-
specific schools of mature adults of Mugil platanus are abundant during
the late summer and fall (Vieira and Scalabrin 1991). Both juveniles and
adults of Mugil platanus feed on diatoms, bacteria, and associated inor-
ganic matter (Vieira 1991b). Species of the Atherinidae are present
throughout the year and total about 30% of the catch. The peak abundance
60 J.P. Vieira and J.P. Castello
100 -
80 - - ENGRAU
e=a PORPOR
- 60 -
= MENAME
!l ~o -
:- =t UMBCAN
... 10
.::... . . PARBRA
;/)
0 =CTEGRA
Z
<t: 20
= MACANC
z = CYNGUA
40
~ - NETBAR
;::::
bO - MICFUR
30
100

100
-;;- 80
~'

;/) - ENGRAU
~ 60 ",. POGCRO
~ 40 =GENGEN
:n 20
- NETBAR
Z - MICFUR
<t: 0
~
~ 20
0 40
0
60
80
100

-;;-
-
0'
100
80
60
-
=
c::::rI

=
GOBSHU
CLUPEI
EUCGUL
ODOARG
=

--
; /)
IENLIN
40
=
:.:.J
:.J MUGPLA
iJ 20 AIHBRA
:n MICFUR
0
Z
<: 20
~
40
~~ 60
SO
100
0 20 -to 60 80 lOO 120 140 160 180 200 220 24~250
TOTAL LENGHr ClASS (mm)

Fig. 4.13.2.
Fish Fauna 61

of Atherinella brasiliensis (22.6%) and Odontesthes argentinensis (7.7%)

occurs during the fall/winter and winter/spring, respectively, the first
always preceding the second by 1 or 2 months. The size classes of these
species rarely coincide in time and space (Bemvenuti 1984), probably
owing to similar feeding habits. Juveniles «70 mm TL) of both species are
plankton feeders, with a preference for phytoplankton followed by zoo-
plankton, whilst adults graze upon benthic organisms (Bemvenuti 1990).
The two abundance peaks of Jenynsia lineata (20.9%) coincide with the
species' reproductive cycle between the spring and fall (Chao et al. 1985). This
small omnivorous species appears to lack ontogenetic changes in diet
preference and grazes principally upon benthic vegetation (Betito 1984).
Species of the pelagic association swim freely through the entire water
column and are sampled by mid-water trawls, although they are also
frequently caught by bottom trawls and beach seines. The engraulidids
«120 mm LT) Lycengraulis grossidens, Anchoa marinii, Engraulis
anchoita, and juveniles of Peprilus paru and Selene ~et-apinis total 40% of
mid-water trawl catches.' Young individuals of Trichiurus lepturus also
form an important component of the pelagic association. Plankton-feeding
juveniles of the clupeids Brevoortia pectinata, Ramnogaster arcuata, and
Platanichthys platana are abundant in somewhat shallower areas.
Engraulidids are plankton feeders and partition the available resources
through differential occupation of space during the year. Juveniles of
Engraulis anchoita enter the estuary during the fall and winter, but Anchoa
marinii and Lycengraulis grossidens are present throughout the year.
Salinity distributions in the estuary are largely responsible for the vertical
and horizontal segregation of the two latter species (Buckup 1984). Salinity
also appears to segregate different age cohorts of Lycengraulis grossidens,
with larvae preferring oligohaline waters for their development (Castello
and Krug 1978). Top predators, such as Pomatomidae, Trichiuridae, and
several Sciaenidae, are often abundant members of the pelagic association
in large estuaries (Vieira and Musick 1994). Since these species are scarce
or absent from the pelagic association in the Patos Lagoon estuary, it is
likely that the spatial and temporal distribution of the local ichthyofauna is
controlled by environmental factors and competition for available food
resources, rather than by predation.
4
Fig. 4.13.2. Contribution of dominant species at different total length classes in A bottom
trawls at more than 4 m depth; B bottom trawls at less than 4 m depth; and C shallow water
«2 m) beach seine catches. ATHBRAAtherinella brasiliensis; CLUPEI Brevoortia pectinata,
Ramnogaster arcuata, Platanichthys platana; CTEGRA Ctenosdaena gradlidrrhus; ENGRAU
Anchoa marinii, Lycengraulis grossidens; EUCGUL Eudnostomus gula; GENGEN Genidens
genide1fs; GOBSHU Gobionellus shufeldti; JENLIN Jenynsia lineata; MACANC Macrodon
ancylodon; MENAME Menticirrhus americanus; MICFUR Micropogonias jurnieri;
MUGPLA Mugil platanus; NETBAR Netuma barba; ODOARG Odontesthes argentinensis;
PARBRA Paralonchurus brasiliensis; POGCRO Pogonias cromis; PORPOR Porichthys
porosissimus; CYNGUA Cynoscion guatucupa; UMBCAN Umbrina canosai
62 C.M. Vooren: Bird Fauna

4.14
Bird Fauna

C.M. Vooren

The open waters and protected shallow bays of the estuary and the wet-
lands and beaches at their margins comprise habitats which in the last
century contained a bird fauna so abundant and diverse as to draw the
attention of visiting scientists and explorers (Appendix). Von Ihering
(1887) states that highly abundant breeding birds such as the black-necked
swan (Cygnus melancorhyphus), the roseate spoonbill (Ajaia ajaja), gulls
and terns of the genera Larus and Sterna, carao (Araunus guarauna), grebes
of the genus Podiceps, and several species of Anatidae and Threskiornitidae
were exploited commercially for feathers, eggs, and meat, and that already
in those days a great decline in their abundance was noted, especially of the
black-necked swan. In spite of all this, the bird" fauna of the myxohaline
waters and shores of the Patos Lagoon estuary has never been studied
systematically. Some general features may be deduced from the studies on
estuarine and marine birds in general (Escalante 1970, 1983; Barattini and
Escalante 1971), from the reviews by Belton (1984, 1985) and Sick (1984),
from the study on the shorebirds and coastal marine birds of Cassino Beach
(Vooren and Chiaradia 1990), and from unpublished personal observations
in the estuary. Thirteen species of birds are common or otherwise of interest,
and are discussed below.
The area supports six species of piscivorous birds. The bigua cormorant
Phalacrocorax olivaceus, one of the most common birds in the estuary,
fishes by diving in shallow water. No breeding places of this bird are known
in or near the area. Trudeau's tern (Sterna trudeaui), common tern (Sterna
hirundo), and black skimmer (Rynchops niger) prey on fishes at the water
surface. Sterna trudeaui probably breeds in the nearby inland marshes,
although Belton (1984) was unable to confirm this, while Sterna hirundo is
a summer visitor from North America (Vooren and Chiaradia 1990). Both
these terns feed mostly at sea but fish in the estuary when the sea is rough.
Sterna hirundo roosts in numbers of up to 5000 birds on a beach in the
estuarine inlet. Rynchops niger roosts in flocks of up to 300 birds on sand
banks in the same area. This species does not breed in the area, and it is not
known from where the birds come. The birds fish mostly at night
(Escalante 1970), probably in the estuary. The snowy egret (Egretta thula)
and the green heron (Butorides striatus) breed in nearby marshes and fish
habitually in the shallow water of estuarine margins and creeks. Cygnus
melancorhyphus breeds in marshes south of the estuary (Taim ecological
reserve), and groups are occasionally seen in the estuary. The species is
herbivorous and may graze on beds of Ruppia maritima, but this remains
to be confirmed. The latter is also true for the red-gartered coot (Fulica
Marine Mammals 63

armillata), which is common in the estuary, probably breeds in the area,

and is mostly herbivorous (Escalante 1983). The brown-hooded gull (Larus
maculipennis) breeds in nearby inland marshes. This gull scavenges in
large numbers in the fishing harbours and at sewage outlets in the estuary,
and roosts at night in protected bays of islands. The kelp gull (Larus
dominicanus) is also a scavenger but feeds mostly on the seashore and only
enters the lower reaches of the estuary to roost in large numbers on the
beaches. The species breeds on the coast of Uruguay and is a non-breeding
visitor to this area. The muddy shore and banks of the inner reaches of
lower estuarine bays are used by waders. The hudsonian godwit (Limosa
haemastica) from North America, the rufous-chested dotterel (Charadrius
modestus) from Patagonia, and the common stilt (Himantopus himantopus)
from nearby inland breeding areas have been seen there. These and other
waders may also use such areas in other bays of the estuary. The small
islands of the estuary are important as roosting places for gulls and herons.
Rynchops niger roost in large numbers on the sandy peninsula of the inlet.
Adjacent salt marshes and the sandy beaches which border them are
important roosting places for Sterna trudeaui and Larus dominicanus and
are at high tide and rough weather a refuge for sandpipers and plovers
from the seashore. These critical areas need to be studied in detail so that
measures for their conservation can be taken.

4.15
Marine Mammals

M.e. Pinedo

Species Composition and Movements

The bottlenose dolphin (Tursiops truncatus; Pinedo et al. 1992), previously

referred to as Tursiops gephyreus (Castello and Pinedo 1977a; Pinedo 1982,
1986), is common in the Patos Lagoon estuary. Between 31 (Moller et al.
1994) and approximately 100 individuals (generally in groups of 3-4) enter
the estuary throughout the year (Castello and Pinedo 1977a; Pinedo 1982),
though they are more frequent in May and November (Pinedo, unpubl.).
Since the species tolerates low salinities (2-3; Castello and Pinedo 1977a),
specimens may reach the limits of the upper estuary near the Sao Gon<;:alo
Channel. Individuals of the sea lion (Otaria flavescens) only occasionally
enter the estuary, but in contrast to the bottlenose dolphin, specimens of
Otaria flavescens may occasionally reach the upper limits of the lagoon
near Porto Alegre, probably in search of food (Pinedo 1990). Although
Pontoporia blainvillei has been reported in the estuary (von Ihering 1892;
Cabrera and Yepes 1940; Carvalho 1961, 1975) the species has not been
seen since 1976.
64 M.e. Pinedo: Marine Mammals

Feeding and Behaviour

Various types of behaviour, like leaps out of the water, tail beating of the
surface, belly exposure, or vertical body positioning are displayed by
Tursiops truncatus (Castello and Pinedo 1977a). Rapid displacements and
occasional leaps always precede the capture of fish. The most common prey
is Micropogonias jurnieri, followed by Trichiurus lepturus and Trachinotus
spp. These species are first bitten and, after floating on the surface, ingested
(Pinedo 1982). Their abundance is likely to influence the occurrence of
dolphins in the estuary (Castello and Pinedo 1977a). A cooperative fishing
method between bottlenose dolphins and fishermen, using a circular nylon
throw-net, is practised at 28° 29' S (Pryor et al. 1990) but has not been
observed in the Patos Lagoon. On rare occasions Tursiops truncatus may be
caught in gill nets set for shrimp fishery in the estuary (Castello and Pinedo
1977a).
5 Energy Flow and Habitats
in the Patos Lagoon Estuary

5.1
Primary Production Cycles

U. Seeliger, C.S.B. Costa, and P.C. Abreu

The functional primary producers in the Patos Lagoon estuary include

emersed and submersed macrophytes, benthic and floating macro algae,
cyanobacteria, and epibenthic, epiphytic, and planktonic microalgae. All
supply food for estuarine and coastal consumers and interact with estuarine
chemical cycles; some perform critical habitat functions during different life
stages of inumerous other organisms (Kennish 1986; Day et al. 1989). In the
Patos Lagoon estuary, the various primary producers display different
production patterns, and the rates of production are likely to vary in
response to biotic and abiotic conditions; thus, the availability of live organic
matter and detritus may fluctuate considerably within and between years.

1966
JANUARY 100 68 66
FEBRUARY 100 66 91
MARCH 135 51 75
APRIL 65 ,~ 66
MAY fID" 106 ' 83
JUNE , I

JULY " 66,' 70

AUGUST 125 146 wf 81
SEPTEMBER 100 136 110 66 126 Ql0
OCTOBER 95 111 66 40 35 113 \Q]
NOVEMBER 55 50 93 75 37 75 83
DECEMBER 135 35 100 50 11 6 70 60
TOTAL 1248 1088 864 {Z92l 1033 ~ 1102 1067 978

Fig. 5.1.1. Minima and maxima of mean monthly precipitaton between 1966-1975 in the
Patos-Mirim Lagoon watershed

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
66 U. Seeliger, C.S.B. Costa, and P.C. Abreu

In most estuaries physico-chemical factors vary predictably on an hourly

to seasonal scale because they are largely controlled by regular tidal and
seasonal flushing cycles (Wolfe and Kjerfve 1986). In contrast, the choked
lagoon characteristics and negligible tidal amplitude make physico-chemi-
cal parameters in the Patos Lagoon estuary largely dependent on wind and
rainfall. Precipitation patterns also tend to vary from year to year
(Fig. 5.1.1; Castello and Moller 1978), owing to the influence of EI Nino and
anti-El Nino phenomena on the climate in the southwestern Atlantic (Arntz
and Fahrbach 1991; IAI 1992). As a consequence, the unpredictable short-
term variations of physico-chemical factors with their irregular amplitude
and timing, their reduced seasonal oscillation, and their pronounced inter-
annual variability become important abiotic forcing factors in the Patos
Lagoon estuary. Further-more, the physiographic heterogeneity of the estu-
ary with deeper open waters and protected, semi-open, and open shallow
marginal bays induces different site-specific environmental characteristics.
Common to all sites are seasonal cycles of surfacewater temperature and
incident light (Coutinho 1982; Cafruni 1983) and generally reduced and
highly unstable underwater light conditions, owing to frequent resuspen-
sion offine sediments by winds (Costa et al. 1988a; Moreno 1994). Differ-
ences between sites are largely due to hydrology, degree of wind exposure,
and sediment characteristics.
Phytoplankton biomass and production in the estuary, with a mean con-
centration of 0.07 g C m- 2 (0.03-7.0 g C m-2; Abreu, unpubl.) and production
rate of 0.072 g C m-2 h-' (0.004-0.320 g C m-2 h-'; Abreu et al. 1994a), are
largely determined by the contribution of flagellates «20 !lm). Peak bio-
mass occurs in the spring and summer, while values close to zero are com-
mon in the winter and fall when heterotrophic processes prevail. Seasonal
variations of phytoplankton biomass and production are principally
related to insolation and temperature. The lowest prima2 production
values tend to coincide with water temperatures below 20 C and mean
water column irradiance below the critical light intensity (Riley 1967) of
40 gcal cm-2 day' (Abreu et al. 1994a). Furthermore, especially during the
summer, the phytoplankton biomass is influenced by zooplankton grazing.
Although grazing will reduce the biomass, it seems to stimulate phyto-
plankton productivity per unit of biomass, owing to the increase in avail-
able light and higher nutrient concentrations (Abreu et al. 1994b).
In contrast to tide-controlled estuaries, both the short-and long-term
phytoplankton dynamics in the Patos Lagoon estuary are directly related to
hydrological and meteorological events. Since short-term chlorophyll a
variability is a function of the residence time of water in the estuary, which
is controlled in turn by wind direction and velocity, high chlorophyll a
biomass arises during runoff after prolonged periods of salt water penetra-
tion (Abreu 1987). Long-term interannual chlorophyll a oscillations
depend on the precipitation patterns in the watershed and estuarine
Primary Production Cycles 67

region. Phytoplankton biomass and production peaks are closely related to

the input of high nitrogenous estuarine nutrient concentrations after ele-
vated rainfall. Since large-scale El Nino events augment precipitation in
southern Brazil, they are likely to promote increased estuarine phyto-
plankton biomass and production (Abreu et al. 1994a; Proenc;:a et al. 1994;
Bergesch et al. 1995)).
The considerable biomass (6-22 g C m-2 ) of benthic microphytes is
dominated by pennate diatoms. After resuspension in the water column
during high incident light, the photosynthesis of benthic cells is inhibited,
suggesting their adaptation to low light conditions (Abreu 1992). The vari-
ability of mean light intensities seems to be the key controlling factor for
these organisms and might explain why phytobenthic cell peaks occur
during the winter, contrary to maximum phytoplankton biomass in the
spring and summer (Bergesch et al. 1995).
Benthic macro algae are responsible for most of the biomass production
in subtidal areas during the winter and early spring. The highest mean
monthly biomass of the dominant Enteromorpha sp. (34.3 g C m-2 ), Rhizo-
clonium riparium (308 g C m-2 ), Ulothrix flacca (2.2 g C m-2 ), and Vaucheria
longicaulis (70.4 g C m-2 ) is linked to regional light and temperature
regimes and to the site-specific availability, permanence, and exposure of
suitable substrate (Coutinho and Seeliger 1986). Throughout the year
cyanobacteria represent the largest single component of the subtidal flora,
with dominance ofLyngbya confervoides in the summer (peak = 55.7 g C m-2)
and Microcoleus chthomoplastes in the fall (peak = 19.8 g C m-2; Coutinho
1982; Coutinho and Seeliger 1986). Although the primary productivity
values of cyanobacteria in saltmarsh environments are lacking, carbon and
nitrogen fixation levels are probably high (Yunes et al. 1990).
Despite the local occurrence of Zannichellia palustris during prolonged
periods of low salinity (Cafruni et al. 1978; Cafruni 1983; Moreno 1994),
Ruppia maritima is responsible for most of the submersed spermatophyte
biomass formation. Rather than being a stable feature of the benthos,
Ruppia maritima may exhibit either perennial growth or annual growth
cycles with massive die-off during the fall, though approximately seasonal
cycles of biomass formation are most common (Cafruni 1983; Asmus et al.
1984; Moreno 1994). Biomass peaks may reach 82.8 g C m-" but the average
value in the estuary is approximately.1O.8 g C m-2, though it may vary
greatly from site to site (Cafruni 1983; Moreno 1994). Data on Ruppia pro-
ductivity are still lacking, but preliminary estimates indicate a net annual
production of 39.6-43.2 g C m- yeaf 1 (Seeliger, unpubl.). Ruppia maritima
also serves as the principal semi-permanent substrate for benthic micro-
and macro algae in the estuary (Ferreira and Seeliger 1985). Since abiotic
conditions, the availability of colonizer species, size and density of Ruppia
beds, and age and turnover of leaves influence epiphyte biomass, produc-
tion patterns are likely to be highly variable. Epiphyte biomass (113.1 g C m-2 )
68 U. Seeliger, C.S.B. Costa, and P.C. Abreu

grows towards the summer and fall and represents between 50 and 70% of
the total Ruppia maritima biomass (Coutinho 1982; Seeliger, unpubl.),
though occasionally it may exceed 100%.
The species-specific seasonal growth cycles (Fig. 5.1.2) of dominant
marsh species with either C3 (Scirpus maritimus) or C4 (Spartina alterniflora,
Spartina densiflora) carbon metabolism are likely to be a function of
different air temperature optima (Rozema et al. 1991). Lower air tempera-
tures «15°C) favour rapid vegetative growth of Scirpus maritimus during
the early spring, followed by peak live aerial biomass at the end of spring
and maximum dead and total aerial biomass at the end of summer (Costa,
in press). In contrast, the highest vegetative growth ofleavesand shoots of
Spartina alterniflora (Cunha 1994) and Spartina densiflora (Silva et al.
1993) coincides with the end of spring when temperatures surpass 20°C,
whilst peak dead and total aerial biomass occur in the early fall and during
fall-winter, respectively. In addition, interannual temperature differences
may decisively affect primary production of the three dominant marsh
species (Fig. 5.1.2). A mean temperature decrease of2.8°C during the growing
season results in 11 % and 33% reduction of aboveground and total Spartina
alterniflora primary production, respectively (Cunha 1994). Mean net above-
ground production ranges from 287.7 to 808.4 g C m o2 year"1 and is

SPECIES MONTHS NAPP

--,-_",---"",--....o...--",ML-L
J_ ....l_"'O""---"'L-.~---"'_.l..L- (gC ai' year' )

SCIRPUS MARCIlMUS

SPAlUINA ALTERNIFLORA
288-412
(SHOn FORM)

SPARTINA DENSIFLORA
1111111111111:1 669
(FREQUENTLY FLOODED)

SPARTINA DENSIFLORA
(OCCASIONAllY FLOODED) I 111~111~1111 714

:::::J VEGETATIVE GROWIH IS DEAD BIOMASS PEAK -TOTAL BIOMASS PEAK

-HIGH GRowm RATE illIIJll LIVE BIOMASS PEAK

Fig. 5.1.2. Aerial growth and biomass cycles and net aerial primary production (NAPP)
estimates based on dry weight to carbon conversion (0.43) of dominant salt marsh
plants in the Patos Lagoon estuary
Primary Production Cycles 69

comparable to highly productive, tide-controlled Spartina alterniflora

marshes elsewhere (Turner 1976; Long and Mason 1983; Adam 1990). Net
beloW'ground production of 727.9-2095.4 g C m-2 year-1 and 1510.2-2282.9 g C
m-2 year- 1 of short Spartina alterniflora (Cunha 1994) and Scirpus maritimus
(Costa, in press), respectively, represent more than 70% of their total net
primary production.
Conservative estimates of net primary production based on maximum and
minimum biomass difference (Roberts et al. 1993) suggest that the dominant
salt marsh plants, macroalgae, and cyanobacteria together are responsible for
as much as 86% of autochthonous carbon addition to the estuary, though
much of the annual marsh production may be recycled in situ,owing to
irregular flooding events (Costa, unpubl). If the annual biomass turnover rates

A S o N o J F M A M J J
MONTHS

Fig. 5.1.3. Total monthly organic carbon contribution by marsh plants (A), epibenthic
micro algae (B), cyanobacteria (C), benthic macroalgae (D), epiphytic micro algae (E),
phytoplankton (F), and Ruppia maritima (G) in the Patos Lagoon estuary
70 C.E. Bemvenuti: Trophic Structure

reported for salt marsh plants (Schubauer and Hopkinson 1984; Costa, in
press), seagrasses (Hillman et al. 1989), and benthic macro algae (Luning
1990) are applied, net annual carbon contributions increase even more. In
particular, the net annual contributions of phytoplankton will be enhanced
by turnover rates of up to 50 times (Abreu et al. 1994a).
Carbon contributions to the Patos Lagoon estuary are the result of both
year-round supply by some producer groups (Le. salt marshes) and
sequential biomass pulses by others (Le. macroalgae, Ruppia maritima, and
epiphytes; Fig. 5.1.3). The apparent seasonal programming of carbon fuca-
tion is one of the functional processes that sustains high estuarine produc-
tion (Rojas-Galaviz et al. 1992). Some producer groups (Le. Ruppia maritima
and salt marsh plants) will also enhance biomass generation by other
primary producers via substrate formation, reduction in hydrodynamics,
and sediment-water column modifications. The combined influence of
different primary producers on biomass formation and decomposition
processes with different duration leads to a highautochthonous carbon
supply throughout the year. Large interannual changes of total annual
primary production and subsequent detrital production may have pro-
nounced effects on consumers and the ecology of the estuary as a whole.

5.2
Trophic Structure

C.E. Bemvenuti

Different primary producer groups play an important role in the trophic

structure of estuaries, owing either to their supply of diverse grazing options
or to their production of detrital matter. Therefore, the trophic web in
estuaries is principally composed of interrelated grazing and detrital food
chains. The discrimination between food chains is more apparent at the
primary consumer level because the food of secondary consumers is often
derived from either source. In estuaries, detrital food chains appear to be
more important (Kennish 1990), and organic detritus particles with attached
microbial organisms typically constitute the lowest trophic level. Large
quantities of detrital material at different stages of decomposition are supplied
by emersed and submersed macrophytes, benthic and floating macroalgae,
cyanobacteria, and epibenthic and epiphytic micro algae. Additionally, part of
the net phytoplankton production is incorporated into the bottom detritus
pool. Although energy contributions by the different primary producers and
detrital fractions to the various animal groups have not yet been quantified in
the Patos Lagoon estuary, studies on food chain partitioning indicate their
differential role in different estuarine habitats (Fig. 5.2.1).
Organic detritus particles represent a large portion of the diet of macro-
benthic invertebrates and the meiobenthos, and thus these organisms are
Trophic Structure 71

likely to provide the major link of energy flow in an oligohaline estuary. As in

estuaries elsewhere (Day et al. 1989), the meiobenthic community in soft-
bottom substrates of the Patos Lagoon estuary is dominated by nematodes
and ostracods, followed by harpacticoid copepods and turbelarians. Some
nematode species feed on detritus and microbial organisms in deeper
anaerobic layers and therefore facilitate the energy flow to upper aerobic
sediments. Most meiobenthic organisms, however, feed on detrital particles
in superficial sediments and may be consequently ingested by deposit feeders
(Coull and Bell 1979) or selectively preyed upon by epibenthic crustaceans
and fishes (Castel 1992). Because meiobenthic organisms are very abundant
and have high population turnover rates, they assume an important function
at specific trophic levels in the estuarine detritus food chain (Day et al. 1989;
Kennish 1990; Castel 1992).
Macrobenthic invertebrates display diverse feeding habits (Bemvenuti
1983, 1992; Asmus 1984), but detritus appears to be an obligate food item
for most species. Infaunal deposit feeders of the first-(:onsumer level, like
Laeonereis acuta, ingest non-selectively detritus particles of surface layers,
whilst the burrower Heteromastus similis feeds on organic matter in deeper
«15 cm) galleries (Bemvenuti 1988). Similarly, the burrowing suspension
feeders Tagelus plebeius and KaUiapseudes schubartii consume detritus and

HERBIVORES

11111 PHYTOPLANKTON
~

1
AND EPIBENTHIC
MICRO-ALGAE
4t:tiR~CI
...
"
MACROALGAE
II ,..~

~~
2"' CONSUMERS
_.: ... 1'r~
" ,,1',,1.'1,:- ,
II

t
Fig. 5.2.1. conceptual diagram of trophic relationships in the Patos Lagoon estuary
72 C.E. Bemvenuti: Trophic Structure

epibenthic micro algae in soft sediments (Capitoli et al. 1978). Predation of

the infaunal polychaete Nephtys fluviatilis on Heteromastus similis repre-
sents an important intermediate link between the non-predatory infauna
and epifaunal macropredators, which aids in the transfer of energy to
higher trophic levels (Bemvenuti 1994). Infaunal predators frequently
regulate the abundance of other non-predatory infaunal organisms, leading
to the establishment of multiple trophic levels in soft-bottom sediments
(Ambrose 1984a,b; Commito and Schrader 1985). In the Patos Lagoon
estuary, however, low infaunal diversity is likely to be responsible for
abbreviated food chains in soft-bottom substrates (Bemvenuti 1994).
The first consumer level of the epifauna is dominated by Heleobia
australis, which feeds on microbes in superficial sediments and on sub-
mersed macrophyte leaves. Tanais stanfordii largely ingests epibenthic
diatoms, but the diet of the amphipod Melita mangrovi also includes
macro algae and copepods (Asmus 1984). The infaunal Kalliapseudes
schubartii (Araujo 1984; Asmus 1984; Bemvenuti 1990) and epifaunal
isopods, amphipods; and tanaidaceans (Reise 1985) are an important food
source for secondary consumers. In particular, the abundance of epifaunal
organisms, owing to their exposure, may provide a measure of predation
pressure by fish and decapods in the estuary (Bemvenuti 1987b). Among
the fishes, primary consumers are represented by deposit-feeding species
of the genus Mugil which ingest large quantities of epiphytes and detrital
matter {Vieira 1985; Fig. 5.2.1}.
Several highly motile decapods with diverse diet preferences are able to
act at different consumer levels. The marsh crabs Chasmagnathus granulata
and Metasesarma rubripes either share detrital food resources with
infaunal and sedentary epifaunal species or alternatively prey on these
organisms (Capitoli et al. 1977; Asmus 1984). Other decapods, like
Cyrtograpsus angulatus, Callinectes sapidus, and Penaeus paulensis, may
shift their food preferences as they grow. After intense recruitment into
shallow vegetated bottoms, their early life stages feed on detrital matter and
attached microbes (Bemvenuti 1983; Asmus 1984; Capitoli and Ortega
1993). As they grow, juveniles (5-15 mm) of Callinectes sapidus, Cyrtograpsus
angulatus, and the small crab Rhithropanopeus harrissi will also ingest
ostracods and nematodes of the meiofauna, as well as epifaunal peracarids
and small infaunal organisms (Bemvenuti 1992). Larger individuals of
young decapods increasingly prey on macro benthic organisms and, like
Callinectes sapidus, may ingest other decapods or small fish. In general,
pre-adults and adults of decapods are typically omnivorous and opportun-
istic feeders, which exploit different trophic levels (Asmus 1984) as food
it~ms become available (Bemvenuti 1983, 1992).
Open water estuarine areas, with a deeper (2-6 m) water column and
strong hydrodynamic activity, are comprised of soft bottoms with low
organic matter concentrations. Under these conditions, the dominance of the
Irregularly Flooded Marginal Marshes 73

suspension feeders Erodona mactroides and Balanus improvisus suggests an

increasing importance of the grazing food chain. Owing to its high abundance
and wide distribution, Erodona mactroides is the most important infaunal
phytoplankton grazer (Bemvenuti et al. 1992), whilst fish of the clupeid and
engraulidid families are prominent zooplankton feeders (Fig. 5.2.1).
In limnic areas of the Patos Lagoon, Erodona mactroides occurs in all size
classes and represents the greatest infaunal biomass (Bemvenuti 1987a). The
larger individuals are the major diet items of adult Callinectes sapidus and
larger fish, like Micropogonias furnieri, Netuma barba, Netuma planifrons,
and Genidens genidens (Araujo 1984). The larvae and juveniles of Erodona
mactroides assume an important trophic role in the shallow estuarine shoals
after intense recruitment in the summer (Bemvenuti et al. 1978). Here they
form, together with Balanus improvisus, the principal food source for fish
and decapods which use the shoals as nursery grounds (Asmus 1984).
In conclusion, the typical environmental stresses of oligohaline estuarine
systems (McClusky 1981; Day et al. 1989) are especiall}' enhanced in the
Patos Lagoon estuary (Capitoli et al. 1978; Castello 1985; Bemvenuti et al.
1992) and may have contributed to the development of a soft-bottom
community with wide trophic niches and abbreviated food chains.

5.3
Irregularly Flooded Marginal Marshes

C.S.B. Costa

Spatial Heterogeneity

Much of the extensive salt marsh surface (40 km2 ) surrounding the Patos
Lagoon estuary (Fig. 5.3.1), particularly that of the southern margin and of
the small islands, displays a considerable variety of physiography, includ-
ing small pools, a dense web of drainage channels, and marginal levees
(Fig. 5.3.2). Owing to the choked characteristics of the estuary, reduced
tidal oscillation, and highly variable seasonal and annual precipitation, the
marshes are irregularly flooded by waters of varying salinity. Euryhaline
conditions tend to prevail in the summer which, during periods of extreme
water deficit, may occasionally approach hypersaline conditions. Oligo-and
mesohaline waters are most commonly found in the winter and spring,
when elevated precipitation causes intense runoff (Castello and Moller
1978; Costa et al. 1988a; Vieira and Rangel 1988). In consequence, the
salinity regimes of the surface and interstitial water in the marshes are
seasonally predictable (Fig. 5.3.3). The interstitial water of the more
frequently inundated lower marshes tends to be more saline than that of
the rarely flooded upper marshes (Bastos et al. 1993a; Silva et al. 1993),
where salts are rapidly leached from increasingly sandy sediments, owing to
74 C.S.B. Costa

... 1 0 1 234 5 6 789

km

PATOS LAGOON
ESTUARY

ATLANTIC
OCEAN
N

Fig. 5.3.1. Distribution of marsh habitats in the Patos Lagoon estuary

Irregularly Flooded Marginal Marshes 75

FINE
SAND

a-\( \')
02.0-2.9
-3.0 - 3.9
.4.0 - 5.0

+0.60
:!:
"'ED IUM~~",=,:----==-_~o--~sn.T. CLAY
SAND ~
+0.40 z
~
~
m
;a
+0.20
r
m
<
m
0.00 r
]:
a I b I C I d ..().20

-10 0 10 20 30 40 50 60 70 80 90 100 110 120

DISTANCE FROM MEAN WATER LEVEL (m)

+
a-SANDFlAT
b - M1CROCUFF
c-RIOGE

f
d - MARSH FlAT
e-CREEK
f -CREEK LEVEE
s. maritimus S. attemiflora S. densiflora g-SALTPOOL

Fig. 5.3.2. Physiographic features of a typical Patos Lagoon estuary salt marsh with
percent organic matter (OM) content and texture of sediments

high mean annual precipitation (1350 mm; Vieira and Rangel 1988). Even in
marshes without pronounced physiographic diversity, the vertical and hori-
zontal influence of different flooding patterns on marsh sediment generates
environmental heterogeneity. Some of the dominant species (i.e. Spartina
densiflora) act as important geomorphological agents. They favour accretion
of the marsh surface and are able to modify drainage patterns and local
physiography (Figueroa et al. 1985) because their canopy promotes deposi-
tion of fine sediments from the water column and their dense root system
resists erosion of marsh creek sediments. In conjunction, these factors open
new niches and provide for pronounced spatial heterogeneity with a marked
influence on diversity, abundance, and distribution of plants and animals.
76 C.S.B. Costa

40
(A)

30

~
~20
~
10

40
(B)
.
....... FREQUEN11...Y FLOODED
. . . . OCCASIONALLY FlOODED *
30 e-e RARELY FlOODED I

J ~.
" ""'-.
~. J
\J
10

o
J AO 0 I F A J A OO IF A J AOO I F A J A 0 0 I F A J AO
1987 1988 1989 1990 1991

Fig. 5.3.3. Salinity of estuarine surface water (A) and of interstitial water in the rooting
zone at different salt marsh levels (B) at the Patos Lagoon inlet. Curves are based on a
simple moving average of three observations

Habitat Function

The fauna of the estuarine marshes is composed of species of aquatic and

terrestrial origin. Since large extensions of the upper marshes are infre-
quently and unpredictably flooded, terrestrial organisms tend to be more
common. However, aquatic organisms have received more attention, espe-
cially communities dominated by the crabs Chasmagnathus granulata
(Capitoli et al. 1977, 1978; D'Incao et al. 1992) and Metasesarma rubripes
(Capitoli et al. 1977). Chasmagnathus granulata is a burrowing omnivore of
the lower marshes (D'Incao et al. 1990, 1992) with larval dependence on
nearshore coastal waters. Although the smaller Metasesarma rubripes is a
herbivore and, rather than burrowing, hides among the roots and shoots of
Irregularly Flooded Marginal Marshes 77

Spartina alterniflora, Spartina densiflora, Scirpus maritimus, and Salicornia

gaudichaudiana, it competes for space and food with Chasmagnathus
granulata (Capitoli et al. 1977). Both species act upon the fragmentation and
remobilization of the belowground macrophyte biomass, thus decisively
influencing the recycling of organic matter in the marshes.
Owing to the origin of sediments and local hydrodynamics, substrate
characteristics vary among and within estuarine marshes. Different silt-clay
ratios in the lower (3.1-72.4%) and upper (0.6-10.5%) marshes are likely to lead
to changes of infaunal composition. Especially in creeks and pools, nematodes
and annelids are common inhabitants of superficial sediments, some being
detritivorous (e.g. Heteromastus similis, Laeonereis acuta) and others predators
(e.g. Nephtys fluviatilis). Since the plant biomass and organic matter
concentrations are considerably higher in marshes (up to 20%) than in
estuarine sediments (1-3.5%; Martins 1966; Moreno 1994), epi- and infaunal
estuarine organisms are attracted to the marshes during flooding. Under
these conditions, the gastropod Heleobia australis f1:equently feeds on
epiphytic and epipsamic micro algae in the lower marshes. Detritivorous
aquatic (Bathyporeiapus bisetosus) and terrestrial (Balloniscus spp.) isopods,
amphipods (Orchestia platensis), spiders, and insects are abundant through-
out the marsh. The abundance of deposit feeders in particular is likely to
correlate with sites of organic matter accumulation in depressions and areas
with dense vegetation cover. Most of these organisms are an important food
source for juvenile fishes (Odontesthes argentinensis, Atherinella=
Xenomelaniris brasiliensis, Micropogonias jurnieri, Mugil platanus; Bemvenuti
1990; Vazzoler 1991; Vieira and Scalabrin 1991) and birds which rest in the
estuary (see Vooren Sect. 4.14).

Organic Matter Contribution

Salt marshes have been described as producer systems that subsidize estu-
arine and nearshore waters (Odum 1971). Their ability to fix carbon at
remarkable rates remains unchallenged, but since marshes display a large
degree of individuality, the ecological fate of the fixed organic matter appears
to be highly variable (Haines 1979; Dixon 1980). In regions with a rapid sea-
level rise, marshes may accumulate plant litter in the sediments. Estuaries with
elevated river flow are likely to export large amounts of net primary marsh
production during runoff, whilst small semi-protected marshes may recycle
large portions of fixed organic matter. In consequence, the fate of organic
matter production by the physiographically diverse and irregularly flooded
marshes of the Patos Lagoon estuary is likely to be manifold. In occasionally
(25% of the time) flooded marshes, up to 20.8% of the annual marsh detritus
pool (2290 g m"2) may be exported, whilst 25.5 and 31% are more likely during
autolysis/leaching and microbial decay, respectively (Costa et aI., unpubl.).
78 C.E. Bemvenuti

5.4
Unvegetated Intertidal Flats and Subtidal Bottoms

C.E. Bemvenuti

Habitat Characteristics

The majority of the Patos Lagoon estuary is comprised of unvegetated soft

bottoms (300 km2); their environmental characteristics and the structure of
the associated benthic community change along depth gradients. Shallow
soft-bottom habitats are typically represented by intertidal flats between
sandy shorelines or marshes of marginal bays and both vegetated and
unvegetated shallow subtidal zones. Since estuarine water levels are largely
dependent on wind and precipitation, rather than on tides (Kantin 1983),
the intertidal flats are irregularly inundated and may be exposed during
prolonged periods in the summer. The larger pi!rt of the estuarine shoals is
dominated by shallow «1.5 m) subtidal soft bottoms, either with or with-
out the occurrence of Ruppia maritima beds, but with epibenthic micro-
algal growth and occasional aggregations of macroalgae. Deeper «6 m)
subtidal soft bottoms are void of plant growth and extend to the edge of the
natural and artificial estuarine channels. The channel bottoms (6-11 m) are
predominantly muddy, with the exception of the estuarine inlet (15 m),
where sediments become increasingly sandy.

Macrobenthic Community

The motile epibenthic organisms of subtidal soft bottoms in the estuary are
largely decapods (Capftoli et al. 1978; Bemvenuti 1987b). Shallower areas are
dominated by the small crab Rhithropanopeus harrissii and juveniles of
Cyrtograpsus angulatus, Callinectes sapidus, and Penaeus paulensis (Fig. 5.4.1).
Between the end of spring and beginning of fall, juveniles use shallow subtidal
shoals as nursery grounds and together with juvenile fish (Sciaenidae, Bothidae,
Jenynsiidae, Atherinidae) exert a severe predation pressure on the macro-
benthic community (Bemvenuti 1987b). At the end of the fall, as is typical for
decapods of temperate regions (Reise 1985), Penaeus paulensis, Callinectes
sapid us, and Cyrtograpsus angulatus migrate to deeper subtidal areas.
The sedentary epifauna of soft bottoms is principally represented by the
widely distributed gastropod Heleobia australis (Hydrobiidae) and by per-
acarid crustaceans, with a dominance of the tan aid Tanais stanfordi, the
amphipods Melita mangrovi, Ampithoe ramondi, Leptocheirus sp., and the
!sopods Dies fluminensis and Munna peterseni (Fig. 5..4.1). Small sedentary
organisms, like peracarids, are often under severe predation pressure (Nelson
1979b; Virnstein et al. 1984; Reise 1985), and their survival depends on the
structural diversity of the habitat, which is often provided by submersed
 SHALLOW WATER c:
~
MARSHES INTERTIDAL FLATS SUBLITTORAL BOTTOMS CHANNELS I~
II>
FLATS (UNVEGETATED OR ......
Ruppla BEDS) ~
0.-
Orchestia platensis : g
II>
- - < 500 ind. m-2
chas'ii~.i7~~~ 2r~"iJ.u.!.a.J.~ 1 __ _ ~
Metasesarma rubripes : - 500-5000 ind. m-2 s:~
- - - - - - - - - - - ~: Laeonereis acuta i ___ : 'TI
- > 5000 ind. m-2 [
• I rage/us p/ebeius :
----t --------- ------
I Melita mangrovi : '"
::l
'"
0.-
~ -=--=- -=-t I~r!.a§~[aDfQijtC == === en
: Munna peterseni : §-
- - - -: Kaiiiapsiudesschubafti/: ......
Nephtys fluviatilis ~
t:J::I
Heteromastus simi/is : o
: Heleobia australis - - - - - - S
i Erodona mactroides S
: Balanus improvisus
'"
I •
• ,

: Cyrtograpsus angulafus :
-------~
___ L----- ------------
_ ______ ~!1 /!fn_e2~~ ~aRigl!.s
- -_----- ------------~
_ ___________ ____ ~ __ --
: Penaeus paulensis :
---------- ----~--- -------- -
:·--- -------- ----T------ : . . ___ L Q'lupt1i§. §.ftfQ.SJd§.. _
: . __ _ LIj~n]ieo......d!!~ ~.'iX~rJ.

Fig. SA.I. Zonation of principal macrobenthic organisms in soft-bottom estuarine habitats

'-l
\0
80 C.E. Bemvenuti

vegetation (Orth et al. 1984; Summerson and Peterson 1984). As a conse-

quence, the densities of epifaunal peracarids, which are typically restricted
to the lower reaches of intertidal flats and shallow subtidal soft bottoms in
the estuary, grow as a result of macroalgal aggregations in shallow subtidal
areas, which supply protection against predation (Bemvenuti 1987b).
The distribution of Heleobia australis ranges from tidal flats to the soft
bottoms of deep channels (Capitoli et al. 1978). Densities may exceed
40 000 indo m-\ though pronounced spatial and temporal changes in density
are common (Bemvenuti et al. 1978, 1992; Lana 1986) since gastropods of the
Hydrobiidae are generally subject to intense predation (Reise 1985). In
shallow inter-and subtidal areas, elevated densities are associated with the
occurrence of fIlamentous algae (Bemvenuti et al. 1978; Orth et al. 1984;
Bemvenuti 1987b). Drastic density fluctuations can also be attributed to low
salinities, which cause osmoregulatory stress (Chomenko and Schaffer 1984;
Bemvenuti et al. 1992). Under this condition, Heleobia australis moves from
the epistratum to the water surface and, utilizing. the water surface tension,
migrates to regions with higher salinity, like deeper subtidal areas and
channels. The small number of competitors and the access to an ample
trophic and spatial niche of the epistratum result in elevated densities of
Heleobia australis in the deeper subtidal areas.
Infaunal polychaetes, pelecipods, and peracarids are characteristic and
abundant inhabitants of environmentally stressful intertidal flats (Reise
1985), though they also occur in subtidal areas. The polychaete Laeonereis
acuta is common to estuaries in the southwestern Atlantic (Orensanz and
Gianuca 1974) and is found in soft bottoms of the Patos Lagoon estuary
down to about 1 m depth (Fig. 5.4.1). As is usual for infaunal burrowing
species (Holland et al. 1980), adults of Laeonereis acuta escape predation in
deeper (approx. 20 cm) sediment strata. However, in the superficial sub-
strate their juvenile stages are under strong predation pressure, which leads
to high mortality and controls the density of the population (Bemvenuti
1992). Dense recruitment, either in shallow areas of the intertidal flat
(Bemvenuti et al. 1978; Capitoli et al. 1978) where preying fish and deca-
pods have limited access or during the winter (Asmus 1984) when several
predators abandon estuarine shoals (Bemvenuti 1987b), counteracts the
otherwise severe impact of predation and guarantees a larger number of
adult individuals. The polychaetes Nephtys fluviatilis and Heteromastus
similis have a wider vertical distribution (Capitoli et al. 1978) than
Laeonereis acuta, with higher densities in shallow subtidal areas but lower
densities in the deeper subtidal (>2 m) (Bemvenuti et al. 1978). Similar to
Laeonereis acuta, Heteromastus similis seeks refuge from predation by
b!lrrowing into deeper sediment strata of shallow subtidal zones
(Bemvenuti 1987b).
Individuals of the burrowing pelecipod Tagelus plebeius characteristi-
cally form patches in silty and clayey areas along the lower edge of
intertidal flats, whilst the isopod Pseudosphaeroma mourei frequently
Unvegetated Intertidal Flats and Subtidal Bottoms 81

occupies increasingly sandy sediments of more unstable, shallow, subtidal

zones (Capitoli et al. 1978). The distribution of the tube-dwelling tanaiid
Kalliapseudes schubartii, which lives in deeper (15 em) strata of sandy mud
bottoms, ranges from the lower reaches of intertidal flats to the margin of
channels at about 6 m depth (Fig. 5.4.1). In spite of intense predation
(Bemvenuti 1983, 1987b, 1992) by fish and decapods (Araujo 1984; Asmus
1984; Bemvenuti 1990), this peracarid is the dominant species (> 10 000 indo m-2)
in shallow subtidal areas. Apart from seeking shelter against predators in
sub-surface sediment strata, Kalliapseudes schubartii is a typical r-strategist
(sensu McCall 1977), with intense reproductive activity, marsupial
spawning, and recruitment, which maintains elevated population densities
(Bemvenuti 1987b, 1992) in shallow subtidal areas. Lower densities
(1000 indo m-z) in deeper subtidal zones (Bemvenuti et al. 1978) may partly
reflect the substrate and water column instability of these areas (Capitoli et
al. 1978; Kantin 1983). The largest macrobenthic biomass is produced by
the pelecipod Erodona mactroides. In the upper estuary, where all size
classes are present, larger individuals (up to 30 mmlength) at lower
densities (461 indo m-z) form an average biomass of 281 g m-2 • In the lower
estuary, the high mortality of Erodona mactroides during the first year
results in individuals of reduced size « 13 mm), and despite high densities
(3722 indo m- z), the mean biomass rarely exceeds 105 g m- (Bemvenuti et al.
1978). Especially at the end of the summer, high mortalities of Erodona
mactroides appear to be related to elevated deposition of fine particles
which may obstruct the filtration system of the organism. The density and
size of Erodona mactroides also influence in part the estuarine distribution
of the cirriped Balanus improvisus which, in the absence of any other hard
substrate, tends to attach to the shells of Erodona. Since small individuals
of Erodona mactroides are unsuitable as substrate, the density of Balanus
improvisus tends to be higher (719 indo m- 2) in the upper estuary.
The abundance and diversity of the macrobenthic community drastically
decrease between the lower reaches of deeper (>6 m) subtidal areas and
channel bottoms (Bemvenuti 1987a), though in the entrance channel diversity
may temporarily increase during periods of inflowing seawater, owing to the
addition of peracarids and small polychaetes (Fig. 5.4.1; Bemvenuti et al. 1992).

Habitat Function

The high production by different primary producer groups and the

continuous supply of detrital matter make inter-and subtidal bottoms ideal
nurseries for invertebrates and fish. The benthic communities of the inter-
tidal flats, shallow and deeper subtidal zones, and channel bottoms are
interdependent and linked by continuous feedback processes. Although
intertidal flats generally lack any conspicuous vegetation cover and, owing
to the dominance of burrowing species, appear to be faunistically poor,
82 U. Seeliger

they are areas of elevated secondary production with high rates of energy
transfer between infaunal organisms and highly motile predators (Reise
1985). In shallow subtidal zones, the deposit feeders represent an impor-
tant link between large amounts of detritus and higher trophic levels. The
deeper subtidal areas are dominated by suspension and deposit feeders,
which largely depend on allochthonous detritus and, in the absence of any
protective vegetation, are exposed to intense predation. These areas are
therefore important feeding sites for fish and decapods (Summerson and
Peterson 1984) and serve as a refuge for decapods during the colder
months (Reise 1985). Even the otherwise inhospitable deep channels con-
stitute an important migratory route for many estuarine organisms, and
the large water column favours the presence of pelagic species.

5.5
Seagrass Meadows

U. Seeliger

The size and physiographic heterogeneity of the Patos Lagoon estuary pro-
vides for diverse and extensive habitats. Rather than being isolated, habi-
tats interface along abiotic gradients via detrital matter transport and
migration of organisms. About 120 km2 of Ruppia maritima beds in the
Patos Lagoon estuary (Fig. 5.5.1) serve as a fishery habitat because they
provide substrate and shelter and offer nursery and feeding grounds for a
diverse assemblage of organisms, which may have adapted their life-cycles
accordingly. The leaves, stems, roots, and rhizomes of Ruppia form a
structurally complex habitat of calm water and stable substrate. The abun-
dant plant and detrital matter supports dense populations of motile and
sessile organisms, and the high in faunal and epifaunal abundance attracts
predators. Additionally, drift algae may form an associated habitat of
potential importance as a nursery.

Seagrass-Epiphyte Complex

In the absence of consolidated sediments, the leaves and stems of Ruppia

maritima provide the principal semi-permanent substrate for many micro-
and macro algae in the estuary, although none of the species appears to be
host-specific. As the leaves of Ruppia emerge from the sediments, epiphyte
colonization commences with a dense cover of the pennate diatom
C;occoneis placentula. Its flat morphology and copious mucus production
enable this species to cling firmly to the leaves, thereby becoming a mono-
specific pioneer stage. A secondary, still selective colonization by Synedra
fasciculata takes place over the living Cocconeis cells and associated detritus.
Seagrass Meadows 83

The formation of dense Synedra populations in the median region ofleaves

favours attachment of Amphora, Nitzschia, and naviculoides. During the
final stage of colonization, the apical region of the leaves is completely
covered by a diverse assemblage of diatoms and macroalgae (Enteromorpha
sp., Rhizoclonium riparium, Achrochaetium flexuosum; Ferreira and
Seeliger 1985). Since the shading effect of epiphytes is likely to decrease the
lifespan of individual leaves (and possibly of the entire population),
epiphytic community turnover must be relatively rapid.

10 km

PATOS
LAGOON
ESTUARY

r
ATLANTIC
OCEAN

52°

Fig. 5.5.1. Distribution of Ruppia maritima beds in the Patos Lagoon estuary
84 U. Seeliger

Faunal Relations

Polychaetes, molluscs, amphipods, the crabs Callinectes sapidus and

Cyrtograpus angulatus (Bemvenuti 1987a,b; Asmus 1989), the pink shrimp
Penaeus paulensis, and juvenile and sub-adult fish live either on and
between the leaves or in burrows between rhizomes of Ruppia (Asmus
1984); thus, vegetated areas tend to exhibit higher faunal abundance than
adjacent unvegetated bottoms. Since the size distribution of particles and
the organic matter contents of sediments do not vary (Moreno 1994), dif-
ferences in animal densities can be directly attributed to macrophyte bio-
mass. As the Ruppia meadow develops, increasing plant density, epiphyte
colonization, and litter production favour a higher abundance and biomass
of consumer populations. The gastropod Heleobia australis, for example,
which normally lives on the sediment surface, preferentially feeds on dense
bacterial fIlms on Ruppia leaves, and reaches its highest abundance during
peak plant growth (Asmus 1989). Many of !he invertebrates feed on
epifauna attached to the leaves, but they probably ingest a mixture of epi-
phytic algae concurrently. In consequence, most of the invertebrates are
opportunistic omnivores, like Cyrtograpus angulatus, which occasionally
show a preference for plant tissue (Capitoli and Ortega 1993). Also water-
fowl, like Fulica armillata and Cygnus melancorhyphus, appear to be
important herbivores of Ruppia tissue and seeds. The abundant plant
detritus in the meadows is likely to provide food for juvenile and sub-adult
fish (Odontesthes bonariensis, Atherinella brasiliensis). Predator-prey
relationships in the beds are influenced by interactions between the
behavioural, physiological, and morphological characteristics of animals
and variations in shoot density, biomass, leaf area, and root and rhizome
depth of the Ruppia plants. The free-living and less motile epifaunal Heleobia
australis is more susceptible to predation than the infaunal Erodona
mactroides and the tube-dwelling Kalliapseudes schubartii. While Kalliapseudes
schubartii appears to be the main diet item of the predator Callinectes
sapidus during the absence of Ruppia in the winter, the protective cover of
plants during the summer impedes the predator'S motility and Heleobia
australis may become the principal food source (Asmus 1989). Patchy
Ruppia beds may provide favourable foraging areas for fish, like Mugil
platanus, Odontesthes argentinensis, and Atherinella brasiliensis, and for
the highly mobile Callinectes sapidus, Cyrtograpsus angulatus, and Penaeus
paulensis, which may exert a strong predation pressure over several
macrobenthic juveniles seeking protection here (Bemvenuti 1992).
Seagrass Meadows 85

Nursery Grounds

One of the most important roles of Ruppia beds in the estuarine ecosystem is
that of a nursery ground in which postlarval stages of many invertebrate and
some fish species concentrate and develop (Asmus 1984). The shrimp
Penaeus paulensis and the blue crab Callinectes sapidus are an important
local fishery resource, and both are estuarine dependent, since they spawn in
coastal water and their post-larvae or megalopi are carried back into the
estuary during the spring, where they find protection in the Ruppia beds.
Although shrimps and crabs are found throughout the estuary, many
continue to forage in the Ruppia beds. Furthermore, Ruppia beds may also be
important nursery grounds for numerous species of commercial fish, and for
small forage species that serve as food for the larger carnivores. At some stage
in their life-cycle, several species of the families Sciaenidae, MugiIidae, Ather-
inidae, and Syngnathidae are associated with the beds. While all large shallow
estuarine areas could potentially be occupied by Ruppia, their size may natu-
rally vary as much as 40% between years (Moreno 1994), and growth may
commence at any time between the early spring and early summer and cease
between the late summer and early winter. In consequence, periods of high
faunal recruitment into the estuary may not coincide with years or periods of
optimal Ruppia bed size and permanence, which may impose habitat limita-
tions for estuarine fisheries stock (D'Incao 1991).

Energy Contribution and Export

Most Ruppia tissue only becomes an available food source as detrital matter.
During the growth period, leaves are continuously shed and tend to
accumulate within the beds, where they undergo rapid decomposition and
serve as food for detritivore organisms (Bemvenuti 1987a, 1992). At the end
of the growth period, whole Ruppia plants and the remaining detritus are
removed by winds and water currents and accumulate in deeper subtidal
areas and channels. During periods of flushing, the detritus along with
recently detached green leaves and uprooted plants is discharged into coastal
waters and transported by NE winds and longshore currents up to 50 km
southward along the beach (Seeliger, unpubl.). Therefore, in addition to
being an important food source in the estuary, the export of live and dead
plant material and detrital matter represents an important trophic link and
energy source for organisms in adjacent and distant coastal nearshore waters.
86 R.R. Capftoli

5.6
Rubble Structures and Hard Substrates

R.R. Capitoli

Colonizing Organisms

In spite of the overwhelming dominance of unconsolidated sediments in

the Patos Lagoon estuary, a variety of artificial structures (approx. 0.18 km 2 )
provide for hard sub-and intertidal substrates. Among the different types
inside the estuary are revetments, piers of port installations, and wooden
pilings which contribute considerably to the total area. The 4-5 km long
jetties of the Patos Lagoon inlet (Fig. 5.6.1) represent the largest continuous
extension of permanent substrate.
Permanent substrates in the estuary and the channel side of the jetties
are under the direct influence of turbid oligo-and mesohaline estuarin~
waters with pronounced salinity variations and reduced tidal oscillation.
These environmental conditions account for an encrusting community of
low floristic and faunistic diversity, dominated by a few tolerant species
(Fig. 5.6.1), like Balanus improvisus, as has been observed in estuaries else-
where (Cory 1967; Dean and Bellis 1975). Balanus improvisus larvae settle
on permanent substrates throughout the year, but their densities are lowest
in the winter. Winter water temperatures of 12°C retard the growth of
recently attached larvae, sexual maturation, and hatching of new larvae,
and thus the abundance of planktonic larvae in the winter and spring tends
to be low (Montli 1980; Capitoli 1983; Baumgarten and Niencheski 1990b).
Intense colonization occurs in the summer and fall together with larvae of
Stylochus sp., Obelia sp., and Boccardia hamata. However, owing to
crowding and high predation pressure, principally by Stylochus sp., the
initially high abundance of Balanus improvisus is rapidly reduced (Capitoli
1983). The dead shells of the cirripeds supply new spatial niches which are
occupied by the polychaete Boccardia hamata, the amphipod Amphithoe
ramondi, and the tanaiid Tanais stanfordi. As a result of strong physical
and biological disturbances within the estuary and, similar to the model for
"unstable environments" proposed by Wood (1987), the community devel-
opment appears to lack seasonal progression and biotic succession. Turbid
estuarine waters also limit algal colonization to the upper 15 cm (Lyngbia
confervoides, Ulvaria oxysperma, Polysiphonia sp., Enteromorpha sp.) or 50 em
(Melosira moniliformis, Biddulphia sp.) of permanent substrates (Coutinho
1982).
Environmental conditions and species composition in the estuary and
inlet channel differ from those on the seaward side of the northern and
southern jetties, each of which display specific habitat conditions with dis-
tinct benthic communities. The seaward side of the southern jetty is
Rubble Structures and Hard Substrates 87

2
5m

ATLANTIC
OCEAN

10'
o 1 2 3 4 6 6 7 8 II 10
km r

Fig. 5.6.1. Vertical distribution of principal macro-invertebrates and algae on rocky

substrate in the Patos Lagoon estuary
88 R.R. Capitoli

protected from swell and wave-overwash during NE winds. Since estuarine

runoff flows southward, the generally turbid freshwater tends to seep
through the gaps of the jetty, and thus the intertidal zone is dominated by
low salinities, whilst salinities between 25 and 30 occur in the subtidal (> 1 m;
Coutinho 1982). In consequence, the benthic community of the intertidal
(cyanophytes, diatoms, Bangia atropurpurea) and upper subtidal «50 cm)
zones (Chthamalus bissinuatus, Balanus improvisus, Enteromorpha
compressa, Bryocladia thyrsigera, Grateloupia cuneifolia) has a low diver-
sity. During prolonged drought periods, when seawater penetrates into the
estuary, Brachydontes darwinian us, Perna perna, and algae (Ulva lactuca,
Cladophora flexuosa, Gelidium crinale, Gymnogongrus griffthsii) colonize
the intertidal zone temporarily. More stable polyhaline salinities below 1 m
depth promote increased diversity and the establishment of Ostrea eques-
tris and associated species (Fig. 5.6.1).
In contrast, the seaward side of the northern jetty is characteristically
exposed to swell and waves, and water transparency and salinity (30) are
high (Coutinho 1982). The colonization of typical marine organisms fol-
lows a clear pattern of zonation (Fig. 5.6.1). Littorina ziczac occupies the
upper and Siphonaria lessoni and Collisella subrugosa, the lower spray
zone, followed by a sparsely populated zone of Chthamalus bissinuatus and
Brachidontes darwinian us. The extensive intertidal is dominated by Perna
perna, Megabalanus vesiculosus, Thais haemastoma, and a diverse algal
flora (Coutinho 1982). Owing to high wave impact, observations of the
subtidal zone are still lacking.

Habitat Function

Although the total area of hard substrate is reduced, it assumes an

important habitat function in this coastal environment, which otherwise is
devoid of permanent substrate. The presence of consolidated substrate is
an essential requirement for the growth and reproduction of cirripeds.
Especially in the summer, a dense layer of adult Balanus improvisus on
rocks and pilings means a continuous high hatching rate and consequently
an important contribution oflarvae to the water column. These will succes-
sively colonize the hard substrate and are likely to assure an ample food
supply for larvae of decapods and juvenile fish. Furthermore, the diverse
types of hard substrate provide physical and trophic niches for resident
and transitory fish. Hypleurochilus fissicornis is typically associated with
hard substrates in the estuary and the inlet channel, where it feeds on soft
parts of Balanus improvisus and amphipods (Capitoli 1983). Other species
are likely to visit the jetties, where they supply the resources for sport fish-
ing activities. Finally, the jetties have biogeographic significance, because
they provide settlement conditions for northern tropical and southern cold
Rubble Structures and Hard Substrates 89

temperate sedentary organisms, which are otherwise unable to bridge the

400 km long barrier of unconsolidated sandy substrate between 30° and 34° S
(Coutinho and Seeliger 1986).
6 Coastal and Marine Environments and Their Biota

6.1
Geomorphological Setting

1.J. Calliari
Continental Margin

The southern Brazilian continental margin is characteristic of low intensity

tectonic evolution processes (Villwock 1987), which levelled the basement,
formed thick sedimentary layers, and led to the development of an exten-
sive coastal plain (Fig. 6.1.1). The gently sloping continental shelf (2 m km-I)
between Santa Marta Grande Cape (28 0 40' S) and Chui (34 0 40' S) has a
width of 100-180 km (Martins et al. 1972) and reaches its largest extension
between 31 0 and 33 0 30' S. Remnant valleys of ancient fluvial channels typi-
cally cut the shelf in this region. Characteristic morphological features of
the shelf include marine terraces (Koswmann et al. 1977) and beachrock
outcrops (Calliari and Abreu 1984), whilst linear sand ridges typically occur
on the inner shelf (Figueiredo 1975). The transition between shelf and
upper slope starts at 160-190 m depths at a rate of 20 m km- I • The consider-
able width of the slope (up to 250 km near Cabo Santa Marta) as well as the
gradual drop to approximately 3000 m depth suggest the dominance of
depositional processes during its formation. At the central part of the
Pelotas Basin, the slope resembles the shape of a cone, which extends from
the outer shelf to about 4000 m depth. A well-developed continental rise
with a maximum width of 400 km (30 0 S) extends between 2700 and 4000 m
depth (Zembruscki et al. 1972). An important feature of the oceanic basin is
the Rio Grande Rise, which extends in an east-west direction between the
continental rise and the flanks of the Mid-Atlantic Ridge and separates the
ocean basins of Brazil and Argentina (Palma 1984).
Since the early Pleistocene (1.8 million B.P.) sea-level changes have been
responsible for the sequential deposition of several barrier islands over the
coastal plain. However, only sea-level changes after the Wisconsin glacia-
tion in the late Pleistocene (18 000 B.P.) have decisively influenced the
sedimentary characteristics of shelf and slope. At that time, sea-levels were
l20-130 m lower and approximately corresponded with the shelf break

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
92 1.J. Calliari

Fig. 6.1.1. Main physiographic features of the continental margin between 29 and 35° S

(Correa 1990). The subsequent phases of rising sealevel during the Holocene
transgression stabilized around 100-110, 60-70, 32-45, and 10-25 m and
imprinted erosive and depositional features of old shorelines over the shelf
(Koswmann et al. 1977; Correa 1990). At the maximum of Holocene trans-
gression (5000 B.P.), the sea was 4 m above present levels and inundated the
older Patos-Mirim Lagoon system. The following regression phase,
interrupted by small trangressions, formed a sequence of beach ridges and
depressions with small lagoonal systems. At the end of the regression phase
(2000 B.P.), the sea-level was 1-2 m below the present level, followed by a
gradual increase to its current height (Villwock and Tommazelli 1989).
The sedimentation patterns of the shelf and upper slope are predomi-
nantly terrigenous and, based on the sand/silt/clay contents, define eight
distinct facies (Correa 1987; Fig. 6.1.2). The inner and outer relict sand
facies of the shelf, composed of medium and fine quartz sand similar to
that of today's beaches and dunes, were deposited during the Pleistocene
regression and reworked during the Holocene transgression (Martins et al.
1972). Relict clayey silt and silty clay facies dominate over the middle and
outer shelf. Continental runoff during the Wisconsin transgression phase
was probably responsible for their deposition. Palimpsest sediments are
represented by transitional facies, composed of similar proportions of
terrigenous sediments and mixtures of biodetritus with sand or mud. The
deposition of recent silty clay and clayey silt facies was a result of sediment
transport from the La Plata River and the Patos Lagoon.
Surface carbonate deposits of shell fragments (Martins et al. 1972; Correa
1983) take up approximately 1000 km2 of the shelf area, and coarse sand
deposits exist near beachrock outcrops south of the Patos Lagoon inlet
(Calliari and Abreu 1984; Calliari et al. 1994). Glauconitic sands are common
sediments of the outer shelf and upper slope, and phosphate concretions (up
to 16% p,os) have been dredged from the upper slope (Klein, unpubl.).
Geomorphological Setting 93

- --
100 km

MIRIM
LAGOON

Fig. 6.1.2. Superficial sediments of the continental shelf and upper slope. (Modified after
Correa 1987)
94 C.A.E. Garcia

Beaches

Except for sandstone, basalt, and volcano-clastic rock formation near 29° S,
which offer a certain degree of shelter, the 653 km long coastline between 29°
and 35° S is totally exposed. Beaches are composed of fine quartz sand, and
most (95%) have a low gradient (z") with few inexpressive beach cusps. Only
a 45 km stretch of the shore south of Albardao lighthouse (33° S) is composed
of coarse shell fragments and fine quartz sands. These beaches display a
higher gradient (4"), erosional scarps at the backshore, and frequently well-
developed cusps (Calliari and Klein 1993). Based on morpho dynamic char-
acteristics (Wright and Short 1984), the beaches are considered generally
intermediate, but immediately south of 3z" and 33° S the beaches are dissipa-
tive and intermediate/reflective, respectively. Beach profiles demonstrate a
seasonal behaviour, owing to wave-induced sand accretion in the late spring
and summer and erosion during the remainder of the year. During cold front
passages, the sub-aerial portion of the intermediate/reflective and dissipative
beaches may suffer volume changes of 40 m 3 m- l and 4.9 m 3 m-\ respectively.
During feriods of high rainfall, a large number of ephemeral creeks (approx.
1.5 km-) break the frontal dunes and drain landward depressions, with
concomitant erosion between frontal dunes and the inner foreshore. The
considerable sand transport to the surfzone is likely to determine the
sediment balance between beach and frontal dunes.

6.2
Physical Oceanography

C.A.E. Garcia

Water Masses and Transport

The southern Brazilian continental shelf is influenced by the Subtropical

Convergence (STC). The Convergence is formed by the southward flowing
Brazil Current (BC), transporting Tropical Water (TW; T >20°C, S >36.00)
(Emilsson 1961), and the coastal branch of the northward flowing Malvinas
(Falkland) Current (MC), transporting Subantarctic Water (SAW; T 4-15°C, S
33.70-34.15; Sverdrup et al. 1942; Thomsen 1962). The convergence and
mixture of TW and SAW form Subtropical Water (STW; T 1O-20°C, S 35.00-
36.00; Emilsson 1961), also called South Atlantic Central Water (Sverdrup et
al. 1942), which flows northward in deeper layers in a counter-clockwise gyre
and may reach the continental shelf (Miranda 1972; Hubold 1980a). In
contrast to oceanic zones, water transport over the continental shelf is largely
a consequence of the impact of offshore currents on the shelf's topographic
features, though the wind regime also plays an important role in circulation.
Long-term current velocity changes follow seasonal patterns and spatial
Physical Oceanography 95

distribution of wind regimes along the continental margin (Luedemann 1971,

1978; Pereira 1977), whilst short term changes of current velocities are
associated with atmospheric conditions as a result of cold front passages in 2-
10 day intervals.
Owing to the influence of the Atlantic gyre, the southward flowing BC
becomes deeper and wider between 24° and 31° S. The convex curvature of
the bathymetry at 24° S and the smooth topography of the shelf cause
instability of the flow and oscillations of the front along the shelf break
(Garfield 1990). The geostrophic circulation and the transport of the water
mass associated with BC vary (Cacciari 1986), but the flow is generally
directed southward. However, although TW and STW masses of the upper
water column flow south, water masses below the thermocline move in
opposite directions. Antartic Intermediate Water (AIW) below the thermo-
cline flows north (Deacon 1933; Wiist 1935), whilst deep North Atlantic
Water (DNAW) flows south. It is likely that the BC extends vertically down to
the interface between DNAW and northward flowing Antartic Bottom Water
(ABW). Due to the anti-cyclonic bottom circulation, some of the AIW may be
transported southward along the slope (Reid et al. 1977; Piola and Gordon
1989). Computations of the southward transport of the BC at 32-33° S (Evans
et al. 1983; Stramma 1989) suggest that during most of the year tropical water
is transported over the shelf (Garfield 1990).
Two types of upwelling processes are common along the southern
Brazilian coast. Coastal upwelling is more frequent during the spring and
summer, owing to the strong influence of NE winds, which provoke a
divergence of superficial waters and cause advection of STW over the shelf
between 28° and 32° S (Miranda 1972; Hubold 1980a). Upwelling at the shelf
break is more common in the winter and spring. The BC and the coastal
branch of the MC form frontal vortices of cyclonic circulation which cause
upwelling of bottom water, principally STW (Hubold 1980a,b).

Thermohaline Structure

The seasonal migration of the Convergence (STC) causes the advection of

TW, SAW, and STW, with contrasting thermohaline characteristics over the
shelf (Olson et al. 1988; Gordon 1989). Furthermore, the thermohaline
properties of the coastal water (CW) vary between seasons and years as a
function of freshwater runoff from the Patos Lagoon and the La Plata River
(Miranda 1972; Miranda et al. 1973; Tseng 1976; Castello and MOller
1977,1978; Miranda and Castro Filho 1979; Godoi 1983; Cacciari 1986; Ciotti
1990; Fillmann 1990). In consequence, the seasonal and interannual
variabilities of the thermohaline properties of the shelf waters are pro-
nounced, with a predominance of TW and SAW during the summer and
winter, respectively (Miranda 1972).
96 L.F. Niencheski and G. Fillmann

The centers of warm anticyclones and cold cyclones directly control the
depth of the thermocline (100-150 m) of the oceanic region (Silva 1981;
Godoi 1983). In contrast, over the shelf the vertical thermohaline structure
varies with season, continental runoff, and direction and velocity of the
winds. During the summer, the shelf (>50 m depth) and the slope are cov-
ered by the tropical surface water of the Be. Owing to reduced continental
freshwater runoff, the vertical thermohaline structure is moderately strati-
fied, and the thermocline occurs at a depth of about 40 m. During the
winter, the BC flows along the outer shelf, and the advection of the north-
ward flowing, cold, and less saline MC water mass results in a strong hori-
zontal thermohaline gradient. Horizontal current sheer may cause upwel-
ling at the front between the two water masses over the shelfbreak. Under
these conditions, the shelf waters become weakly stratified (Lima 1992).
High freshwater discharge by the Patos Lagoon and the La Plata River
(Castello and Moller 1978) and less frequent south winds correct this situa-
tion during the spring. Coastal waters over the-shelf with less than 50 m
depth become strongly stratified, with vertical sigma-t gradients of 2.5 m- 1
(Brandhorst and Castello 1971; Podesta 1990; Lima 1992).
Shelf water surface temperatures (Miranda 1972; Castello and Moller
1977; Hubold 1980a,b) are largely a consequence of the different water
masses. The seasonal variation of mean surface temperature, with 23°C in
February and 12.5°C in August, is most pronounced over the southern
shelf. The presence of tropical water over the shelf during the summer
causes high monthly mean surface temperatures with a low standard
deviation, whilst high monthly surface temperature deviations are typical
for the spatial heterogeneity of water masses in the winter (Lima 1992;
Belem, unpubl.).

6.3
Chemical Characteristics

L.F. Niencheski and G. FiHmann

The chemical characteristics of shelf and slope water largely depend on

continental runoff and on the influence of the different water masses at
each season (Magliocca 1971, 1973; Castello and Moller 1977; Hartmann et
al. 1980; Hubold 1980a,b; Kantin 1982, 1983; Kantin et al. 1982; Magliocca
et al. 1982; Dohms 1983; Carreto et al. 1986; Fillmann 1990). Sediment-
water column exchange, assimilation, remineralization, and enrichment
processes define the chemical characteristics of each water mass.
The Brazil Current (BC) is characterized by oligotrophic Tropical Water
(TW) which reaches the southern Brazilian shelf most of the year. Low
average dissolved nutrient concentrations (phosphate 0.31 ~, nitrite 0.09 ~,
nitrate 1.43 11M, ammonium 0.78 ~, silicate 4.83 11M) and low mean total
Chemical Characteristics 97

suspended matter levels (4.45-5.96 mg r') of their water indicate a lack of

enrichment from continental runoff. In contrast, the Subantartic Water
(SAW) mass represents an important source of nutrients for shelf waters.
Especially during the winter, the addition of high dissolved nitrate, ammo-
nium, phosphate, and silicate, with maximum concentrations of 11.1 1lM,
6.6 11M, 1.1 11M, and 10 11M, respectively, fuels the primary production over
the shelf (Fillmann 1990; Ciotti 1990; Sales Dias, unpubl.; Fig. 6.3.1).
Continental runoff from the La Plata River and the Patos Lagoon influ-
ences the chemical characteristics of coastal water (CW) over the shelf. In
consequence, both short-term and seasonal changes in volume and compo-
sition of runoff cause high variability of chemical parameters in CWo Dis-
solved mean oxygen concentrations (8.18 mg r') and saturation levels
(96.06%) are higher than in the other water masses, though extremely low
values (3.06 mg r'; 38.15%) may occur during the spring, and the pH (7.78 -
8.80) may also fluctuate considerably. Average total suspended matter
(> 16 mg 1"') and silicate (23.10 1lM) are highest in CWo The generally low
nitrate (2.61 1lM) concentrations and N:P ratios indicate that nitrogen may
limit primary production here (Fig. 6.3.1).

i'
:l.
W 1
~
::r::
B.
(f)
o::r::
Q.

i'
:::l.0.6

~ 0.4
it:
!:: 0.2
z

TW SAW CW STW TW SAW CW STW

o SPRING • WINTER • SUMMER 0 FALL

Fig. 6.3.1. Mean seasonal nutrient concentrations and N:P ratios (CV=95%) of different
water masses over the continental shelf between 29 and 35° S
98 U. Seeliger

Subtropical Water (STW) is a major source of dissolved nitrate, with a

mean concentration of 9.29 ~M and a range between 0.22 and 26.28 ~M.
This water is also rich in dissolved inorganic phosphate (mean 0.73 ~M,
maximum 1.81 ~), while mean nitrite (0.08 ~M) and ammonium (0.74~)
concentrations are low. The suspended matter concentration (4 mg r1) of
the deep STW is low. The oxidation of organic matter during transport
leads to low dissolved oxygen (mean 6.6 mg }"1) and oxygen saturation lev-
els (approx. 79%), which also results in increased nitrate values. The influ-
ence of STW on the continental shelf strongly depends on the direction of
dominant winds since the great depth of these waters obstructs input
processes. Nutrients rarely reach the euphotic zone (Fillmann 1990) and
only cause elevated productivity and chlorophyll a levels during coastal
and shelfbreak upwelling (Hubold 1980a; Ciotti 1990).

6.4
Coastal Foredune Flora

U. Seeliger

Floristic Composition and Affinities

Coastal foredunes between 32° and 34° S are exclusively colonized by herba-
ceous plants, whilst arboreous climax vegetation is restricted to older
inland dunes. Owing to the recent formation of the coastal plain, most
species are migrants from neighbouring provinces, and endemic species are
rare (Waechter 1985). The composition of the foredune flora changes along
latitudinal gradients (Legrand 1959; Reitz 1961; Eskuche 1973; Pfadenhauer
1980; Cordazzo and Seeliger 1987). The floristic composition of the fore-
dunes of Brazil's extreme south (30-34° S) and northern Argentina (37-39° S)
is much alike, but shows little similarity with northern tropical (28° S) or
southern cold-temperate (42° S) regions (Fig. 6.4.1). These distributional
discontinuities correspond to the northern and southern latitudinal limits
of the Subtropical Convergence during the winter and summer, respec-
tively (Legeckis and Gordon 1982), thus characterizing this coast as a
warm-temperate transition zone (Coutinho and Seeliger 1986; Cordazzo
and Seeliger 1988a).
The 71 foredune species (Appendix) exhibit perennial, as well as sum-
mer and winter annual life-forms (Cordazzo and Seeliger 1987, 1988a;
Seeliger 1992a). Summer annuals complete their life-cycles between the
spring and fall, while winter annuals germinate in the fall, persist vegeta-
tively during the winter, and set flowers and seeds the next year. All peren-
nials exhibit drastically reduced growth vigour during the winter (Cordazzo
and Seeliger 1988b), owing either to suboptimal environmental conditions
or endogenously induced resting periods, as has been observed for Hydro-
Coastal Foredune Flora 99

cotyle bonariensis, Gamochaeta americana, and Blutaparon portulacoides

in the Brazilian tropics (Hueck 1955; Ormond 1960). True dormancy only
occurs in Oenothera affinis, Pterocaulon purpurascens, and Achyrocline
satureoides, which overwinter through underground organs. Germination
and flowering of all species appear to be largely controlled by local soil
moisture conditions or regional temperature and light regimes, respectively
(Cordazzo and Seeliger 1988b).

25

--

100
N
C»
to>
""
to>
U)
""
N
0
(Jl
to> to>
N
0 .....0
~ 0
(Jl (Jl (Jl

Sporobulus rigens
Panicum urvilleanum
Hyalis argentea
Senecio quequensis
Spartina coarcta
• Calycera crassifolia
Cakile maritima
Blutaparon portulacoides
Panicum racemosum
Senecio crassiflorus
Gamochaeta americana
Hydrocotyle bonariensis
Spartina ciliata
Ipomoea pes-caprae
Canavalia maritima
Acicarpha espathula
Remlrea maritima
Scaevola plumieri
Sporobulus virginicus

Fig. 6.4.1. Similarity and species composition of dominant coastal foredune vegetation
between southern Argentina (420 S) and St. Catarina, Brazil (260 S)
100 U. Seeliger

Biology of Dominant Species

Their biomass and abundance classify Blutaparon portulacoides, Panicum

racemosum, Spartina ciliata, Hydrocotyle bonariensis, Andropogon arenarius,
and Androtrichum trigynum as foredune dominants. Their distribution and
abundance are largely a function of substrate stability and water table
distance (Cordazzo 1985; Cordazzo and Seeliger 1993). Sand accretion and
erosion are particular controlling factors for the perennial rhizomatous
Blutaparon portulacoides, which is common to the Atlantic ocean beaches
between northern Brazil and Argentina (Hueck 1955; Pfadenhauer 1978).
Succulent and frequently shed leaves, foliar salt excretion glands, and a
reduced number of stomata enable this species to tolerate salt-stressed
backshore conditions (Dillenburg et al. 1986). The clonal growth favours
the occupation of the unstable substrate, and the leaves and rhizomes aid
in the capture of windblown sand (pfadenhauer 1980; Bernardi and Seeliger
1989). Although cyclic events like extreme storm tides drastically reduce
the biomass and density of the Blutaparon population (pfadenhauer, 1978),
the seasonal recuperation (Cordazzo 1994) from rhizome fragments is
rapid (Bernardi et al. 1987; Bernardi and Seeliger 1989; Cordazzo and
Seeliger 1993).
Stands of the perennial rhizomatous grass Panicum racemosum extend
from northern Brazil to Argentina (Smith et al. 1982). This species is domi-
nant on foredunes with variable substrate stability and nitrate «0.5 mg kg-I),
phosphate (0.2-1.2 mg kg-\ and potassium (3-35 mg kg-I) deficient sands
(Costa 1987). Occasionally, Panicum racemosum occurs together with the
strandline species Cakile maritima, Calycera crassiflora, and Calystegia
soldanella (Seeliger 1992b; Cordazzo and Seeliger 1993). The highest flow-
ering and growth response occur under optimal temperature and precipi-
tation during spring in sand accreting areas. Although cutting experiments
with Panicum tillers show that sand-accretion accompanies plant growth,
the mechanical deposition of sand itself does not stimulate growth, but
rather provides a substrate and a source of nutrients for continuous verti-
cal growth of rhizomes and tillers. Over time and with distance from the
sea, Panicum racemosum populations change from "invader", through
"mature", to "regressive" stages, expressed by a gradual reduction in popu-
lation density and tiller vigour, in response to the spatial patterns of sand
deposition and salt spray input (Costa et al. 1984, 1991). On dunes with
reduced sand accretion, Gamochaeta americana and Senecio crassiflorus
may locally become dominant (pfadenhauer 1978, 1980; Cordazzo and
Seeliger 1987, 1988a). The distribution of Senecio crassiflorus can be
explained by the species' germination success and the growth capacity of
seedlings in response to sand accretion (Cordazzo and Souza 1993).
Coastal Foredune Flora 101

The distribution and biology of the other dominant species are largely
controlled by the groundwater table distance and inundation stress (Cordazzo
and Seeliger 1988a). In areas close to the groundwater table, with reduced
sandflow and low levels of nutrients, Spartina ciliata assumes dominance.
Although both Spartina and Panicum maintain their populations principally
via clonal growth, each species adopts different colonization strategies (Costa
et al. 1984, 1991). While rhizome fragments are the principal dispersal units
of Panicum, a large number of salt-tolerant seeds permit Spartina ciliata to
rapidly occupy disturbed sites, owing to high germination success and
efficient seedling establishment (Cordazzo 1994). At drier sites, the number
of live Spartina leaves/tillers relates to precipitation. High death rates occur
during dry spring and summer, and high birth rates during the fall. At humid
sites, birth and death peaks alternate, and Spartina tillers reach their
maximum height and leaf number. These differences suggest that the density
of leaves/tillers depends on the carrying capacity of each site (Costa and
Seeliger 1988a).
The pronounced phenotype plasticity and the extensive and fast-growing
integrated rhizome system, which compensates for nutrient limitation or
competition (Costa and Seeliger 1988b, 1990), enable Hydrocotyle
bonariensis to opportunistically exploit seasonally flooded humid areas
(Costa 1987). Similar distribution patterns of Hydrocotyle bonariensis have
been observed in dune systems elsewhere (Moreno-Casasola and Espejel
1986; Barbour et al. 1987). The local co-dominance of humid sand indicators,
like Paspalum vagina tum (pfadenhauer 1978, 1980; Pfadenhauer and Ramos
1979; Cordazzo and Seeliger 1988a, 1993; Costa et al. 1988b), confirms this
patterns for the southern Brazilian coast. The leaf demography and forma-
tion of reproductive structures of Hydrocotyle bonariensis depend on the
carrying capacity of each site as well as on resources in belowground organs.
A high below/above ground biomass ratio indicates that Hydrocotyle depends
principally on rhizome-propagated growth. Groundwater table fluctuations
promote different leaf morphologies, with the rapid sprouting of small leaved
and long stemmed plants after flooding (Costa and Seeliger 1988b, 1990).
Similar to Hydrocotyle bonariensis, areas with a seasonally fluctuating
watertable are dominated by the perennial grass Andropogon arenarius. This
species is endemic to stable dunes of southern Brazil and Uruguay and grows
in association with Spartina ciliata, Imperata brasiliensis, Androtrichum
trigynum, and Hydrocotyle bonariensis (Cordazzo 1985). Within distribu-
tional limits, the growth of Andropogon appears to be influenced by com-
petition for nutrients and/or space with Spartina ciliata. Site-specific abiotic
factors favour different strategies of spatial occupation. At dry sites, with
desiccation stress and/or sand accretion or erosion, Andropogon seedlings
rarely become established, and spring growth initiates from clonal budding of
rhizomes, which results in dense clusters of plants. In contrast, wet areas
promote seed germination and seedling survival, which leads to the random
102 N.M. Gianuca

growth of single shoots with low density. Low aerial biomass during winter
suggests a pre-adaption mechanism to seasonally flooded conditions (Costa
1987; Costa et al. 1988a).
Winter-inundated sites are typically dominated by the perennial Cyper-
aceae Androtrichum trigynum, which is endemic to southern Brazil and
northern Argentina (Costa 1987; Costa et al. 1988b). Associates include Phyla
canensis, Bacopa monnieri, Pluchea sagittalis, and Paspalum vaginatum,
inundation-tolerant species of drier sites (Andropogon arenarius, Hydrocotyle
bonariensis, Spartina ciliata), and marsh remnants, like funcus acutus and
Typha domingensis (Cordazzo and Seeliger 1987, 1993). The change of domi-
nance between Andropogon and Androtrichum can be attributed to different
growth and reproductive responses of these species to inundation stress. The
aerial biomass of Androtrichum is positively related to the watertable dis-
tance, and plants are absent from areas with a deeper (>2 m) mean annual
groundwater table. The rapid occupation of wet areas during spring seems to
be related to an efficient uptake of nutrients which become available after
drainage (Lacerda et al. 1986). A high proportion of monocarpic flowering
shoots induces high mortality in the fall after flowering. In contrast, a low
proportion of flowering shoots in dry areas results in low and constant mor-
tality rates throughout the year (Costa et al. 1988b).

6.5
Coastal Foredune Fauna

N.M. Gianuca

As has been reported for dune systems elsewhere (van Heerdt and Morzer
Bruyns 1960; Callan 1964; CaussaneI1970), insects are the dominant fauna in
the extensive coastal foredunes of Brazil's extreme south and represent the
most diverse group of organisms, though some vertebrate species are also
important (Gianuca 1985, 1988; Appendix).

Insects

Ants, with nine species, are among the most conspicuous organisms
throughout the year, although in the spring and summer elevated noon sur-
face sand temperatures drastically reduce their activity on the surface.
Despite an apparently low diversity of food sources, diet preferences vary
between species. Pheidole spininodis, Monomorium sp., and two species of
the genus Conomyrma are omnivorous, whilst Camponotus punctulatus and
Solenopsis geminata are essentially herbivorous and carnivorous, respec-
tively. The basic food of Mycetophylax simplex and Brachymyrmex sp. are
fungi on dead organic matter and faecal pellets (Bicho, unpubl.).
Coastal Foredune Fauna 103

The coleopterans (>40 species) are the most diverse group. Sand bur-
rowing scarabs like Thronistes rouxi, Ligyrus gianucai, Plectris bonariensis,
Athyreus chalybeatus, and Psammodius sp. are typical of the dunes. Their
adult forms are largely restricted to the summer and only surface from the
sandy sediments on wind-still nights with high temperature and humidity.
Both larvae and adults of scarabaeids feed on detrital matter of roots and
rhizomes of resident vegetation. In contrast, all species of carabids are
predators. Tetragonoderus variegatus remains buried at night and preys
during the day on collembolans and delphacids, whilst Tetragonoderus
undatus surfaces and feeds at night. The carabids Sphizogenius costiceps,
Bembidion sp., and Scarites spp. are usually restricted to slacks, where they
prey on larvae and adults of other insects, preferentially the rove beetles
Bledius bonariensis, Bledius microcephalus, Bledius fernandezi, the hetero-
cerid Efflagitatus freudei, the hydrophilid Paracymus rufocinctus, and the
orthopteran Neotridactylus carbonelli, which reaches high densities in the
spring and summer. Sand micro algae (Garcia-Baptista 1993) and organic
matter derived from the decomposition of plant litter make up the basic
diet of these slack dwellers. Several coleopterans are associated with the
aerial parts of dune plants. The anticid Lagrioida nortoni is abundant on
the leaves and spikes of the grasses Panicum racemosum and Spartina
ciliata. The weevil Listroderes uruguayensis lives between and feeds on
plants of Hydrocotyle bonariensis. Caterpillars of the moth Ecpantheria
indecisa graze upon the leaves of Senecio crassijIorus (Pereira 1957) as well
as on Blutaparon portulacoides, which is also mined by larvae of the small
fly Hapopleudes vogti (Agromyzidae). Other species associated with the
vegetation of frontal dunes include the cicadas Proarna uruguayensis and
Proarna sp. and the delphacid Delphacodes kuscheli. The scavenging mole
cricket Scapteriscus riograndensis and the predator earwig Labidura riparia
(Labiduridae) are common species throughout the year.
Some flies (i.e. Eccritosia rubriventris, Asilidae) and several wasps are
well adapted to the sand dune environment owing to both their ability to
burrow in loose sand and to locate and catch either flying or buried insects.
The spider wasp Anoplius bilunulatus attacks the white spider Moenkhausiana
halophila (Zoodaridae), which serves as food for the larvae in sub-surface
galleries. The larvae of one of the largest species (Campsomeris cineraria,
Scoliidae) parasitize the white grubs of the scarabs Thronistes rouxi and
Ligyrus gianucai. Eucoila sp. (Cynipidae) attacks larvae of flies. Under-
ground nests of Tachytes ornatipes (Larridae) are most common in the slacks,
where adults paralyze and store the pigmy mole cricket Neotridactylus
carbonelli to feed the larvae.
104 P. C. Abreu: Bacterioplankton

Vertebrates

The frontal dunes are the preferred habitat of several vertebrate species. At
night, the sand toad Bufo arenarum arenarum (Bufonidae) and the sand frog
Pleurodema darwini (Leptodactylidae) are the dominant predators of differ-
ent insects and spiders. During daylight, the lizard Liolaemus occipitalis
(Iguanidae) feeds opportunistically on the available insects. Small toads,
frogs, and especially lizards are preyed upon by the hognose snake
Lystrophis dorbignyi (Colubridae). Two rodents, the mouse Calomys Zaucha
and the tuco-tuco Ctenomys flamarioni (Ctenomyidae), are common to the
drier frontal dunes, where they ingest seeds and plant tissue of foredune
vegetation. During most of the day, Ctenomys flamarioni hides in an exten-
sive network of tunnels, and storage food and building material for the nest
is almost always collected at night. The sheltered areas and pastures behind
the foredunes are inhabited by species like the burrowing owl Athene
cunicularia (Strigidae), the skunk Conepathus chinga (Mustelidae), the
armadillo Dasypus hybridus (Dasypodidae), and the fox Dusicyon gymnocercus
(Canidae), all of which undertake feeding incursions into the foredunes at
night.

6.6
Bacterioplankton

P. C. Abreu

Both autotrophic picoplankton «2 ~) and heterotrophic bacteria are

important components of oligotrophic oceanic water masses because of
their contribution to primary production (Platt and Li 1986) and their
influence on remineralization processes and importance as trophic links
(Azam et al. 1983), respectively. In spite of their relevance, little is yet
known about the biology of bacterioplanktonic organisms and their ecol-
ogy in coastal and oceanic waters of this region. In superficial shelf and
slope waters (31 0 30' - 340 30' S) the number of free bacteria (length 0.5-1 ~;
width 0.3-0.5 ~) oscillates between 0 and 1.35 xlO s cells r\ whilst the
number of bacteria (length 1-2~; width 0.3-0.5 ~) attached to particles
varies between 0 and x5.25 105 ceils rl. Free and attached bacteria are more
abundant during the spring and distribution patterns generally follow
those of phytoplankton biomass (chlorophyll a), being highest in coastal
water and during the influence of subantartic water. The biomass of total
free and attached bacteria and phytoplankton is of similar magnitude
(Fig. 6.6.1). The present data indicate that bacteria efficiently metabolize a
large part of the organic matter produced by phytoplankton, which is an
important carbon source for higher trophic levels.
Phytoplankton 105

80
EJ PHYTOPLANKTON
BACTERIA
60
:...
u
Cl
.:
III 40
III
«
:;;
o
iii
20
Fig. 6.6.1. Mean bacterial
and phytoplankton
biomass with standard
deviation in southern
SEPTEMBER FEBRUARY JULY Brazilian coastal waters

6.7
Phytoplankton

C. Odebrecht and V. M. T. Garcia

Taxonomic Composition, Cell Abundance, and Diversity

Descriptions of phytoplankton species of the southern Brazilian shelf were

first done early this century (Cunha and Fonseca 1918; Peters 1932; Kasler
1932; Gemeinhardt 1932). These and subsequent studies showed that a
large number of the diatom species belong to the genera Chaetoceros,
Coscinodiscus, Rhizosolenia, and Nitzschia (= Pseudonitzschia), and that
Ceratium, Dinophysis, Prorocentrum, Scrippsiella, Noctiluca, Oxytoxum,
and Gymnodiniales are important dinoflagellates (Rosa 1979; Kremer and
Rosa 1983; Dohms 1983; Ciotti 1990). Early estimates of total phytoplank-
ton abundance (diatoms, naked and thecate dinoflagellates, nanoflagellates,
coccolitophorids) at various depths (Hentschel 1932) reported con-
centrations of 104 _10 5 cells r\ though recent studies have shown concentra-
tions of 106 cells r' in coastal and offshore waters (Barth 1968; Ciotti 1990),
and 107 _10 9 cells r' in the surfzone (Odebrecht et al. 1995a).
The convergence between the Tropical Water (TW) of the Brazil Current
(BC) and Subantartic Water (SAW) of the Malvinas Current (MC), as well
as the freshwater input from the La Plata River and the Patos Lagoon, all
with distinct phytoplankton communities, results in high phytoplankton
diversity over shelf and slope (Appendix). At the southern end of the shelf
106 C. Odebrecht and V. M. T. Garcia

during the winter, the Shannon-Weaver index (3.4-4.6) for diatoms and
silicoflagellates compares favourably to values of all phytoplankton taxa
from highly diverse marine ecosystems elsewhere (Lange and Mostajo
1985). Of the 131 diatoms and silicoflagellates species identified by these
authors, 45% are of neritic and 16% of oceanic origin. The dominant spe-
cies, like Rhizosolenia setigera, Thalassiosira rotula, Thalassionema
nitzschioides, Dictyocha fibula, and Dictyocha speculum, have a cosmo-
politan and temperate distribution. During the late winter and spring ele-
vated phytoplankton concentrations (10 5 _10 6 cells r') occur both nearshore
and offshore. Under the influence of continental runoff, the diatom Skele-
tonema is abundant in coastal waters near the mouth of the Patos Lagoon
estuary, while Rhizosolenia and pen nates like Pseudonitzschia are abun-
dant in offshore regions under the influence of SAW. The influence of
Coastal Water (CW) also favours an abundance of thecate dinoflagellates
(Scrippsiella cf. trochoidea, Prorocentrum spp., Dinophysis spp., Ceratium
spp.) in the northern central shelf regions. The_ autotrophic ciliate Meso-
dinium rubrum is common along the entire southern Brazilian coast. Nan-
oflagellates largely dominate oceanic areas under the influence of TW but
share dominance with diatoms and dinoflagellates over the shelf and in
coastal regions in the winter and spring (Ciotti 1990; Castello et al. 1990).
Under the strong influence of the BC in the summer, phytoplankton abun-
dance is usually low, and warm water diatoms (Palmeria hardmanniana,
Pseudosolenia calcar-avis, Rhizosolenia robusta, Rhizosolenia imbricata,
Proboscia indica, Chaetoceros decipiens, Chaetoceros peruvian us, Chae-
toceros coarctatus) extend into nearshore waters (Dohms 1983).

Chlorophyll a and Primary Production

High chlorophyll a concentrations and primary production rates occur off

the coast of southern Brazil. Based on the data of six cruises (Table 6.7.1),
the mean annual particulate primary production is approximately
160 g C m-2 year-I. High productivity rates result mainly from northward
transport of nutrient-rich SAW, continental runoff, and upwelling of Sub-
tropical Water (SACW; Fig. 6.7.1). As a consequence, the distribution
patterns of phytoplankton chlorophyll a concentrations and primary
production rates are directly a -function of the temporal and spatial
variability of the different water masses. Large-scale meteorological
processes (El Nino-Southern Oscillation) may cause pronounced
interannual variations of hydrographic characteristics as a result of
differences in the total annual precipitation. Therefore, chlorophyll a
concentrations are higher in CW during El Nino years (warm periods in the
Pacific Ocean) than in La Nina years (cold periods; Ciotti et al. 1995).
Phytoplankton 107

26
Spring

22
CW
18 ••
.........
U
0
14
'-'
~ 10
....
::s

.
~ 26 Summer
....
~ 22
0..
El Chlorophyll a (mg m- 3)
~ 18 . < 0.5 .0.5 - 2.0 • 2.0 - 4.0 • > 4.0
E-4
14

10

28 30 32 34 36
Salinity
Fig. 6.7.1. T-S diagrams with water masses and chlorophyll a concentrations off southern
Brazil. Subantarctic Water (SA W); Tropical Water (TW); Coastal Water (CW); and South
Atlantic Central Water (SACW = STWj

In late winter and spring, the greatest chlorophyll a values and primary
production rates are related to nutrient input from SAW and continental
runoff, with sub-surface and surface nuclei of high biomass, respectively
(Ciotti et al. 1995). In the summer, the low chlorophyll a concentrations and
primary production rates are due to nutrient limitation during oligo-trophic
TW influence. Furthermore, phytoplankton concentrations, production
rates, and chlorophyll a concentration may be limited by light in the winter.
Shelf break and coastal STW upwelling favour high biomass concentrations
in slope and coastal waters, respectively, the extent of STW intrusion over the
shelf being an important factor. Large-scale coastal upwelling events lead to
high chlorophyll a values (up to 7 mg m'3 ) in surface waters (Hubold 1980a).
In contrast, incomplete upwelling and mixing result in sub-surface
chlorophyll a maxima (approx. 3 mg m'\ owing mainly to the growth of
large diatoms with an increased cell chlorophyll a content, rather than to an
increase-in cell number (Odebrecht and Djurfeldt 1996).
108 C. Odebrecht and V. M. T. Garcia: Phytoplankton

Table 6.7.1 Particulate primary production rates (mg C m-2 h- 1 ) in different

seasons off the coast of southern Brazil.

Season mean min. max.

Winter' 12.3 8.5 17.2
Late winter'
, 78.6 44.8 176.7
Spring 120.3 52.6 344.3
Summerd 27.3 9.0 56.7
Fall'" 36.0 3.4 98.1
'Garcia (unpubl.). bCiotti (1990). 'Teixeira et al. (1973). dOdebrecht (unpub!.).
'Gonzalez (1994).

Bloom Events

Under favourable climatic, hydrographic, and physico-chemical condi-

tions, phytoplankton blooms are a common phenomenon along the south-
ern Brazilian coast. Unidentified red-coloured surface patches have been
observed since the beginning of the century (Cunha and Fonseca 1918), and
high concentrations of the diatoms Coscinodiscus, Rhizosolenia, and Nitzschia
(Barth 1968), of the cyanophyte algae Anabaena and Trichodesmium, and of
the dinoflagellate Ceratium massiliense (Rosa 1979) occurred at several
occasions in nearshore and coastal waters. Especially in the surfzone,
environmental conditions appear to favour blooms of some phytoplankton
species.
Large concentrated patches (109 cells fl) of the diatom Asterionellopsis
(=Asterionella) glacialis, with chlorophyll a concentrations up to 1647 mg m-3,
are a common feature of the surf zone in southern Brazil. Patch formation
is related to cycles of cold front passages every 6 to 10 days, which force
coastal waters onshore, and diurnal cycles (Odebrecht et al. 1995a). Since
mucus production of Asterionellopsis glacialis cells in unialgal cultures is
stimulated by the addition of fine sediments, diurnal chlorophyll a
oscillations in the surf zone may be a result of cyclic mucus production
which favours attachment of suspended particles to the cells and causes
their sedimentation. Cells are then likely to survive in low light on bottom
sediments, because under controlled conditions at low light (30 J..lE m-2 S-I)
the cellular chlorophyll a content (6.18 pg cell-I) of Asterionellopsis glacialis
exceeds that of cells (1.52 pg celn cultured under high light (300 J..lE m-2 S-I;
Garcia and Rorig, unpubl.).
The presence of potentially toxic dinoflagellate blooms in the surf zone,
such as Gyrodinium cf. aureolum, Dinophysis acuminata, and Noctiluca
scintillans have been observed on several occasions and may be largely a
result of rapid offshore cell growth during calm weather and their concen-
tration in the surfzone during cold front passages (Odebrecht et al. 1995b).
Protozooplankton 109

These conditions may have led to the massive mortality of the intertidal
molluscs Mesodesma macroides and Donax hanleyanus and the crustacean
Emerita brasiliensis along several hundred kilometers of southern Brazilian
beaches (Machado 1979; Rosa and Buselato 1981; Tommasi 1983; Garcia et
al. 1994; Odebrecht et al. 1995b).

6.8
Protozooplankton

C. Odebrecht

Proto zooplankton is composed of apochlorotic flagellates, ciliates, and

ameboid organisms. These heterotrophic protista are important ecological
intermediates of the microbial loop because they act on nutrient recycling,
as predators of other micro-organisms, and serve as prey for larger con-
sumers (Sieburth et al. 1978). Early citations of protozooplankton organ-
isms in southern Brazili'an shelf waters included tintinnid ciliates
(Tintinnopsis campanula, Tintinnopsis beroidea) and heterotrophic dino-
flagellates (Noctiluca scintillans, Peridinium = Protoperidinium divergens;
Cunha and Fonseca 1918). Estimates of abundance in formaldehyde-fixed
bottle samples from various depths demonstrated that zoo flagellates and
naked apochlorotic dinoflagellates (Gymnodiniales) were the most abun-
dant forms (103 cells rl; Hentschel 1932).
Proto zooplankton organisms appear to be an important component of
the pelagic shelf system (Appendix). Both the vertical and horizontal
distribution patterns of their cell concentrations tend to coincide with
phytoplankton and chlorophyll a concentrations, and are higher in either
surface layers or under the influence of nutrient-rich SAW, STW, and CWo
Because of CW, cell concentrations of Noctiluca scintillans, Gyrodinium
spp., Polykrikos schwartzii, unidentified dinoflagellates, and naked ciliates
tend to be high (>10' cellsn near the Patos Lagoon mouth (Castello et al.
1990). Because of SAW, large (40-80 J..Illl) naked ciliates (Strombidium),
tintinnids (Tintinnopsis, Eutintinnus), and large heterotrophic dino-
flagellates (Gyrodinium, Protoperidinium) exceed cell concentrations of
10' cells r1 in shelf waters (Ciotti 1990). In contrast, in oceanic regions,
protozooplanktonic organisms are large\y represented by small «20 11m)
n
naked ciliates (103 -10' cells in oligotrophic warm waters of the Brazil
Current (Dohms 1983; Ciotti 1990).
110 M. Montu, I. M. Gloeden, A. K. Duarte, and C. Resgalla Jr.

6.9
Zooplankton

M. Montu, 1. M. Gloeden, A. K. Duarte, and C. Resgalla Jr.

Taxonomic Composition and Distribution

Coastal and marine zooplankton is represented by foraminiferans (37),

medusae (53), siphonophores (32), polychaetes (23), molluscs (29), clado-
cerans (7), ostracods (67), copepods (152), mysids (5), euphausiids (26),
chaetognaths (20), and tunicates (39; Appendix). The species composition
and the spatial and temporal distribution of the zooplankton organisms are
directly related to the dominance of different water masses over the south-
ern Brazilian shelf and slope. Although planktonic foraminiferans are good
indicators of different water masses, they are generally less abundant,
except for the more frequent Globigerinoides ruber, Globorotalia inflata,
Globorotalia menardii, and Globoquadrina dutertrei (Boltovskoy 1959,
1970, 1981; Closs and Barberena 1960, 1962; Roettger 1970; Be and Tolderlund
1971).
The abundance and diversity of medusae increase during the spring and
summmer, with Liriope tetraphylla and Proboscidactyla ornata being
dominant (Meneghetti 1973). While some species (Aglaura hemistoma,
Rhopalonema vela tum, Aegina citrea, Liriope tetraphylla, Proboscidactyla
ornata) have a widespread distribution (Ramirez and Zamponi 1981), others
are more restricted to either coastal (Obelia spp., Eucheilota ventricularis)
(Navas-Pereira 1973) or shelf regions (Zanclea costata, Cytaeis tetrastyla,
Bougainvillia ramosa, Bougainvillia platygaster, Sarsia eximia, Turritopsis
nutricula, Proboscidactyla ornata, Cunina peregrina). Both Aglaura
hemistoma and Cunina octonaria are typical of the Brazil Current
(Vannucci 1957, 1963; Navas-Pereira 1973).
Like medusae, siphonophores (Chelophyes appendiculata, Diphyes bojani,
Bassia bassensis, Abylopsis eschscholtzi, Muggiaea kochi, Enneagonum
hyalinum) are most abundant in high temperature and salinity waters
during the spring and summer. Muggiaea kochi and Enneagonum
hyalinum occur preferentially in nearshore waters, whilst Chelophyes
appendiculata, Eudoxoides spiralis, Diphyes dispar, Diphyes bojani, and
Bassia bassensis appear in the fail. Under the influence of different water
masses in winter, Lensia subtilis, Diphyopsis mitra, Sulculeolaria quadrivalvis,
and Abylopsis tetragona occur. Only Hippopodius hippopus can be found
throughout the year in oceanic regions (Gloeden, unpubl.). In addition,
shelled planktonic pteropod molluscs (Limacina rrochiformis, Creseis
virgula) , some characteristic of the Brazil Current (Limacina lesueuri,
Limacina bulimoides) and others of wider distribution (Cavolinia inflexa,
Clio pyramidata, Hyalocylis striata), are most abundant during the
Zooplankton 111

summer. Species of the genera Creseis and Cavolinia as well as Limacina

inflata, Limacina bulimoides, Limacina lesueuri, and the predominant
Limacina f. retroversa are restricted to southern shelf waters in the winter
(Resgalla and Montu 1994).
The cladocerans are essentially neritic, generally with higher concentra-
tions close to the coast (Navas-Pereira 1973). The species richness rises in
the winter when Penilia avirostris, Pseudoevadne tergestina, Evadne spinif-
era, and Podon intermedius become common. Species like Pleopys
polyphemoides and Evadne nordmanni are indicators of cold and Podon
schmackeri of warm coastal waters, whilst Penilia avirostris is characteristic
of the Brazil Current (Resgalla and Montu 1993).
As elsewhere, copepods (up to 2000 org. m-3 ) constitute the most abun-
dant group of zooplankton in southern Brazilian neritic and oceanic
waters. The dominance of SAW during the winter favours the presence of
cold water species (Calanus simillimus, Ctenocalanus van us, Calanoides
carinatus, Eucalanus elongatus, Centropages brachiatus, Pontella marplat-
ensis, Oncaea conifera, Oithona similis) over the shelf and the occurrence of
Rhyncalanus nasutus, Conea rap ax, and Acartia danae in oceanic regions.
During the summer, typical TW species (Paracalanus quasimodo, Paraca-
lanus parvus, Calocalanus pavoninus, Clausocalanus furcatus, Temora
stylifera, Oithona nana) occur. During this period Paracalanus campaneri,
Mecynocera clausi, Calocalanus pavo, Lucicutia flavicornis, Acartia negligens,
Oncaea venusta, Corycaeus furcifer, Corycaeus latus, Sapphirina auronitens,
Sapphirina ovatolanceolata, and Sapphirina scarlata are found more
frequently in oceanic regions, and other species like Acartia tonsa,
Paracalanus parvus, Temora stylifera, Eucalanus sewelli, Euterpina acutifrons,
and Clytemnestra rostrata are common to the shelf (Bjornberg 1963; Duarte
and Montu, unpubl.).
Species of mysids tend to be abundant in coastal waters, and Metamysi-
dopsis ?longata and Neomysis americana may constitute up to 90% of the
zooplankton (Bersano 1994). In contrast, euphausiids are generally more
common in slope and oceanic regions, and especially the adult forms of
some dominant species (Euphausia similis, Euphausia recurva, Euphausia
lucens, Thysanoessa gregaria) appear to have a preference for these waters
(Gorri 1995). Several Euphausiacea (Euphausia lucens, Nematoscelis megalops,
Thysanoessa gregaria) occur in SAW, whilst others (Stylocheiron
abbreviatum, Stylocheiron affine, Thysanopoda aequalis, Thysanopoda
monacantha, Thysanopoda obtusifrons, Thysanopoda orientalis, Euphausia
brevis, Euphausia hemigibba, Euphausia recurva, Euphausia spinifera,
Nematobrachion flexipes) are present in TW (Seguin 1965; Barth 1966;
Mauchline and Fischer 1969; Ramirez 1971; Brinton 1975; Mauchline 1980;
Gorri 1995).
112 M. Montu, I. M. Gloeden, A. K. Duarte, and C. Resgalla Jr.

30· 30·

SUMMER FALL
x=O.42 x:0.30
55' 50' 55· 50·

30' 30°

WINTER SPRING
x:0.31 )(:0.27

50· 55°

00.01-0.25 00.25~.S D O.s~.75 .0.75-1.0

Fig. 6.9.1. Distribution of zooplankton biomass in volume (ml m·') over the continental
shelf

Several species of the Chaetognatha are indicators of TW (Sagitta serra-

todentata), STW (Krohnitta subtilis, Sagitta decipiens), CW (Sagitta tenuis),
and SAW (Sagitta tasmanica; Resgalla 1993). Other common cold water
species include Sagitta planctonis, Sagitta maxima, and Eukrohnia hamata
(Boltovskoy and Jankilevich 1981; Duarte and Montu, unpubl.). Through-
out the year, Sagitta tenuis is the most abundant species close to the coast
and over the shelf, whilst Sagitta enflata, Sagitta serratodentata, Sagitta
minima, Sagitta bipunctata, Sagitta hispida, and Pterosagitta draco occur in
shelf and slope water. Similarly, different species of tunicates (Appendicularia)
are more abundant in nearshore water (Oikopleura dioica), shelf and slope
Zooplankton 113

water (Oikopleura longicauda, Oikopleura fusiformis, Oikopleura rufescens,

Fritillaria borealis, Fritillaria pellucida), or occur predominantly in oceanic
water (Oikopleura albicans, Oikopleura cophocerca, Stegosoma magnum,
Fritillaria formica). Species of the Thaliaceae (Ihlea punctata, Ritteriella
retracta, Salpa aspera, Salpa fusiformis, Cyclosalpa polae, Brooksia rostrata,
Thalia democratica) are of tropical or subtropical origin and abundant
during the spring and summer, whilst the subantartic Ihlea magalhanica
and Salpa thompsoni are common to epipelagic layers (Amaral, unpub!.).

Migration

The thermocline appears to control the vertical migration patterns of

several zooplankton species, such as the migration of medusae to layers
above (Bougainvillia platygaster, Euphysora furcata) and below (Cunina
peregrina) the thermocline (Correia 1983) as well as the migrations of
Siphonophora (Lensia campanella, Lensia cossack, Sulcoleolaria quadri-
valvis, Sulcoleolaria turgida, Sulcoleolaria chuni) from deeper layers to the
surface. Although Pteropoda are known for their pronounced vertical
migration (Haagensen 1976), short distance migration of Limacina species
between epipelagic and meso pelagic layers are common in southern
Brazilian waters. During the winter, nictimeral migrations of Chaetognatha
over the slope result in high concentrations in surface (Sagitta tenuis,
Sagitta tasmanica) and epipelagic layers (100 m; Eukrohnia hamata,
Krohnitta subtilis, Sagitta decipiens) at night, whilst Sagitta enflata, Sagitta
hexaptera, Sagitta lyra, and Pterosagitta draco concentrate in surface layers
during the day. In contrast, direct migration of the three latter species
during the summer results in high concentrations at night (Resgalla 1993).

Biomass

Zooplankton biomass evaluations provide important information to help

clarify community structure and energy flow among trophic levels. Just as
for species composition, different water masses control the spatial and
temporal distribution patterns of zooplankton biomass. Generally, biomass
values tend to decrease northward and with increasing distance from the
coast. High carbon values over the shelf are related to the influence of SAW
and CW, with an important contribution from continental runoff from the
La Plata River and the Patos Lagoon during the coldest months. However,
the highest carbon values (up to 98,47 mg C m-3 ) occur during the spring
and summer, principally in coastal surface waters. Especially in the surf
zone, mysids may achieve concentration peaks of 8142 mg C m- 3 (Bersano
1994).
114 N. M. Gianuca

Annual variations of estimated zooplankton volumes (Bongo net sam-

ples) are highest (> 1 ml m-3 ) during the summer and fall in areas close to
the coast, owing to the abundance of Thaliacea species which are large
organisms with a low energetic content (Meneghetti 1973; Navas-Pereira
1973; Hubold 1980a, b). In contrast, during the winter and spring, the high-
est values (0.25-1 ml m-3 ) occur in offshore areas under the influence of
SAW and Subtropical Convergence (Fig. 6.9.1). Owing to the diversity of
the zooplankton community, the summer and fall biomass peaks are fre-
quently associated with gelatinous plankton, whilst the winter and spring
biomass is largely a function of copepods and occasionally of euphausids.

6.10
Benthic Beach Invertebrates

N. M. Gianuca

The extensive, gently sloping, fine grained sandy beaches of Brazil's

extreme south favour the occurrence of an abundant and diverse macro-
benthic fauna (Appendix). Their vertical distribution follows a typical
zonation pattern, including a backshore or supratidal zone, an intertidal
zone, and inner and outer surfzones (Fig. 6.10.1). Early studies of beach
invertebrates were concerned with the assessment of commercially valuable
stocks (Gianuca 1971) or concentrated on taxonomic and qualitative sur-
veys (Orensanz and Gianuca 1974; Escofet et al. 1979). More recent quali-
quantitative studies characterized the fauna of the intertidal zone (Gianuca
1983, 1985, 1987; Santos 1990) and the inner (Paes 1989; Soares 1992) and
outer (Borzone and Gianuca 1990) surfzone, revealing that they are among
the richest and most productive sandy beach environments anywhere.

Supratidal Organisms

The supratidal zone, or backshore, is only flooded by seawater during

extreme storm tide events. Among the organisms that inhabit this zone,
Orchestoidea brasiliensis is restricted to the lower fringe, whilst Ocypode
quadrata occupies the entire upper part of the beach (Gianuca 1985, 1988).
However, most of the resident species are of terrestrial origin. Insects con-
stitute the dominant fauna and are represented by well-adapted coleopter-
ans, like the beetles Phaleria testacea, Bledius bonariensis, Bledius micro-
cephalus, Bledius Jernandezi, Cicindela nivea conspersa, and Cicindela
patagonica. Several predators (Carabidae) are especially attracted by the
high densities (2350 indo m-2 ) of Bledius spp. (Gianuca 1987). A particular
association of insects concentrates in moist sands around freshwater
creeks, which drain marshes behind the fore dunes. The most conspicuous
Benthic Beach Invertebrates 115

MdJita quinquiesperforata
Magdona riojai
OliYanciliaria deshayesia.n a
Mactra isabelleana
Syniljotea marpiatensis
AnClnus gaucho
Diastylis sympterigiae
-
PhoxocephaJopsis zi=ri ___ I
Arcnaeus cribarius
OJivancilJaria auricularia
Macrochiridothea lilianae
Buccinanops duartei
$jgaJion Clrriferum
------
___ I

ITonax gemmula
EuzolJuS fUEi.f.eruli
~

-
Emerita brasiliellSJS

~
Spio gaucha
Donax hanJeyanus
Mesodesma mactroides
ExciroJana armata ~I
Orchestoidea brasiliensis ~~
BJedius bonariensis
Ocxpode quadma .:::'
...
Cicindda patagonica
~ ~
r--....:: ~ ~

Ft ST IT ---- ISZ----~
'--- ---osz--
Fig. 6.10.1. Zonation of principal invertebrates between foredunes and surf zone: fore-
dune (FD); supratidal (ST); intertidal (IT); inner surf zone (lSZ); outer surf zone (OSZ)

species are the beetles Efflagitatus freudei and Paracymus rufocinctus and
the orthopteran Neotridactylus carbonelli. All feed on organic detritus,
microscopic psammic algae, and associated protozoans in the surface layer
of the sand. The principal predators of most supratidal insects are the resi-
dent birds Charadrius collaris and Anthus correndera.

Intertidal Organisms

A wide intertidal zone is characteristic of intermediate and dissipative

beaches. Most resident species exhibit vertical distribution patterns, but
since their vertical migration depends on tidal flooding, their distributional
range follows seawater level and beadi profile changes (Gianuca 1987).
Along dissipative beaches, some of the subtidal species, like Neocallichirus
mirim and associated crustaceans (Pinnixa patagoniensis, Hemicyclops
subadhaerens, Yoninae sp.), polychaetes (Grubeulepis bracteata, Leocrates
sp.), the turbelarian Stylochoplana sp., and an actiniarian which inhabit its
tunnels, extend into the lower intertidal zone (Gianuca 1985).
The biomass and production of benthic invertebrates are highest in the
intertidal zone (Gianuca 1983, 1985). Accumulated annual production
(ash-free dry weight) of Emerita brasiliensis and the bivalves Mesodesma
116 N. M. Gianuca

mactroides and Donax hanleyanus equals 238.5, 185.3, and 13.6 g m-2 ,
respectively. In contrast, in aIm wide transect, the highest annual pro-
duction was associated with Mesodesma mactroides (3251.76 g m- I ), owing
to the wider vertical distribution range of this species, followed by Emerita
brasiliensis (499.32 g m- I ) and Donax hanleyanus (55.57 g m- I ). Together,
these species are responsible for approximately 95% of the total intertidal
invertebrate biomass, and probably also for the total annual production
(Gianuca 1985). Other abundant species are the suspension-feeding poly-
chaete Spio gaucha, with a biomass of 35 g m-2 and an annual production of
114 g m-\ and the isopod Excirolana armata (8 g m-2 ) with densities of up to
7075 indo m-2 (Gianuca 1985; Santos 1990). During high tide, the consider-
able density and biomass of intertidal invertebrates attract several preda-
tors, like fishes, gastropods, and swimming crabs, whilst at low tide resi-
dent and migrating birds become the main predators (Gianuca 1983).

Surfzone Organisms

The shallow and turbulent sandy bottoms of the inner surfzone, between
the beach and about 2 m depth, are under the influence of regular wave
action and strong currents. Frequent winter storms further increase the
stressful hydrodynamic and substrate conditions. As a result, only 10 of the
60 benthic inner surfzone species occur during the entire year. The bivalve
Donax gemmula is the most abundant species (Gianuca 1985; Paes 1989).
Other common organisms are the crustaceans Pinnixa patagoniensis,
which inhabits the galleries of Neocallichirus mirim, Macrochiridothea
lilianae, Macrochiridothea giambiagiae, Arenaeus cribarius, and Phoxo-
cephalopsis zimmeri, gastropods (Buccinanops duartei, Olivancillaria
auricularia, Olivancillaria uretai), and polychaetes (Sigalion cirriferum,
Nephtys simoni; Gianuca 1983, 1985, 1988). Benthic macrofaunal densities
of up to 431000 indo m- I near Cassino Beach (3l14' S) are among the high-
est ever recorded for inner surfzones (Soares 1992).
The outer surfzone (2-10 m depth), although exposed to high wave tur-
bulence during storms, is inhabited by a total of 126 species (Borzone and
Gianuca 1990). Crustaceans make up 39%, with the isopods Ancinus gaucho
and Synidotea marplatensis and the cumacean Diastylis sympterigiae being
the most abundant species. Among the molluscs, which contribute 32%,
small species «15 mm) like Donax gemmula and Parvanachis isabellei and
larger species (15-70 mm) such as Olivancillaria deshayesiana, Mactra
isabelleana, and Dorsanum moniliferum are predominant. Thirty species of
polychaetes represent 26% of the benthic community. The most abundant
ones are Stheneloplis oculata, Magelona riojai, Parandalia americana,
Sthenelais limicola, Kinbergonuphis difficilis, Nephtys simoni, Odontosyllis
hetero-falcheata, and Owenia fusiform is. Mellita quinquiesperforata and
Continental Shelf Benthos 117

Amphioplus lucyae are the dominant echinoderms, which represent 3% of

the total number of benthic species.
The vertical distribution of the benthic macrofauna in the surf zone is
largely a function of depth. Cluster analysis identified three associations
(Borzone and Gianuca 1990). The first corresponds to shallower areas (2-5 m)
and is characterized by Olivancillaria auricularia, Buccinanops duartei,
Corbula caribaea, Donax gemmula, Macrochiridothea lilianae, Sigalion
cirriferum, Diopatra viridis, and Mellita quinquesperforata. A second asso-
ciation, with higher diversity but few exclusive species, like Olivancillaria
carcellesi, Olivancillaria urceus, Acteon pelecais, Epitonium angulatum,
Apoprionospio pygmaea, and Amphicteis sp., occurs at depths between 5 and
8 m. In deeper waters (8-10 m) of the outer surfzone, a third association is
dominated by Kinbergonuphis difficilis, Paraprionospio pinnata, Mactra
isabelleana, Tellina petitiana, Tellina sandix, and Terebra gemmulata.
However, several species (Diastylis symppterigiae, Synidotea marplatensis,
Ancinus gaucho, Abra lioic:a, Parvanachis isabellei, Parandalia americana)
have a wider distribution and can be found in high densities throughout the
outer surfzone.

6.11
Continental Shelf Benthos

R. R. Capitoli

Inner Shelf Fauna

Shallower inner shelf regions (10-50 m depth), which are under the
influence of highly turbid freshwater runoff from the Patos Lagoon, suffer
periodic perturbations because of elevated silt and clay deposition (Calliari
and Fachin 1993). The resulting environmental instability induces a state of
disequilibrium (Borzone 1988) since recolonization by species without
pelagic life-cycle phases is frequently unsuccessful. Common species in this
area include molluscs (Mactra isabelleana, Natica limbata, Parvanachis
isabellei, Olivancillaria urceus, Abra lioica), polychaetes (Kinbergonuphis
difficilis, Paraprionospio pinnata, Neanthes bruaca, Parandalia americana),
cumaceans (Diastilis sympterigiae), and isopods (Synidotea marplatensis,
Ancinus gaucho). Based on depth and sediment type the 93 molluscs form
five associations between 10 and 50 m depth (Appendix; Absalao 1986). At
a depth of 10-20 m, the omnivorous Parvanachis isabellei and opportunists
like Mactra isabelleana and Solen tehuelchus dominate an association with
low diversity in depositional areas where physical processes tend to rework
the muddy (30%) sediments. A deeper placed (20-40 m) association in pre-
dominantly fine and very fine sandy substrate is characterized by Tellina
gibber, Cadulus tetrachistus, and Olivella puelcha. Co-occurring species
118 R.R.Capitoli

}PO 500
.~ .........

100 km

Ht::::H !~Oe~ ~~ assoc~tion

1

D inner shelf association

(15 -25m)

D inner shell transitional

bottoms (25 - 5Om)

Sha low intermediate shelf

association (50 - BOrn)
~ ,,.: , Intermediat. she~
,:;:,. association (80 -130m)
.:o:})V· .~
.' '-~~::;.:- ... - Ollter she~ blodetrrtal
••• • :.> bottoms (130 - 220m)

;:l:!i~:;:'<'
n
1::':['::1:,:1 f,~r_ ~m')ssOOation
.... 50
1 00
Slope association
.·····50···
....... ... (200 - 500m)
20'0 500

Fig. 6.11.1. Macrobenthic invertebrate associations of the continental shelf and slope

include Pitar rostatus, Mactra petiti, Adrana electa, the predators Natica
limbata and Buccinanops lamarcki, the polychaete Owenia sp., and the
echinoderms Astropecten armatus, Olivancillaria articulatus, and Encope
emarginata. Nucula puelcha, Corbula patagonica, and Dentalium infractum,
together with ophiuroids (Amphiura joubini) represent an association in
sandy mud at approximately 40 m depth. Two less extensive areas of variable
sandy substrate types, proximate to beachrock outcrops, are occupied by
Halistylus columna and Crassinella lunulata and by Olivancillaria urceus and
Glycymeris longior, respectively (Absalao 1986).
Continental Shelf Benthos 119

More distant inner shelf areas south of 3i S are not affected by runoff
from the Patos Lagoon. The shallower (10-15 m) sandy areas (Fig. 6.11.1)
are characterized by the presence of cumaceans, isopods, the penaeid
Artemesia longinaris, and the molluscs Dorsanum moniliferum, Olivancil-
laria urceus, Adelomelon brasiliana, and Mactra isabelleana, the last
becoming more abundant with increasing silt concentrations. Deeper (15-
25 m) areas with fine sand (Fig. 6.11.1) are occupied by Encope emarginata,
Buccinanops lamarcki, Loxopagurus loxochelis, Astropecten armatus,
Adrana electa, isopods of the genus Serolis, and polychaetes of the family
Owenidae. Transitional sandy mud bottoms (25-50 m; Fig. 6.11.1) between
the inner and intermediate shelf regions are characterized by Pitar rostratus,
Corbula patagonica, Buccinanops gradatum, Mactra petiti, Dardanus arrosor,
Persephona punctata, Hepatus pudibundus, and ophiurids. Phyllochaetopterus
socialis, Astrangia rathbuni, serpulid polychaetes, and Crepidula pro tea are
conspicuous organisms in areas with frequent beachrock outcrops,
consolidated substrates, and accumulation of shell fragments. Chlamys
tehuelcha, Acroscalpellum bubaloceros, Crepidula aculeata, Atrina seminuda,
and Pteria hirundo may co-occur in less extensive areas.

Intermediate and Outer Shelf and Slope Fauna

Surveys of the benthic macrofauna of the southern Brazilian intermediate

and outer shelf and slope (50-500 m) have largely focused on the general
distribution of species (Tommasi 1969; Tommasi et al. 1973; Rios 1994).
South of 32° S an association, represented by Pitar rostratus, polychaetes
(Asychis sp., Owenia sp.), amphipods, and ophiurids, is relatively abundant
in muddy sand bottoms of the shallower (50-90 m) intermediate shelf
(Capitoli, unpubl.). These areas also display the highest relative abundance
of demersal and benthic fish among all shelf areas (Haimovici et al. 1994a,
1995). The association of the extensive sandy mud bottoms of the interme-
diate shelf (80-130 m; Fig. 6.11.1) displays the highest faunal abundance,
with densities and biomass of 500-1000 indo m-2 and 10-100 g m-2 , respec-
tively. The macro benthic assemblage is largely represented by Diopatra
tridentata, Chasmocarcinus typicus, Terebellides sp., Onuphis tenuis,
Astropecten cingulatus, Squilla brasiliensis, and Portunus spinicarpus. In
muddy bottoms (100-130 m) the abundance of characteristic assemblage
species decreases, and the polychaetes Eupanthalis rudipalpa, Panthalis
oerstedi, Spiochaetopterus sp. and the sipunculid Nephasomma sp. become
more abundant. Low faunal abundance coincides with regions of biodetrital
bottoms (Tommasi et al. 1973), which occupy large areas between 130 and
220 m depth north of 32° S but become narrower around shelf break areas
(170-200 m) further south (Fig. 6.11.1). Biodetrital bottoms are composed
of a mixture of sand/silt with either exoskeleton debris of ahermatipic
120 C. Sinque and J. H. Muelbert

corals (Cladocora debilis, Trochocyathus sp.) in the north or shell frag-

ments and debris of the coral Rhizopsammia manuelensis in the south. The
macrobenthic invertebrate association of outer shelf (130-200 m) bottoms
(Fig. 6.11.1) is poor (100-500 org. m-\ <10 g m- 2) and is largely composed of
gorgonians (Primnoella bisseialis, Primnoella compressa, Elisella barbadensis,
Elisella elongata, Muriceides hirtus, Bebryce cinerea, Acanthogorgia
scharammi, Muricea midas), echinoderms (Anthenoides brasiliensis,
Ophiomisidium pulchellum, Brissopsis atlantica, Stylocidaris affinis,
Crinometra sp.), isopods (Cirolana exigua), barnacles (Rostratoverruca
nexa), and serpulid polychaetes. Slope areas (200-500 m) display faunal
heterogeneity and low density «100 indo m-2 ) and biomass «10 g m-2 ).
However, different depth strata (200-300 m and 300-500 m) with varying
substrate composition appear to select for different species dominance.
Consolidated substrates of upper slope regions are typically occupied by
Eunice frauenfeldi, Zoanthidea sp., and species of Porifera, Actinaria, and
ophiurids. Co-occurring species include Myriopsis quinquespinosa, Stylocidaris
affinis, Verruca carlbea, Brissopsis atlantica, and Spiochaetopterus sp.
Lower slope regions are characterized by Acanella eburnea. Flabellum bra-
siliensis (coral), Ophiurolepis sp. (ophiurid), and Priapulus sp. are relatively
constant components. Molpadia sp. (holothurian), Batyplax typhla
(brachyurid), Bathynomus giganteus (isopod), Antipathes sp. (black coral),
and Cephalodiscus (pterobranchia) are occasionally present.

6.12
Ichthyoplankton

C. Sinque and J. H. Muelbert

Taxonomic Composition

Initial ichthyoplankton surveys over the southern Brazilian shelf and slope
(Aboussouan 1969) aimed to elucidate aspects related to fishery biology.
Eggs and larvae of about 88 fish species occur in the plankton community
and are distributed among 48 families (Appendix). Three species of
clupeids (Sardinella brasiliensis, Harengula jaguana, Brevoortia pectinata;
Hubold and Ehrlich 1981) and of Engraulididae (Lycengraulis grossidens
=L. olidus, Engraulis anchoita, Anchoa marinii; Weiss et al. 1976; Weiss and
Souza 1977a,b; Phonlor 1984a,b) occur in this region. Stomiiformes are
recognized as the most important component of oceanic mid-water
ichthyofauna (Bonecker and Hubold 1990). They are represented by the
families Sternoptychidae (Valenciennellus tripunctulatus), Photichthyidae
(Vinciguerria nimbaria, Vinciguerria poweriae, Vinciguerria attenuata,
Pollichthys mauli), Stomiidae (Stomias sp.), Idiacanthidae (Idiacanthus sp.)
and Gonostomatidae. The Gonostomatidae are represented by five species of
Ichthyoplankton 121

Cyclothone, Diplophos taenia, Gonostoma spp., Magarethia obtusirostra, and

Maurolicus muelleri which is the most abundant species among the
Stomiiformes (Weiss et al. 1988; Bonecker and Hubold 1990).
Larvae of Myctophidae occur throughout the year and are occasionally the
dominant component of the ichthyoplankton. The most important species
are Diogenichthys atlanticus, Hygophum reinhardtii, Myctophum spp., and
Diaphus spp. which may account for up to 60% of myctophid larvae. Among
the Gadiformes, Bregmacerotidae is the most abundant family, with
Bregmaceros sp. making up 44% of the larvae. Although commercially
important, planktonic stages of Urophycis spp. (Gadidae) and Merluccius
hubbsi (Merluccidae) are not abundant, and larvae of Coelorhynchus sp.
(Macrouridae) are rare.
Eight species of Sciaenidae occur in the ichthyoplankton (Weiss 1981;
Muelbert and Weiss 1991; Ibagy and Sinque 1995), and the abundant
Micropogonias jurnieri, Paralonchurus brasiliensis, and Macrodon ancylodon
are important local fishery resources. The suboide-r Scombroidei is
represented by larvae of Gempylidae (Diplospinus multistriatus, Thyrsitops
lepidopoides, Neolatus tripes) and Scombridae (Scomber japonicus, Sarda
sarda, Katsuwonus pelamis, Euthynnus alletteratus, Auxis sp.) and by the
eggs and larvae of Trichiuridae (Mafalda 1989). Among the Trichiuridae, the
eggs and larvae of Trichiurus lepturus are the most abundant ones, while
larvae of Lepidopus caudatus are less common. Pleuronectiformes,
represented by eggs and larvae of Pleuronectidae and larvae of Bothidae,
Cynoglossidae, and Soleidae, also comprise an important fisheries resource.
Larvae of Anguilliformes, Exocoetidae, Atherinidae, Zeidae, Caproidae,
Triglidae, Serranidae, Carangidae, Mugilidae, Gobiidae, and Balistidae are
less abundant in the ichthyoplankton (Weiss et al. 1976, 1987).

Spatial and Temporal Distribution

Although correlations between feeding, growth, predation, transport, and

retention of eggs and larvae and primary and secondary production
patterns are still lacking, it is clear that the spawning periods of different
fish species and the dynamics of different water masses influence both
species composition and large scale spatial-temporal distribution patterns
of eggs and larvae.
With the exception of the winter months, clupeid eggs are present
throughout the year. Their abundance is highest along the 30 m isobath and
decreases towards the open ocean. Brevoortia pectinata spawns in coastal
areas, and larvae of different developmental stages enter the nursery areas of
the Patos Lagoon estuary (Weiss et al. 1976). Engraulis anchoita occurs in
subtropical waters (STW) and in mixed waters of the La Plata River between
29° and 35° S (Weiss and Souza 1977b), but the greatest amount of spawning
122 c. Sinque and J. H. Muelbert: Ichthyoplankton
occurs at the southern limits of this area under optimal temperature (9-20°C)
and salinity (30-36) conditions (Phonlor 1984b). A dominance of Engraulis
anchoita during the winter and spring can be attributed to the northward
displacement of the Malvinas Current (MC), whilst in the summer and fall,
owing to the influence of the warmer Brazil Current (BC), other species
dominate. Lycengraulis grossidens spawns in shallower coastal and less saline
waters between 12.9 and 18.4°C (Hubold and Ehrlich 1981) south of 28° 58' S;
the planktonic stages then grow further in estuarine areas (Weiss 1981;
Muelbert and Weiss 1991). Anchoa marinii is a typical marine species which
is virtually continuously present and widely distributed geographically,
though the eggs and larvae are concentrated over the southern region of the
shelf (Weiss et al. 1976; Hubold and Ehrlich 1981).
The eggs and larvae of Stomiiformes are distributed in tropical and tem-
perate pelagic oceans (Bonecker and Hubold 1990). The eggs of Maurolicus
muelleri occur throughout the year in deeper waters «250 m) but are most
abundant in the fall, whilst larvae are common between 100 and 150 m depth
(Weiss et al. 1988) and reach their highest densities during the winter and
spring at mean temperatures between 15 and 25°C and salinities of about 36.
The larvae of Cydothone spp. and Vinciguerria spp. are confined to oceanic
waters of the BC below 200 m depth. Their highest concentrations are
reached in the winter and spring between 18 and 23°C and a salinity of 36.
Because of their low abundance, the distribution pattern of other
Gonostomatidae is less well defined. With the exception of Pollicthys mauli in
shelf waters of Cabo st. Marta (29° S), the species are always distributed in
oceanic waters (Bonecker and Hubold 1990).
The larvae of Myctophidae also have an oceanic distribution and occur
throughout the year along the shelfbreak. However, they are most abundant
over the southernmost Brazilian shelf region along the 100 m isobath in the
winter and summer (Diaphus spp.) when they may account for 18-25% of the
ichthyoplankton. The larvae of Bregmaceros sp. are most abundant in the fall.
Bregmaceros can tori and Bregmaceros atlanticus dominate over the outer
shelf north of 29° S, and the highest densities of Bregmaceros can tori larvae
are associated with the maximum chlorophyll layer (Matsuura et al. 1993).
The distribution of Merluccius hubbsi, which is highly influenced by
temperature (l2-13°C), salinity (26-28), and depth (60-90 m), is determined
by the northward movement of the MC over the shelf during the winter and
spring (Pereira 1983).
Sciaenids are demersal fish that inhabit coastal waters. In general,
spawning occurs close to the shore, and several species use the Patos
Lagoon estuary as a nursery area (Weiss 1981; Muelbert and Weiss 1991;
Ieagy and Sinque 1995). Although eggs and larvae are present throughout
the year, the highest densities are registered during the spring and summer.
Reproduction of Antigonias capros and Zenopsis conchifer occurs over the
shelf at about 200 m depth under the influence of the warm (l7.S-23.4°C)
Pelagic Teleosts 123

and saline (32.2-36.3) waters of the BC during the fall and spring, respec-
tively. Eggs and larvae of the eurythermal and euryhaline species Trichiurus
lepturus are present throughout the year, with the highest densities
appearing during the summer and fall in TW, STW, and CW (Mafalda
1989). During the summer and fall they occur closer to the shore and also in
the estuaris (Weiss 1981; Pereira 1986; Muelbert and Weiss 1991). Lepidopus
caudatus larvae concentrate during the spring and fall in TW and STW
north of this region.
The highest larval densities of the Gempylidae (Diplospinus multistriatus,
Neolatus tripes, Thyrsitops lepidopoides) occur during the winter in oceanic
waters and during the summer in coastal waters (Mafalda 1989). Larvae of
Scombridae (Scomber japonicus, Sarda sarda, Katsuwonus pelamis, Euthynnus
alletteratus, Auxis sp.) tend to occur during the summer in coastal regions
under the influence of TW. Similarly, Pleuronectiformes are most abundant
during the summer, when their eggs may account for 7,% of planktonic fish
eggs and their larvae for 17% of the ichthyoplankton. Larvae of Cynoglossidae
appear during the fall. Eggs and larvae of less abundant species of Exocoeti-
dae and Bregmacerotidae are noted in the summer, those of Triglidae in the
summer and fall, whilst Atherinidae larvae are common in the winter. Coastal
waters of the southern part of this region favour the presence of larvae of
Carangidae (Trachurus lathami =T. picturatus, Parona signata, Chloroscom-
brus crysurus) and Mugilidae during the summer and fall, respectively (Weiss
et al. 1976).

6.13
Pelagic Teleosts

J. P. Castello
Based on their distribution and ecological and economic importance, pelagic
fish (Appendix) of the southern Brazilian shelf and slope regions can be clas-
sified as (1) inhabitants of nearshore and shelf waters down to approximately
30 m depth, (2) inhabitants of the shelf between the coast and the break at
about 200 m depth, and (3) oceanic inhabitants of the slope and deeper
waters.

Shallow Coastal Water Inhabitants

Typical inhabitants of shallow waters are Trachinotus marginatus, Anchoa

marin ii, Lycengraulis grossidens (=L. olidus), Brevoortia pectinata, and
juveniles of Engraulis anchoita (Cunha 1987), whilst Centengraulis
edentulus and Anchoa tricolor are found rarely (Buckup 1984). Among the
different Trachinotus species (T. marginatus, T. goodei, T. carolinus,
124 J. P. Castello

T. falcatus), characteristic of turbulent surfzones with variable salinity,

Trachinotus marginatus, with a total length (TL) of 15-60 mm, is the most
abundant one (Cunha 1987). During their first year, the fast growing juve-
niles feed on molluscs (Mesodesma mactroides, Donax hanleyanus) and the
crustacean Emerita brasiliensis and then migrate to deeper waters. The two
engraulidids Anchoa marinii «150 mm TL) and Lycengraulis grossidens
(60-100 mm TL <180 mm; Castello and Krug 1978) and the clupeid
Brevoortia pectinata «350 mm TL; Figueiredo and Menezes 1978) occur in
coastal waters of less than 25 m depth, though the distributional range of
Lycengraulis grossidens is narrower than that of the other two species. Both,
Lycengraulis grossidens and Brevoortia pectinata reproduce principally
from the spring to fall and the early developmental and juvenile stages of
their life-cycle are associated with estuarine waters (Weiss and Krug 1977;
Castello and Krug 1978; Weiss 1981). During the spring, Anchoa marinii
commonly occurs together with Pomatomus saltatrix, Balistes spp.,
Peprilus paru, Sarda sarda, and Stromateus spp. (Mello et al. 1992).

Shelf Inhabitants

Pomatomus saltatrix, Sarda sarda, Scomber japonicus, Trachurus lathami,

Katsuwonus pelamis, three species of Mugil, and adults of Engraulis
anchoita are abundant pelagic species found over the shelf. Owing to its
wide distribution, great abundance, and role as a secondary consumer,
Engraulis anchoita is an important pelagic species in the Southwestern
Atlantic (23-45 0 S; Castello and Habiaga 1982; Habiaga and Castello 1986;
Acuna and Castello 1986). The distribution of adult shoals follows the
coastal branch of the Malvinas Current and is limited to the east by the
thermohaline front with the Brazil Current. Freshwater runoff from the
Patos Lagoon and La Plata River during the winter (Castello et al. 1990;
Lima 1992) provides stable water column conditions in the coastal zone for
the development of larvae which are retained in this region by Ekman
transport mechanisms (Lima and Castello 1995). Coastal zones north of 31 0 S
are preferentially occupied by larvae and juveniles, but further south larvae,
juneniles, and adults are associated with low-temperature waters of sub-
antartic origin (Castello and Habiaga 1982; Lima and Castello 1995). Near
sub-surface upwelling areas (290 S), the daily growth rates (0041-0.73 mm dail)
oflarvae (8-32 mm CP; Castello and Vasconcellos 1995) are similar to those
in Argentine waters (41 0 S) (Sanchez et al. 1991). Since males and females
abandon shallower coastal waters after reaching first maturation at 70-85 mm
TL (Phonlor 1984b; Acuna and Castello 1986), the size of individuals tends
to increase towards deeper waters. The species grows rapidly, and more
than 50% of the asymptotic length (175 mm) is achieved during the first
year. Adults congregate over the inner and intermediate shelves during
Pelagic Teleosts 125

reproduction in the winter and spring, and at least four spawning groups
have been identified for southeastern and southern Brazilian shelf regions
(Weiss and Souza 1977b; Nakatani 1982). Batch spawning occurs at night
(22.00-4.00 h) in near-bottom water (Acuna and Castello 1986) every 12-17 days
(Christiansen and Cousseau 1985). In the late spring and early summer,
adults migrate south to Uruguayan and Argentine waters, returning in the
winter. Adults of Engraulis anchoita feed during the day, and even though
their diet preference may overlap with Trachurus lathami and Scomber
japonicus, their diet diversity is considerably larger. About 90% of the food
is composed of calanoid copepods, euphausids, and hyperid amphipods,
though occasionally phytoplankton is also ingested (Schwingel 1991;
Schwingel and Castello 1994). The production of the stock, with up to 90%
during the first year onife, varies between seasons. A winter biomass of 60 g m-2
results in 44.2 g m-2 production, whilst the spring biomass (14 g m-2 )
produces 9.3 g m-2 , the natural mortality of Engraulis anchoita being the
principal limiting factor. Catches are largely composed of 1- and 2- year-
old individuals and equal ratios of males and females, though females tend
to dominate in mid-water trawls during peak production (Acuna and
Castello 1986). This considerable stock which, owing to differential
recruitment and fluctuating oceanographic conditions (Lima and Castello
1995), may vary interannually between 35 000 and 1 900000 tons, is still not
being commercially exploited.
Of the three Mugil species, only M. curema and M. gaimardianus are
essentially warm water representatives and are seen during the summer and
fall, when high water temperature and salinity conditions prevail. In contrast,
juveniles of the more common Mugil platanus persist throughout most of the
year, though they are more abundant in the winter and spring in nursery
areas of the Patos Lagoon estuary and adjacent coastal waters (Vieira 1985).
In the fall, which is the major period of catch, adults of Mugil platanus leave
the estuary, owing to gradually decreasing water temperatures and frequent
seawater penetration, and initiate their reproductive migration. Spawning
occurs in warmer (19°_21°C) offshore waters at about 2t S between the end of
the fall and winter (Vieira 1985; Viera and Scalabrin 1991). Eggs and larvae
are generally transported by currents towards the surfzone, followed by long-
shore migration to the Patos Lagoon estuary (Vieira 1991b).
The bluefish Pomatomus saltatrix has a wide biogeographic distribution
and is present throughout the year (Krug 1984). Decreasing temperatures
in Argentine waters (38° S) in late fall appear to stimulate migration to the
southern and southeastern Brazilian shelf, where the highest abundance is
reached in the winter. Approximately 50% of growth occurs during the first
2 years,. although males and females may reach a maximum age of 7 and
8 years, respectively. Batch spawning occurs during the late spring and summer
after males and females (35-40 cm TL) reach first sexual maturity at an age of
approximately 2 years. The period of reproductive activity coincides with a
126 J. P. Castello

second peak of abundance at approximately 3t S (Krug and Haimovici

1991). While juveniles of Pomatomus saltatrix «25 cm TL) feed on small
pelagic fish and crustaceans, adults (>25 cm TL) prefer anchovy, Atlantic
sabretooth anchovy, horse mackerel, and less frequently squids, cutlass fish,
and weakfish (Haimovici and Krug 1992). The scombriform Sarda sarda
(mean 53 cm TL, maximum 78 cm TL), which largely feeds on anchovy, is a
migratory species. Migrations from Argentina (38° S) into southern Brazilian
shelf waters are limited by superficial isotherms of 15-16°C (Aasen and
Castello 1968a,b). After reproduction in southern Brazilian shelf waters from
November to January, Sarda sarda returns to waters near Mar del Plata (38° S),
where the stocks form a resource for the Argentine purse seine fishery.
Scomber japonicus is present during the entire year and migrates latitudi-
nally in the Southwestern Atlantic. Reproduction occurs in the winter over
the southeastern Brazilian shelf. In spite of the assumption that only one
population exists between 22° and 4t S (Seckendorff and Zavala-Camin
1985), evidence based on morphometric an~ meristic characteristics
(Perrotta et al. 1990) and on reproduction in the spring at 38° S (Perrotta
1987) suggests the presence of two populations. Also, shoals of different size
classes reach the Mar del Plata region (Gagliardi and Cousseau 1970) and the
interannual abundance varies strongly (Castello and Cousseau 1976). In
southern Brazilian waters, the abundance of Scomber japonicus may alternate
with the abundance of the widely distributed horse mackerel (Trachurus
lathami; Saccardo 1980, 1987), thus modifying the composition of the pelagic
association (Mello et al. 1992). Trachurus lathami from Brazilian and Trachu-
rus picturatus australis from Argentine waters (38° S) have been considered as
two different taxa (Cousseau 1967; Ciechomski and Cassia 1980), however,
since they are morphologically identical the presence of two taxa is not justi-
fied (Saccardo 1980). The size of Trachurus lathami (85-210 mm TL) in
southeastern and southern Brazilian shelf waters is similar. Sex maturation
occurs at 115-119 mm TL, and even though the reproductive period is
prolonged peak reproduction is reached in November (Saccardo 1980, 1987;
Castello, unpubl.). The species can live to a maximum age of 8 years, but an
average of 2-3 and 3-4 years is found for the southern and southeastern
Brazilian shelf regions, respectively. During the day, shoals of Trachurus
lathami form at 50-90 m depth in southeastern and at 20-75 m depth in
southern shelf waters (Saccardo 1980, 1987; Castello, unpubl.). At night, the
species disperses in surface water and frequently mixes with Engraulis
anchoita (Mello et al. 1992).
The abundant small to medium-sized skipjack (Katsuwonus pelamis) is
widely distributed in tropical and subtropical oceans and represents an
i.l11portant fishery resource (Matsuura 1982a; Jablonski and Matsuura 1985;
Vilela 1990; Vilela and Castello 1991, 1993). Although Katsuwonus pelamis
undergoes extensive latitudinal migrations, the species is found throughout
the year over the southeastern Brazilian shelf (Maluf et al. 1984). Schools
Pelagic Teleosts 127

migrate to more southern regions in the late spring, where their distribu-
tionallimits coincide with the western margin of subtropical waters (l8°C,
3.5 ml r 1 dissolved oxygen) (Habiaga and Castello 1986). In these areas,
Katsuwonus pelamis feeds from November to May largely on Maurolicus
muelleri, Engraulis anchoita, and Euphausia similis. Other pelagic species of
the southern Brazilian shelf include Thyrsitops lepidopoides, Balistes spp.,
Lagocephalus laevigatus, Peprilus paru, Stromateus maculatus, Stromateus
brasiliensis, Parona signata, Saurida brasiliensis, Decapterus sp. (Castello,
unpubl.), and Auxis thazard (Miyake and Hayasi 1978). The species
Cynoscion guatucupa, Trichiurus lepturus, and Merluccius hubbsi only
become pelagic as juveniles or during nocturnal feeding (Mello etal. 1992).
Mid-water trawls produce a characteristic assemblage of pelagic species,
Engraulis anchoita, Trachurus lathami, Loligo sanpaulensis, Trichiurus
lepturus, Cynoscion guatucupa, and Merluccius hubbsi being the most
frequent ones in the winter and spring. A second assemblage during spring is
composed of Peprilus paru, Balistes sp., Pomatomus saltatrix, Sarda sarda,
Katsuwonus pelamis and Anchoa marinii (Fig. 6.l3.1; Mello et al. 1992).

Slope and Deep Water Inhabitants

Tunafish and similar species, pelagic sharks, and other smaller oceanic fish
occur abundantly over the slope or, under favourable environmental con-
ditions, also over the outer shelf. The tunafish Thunnus albacares, Thunnus
alalunga, Thunnus thynnus, and Thunnus obesus are commercially valuable
species, of which Thunnus albacares is the most important one. The
concentration of shoals is controlled by the influence of the Subtropical
Convergence. Thunnus albacares as well as Thunnus alalunga and Thunnus
obesus occur in southern Brazilian waters (27-35° S) from August to Febru-
ary and from April to September, respectively (Zavala-Camin 1974, 1978;
Zavala-Camin and Antero da Silva 1991). Since planktonic eggs and larvae
are absent from southeastern regions (Matsuura 1986; Mafalda 1989),
reproduction of Thunnus albacares occurs in the tropics (Zavala-Camin
and Antero da Silva 1991). In southern Brazilian waters, the age structure
of Thunnus albacares varies between 1 and 6 years (Vaske 1992), probably
as a result of the migration of two or three cohorts twice a year (Zavala-
Camin 1978). Juveniles form large schools in surface waters and often mix
with other species, like Katsuwonus pelamis, whilst adults occur in sub-
surface and upper mesopelagic waters (Zavala-Camin 1981). Both
Thunnus albacares and Thunnus alalunga prey during the winter and spring on
fish, principally Brama sp., cephalopods (Ommastrephidae and Ornitotheuthis
antillarum), and hyperid crustaceans (Zavala-Camin 1981; Vaske 1992). Other
tunafish species (Thunnus atlanticus, Thunnus maccoyii) are rare.
128 J. P. Castello: Pelagic Teleosts
ENGRAULIS ANCHOITA 87.5 rI

L~~I:~H~:~A~';= ~:::::::::}t:Jf:Jf):: ':::"::'::::::':::~~~", WINTER

i~a1ii~~~=il; :'-----
:::

~~=~=; ~: t-------------+---,
TRICHJURUS lEPTURUS 50.0 _
PEPRILUS PARU 12.5
PARONA SIGNATA 8.3 ..........--...
GALEORHINUS GALEI 16.7

POMATOMUSMACULATUS
STROMATEUS SALTATRJX 12.5
8.3---.1 ~--=~:::::::::J~---l
ANCHOA MARINII 16.7

THYRSITOPS LEPJDOPOJDES 8.3 !;;::::;;;=========j:=:::::==~

727667 559 471 360 300 183 099
ENGRAULIS ANCHOITA 91.8
TRACHURUS LATHAMI 42.6
LOLJGO SANPAULENSIS 42.6 SPRING
TRICHIURUS LEPTURUS
42.6
THYRSITOPS LEPIDOPDIDES 14.8
-r-
CYNOSCJON GUATUCUPA 23.0
MERLUCCIUS HUBBSJ 23.0 ~
MACRODONANCYlODON 6.6
KATSUWONUS PELAMIS 3.2 tr-
ANCHOA MARIN II 13.1
' ::::.
PEPRllUS PARU 9.8 ~
BALISTES SP. 8.2
:':':::"
POMATOMUS SALTATRIX 4.9
SARDASARDA 3.2
STROMATEUS SP. 6.5 ~

GALEORHINUS GALEI 4.9

PARONA SIGNATA 3.2 It-
SCOMBER JAPONICUS 3.2
SPHYRNASP. 3.2

667 600 417 309 174087 021

Fig. 6.13.1. Winter and spring dendrograms of pelagic species assemblages (shaded
areas) with frequency of occurrence

Several commercially valuable tunafish-like species, represented by bill-

fish, swordfish, and marlin (Euthynnus alletteratus, Coryphaena hippurus,
Xiphias gladius, Isthiophorus albica,!s, Tetrapturus albidus, Makaira nigri-
cans) are caught in southern Brazilian waters (Zavala-Camin 1974, 1978;
Mello et al. 1993). In particular, the swordfish (Xiphias gladius) , which
potentially competes with Thunnus obesus, is an important fishery resource.
The preferential diet of this species is Illex argentinus (Mello 1992). Finally,
smaller fish species, like Maurolicus muelleri, genera of the Mycthophidae
(Diaphus, Symbolophorus, Myctophum, Diogenichthys), and Lampanyctinae,
which undergo large vertical migrations, occasionally reach a high level of
abundance (Castello and Habiaga 1982, Souto, unpubl.).
Demersal and Benthic Teleosts 129

6.14
Demersal and Benthic Teleosts

M. Haimovici

Over the past few decades, the search for new fishery resources stimulated
several surveys of demersal and benthic teleost fish over the southern
Brazilian shelf and slope (Nakamura 1963; Vazzoler and Iwai 1971; Menezes
1971; Roux 1973; Yesaki 1973; Vazzoler 1975; Yesaki et al. 1976; Vooren et
al. 1988). Today, over 200 species from more than 100 families have been
identified (Menezes 1971; Benvegnu-Le 1973; Figueiredo 1977; Figueiredo
and Menezes 1978, 1980; Menezes and Figueiredo 1980, 1985; Haimovici et
al. 1994a; Appendix).

Seasonal Patterns of Species Richness and Abundance

The demersal ichthyofauna of the southern Brazilian shelf, transitional

between tropical and temperate zones, is part of the Argentine zoo-
geographic province, which extends between 22° and 42° S (Figueiredo
1981). The fauna can be divided into species with tropical and temperate
water affinities of both the northern and the southern hemisphere. Most
species are eurybathic and eurythermal, and few endemics are found.
Although the species composition varies little with the seasons, annual
variations of species richness and relative abundance follow annual bottom
temperature variations in coastal waters (> 10°C) and at the shelf break (4-
5°C), as a result of the latitudinal oscillation of the Subtropical Convergence
between the warm Brazil Current (BC) and the cold Malvinas Current (MC;
Miranda et al. 1973; Haimovici et al. 1996). In winter, the highest biomass
at 40-60 m (Micropogonias jurnieri, Umbrina canosai, Cynoscion guatucupa
= Cynoscion striatus) coincides with the northward flowing cold waters of
the coastal branch of the MC, though the relative abundance decreases
from coastal to deeper waters. In contrast, at the end of the summer, abun-
dance increases from the inner shelf towards the shelf break, probably
owing to the existence of vortices and to partial upwelling of subtropical
water (Podesta 1990; Lima and Castello 1995). The total number of species
varies little between seasons. The greatest species richness over the shelf
break is probably a result of both different types of substrate and different
water layers, whilst a pronounced decrease in total species number over the
slope below 350 m appears to be associated with the poor benthic inverte-
brate fauna (Capftoli, Sect. 6.11).
130 M. Haimovici

12 ~ ~ E E E E E
'"....... ...'" '"::J §
E
l!! Jl E E E '"
~ E
~ E
..,'" '"'" ~
2': !:::
g
M

g 0... ~ .... &\.... ...~

&\
~
!.! ::'" ~ 0 0
'"
000
Genera alld Species Family % %

Memicirrllll$ tiIlO,"/i Sciacnidac 25 61 39

Partllonc:lwrlIS brasiliensis Sciacnidac 103 49 51
MtlcrOtlolJ ancylodon Sciacnidac 83 53 47
Ptpr;/us IXln, Slromalcidac 103 89 II
Micropogollh1s!um;pr; Sciaenidac 172 52 48
Urophycis brtlsilicn,fis Gadidae 201 58 42
Netuma SI'. Ariit.!ac 83 M 36
Conger (Jrbign)'(wus Congridae 108 40 60
Cynoscion guatllCUI'XI Sciacnidae 222 50 50
Trichillrrt.f e
l pmrlls Trichuridae 231 64 36
Balisfl!S calJriscrl,f Balislidae 41 ~8 32
Umbrinll canosa; Sciacnidac 190 47 53
p,jonoltls IJlmCUlW ,f Triglidac 1M 57 43
PluaUcJr,Ir.','s fJQmgollicfls BOlhidae 136 55 45
Poric1lthys l>Ortui.uimu.f Batrachoididac 151 48 52
C,rnosci01I jamaicen.'iis Sciacnidac 88 76 24
CIt'lJoscitlena grnc:ilidrrllll,f Sci:lcnidac 87 72 28
TftlChums latlwmi Carangidae 120 42 58
Sallrill" caribbatll Synodontidae 44 74 26
Thyr ilops It!pie/oIJoides Gempylidae 5H 38 62
Pagrtt,r t)(l8nls Sparidae 40 50 50
PriQl1oltl.r 1Ul(ligu/a TrigUdac 60 55 45
Mer/fledu.! IlIIhb.si Mcrluccidac 135 31 69
MlIllus argt!nlinae Mullidac 74 50 50
Para/ic/',Ir.\'s isosceles BOlhidac 56 23 77
Lopllius gClstrophysus Lophiidac 39 45 55
z.enopsis conchifertl Zeidac 59 47 53
Antigon;a capros Caproidac 21 45 55

.
S:'t'IWgroIJ.'f slJinosrls Acropornalic.Jnc M 52 4K
Uro{J/'yt'is mys,m;eu Gadidae 60 36 64
Lnp/,oItj,Uu.f \'illtlr;; Branchiostcgida t 5 100
Gt"Yl1lertlS bra~ilirm';$ Ophidiidac 30 44 56
Bf'lI/brops heten,ms Pcrcophidac 24 30 70
8~1t1"(J(/es"",s tlnnl:llIus Trichiuridac 18 33 67
Ariomma bantU Ariommatidac 31 48 52
SYIF{l8rOjlS bellus Acropom:uidae 16 19 81
IJelic:nit..,rll.'i la/riller $corpacnidae 25 42 58
Argf!lIIi,w slriliUl Argcn tinidac 19 42 58
Po/yprioll americallus Polyprionidae 10 77 23
Diapl11u (1Ilm~rilii Myctophidac 15 40 60
Polymixia JO.... l', Polymixiidac 29 41 59
Ma/(,cocephalus occidentalis Macrouridae 17 47 53
CoelOriflChu marinii Macrouridae 20 SO 50

Fig. 6.14.1. Most common teleost species in bottom trawl surveys (1981-1987) from the
southern Brazilian shelf and upper slope with absolute frequency of occurrence (FO) in
315 hauls. Occurrence of species (black >50%, dark gray 25-50%, light gray <25%)
during northward (cold season) and southward (warm season) shifts of the Subtropical
Convergence (* commercially important species)
Demersal and Benthic Teleosts 131

Composition and Depth Distribution

The wide and gently sloping southern Brazilian shelf is predominantly

covered by soft-bottom substrates (Zembruski et al. 1972; Furtado 1973).
As a result, the gradual substitution of species is largely associated with
depth (Benvegnu-Le 1978; Haimovici et al. 1994a; Fig. 6.14.1). Over the
shelf down to 120 m depth, the families with the larger number of species
are Sciaenidae, Carangidae, Clupeidae, Bothidae, and Serranidae. Based on
percent catch per unit effort (CPUE), Sciaenidae (80.9%) and Trichiuridae
(14.5%) have the highest relative abundance. Four species alone
(Micropogonias furnieri, Cynoscion guatucupa, Umbrina canosai, Trichiurus
lepturus) are responsible for 71.9% of the total biomass and together with
another 11 abundant species (Macrodon ancylodon, Prionotus punctatus,
Merluccius hubbsi, Peprilus paru, Ctenosciaena gracilichirrus, Paralonchurus
brasiliensis, Cynoscion jamaicensis, Trachurus lathami, Urophycis
brasiliensis, Porichthys porosissimus, Balistes capriscus) represent as much
as 95.5% of the biomass (Haimovici et al. 1996).- The generally low
abundance of small Anguilliformes and Pleuronectiformes may be a result
of inefficient sampling (Arena et al. 1992), whilst the low abundance of
Pagrus pagrus, Pogonias cromis, and Netuma barba reflects the depletion of
their stocks between 1970 and 1990 (Haimovici et al. 1989a; IBAMA 1993).
Over the outer shelf between 120 and 180 m, Trachurus lathami, Trichiurus
lepturus, and juveniles of Cynoscion guatucupa, Umbrina canosai, and
Scomber japonicus are the dominant species and together total 83% of the
teleost biomass. Over the shelf break (180-249 m depth), Trichiurus
lepturus and Thyrsitops lepidopoides are abundant, whilst in areas with
irregular coralline substrates Antigonia capros and Priacanthus arenatus
occur, which together represent 63% of the biomass. The demersal fish
fauna of upper slope regions (250-587 m) with extensive irregular bottoms
is largely composed of Serranidae, Macrouridae, Trichiuridae, and Mycto-
phidae. The monospecific families Ariommatidae and Polyprionidae are
most abundant, with Ariomma bondi, Helicolenus dactylopterus, and
Polyprion american us making up 77% of the total biomass (Haimovici et al.
1994a).

Ecology of Principal Species

Macrodon ancylodon occurs in shallow coastal waters between Venezuela

(10 0 N) and Argentina (40 0 S). The species is most abundant over the south-
ern Brazilian shelf, and the local stock mixes little with northern popula-
tions (Yamaguti 1979). Spawning occurs in late spring and summer in
coastal areas adjacent to the Patos Lagoon mouth (3t S), followed by
migration to more northern latitudes (28 0 S) in winter (Santos and
132 M. Haimovici

Yamaguti 1965; Yamaguti 1979). Intense exploitation of Macrodon ancylodon

in southern Brazil has altered the density and consequently the growth and
age structure of the stock. Prior to 1970, first sexual maturity was reached
between 3 and 4 years and at 25 cm total length of maturity (TIm; Yamaguti
1967), with a maximum age of adults around 12 years (45 cm; Yamaguti and
Santos 1966). At the end of the decade, older age classes (>5 years) were
absent, but the length of age classes had increased (Martins Juras 1980;
Haimovici 1988a), and first sexual maturity was reached at 1-2 years (TLm
females 27.4 cm, males 21.5 cm; Juras and Yamaguti 1989). Sub adults and
adults of Macrodon ancylodon feed preferentially on shrimp (Artemesia
longinaris), cephalopod (Loligo sanpaulensis), and fish (Paralonchurus
brasiliensis, Macrodon ancylodon), but fish become more important in the
diet as size increases (Juras and Yamaguti 1985).
Micropogonias furnieri is widely distributed along the Atlantic coast of
South America down to 40° S (Isaac 1988; Vazzoler 1991). Although geneti-
cally distinct populations are absent from southeastern and southern
Brazilian waters (Maggione et al. 1994), the exchange between stocks from
southern (29-34° S), southeastern (22-29° S), and northeastern «22° S)
latitudes is limited (Vazzoler 1971; Valentini et al. 1991a; Haimovici and
Umpierre, unpubl.). However, stocks which reproduce south of this region
(34° S) migrate into southern Brazilian waters during the winter, following
the coastal branch of the MC. Maximum spawning in southern Brazilian
waters occurs in the spring and summer in coastal water under the influ-
ence of freshwater runoff from the the Patos Lagoon (Vazzoler 1971), and
the estuary of the lagoon constitutes an important nursery ground (Weiss
1981; Barbieri 1986; Castello 1986; Pereira 1994). Although some juveniles
reach first sexual maturity at 18-20 cm TLm in the estuary, most mature at
30-40 cm after leaving the estuary (Vazzoler 1971; Haimovici, unpubl.).
Subadults «30 cm TL) of Micropogonias furnieri occur throughout the
year near the coast, whilst adults disperse over the shelf, where they feed on
polychaetes, crustaceans, molluscs, ophiuroids, and fish (Tanji 1974;
Vazzoler 1975). During the fall and winter, adults concentrate in waters of
up to 100 m depth and migrate from Uruguay to southern Brazil. In con-
trast, during the summer they are seen in shallower waters «50 m;
Haimovici 1987; Haimovici et al. 1996) and migrate southward (Vazzoler
1991; Reis 1992). In southern Brazilian waters Micropogonias furnieri
attains over 70 cm TL. Age determinations, based on scales (Vazzoler 1971;
Castello 1986) and transverse cut of otholits (Schwingel and Castello 1990;
Haimovici and Umpierre 1996) show a correspondence between the
number of scale rings and opaque otholit bands for individuals younger
than 5 years (Haimovici 1977). Although the age of individuals frequently
eJl:ceeded 30 years prior (1980) to intensive exploitation (Schwingel and
Castello 1990), today individuals more than 20 years old are rarely found.
Demersal and Benthic Teleosts 133

Netuma barba is a slow-growing species with a calculated life span of

23.1 years and a mortality of 0.13, though adults may occasionally attain
36 years of age and a total length of 98 cm. Sexual maturity of females
(41.5 cm) and males (43 cm) is reached at about 7 years. At the end of the
winter Netuma barba migrates into the Patos Lagoon estuary. Gonad
maturation and reproduction, with low female fecundity (32-272 intraovaric
oocytes of 12-19 mm), take place in early spring in the estuary followed by
spawning in the coastal waters. The males incubate the eggs for up to
2 months in the buccal cavity (Reis 1986a,b). Between spawning seasons,
adults disperse over the entire shelf (Haimovici et al. 1996). The first
juveniles (6 cm TL) appear in late summer near the Patos Lagoon inlet and
also in the estuary, where they feed on pereiopods of Callinectes spp. and
polychaetes (Araujo 1984). The diet of adults is composed of molluscs and
polychaetes in the estuary, epibenthic crustaceans in adjacent coastal
waters, and principally sciaenid fish in both environments.
Although Pogonias cromis ranges from 45° N to 40° S (Fischer 1978),
information on the species from the Southwestern Atlantic is scarce. Over
the southern Brazilian shelf the species occurs at less than 40 m depth near
the inlet of the Patos Lagoon, and juveniles of 1-2 years are commonly
found in the estuary. Between the end of the spring and summer, the
gonads of females reach an advanced stage of maturity, and partial
spawning takes place. Growth appears to be rapid, since a total length of
more than 1 m is reached after 10 years, though the age of adults (TL 140 cm,
40 kg) may exceed 40 years (Haimovici et al. 1989a; Haimovici, unpubl.).
Juveniles feed on Heleobia (Litoridina), Erodona, and Balanus improvisus
in the estuary, whilst the diet of subadults and adults in coastal waters is
composed of molluscs (Mactra patagonica, Mactra isabelleana, Glycymeris
longior, Corbula sp.), gastropods (Buccinanops gradatum) , and crustaceans
(Hepatus pudibundus, Loxopagurus loxochelis, Libina spinosa).
Cynoscion guatucupa (=c. striatus) is a demersal-pelagic sciaenid
between the latitudes 2t and 40° S. The species is most abundant from 29°
to 40° S where it appears to belong to one stock (Vargas-Boldrin 1980; Vieira
1990). Adults of Cynoscion guatucupa occur at depths of less than 50 m,
especially between the spring and fall. In southern Brazilian shelf waters,
multiple spawning peaks coincide with the spring and the end of the
summer or the beginning of the fall. Principally during the summer and
fall, small juveniles ( < 10 cm) are restricted to shallow coastal waters, whilst
larger juveniles and sub adults (10-30 cm) are abundant throughout the
year at depths between 25 and 100 m. First sexual maturity of females (32-
35 cm) and males (29-31 cm) is reached at about 4 years, and the maximum
age approaches 15 and 13 years, respectively (Vieira 1990; Vieira and
Haimovici 1993). The diet preference of Cynoscion guatucupa changes with
age. Juveniles preferentially feed on planktonic crustaceans (hiperiid
amphipods, euphausids, copepods, mysidaceans), whilst adults ingest a
134 M. Haimovici

diet composed of fish (Anchoa marinii, Engraulis anchoita, Paralonchurus

brasiliensis, Macrodon ancylodon, Cynoscion guatucupa) and shrimp
(Artemesia longinaris, Pleoticus muelleri; Vazzoler 1975, Vieira 1990).
The southern Brazilian stock of the demersal sciaenid Umbrina canosai,
which represents a major part of bottom trawl catches during the winter
and spring, exhibits a clearly defined migratory cycle. Partial spawning
initiates with larger females at the end of the winter as far north as 29° S
and continues southward until the end of the spring. The large females are
also the first to migrate to southern feeding grounds in Uruguay and
Argentina, followed by males and smaller females during the summer and
fall (Gonzalez Alberdi and Nani 1967; Haimovici and Cousin 1989). During
their first year, juveniles remain over the southern Brazilian intermediate
shelf (Zaneti Prado 1979; Haimovici 1982). Males and females reach first
sexual maturity at 18.4 and 21.9 cm, respectively. At age II, 60% of the
males and 27.4% of the females are sexually mature. Adult females tend to
be larger than males and occasionally surpass an age of 20 years (Haimovici
and Reis 1984), although intense exploitation of stocks caused a significant
age-dependent average weight increase (Haimovici 1988b). Umbrina canosai
displays its highest feeding activity during the summer (Haimovici et al.
1989b) and feeds principally on benthic organisms. Smaller individuals
prefer a diet of amphipods and mysids, whilst larger individuals feed on
polychaetes, ophiuroids, bivalves, gastropods, decapods, and juvenile fish
(Vazzoler 1975; Haimovici et al. 1989b). Adaptive characteristics like high
fecundity, prolonged spawning over a large area, great longevity, age-
dependent partitioning of feeding areas, and migratory patterns appear to
explain the abundance of Umbrina canosai in southern Brazil (Haimovici
1982; Haimovici and Cousin 1989).
The cosmopolitan Trichiurus lepturus is an abundant species over the
southern Brazilian shelf (Haimovici et al. 1996), and the eggs and larvae of
this species are common (Mafalda 1989). Under the influence of tropical
waters in the summer, spawning females occupy areas near the coast, but
they are also found over the shelf break throughout the year. First sexual
maturity is reached at approximately 70 cm TLm' The diet preference of
Trichiurus lepturus changes from copepods to mysidaceans (Promysys
atlantica), sergestids (Lucifer faxoni), and euphausids (Eufausia similis)
during the early juvenile stages. Feeding diversity augments with increasing
size, including juvenile sciaenids, Engraulis anchoita, benthic macro-crus-
taceans (Artemesia longinaris, Squilla brasiliensis) and cephalopods (Loligo
sanpaulensis, Illex argentinus; Martins 1992).
Although only Prionotus punctatus is of some commercial importance,
both Prionotus punctatus and Prionotus nudigula inhabit the southern
Brazilian shelf and slope throughout the year, becoming more abundant
during the spring. Both species reproduce and spawn from the spring to the
fall. Prionotus punctatus reaches first sexual maturity at 26.2 cm, and adults
Demersal and Benthic Teleosts 135

(40 cm TL) are restricted to depths of less than 135 m between 11 and
22.5°C. Prionotus nudigula matures sexually at 16 cm, and adults occur at
depths of up to 378 m and bottom temperatures of 9.8-21.2°C. Although the
diet of both species is largely composed of crustaceans, food preferences
differ between species and change with size, depth, and season. Prionotus
punctatus displays higher feeding activity during the afternoon and in the
summer and ingests bottom dwellers of reduced motility. With increasing
size, the diet preference changes from cumaceans, amphipods, Leptochela
serratorbita, and mysids, over Portunus spinicarpus, other brachyurans,
and Pleoticus muelleri, to larger brachyurans, stomatopods, and benthic
fish (Porychthys porosissimus). Prionotus nudigula, on the other hand, also
preys on organisms in the water column and, during the summer and fall,
ingests Munida sp. and the fish Bregmaceros atlanticus at depths exceeding
100 m, but in the spring it feeds on Euphausia similis and amphipods in
shallower depths (Teixeira and Haimovici 1989).
Peprilus paru is a less abundant species which seems to follow the dis-
placement of the Subtropical Convergence. Although Peprilus paru is pres-
ent throughout the year, it is less important in bottom trawl catches in the
winter and spring, and juveniles only occur during the summer and fall.
Multiple spawning occurs from the end of the spring to early fall, and
females and males reach first sexual maturity at 120 and 130 mm fork
length, respectively. Juveniles seem to feed principally on gelatinous
organisms, like jellyfish and salps, but also on small crustaceans, poly-
chaetes, and ascidians (Cerqueira and Haimovici 1990).
The distribution of Merluccius hubbsi ranges from the upwelling area of
Cabo Frio (22° S) to the extreme south of the American continent
(Fagundes Netto and Oliveira 1991). The southern Brazilian shelf, where the
species occurs at bottom temperatures of 6.8-21 ° C between 22 and 490 m
depth, appears to serve as a growth region for locally reproducing indi-
viduals and those which reproduce further south. Multiple spawning
occurs during the winter and early spring, and first sexual maturity of both
sexes is likely to be reached at the end of the third year (30-40 cm). Juve-
niles are present throughout the year but tend to be associated with sub-
tropical water of the outer shelf and slope. In contrast, adults occur mainly
in the winter. Both juveniles and adults are most frequently found at depths
between 40 and 120 m. Engraulis anchoita is an important component of
the diet, though juveniles of Merluccius hubbsi also ingest mysids and
amphipods. Larger individuals prey on Trachurus lathami, Merluccius
hubbsi, and cephalopods (Illex argentinus, Loligo sanpaulensis; Haimovici
et al. 1993).
Pagrus pagrus is a hermaphrodite species which is widely distributed in
the Atnmtic and Mediterranean (Manooch and Hassler 1978). Based on
growth rings, two groups of this species have been identified for southern
Brazilian shelf waters. During the winter, individuals with well-defined
136 G. Phonlor and J. c. B. Cousin

rings concentrate over the intermediate shelf south at 33° 40' S, whilst a
group without obvious rings is less abundant and occurs over the entire
southern shelf during the summer. Juveniles «2 years) are rare over the
southern Brazilian shelf. The growth of males and females is similar, but
bisexual individuals tend to be smaller. Advanced stages of sexual maturity
occur from late spring to the end of summer, and adults may exceed an age
of 15 years. The diet of Pagrus pagrus is principally composed of fish
(Engraulis anchoita, Trachurus lathami, Umbrina canosai, Synagrops spinosus,
Cynoscion guatucupa) and squid (Loligo sanpaulensis), but in coastal areas
this changes to predominantly benthic invertebrates (Capftoli and
Haimovici 1993).
Despite slow growth, the serranid Polyprion american us reaches a total
length of 140 cm and up to 80 kg weight. In contrast to other serranids, the
sexes of Polyprion americanus are separate, and reproduction occurs in the
winter. Gonad maturation is common in females and males of more than
80-90 cm, and vitellogenesis of oocytes is restricted to May, July, and
September. The diet of Polyprion americanus is largely composed of
demersal fish (Helicolenus dactylopterus, Merluccius hubbsi, Urophycis
mystaceus), but the squid Illex argentinus and to a lesser extend Lycoteuthis
diadem a, Ancistrocheiros lesueuri, and the deep water crab Chaceon
notialis are also ingested (Peres and Haimovici, unpubl.).
Flatfishes of the genus Paralichthys are represented by P. orbignyanus,
P. patagonicus, and P. isosceles. The first is largely restricted to the Patos
Lagoon estuary, the second is most abundant over the shelf, and the third is
the smallest species which occurs mostly over the outer shelf (Haimovici et
al. 1996). All three species spawn during the spring and summer and are
active predators on shrimp, fish, and cephal ods (Kawakami 1975).

6.15
Early Life History of Silverside Fishes

G. Phonlor and J. C. B. Cousin

Several species of the family Atherinidae are abundant in southern Brazil-

ian coastal nearshore waters (Odontesthes incisa, Odontesthes argentinen-
sis) and the Patos Lagoon estuary (Atherinella brasiliensis, Odontesthes
sp.), as well as in coastal freshwater bodies (Odontesthes bonariensis,
Odontesthes humensis, Odontesthes retropinis; Kleerekoper 1945; Figueiredo
and Menezes 1978; Chao et al. 1982; Bemvenuti 1987, 1993; Prodohl and
Levy 1989; Phonlor and Vinagre 1990). Owing to a considerable
morphological similarity between species, the taxonomy of the family is
confused (Figueiredo and Menezes 1978). Furthermore, since marine
representatives also invade and colonize brackish and freshwater environ-
ments, different phenotypes of the same species are common (Boyd and
Early Life History of Silverside Fishes 137

Fig. 6.15.1. Eggs of marine (A) and estuarine (B) morphotypes of Odontesthes argentinensis

Simmons 1974; Prodohl and Levy 1989). Brackish and marine morphotypes
of Odontesthesargentinensis (Bemvenuti 1987, 1990, 1993) differ in egg
size. They either exhibit a smooth chorion with 3-6 filaments which sur-
rounds the micropyle (estuarine morphotype) or 17-19 looped filaments,
attached by both ends to the egg, and 3 filaments attached at one end to the
chorion (marine morphotype; Fig. 6.15.1). Although karyotype and
electrophoretic studies suggest the presence of one species, differences in
chorion structure and filament number in other species (Atherinops affinis;
White et al. 1983) have been interpreted as either species-specific (Ivankov
and Kurdyayena 1973; Lonning 1981) or as a result of abiotic gradients
(Fundulus heteroclitus; Brummet and Dumont 1981).

Spawning, Fertilization and Hatching in the Genus Odontesthes

The spawning periods of different Odontesthes species vary between

habitat&. In coastal freshwater lagoons, O. humensis and O. bonariensis
spawn between the end of fall and the end of winter, though spawning
periods tend to overlap (Phonlor and Vinagre 1990). In brackish Patos
Lagoon waters Odontesthes sp. spawns from late fall to early summer,
138 G. Phonlor and J. C. B. Cousin

whilst spawning of O. argentinensis in coastal waters lasts from the end of

winter to early summer (Bemvenuti 1987; Phonlor and Vinagre 1989, 1990;
Phonlor and Sampaio 1992). The oxygen consumption of spawned
O. argentinensis eggs is low, probably because of their slow embryonic
development and/or their occurrence in bottom layers with reduced
dissolved oxygen concentrations.
The fertilization success of Odontesthes eggs depends on the salinity tol-
erance of the gametes. Below a salinity of 2.5, fertilization of Odontesthes
argentinensis and Odontesthes sp. eggs is absent; it increases to 12 and 39%
at salinity 5, respectively. Fertilization success exceeds 85% in both species
above a salinity of 5, and the highest sperm motility occurs at salinities of
20-30. Fertilization success is therefore likely to be caused by different
salinity optima of gametes.
Environmental gradients also influence the hatching of eggs (Phonlor
and Vinagre 1989). Similar to fertilization, Odontesthes argentinensis does
not hatch in freshwater, though hatching rates gradually increase to 100%
in salinities between.10 and 30 (Sampaio and Phonlor 1992). The period of
larval survival (6 days) is much lower in freshwater, but virtually all larvae
survive in salinities above 5 (Phonlor and Sampaio 1992). In contrast,
hatching rates of freshwater representatives (0. humensis, O. bonariensis)
increase from 40 to 100% between salinities of 0 and 10, but decrease to
38% at higher salinities (20). In addition, temperature shocks (3°C) increase
hatching of Odontesthes bonariensis significantly. The hatching time of
Odontesthes argentinensis eggs is reduced by low temperature (15°C) and
light intensities. Furthermore, Odontesthes argentinensis eggs tend to be
exposed to prolonged desiccation after their deposition on the beaches
during exceptionally high tides (Phonlor and Vinagre 1989). Under
experimental conditions, desiccation shortens the hatching period, and all
embryos survived 96-120 h of desiccation. However, embryos in early
stages of development (16 cells) are more susceptible than those just prior
to hatching. Since the mortality of hatched larvae is insignificant and their
locomotion and feeding behaviour remain unchanged, survival chances
after desiccation under natural conditions appear to be high. Physiological
adaptations of Odontesthes species of marine origin, principally to different
salinity regimes in brackish and freshwaters (Phonlor and Sampaio 1992),
thus confirm the genetic plasticity of Atherinidae (Bamber and Henderson
1988).

Larval Development and Growth of Odontesthes argentinensis

The euryhaline characteristics of Odontesthes argentinensis not only extend

the distributional limits of the species, but also distinguish it as an excellent
organism for bioassay studies and, together with high larval survival, make it a
Early Life History of Silverside Fishes 139

potential candidate for large-scale culture. Under controlled culture conditions

(20De), larval feeding commences on the first day. Weight increases and aerobic
metabolism of hatched larvae (6.94 mm, 0.6 mg dry weight) are a function of
time. The weight specific growth rate decreases from 0.14 to 0.07 mm dail after
4 weeks (Phonlor and Sampaio 1996). A decrease in food intake per unit larval
weight after 14 days is possibly compensated for by an increased assimilation
efficiency, accompanied by higher oxygen consumption and growth rates.
Organogenesis and processes of cellular differentiation accelerate after
hatching. As the sole energy source, the vitellus (Fig. 6.1S.2a) assures the
formation and function of transient larval organs. Major changes of the
digestive system include the expansion of the intestinal tract and division into
anterior, middle, and posterior parts (Fig. 6. 1S.2b,c), the formation of a
sphincter that separates the oesophagus from the anterior intestine, and the
complete stratification of the oesophagus and pharynx epithelium, with an
increase in number of teeth, taste buds, and goblet cells (Fig. 6.1S.2d). Owing to
the formation of new hepatocytes and capillarization, the Jtver volume increases
rapidly and substitutes for the vitellus (Fig. 6.1S.2c). In the p-ancreas, groups of
exocrine cells assume an acini form, and the basophil cells become replete with
zymogen granules. The gas gland and the rete mirabile differentiate into the
swim bladder, which inflates and remains connected to the digestive tract by a
short pneumatic duct. Since newly hatched larvae increase in body weight and
notocord length during exposure to thyroxine, it is likely that after the
development of the thyroid gland, increased thyroxine secretion influences
tissue differentiation and provokes adaptations to higher salinities (Santos and
Santos 1989).
During a period of starvation, larvae consume most of the vitellus in the first
4.5 days, with some remaining until the 8th day (Fig. 6.1S.2e). At that age, low
mortality (10%) emphasizes the resistance of larvae to starvation, though
cellular atrophy leads to volume reduction and structural changes of tissues and
organs. The digestive system in particular undergoes significant modifications.
Intestinal folds begin to disappear, the entire intestinal wall becomes thinner
(Fig. 6.1S.2f) and cellular basophilia increases, owing to cytoplasmic density.
After 10 days of starvation, the space between the organs in the abdominal
cavity grows, the gall bladder expands, the pancreatic acini begin to
disintegrate, and localized necrosis of cells commences. In spite of these tissue
alterations, surviving larvae (3%) resume growth upon being supplied with
food, which retards the "point of no return" (Blaxter and Hempel 1963) and
establishes a somewhat flexible "critical period" (Fabre-Domergue and Bietrix
1897) for this species. Starvation beyond 10 days causes general cellular
atrophy, peeling of the intestinal epithelium, a virtual absence of zymogen
granules in the pancreas, and increasing necrosis of the liver. Similar
lesions 'have been reported for other species of teleosts after starvation
(Ehrlich et al. 1976; Theilacker 1978; Watanabe 1985; Cousin et al. 1986).
Odontesthes argentinensis larvae do not survive 12.5 days of starvation.
 MI NS

c
- d
-

e
-
Fig. 6.15.2. Histological sections of starved Odontesthes argentinensis larvae at hatching (a)
and after 6 (b), 4 (c,d), 8 (e), and 12 days (f). (AE Atrophied epithelium; AI anterior
intestine; BC buccal cavity; BD bile duct; BV blood vessel; CT connective tissue; E
esophagus; EP epidermis; FT faringeal teeth; GB gas bladder; GA gill arches; GC goblet cells;
H heart; L liver; MI middle intestine; N notochord; NH necrotized hepatocytes; NS nervous
system; PF pectoral fins; P pigment; PI posterior intestine; S spleen; SE stratified
epithelium; SM striated skeletal muscle; TB taste buds; Vvitellus). Bars equal 400 11m (a,b)
and 100 11m (c,d,e,f)
Demersal Elasmobranchs 141

In culture, the survival and growth of Odontesthes argentinensis larvae

are largely a function of their density and of the available food's nutritional
quality. A diet composed of Artemia nauplii (20 org. mr') satisfies nutri-
tional requirements and permits optimal growth (0.11 mm dai') of larvae
(IS org. 1"') up to the 2nd month of life (28.6 mm TL) with less than 5%
mortality. Proportional increases of larval and Artemia densities, or
Artemia supplied ad libitum, alone or in combination with rotifera, do not
improve the growth rates further. Although artificial diets potentially
reduce costs and may simplify fish culture procedures (Tandler 1985) and
larvae of Odontesthes argentinensis larvae ingest this diet immediately after
hatching, their limited nutritional value only leads to suboptimal growth.

6.16
Demersal Elasmobranchs

C.M. Vooren

In terms of abundance and species diversity, elasmobranchs are important

components of the ichthyofauna of the southern Brazilian shelf and upper
slope. Bottom trawl surveys between 1968 and 1972 (Sadowsky 1973;
Vazzoler et al. 1973; Figueiredo 1977) and from 1980 to 1986 (Peres and
Vooren 1991; Vooren 1992) provided mostly information on demersal
species, but occasional catches of pelagic sharks also yielded data on the
genera Carcharhinus, Sphyrna, Alopias, Squaliolus, and Etmopterus. Sharks
of the pelagic environment are frequently caught by the commercial tuna
longline fishery on the continental slope, but their biology and ecology
have not been studied.

Species Composition and Abundance

Fifty-seven species have been recorded over the shelf and upper slope, at
depths from 10 to 500 m between 28° 40' and 34° 34' S (Cape Santa Marta
Grande to Chul; Appendix). The sharks are represented by 32 species
(3 Squatiniformes, 2 Lamniformes, 8 Squaliformes, 16 Carcharhiniformes,
3 Hexanchiformes) and 25 species belong to the rays and skates (l4 Rajiformes,
7 Myliobatiformes, 4 Torpediniformes), though several identifications are
uncertain at the species level. Sadowsky (l973) and Figueiredo (l977) cite
Myliobatis goodei Garman, 1885 in the area, but in later studies of material
which fitted the description of this species, two morphotypes were found,
with different morphometric dimensions of the head and the tooth plates,
which were classified as Myliobatis NT (narrow teeth) and Myliobatis BT
(broad teeth), respectively. Levy and Concei~ao (1989) showed by
biochemical methods that the morpho types represent two separate species,
142 C.M. Vooren

but their identification was not possible because available taxonomic

descriptions are lacking in relevant detail. Of the genus Squatina,
Figueiredo (1977) and Compagno (1984) cite only S. argentina in the area,
but Sole-Cava et al. (1983) and Sole-Cava and Levy (1987) found three
morphotypes which were characterized as separate species through
frequency of isoenzymes. Vooren and Silva (1991) identified these as
S. guggenheim, S. argentina, and the new species S. occulta. Of the genus
Squalus, Calderon (1994) identified S. megalops as common in the area,
while S. mitsukurii and S. acanthias occurred sporadically. The occurrence
of S. cubensis in the area, cited by Sadowsky (1973), Figueiredo (1977), and
Compagno (1984), was not confirmed in later studies.
In the years 1981 to 1983, the abundance of demersal elasmobranchs as a
group on the continental shelf at depths between 10 and 100 m varied
seasonally from 43 000 tons in the summer to 96 000 tons in the late winter
and spring. Twenty-five elasmobranch species account for 94% of the total
biomass. Angel sharks, rays, and skates consJituted about 90% of th,e
demersal elasmobranch biomass. Squatina guggenheim, Squatina occulta,
Sympterigia acuta, Sympterigia bonapartei, Myliobatis NT, Myliobatis BT,
and Raja castelnaui together made up about 80% of the biomass of bottom-
dwelling elasmobranchs, while Galeorhinus galeus and Mustelus schmitti
accounted for 75% of the biomass of free-swimming demersal elasmo-
branchs in the winter. The decrease of the catch of elasmobranchs per unit
effort in the commercial otter-trawl fishery is evidence that since 1985 the
abundance of many species has decreased drastically. Indeed, the popula-
tion of the guitar fish Rhinobatos horkelii is in danger of extinction
(IBAMA 1993; Vooren, unpubl.).

Depth Distribution and Migrations

According to their temporal distribution, species are classified as perma-

nent residents, winter or summer migrants, and sporadically occurring
species. Psammobatis rutrum, Psammobatis glansdissimilis, and Psammobatis
bergi are not included in this classification because they have only been
identified in preserved material and not routinely in the field.
Among the 18 permanent residents, 12 are dominant species which are
seen throughout the year without predictable changes in abundance, and
complete their entire life-cycle in the waters off southern Brazil: Squatina
guggenheim, Mustelus jasciatus, Rhinobatos horkelii, Zapteryx brevirostris,
Raja castelnaui, Raja agassizi, Raja platana, Raja cyclophora, Sympterigia
a,cuta, Myliobatis NT, Myliobatis BT, and Gymnura aitavela. Four species
occur constantly but in small numbers: Carcharhinus plumbeus, Carcharhinus
obscurus, Carcharhinus signatus, and Rhizoprionodon lalandei. Squatina
argentina and Squatina occulta are common all year round, but gravid
Demersal Elasmobranchs 143

females and neonates are rarely found, so gestation and birth cannot take
place in that area. The constant presence of Myliobatis NT and Myliobatis BT
with a peak abundance in the spring and fall is a result of complex migrations
and the passage of different populations through the area. Sympterigia acuta
and the four dominant species of the genus Raja are sedentary, each species
occupying a distinct depth range (Fig. 6.16.1). The gravid females of the
species Squatina guggenheim, Rhinobatos horkelii and Mustelus fasciatus live
at depths greater than 20 m for most of the year but migrate during the
spring and summer to shallow coastal waters to give birth, and the juveniles
remain there during their first year of life. Intense commercial fishing in
these waters interferes with reproduction and recruitment and therefore
represents a threat to the survival of these species (Queiroz 1986, 1995; Lessa
et al. 1986; Vooren et al. 1988; Vooren and Silva 1991; IBAMA 1993).
The 12 winter migrants are the skate Sympterigia bonaparte, the ray
Discopyge tschudii, and the sharks Galeorhinus galeus, Mustelus schmitti,
Mustelus canis, Sphyrna zygaena, Eugomphodus taurus" Squalus megalops,
Squalus mitsukurii, Squalus acanthias, Notorhynchus·· cepedianus, and
Heptranchias perlo. These species reproduce in Uruguayan and Argentine
waters during the summer, and migrate in the winter to the continental shelf
of southern Brazil. All sharks of this group represent important regional
fishery resources during the winter (Barcellos 1957; Souto 1986; Batista 1988;
Vooren et al. 1988, 1990; Ferreira and Vooren 1991; Peres and Vooren 1991;
IBAMA 1993; Calderon 1994; Queiroz 1995).
The six summer migrants are the shark Sphyrna lewini and the rays
Mylio ba tis freminvillei, Dasyatis say, Dasyatis centroura, Rhinoptera bonasus,
and Narcine brasiliensis. These warm water species migrate with tropical
water masses during the summer to the shelf of southern Brazil where
Sphyrna lewini, Dasyatis say, and Myliobatis freminvillei give birth in
nearshore waters. With the exception of Narcine brasiliensis and Rhinoptera
bonasus, all summer migrants represent commercial fishery resources. Since
large juveniles of Carcharhinus plumbeus are fished with beach seines during
the summer in southern Brazil (Vooren, unpubl.) and Uruguay (De Buen
1952), it is likely that this species is a summer migrant to inshore waters of
less than 10 m depth.
Eighteen species occur sporadically: Galeocerdo cuvier, Carcharhinus
brevipinna, Carcharhinus acronotus, Carcharhinus brachyurus, Scyliorhinus
besnardi, Schroederichthys bivius, Alopids vulpinus, Squaliolus laticaudus,
Etmopterus hillianus, Etmopterus pusillus, Centroscymnus cryptacanthus,
Echinorhinus brucus, Hexanchus griseus, Raja leptocauda, Psammobatis
len tigin osa, Gurgesiella dorsalifera, Torpedo puelcha, and Benthobatis sp.
(Barcellos 1963; Barcellos and Pinedo 1980; Vooren, unpubl.). Most species
occur in small numbers at depths from 100 to 500 m. However, the abun-
dance of pelagic sharks such as members of the genera Etmopterus and
Squaliolus may be greater than suggested by from bottom-trawling.
 DEPTH (m)

50 100 150 200 250 300 350 400 450

...........
RHINOPTERA BONASUS
SYMPTERIGIA ACUTA
SYMPTERIGIA BONAPARTE I
ZOPTERYX BREVIROSTRIS
RAJA AGASSIZI
SQUATINA GUGGENHEIM
--
MYLIOBATIS BT
SPHYRNA LEWINI
S~RNA 2YG.AENA
RHINOBATOS HORKELII
RAJA PLATANA
MUSTELUS CANIS
MUSTELUS FASCIATUS
MUSTELUS SCHMITTI
DASYATIS SAY
MYLiOBATIS NT
MYLIOBATIS FREMINVILLEI
GYMNURA AL TAVELA
EUGOMPHODUS TAURUS
RAJA CASTELNAUI
DASYATIS CENTROURA
RAJA CYCLOPHORA
GALEORHINUS GALEUS
SQUATINA OCCUL TA
SQUALUS MEGALOPS
SQUATINA ARGENTINA

o
~
<:
o
Fig. 6.16.1. Depth ranges of dominant demersal elasmobranch species over the continental shelf and upper slope between 28' 40' o
and 34' 34' S ;;l
::l
Demersal Elasmobranchs 145

Reproductive Strategies and Evolution

The elasmobranch fauna of southern Brazil is composed of 13 oviparous

species (Rajidae and Scyliorhinus besnardi) and 44 viviparous forms (all
others). All types of trophic relation between mother and embryo as classi-
fied by Wourms et al. (1988) are observed among the viviparous species,
and these fishes display different reproductive strategies in terms of sexual
development, reproductive cycle, body size at birth, and fecundity. Among
the 10 species that have been studied in detail, lecithotrophic gestation
occurs in Rhinobatos horkelii, Squalus megalops, Squalus mitsukurii,
Squatina guggenheim, Squatina occulta, Squatina argentina, and Galeorhinus
galeus, while Mustelus schmitti, Mustelus canis, and Mustelus Jasciatus have
matrotrophic gestation (Vooren 1992). In most species the reproductive
cycle of the female is annual, with vitellogenesis and gestation occurring
simultaneously. However, the female reproductive cycle of Squatina
guggenheim, Squatina occulta, and Galeorhinus galeus lasts 3 years: 1 year
for gestation, and 2 years for gametogenesis and yolk formation (Peres and
Vooren 1991; Vooren 1992; Sunye 1993; Silva 1996). In matrotrophic
species, the annual production of litter mass as a proportion of maternal
body mass ranges from 11 % in Mustelus Jasciatus to 26% in Mustelus canis.
In lecithotrophic species the values range from 5 to 7% in Rhinobatos
horkelii, Squatina guggenheim, Squatina occulta, and Galeorhinus galeus,
but reach 19% in Squalus megalops and Squalus mitsukurii. If the
production of yolk is included, annual reproductive investment as a
proportion of maternal body mass reaches 32% in Squalus megalops and
Squalus mitsukurii (Vooren 1992; Calderon 1994). Thus, there are
considerable differences between species in the relative magnitude of the
annual investment of metabolic energy in reproduction by the female. The
partition of metabolic investment between reproduction, growth, and
maintenance is genetically determined and results in the mutual adjust-
ment between recruitment and mortality at the population level. Future
research on population energetics and mortality will elucidate the mecha-
nisms and constraints which determine the process.
Rhinobatidae, Squatinidae, and Hexanchidae have existed since the Juras-
sic, Sphyrnidae appeared during the Miocene, while other families such as
Rajidae, Myliobatidae and Carcharhinidae date back to the Cretaceous
(Cappetta 1987). Thus, the elasmobranch fauna of southern Brazil contains
elements of widely different degrees of antiquity. This circumstance
favours the comparative investigation of the biology and adaptation of
elasmobranchs at different evolutionary stages, and permits study of the
evolution of viviparity among the group. For instance, the cloacal gestation
of the genus Squatina is unique among elasmobranchs and may represent
an early stage of the evolution of viviparity, though uncertainty persists as
to whether this mode of reproduction evolved from or gave rise to oviparity
146 M. Haimovici

(Sunye 1993). The elasmobranch fauna of southern Brazil is a "laboratory

of evolution", and there is an urgent need to preserve this fauna, which is
being severely threatened by the fishery industry.

6.17
Cephalopods

M. Haimovici

Early reviews on the taxonomy and zoogeography of Brazilian cephalopods

are based on museum collections of expeditions during the 19th and the
beginning of the 20th century as collated by Palacio (1977). More recent
studies have re-examined the octopus Eledone massyae (Voss 1964) and the
abundant Loligo sanpaulensis (=L. brasiliensis; Brackoniecki 1984), as well as
described a new genus and one member (Vosseledone charrua; Palacio 1978)
and a new species of Eledone (E. gaucha; Haimovici 1988c). Based on the
revision of cephalopod collections (Haimovidet al. 1989c; Perez arid
Haimovici 1991a; Haimovici and Perez 1991a,b), the study of cephalopod
beaks in the stomachs of predators (Santos 1992), as well as macro-
zooplankton samples of paralarvae and juveniles (Haimovici and Perez
1991a; Haimovici et al. 1994b), the knowledge concerning southern Brazilian
cephalopods has improved (Palacio 1977; Juanic6 1980; Roper et al. 1984;
Haimovici 1985; Haimovici and Perez 1991b; Haimovici et al. 1994b).

Faunistic Composition and Ecology of Dominant Species

For much of the cephalopod fauna, the Brazilian coast between 20° and 34° S
represents a biogeographic transition zone. This region appears to be the
centre of temperate coastal species with a relatively restricted distribution,
like the frequent Loligo sanpaulensis, Eledone massyae, Eledone gaucha,
and Octopus tehuelchus. Temperate species with a wide distribution
(tropical species Loligo plei and Semirossia tenera, and the cosmopolitan
Octopus vulgaris) are rare (Palacio 1982). Compared with more northern
regions, the southern Brazilian shelf and slope exhibit an apparently
impoverished cephalopod fauna (Haimovici and Andriguetto 1986;
Haimovici et al. 1989c; Haimovici and Perez 1991a). The low diversity of
benthic shelf habitats and pronounced seasonal water temperature oscilla-
tions, owing to largely sandy bottoms and to the influence of the Brazil and
Malvinas Currents, respectively, do not favour the occurrence of tropical
neritic species like Loligo plei, Pickfordiateuthis, and Lolliguncula brevis,
nor of cold temperate coastal species like Loligo gahi.
Cephalopods 147

Over 40 cephalopod species have been recorded for the southern

Brazilian shelf and slope between Santa Marta Grande Cape and Chui (28°
30'-34° 40' S; Appendix; Haimovici and Andriguetto 1986; Haimovici and
Perez 1991b; Perez and Haimovici 1993; Fig. 6.17.1). Octopus tehuelchus is
the most abundant benthic species in shelf waters down to 50 m depth,
whilst Semirossia tenera and Octopus vulgaris occur in deeper waters.
Between 50 m depth and the shelfbreak (200 m), the Octopodidae Eledone
massyae and Eledone gaucha are the most frequent and dominant benthic
species (Haimovici and Andriguetto 1986; Haimovici and Perez 1991a). The
life-yc1e of Eledone massyae extends approximately 2 years. Males reach
sexual maturity prior to females in the fall, followed by reproduction in the
spring. Intraovaric fecundity with 27-126 ovocytes in Eledone massyae
(Perez and Haimovici 1991b) and 5-58 in Eledone gaucha (Perez et aI.,
unpubI.) is low. In contrast to other Octopodidae, the oviductal gland in
females of both species lacks the function of a spermatheca since sperms
are stocked in filaments of the intraovaric maturing oocytes (Perez et aI.
1990). Adults of Eledone massyae and Eledone gaucha appear to explore
different food sources. While the diet preference of Eledone massyae
changes with age from amphipods to portunid crabs and polychaetes,
Eledone gaucha continues to feed on amphipods and isopods (Perez and
Haimovici 1995). In the deeper slope regions species of subantarctic origin
(Voss 1988), like Pareledone charrua, Pareledone turqueti, and eledonids
occur on sandy bottoms, in contrast to Vosseledone charrua, which is
probably restricted to consolidated bottoms of relict corals.
Loligo sanpaulensis is the most abundant neritic species in shelf waters
during the entire year (Haimovici and Andriguetto 1986; Andriguetto and
Haimovici 1991). The highest densities and largest individuals occur south
of 32° S in waters between 40 and 80 m depth at less than 18°C. Since the
spring biomass may reach 3500 tons, Loligo sanpaulensis stocks are of
potential commercial value. The southern Brazilian shelf waters appear to
be a spawning area of the species, with smaller and larger individuals
spawning in shallower «40 m) and deeper waters, respectively. Smaller
individuals are more common in the summer and fall, whilst mature
females are found throughout the entire year but are less frequent in the
fall (Andriguetto and Haimovici, in press). The availability of food, largely
composed of fish and crustaceans, is likely to influence the distribution
patterns of Loligo sanpaulensis (Juanic6 1979; Costa and Fernandes 1993).
Other pelagic species of the shelf water, especially during the summer, are
Argonauta nodosa and Loligo plei.
Illex argentinus is the most frequent and abundant species over the
southerp Brazilian outer shelf and upper slope (Haimovici and Perez
1991b). The species has a life-cycle of 1 year (Arkhipkin 1990) and repro-
duces essentially during the winter and spring when spawning adults are
commonly found along the shelf break. Based on their size and the absence
 Depth range (m) :;
00
0\0\0\0\00
0\ ..... .... 0
.... ....
0\ 0\ 0\ N «'\ ....-00
C\ ('I""j Lr) f". - I , I I I 0

Species and genera Families I OOOO~~~~~~ Habitat

ON~\oOO N",,-.::rA
Order SEPIOIDEA
Spirula spirula Spirulidae fSr[)}Wi Mesopelagic
Semirossia tenera Sepiolidae d= ¢ @¥i£i! Demersal,muddy and sandy bottoms
Heteroteuthis atlantis Sepiolidae [[IKII Mesopelagic
Order TEUTHOIDEA
Loligo sanpaulensis Lolliginidae Demersal-pelagic
Loligo plei Lolliginidae Demersal-pelagic in the warm season
Illex argentinus Ommastrephidae Demersal-pelagic
Ornithoteuthis volatilis Ommastrephidae Epipelagic
Ornithoteuthis antillarum Ommastrephidae Epipelagic
Ommastrephes bartramii Ommastrephidae Epipelagic
Todarodes angolensis Ommastrephidae Demersal-pelagic
Thysanoteuthis rhombus Thysanoteuthidae Epipelagic
Onychoteuthis banksi Onychoteuthidae Epipelagic
Moroteuthis ingens Onychoteuthidae Demersal-pelagic
Moroteuthis robsoni Onychoteuthidae Demersal-pelagic
Onychia carribaea Onychoteuthidae Mesopelagic
A braliops is pfefferi Enoploteuthidae Mesopelagic-bathypelagic
Abralia veranyi Enoploteuthidae Mesopelagic-bathypelagic
Abralia redfieldi Enoploteuthidae Mesopelagic-bathypelagic
Ancistrocheirus alessandrini Enoploteuthidae Mesopelagic-bathypelagic
Pterygioteuthis giardi Enoploteuthidae Mesopelagic-bathypelagic
Histioteuthis dofleini Histioteuthidae Mesopelagic
H istioteuthis atlantica Histioteuthidae Mesopelagic
~
::r:
~.
3o
:S.
o.
 0\0\0\.0\0 Q
O\f"-..-.:t'~~O
O\C\O'\_ ...... Nt't'\-.:!''OO 'g.
O'\('f"')lf)r--. - t tl I ' 0
~oooo~~g~~\D e:-
o
Species l\lld genera Families ON-.:I"\Ooo NC'f""l.-..:t'A Habitat '0
o
0.-
Liocranchia reinhardti Cranchiidae Mesopelagic-bathypelagic '"
Megalocranchia sp Cranchiidae mJ Mesopelagic-bathypelagic
Chiroteuthis veranyi Chiroteuthidae Mesopelagic-bathypelagic
Lycoteuth is diadema Lycoteuthidae Bathyal, mesopelagic
Order OCTOPODA
Octopus tehuelchus Octopodidae }l~ ~~f..w..t: Benthic, muddy and rocky bottoms
Octopus vulgaris Octopodidae Benthic, rocks and coral reefs
Scaeurgus unicirrhus Octopodidae Benthic,sand and coraline bottoms
Pteroctopus tetracirrhus Octopodidae Benthic, muddy bottoms
Eledone massyae Octopodidae Benthic, sandy and muddy bottoms
........;.:(
"i:-.
Eledone gaucha Octopodidae IIRr:" ~~,:~, . Benthic, sandy and muddy bottoms
Pareledone turqueti Octopodidae Benthic, mud
Pareledone charcoti Octopodidae Benthic, sandy bottoms
Vosseledone charrua Octopodidae Benthic, on rocky and coraline bottoms
Eledonella pigmea Bolitaenidae Bathypelagic
Japetella diaphana Bolitaenidae Bathypelagic
Allopsus mollis Allopsidae Epipelagic - bathypelagic
Argonauta nodosa Argonautidae Epipelagic in warm waters
Tremoctopus violaceous Tremoctopodidae Epipelagic
Ocythoe tuberculata Ocythoidae Epipelagic
Thysanoteuthis rhombus Thysanoteuthidae Epipelagic

Fig. 6.17.1. Depth ranges and habitats of neritic and upper slope cephalopods along the southern Brazil coast (black abundant; dark gray
frequent; light grey infrequent)

......
~
150 M. C. Pinedo

of didymozoid parasites in their digestive tract, spawning adults from

southern Brazil, Uruguay, and northern Argentine may belong to the same
group (Santos and Haimovici, in press). Paralarvae and juveniles of less
than 50 mm mantle length exhibit three growth phases. They concentrate
principally during the winter and spring in shelf waters (> 100 m, >1tc) of
tropical origin and at the convergence with the Malvinas Current
(Haimovici and Perez 1990; Vidal 1994a,b; Haimovici et al. 1995; Vidal and
Haimovici, unpubl.). The diet preference of Illex argentinus changes during
development. Recently hatched paralarvae appear to feed on micro-
organisms, whilst crustaceans are the preferred diet of older paralarvae.
During continued growth, the relative importance of crust~ceans changes
in favour of a diet of fish and other cephalopods, which includes cannibal-
ism (Santos 1992). Illex argentinus represents an important food source for
the tunafish Thunnus obesus, the swordfish Xiphias gladius, and the wreck-
fish Polyprion americanus (Santos 1992; Santos and Haimovici, unpubl.).
Although Illex argentinus is of considerable economic importance world-
wide (Csirke 1987; Haimovici et al., in press), the potential value of the
southern Brazilian stocks still needs to be evaluated (Haimovici and Perez
1991a).Other typical slope species include Abralia verany, Abralia redfieldi,
Lychoteuthis diadem a, and Ancistrocheirus lesueuri (syn. A. alessandrini)
which are abundant during the winter (Haimovici and Perez 1990), as well
as oceanic species with wide distribution in temperate and tropical waters,
like Ornithoteuthis antillarum and Ommastrephes bartrami (Santos 1992;
Santos and Haimovici, unpubl.).

6.18
Marine Mammals and Turtles

M. C. Pinedo

Species Composition and Distribution

Of the approximately 78 known worldwide cetaceans and the 13 endemic

pinnipeds of South America and the Antarctic, 27 and 7, respectively, occur
along the southern Brazilian coast between the latitudes 25° 14' and 33° 45' S
(Castello and Pinedo 1977b; Pinedo 1990; Pinedo et al. 1992; Appendix). The
pinnipeds, some of which (Arctocephalus gazella, Hydruga leptonyx, Lobodon
carcinophagus) have a circumpolar distribution, follow the Malvinas Current
to reach the Brazilian coast (Castello and Pinedo 1977a). The cetaceans
Hyperoodon planifrons, Phocoena spinipinnis, Sotalia jluviatilis, Australo-
phocaena dioptrica, and Pontoporia blainvillei are restricted to the Southern
Hemisphere, the last being endemic to the Southwestern Atlantic. Five
cetaceans (Mesoplodon densirostris, Stenella coeruleoalba, Stenella attenuata,
Steno bredanensis, Kogia simus) and two pinnipeds (Arctocephalus tropicalis,
Marine Mammals and Turtles 151

Arctocephalus gazella) have been cited for the first time in the southwestern
Atlantic (Castello and Pinedo 1977b; Pinedo and Castello 1980; Pinedo 1987;
Pinedo et al. 1992), whilst five others (Megaptera novaeangliae, Phocoena
spinipinnis, Pseudorca crassidens, Kogia breviceps, Australophocaena
dioptrica) have been registered for the first time in the southern Brazilian
region (25° 14'-33° 45' S; Pinedo 1985, 1989; Rosas and Pinedo 1989). Among
the 27 cetaceans, Balaenoptera musculus is an endangered species, and
Megaptera novaeangliae and Eubalaena australis are classified as vulnerable
because overexploitation and degradation of habitats has drastically reduced
the original stocks. The remaining 24 cetacean species are categorized as
"insufficiently known" (IUCN 1991), which impedes their characterization as
endangered or vulnerable. Populations of nine of these species run a serious
risk of being accidentally caught in fishing nets (Perrin 1988). Apart from the
rich marine mammal fauna, four (Caretta caretta, Chelonia mydas,
Eretmochelys imbricata, Dermochelys coriacea) of the eight known marine
turtle species have extended their distribution into southern Brazilian waters
(de Menezes 1972; Frazier 1984; Marquez 1990).

Ecology of Principal Cetaceans

Most of the cetaceans over the southern Brazilian shelf are coastal species,
though 10 (Delphinus delphis, Balaenoptera musculus, Physeter macrocephalus,
Hyperoodon planifrons, Mesoplodon densirostris, Ziphius cavirostris,
Grampus griseus, Globicephala melas, Stenella clymene, Stenella coeruleoalba)
are oceanic and another four (Balaenoptera acutorostrata, Megaptera
novaeangliae, Stenella frontalis, Stenella longirostris) commonly occur in
both environments. The more frequently encountered species are
Pontoporia blainvillei, Tursiops truncatus, Eubalaena australis, and
Delphinus delphis (Pinedo 1985, 1986; Castello and Pinedo 1986).
Pontoporia blainvillei occurs in waters of up to 30 m depth near the
coast «50 km; Praderi et al. 1989) between 18° and 39° S (Perez-Macri and
Crespo 1989; Pinedo 1994a). Male and female adults reach a total length of
158 cm and 177 cm, respectively (Brownell 1989), though geographically
distinct forms of smaller and larger sized adult individuals are seen north
and south of this region at 22-2t Sand 32-38° S, respectively (Pinedo 1991,
1995). First sexual maturation occurs between 2 and 3 years (Kasuya and
Brownell 1979), followed by reproduction in the late spring. Gestation
generally exceeds a period of 9 months, and newly born individuals
measure about 75 cm (Pinedo et al. 1989). Both sexes display early devel-
opment of fins, respiratory system, and sensory organs. The life span of
Pontoporia blainvillei is relatively short since physical maturity occurs
between 4 and 8 years (Pinedo 1991), and the longevity of males and
females is about 16 {Brownell 1989) and 21 years (Pinedo 1994b), respectively.
152 M. C. Pinedo

Pontoporia blainvillei is often caught accidentally in beach seines along

the southeastern and southern Brazilian coast, which represents the princi-
pal cause of mortality (Perez-Macri and Crespo 1989; Pinedo et al. 1989;
Praderi et al. 1989; Pinedo 1994a; Corcuera et al. 1994). It is the most
common species interacting with fisheries between approximately 26° and
34° S (Pinedo 1994a). Since most of the 1085 dead specimens registered
over a 10-year period along the southern coast (29° 20'-33° 45' S) were still
sexually immature «3 years), their mortality is likely to have had an
impact on the age structure of the population (Pinedo 1994a,b). Pontoporia
blainvillei preys on a large variety of benthic organisms and fish. The fish
diet is largely composed of Sciaenidae, particularly juveniles «50 mm) of
Cynoscion guatucupa (Brownell 1975; Pinedo et al. 1989). The cephalopod
Loligo sanpaulensis is ingested especially by females, and young individuals
also feed on the crustaceans Pleoticus muelleri, Artemesia longinaris, and
Penaeus paulensis (Pinedo 1982).
Tursiops truncatus (Castello and Pinedo 1977a; Pinedo 1982, 1986;
Pinedo et al. 1992) is common in coastal shelf w~ters and adults and young
individuals are seen throughout the year in the Patos Lagoon estuary
(Castello and Pinedo 1977a). Natural death appears to be the principal
cause of mortality, followed by accidental capture in fishing nets (Pinedo
1986). Twenty of the 76 strandings (29 0 20' and 33° 45' S) between 1976 and
1993 were attributed to fishery interactions, five being caught two miles
offshore in gill-nets (22 em mesh) set for sharks (Pinedo 1986, 1994a).
Along this coast Tursiops truncatus does not ingest cephalopods or shrimp
but essentially preys on fish (> 150 mm TL). Micropogonias furnieri domi-
nates among the 10 teleost species which compose the diet (79%; Pinedo
1982).
Other species, like Globicephala melas, Steno bredanensis, Sotalia jluviatilis,
Stenella frontalis, and Pseudorca crassidens, are caught by long lines or nets.
The coastal form of Sotalia jluviatilis is common between the equator and 29°
S and is the principal species interacting with fisheries at about 25 0 S.
Pontoporia blainvillei is the most common species along the coast between 26
and 340 S (Pinedo 1994a). Pelagic and oceanic species, like Physeter
macrocephalus, Kogia simus, Globicephala melas, Phocoena spinipinnis, and
Pseudorca crassidens, feed on a diverse diet of cephalopods, shrimps (Clarke
et al. 1980; Pinedo 1987), and Sciaenidae and Serranidae (Pinedo 1989;
Pinedo and Rosas 1989), whilst Orcin us orca sometimes ingests individuals of
the swordfish Xiphias gladius which are hooked on long lines (Pinedo 1994a).
Every year groups of up to ten individuals of Eubalaena australis are seen
during the winter and spring, with the highest incidence in October, over the
s9uthern Brazilian shelf (27 0 23'-3l1O' S) between the coast and about 30 km
offshore (Pinedo 1985; Pinedo, unpubl.). The seasonal occurrence and
behaviour associated with copulation (Pinedo 1985) provide evidence that
the southern Brazilian shelf may have greater importance as a region of
Marine Mammals and Turtles 153

reproduction between populations from northern (28° 29' S) and southern

(42° 30' S) nursing areas in the southwestern Atlantic than previously
considered (Best et al. 1993). It is likely that the Eubalaena australis populat-
ion is recuperating from past overexploitation. In contrast, Delphinus delphis
occurs in groups of up to 500 individuals throughout the year over the
intermediate shelf and slope (29° 22'-340 31' S) between 70 and 2500 m depth
and is occasionally caught by fishing nets (Castello and Pinedo 1986; Pinedo
1994a).

Ecology of Principal Pinnipeds

Otaria flavescens and Arctocephalus australis are the most frequently

encountered species, especially between the fall and spring, although
recordings of Arctocephalus tropicalis have increased over the last 15 years
(Pinedo 1986, 1990). The remaining pinniped species are only occasional
visitors (Castello and Pinedo 1977c). After reproduction along the coast of
Uruguay and Argentina during the summer, starved individuals of Otaria
flavescens migrate to both southern (Vaz-Ferreira 1982a) and northern
feeding grounds. The migratory range of females appears to be more
restricted since they are most abundant at the southern limits of Brazil's coast
(34° S) in the winter (Pinedo 1986). In contrast, subadult and adult males
(3-5 years) migrate further north. The tip of the northern breakwater of the
Patos Lagoon inlet (32° S) serves as a resting place for individuals of Otaria
flavescens throughout the year, though there are more in the spring (Rosas
1989). Sexual maturation of females and males is reached between 3 and 4 years
and after 5 years, respectively (Vaz-Ferreira 1982a). Both sexes attain 95% of the
maximum length after 8 years (females 194.4 cm, males 254.4 cm; Rosas et al.
1993), although females may live to be 14 years and males 16 years. Especially in
the winter and spring, the principal cause of mortality is due to aggression by
fishermen during fishing activities, since approximately 30% of the 292 dead
specimens analyzed over a lO-year period displayed mutilations or perforations
caused by firearms (Pinedo 1986, 1990; Rosas 1989). Otaria flavescens preys on
five different elasmobranchs and on at least 15 different teleost fish of which
Paralonchurus brasiliensis, Macrodon ancylodon, Cynoscion guatucupa,
Micropogonias jurnieri, and Trichiurus lepturus represent the major diet items.
Like Otaria flavescens, the pinnipedArctocephalus australis leaves the
reproductive colonies in Uruguay (35° 10' S; Vaz-Ferreira 1982b) to migrate to
more northern regions. Most of the 106 dead individuals encountered over
10 years between latitudes 290 20' -330 45' S were young males of 80-100 cm
(PinedQ 1986, 1990). Dead specimens do not generally display mutilations since,
unlike Otaria flavescens, Arctocephalus australis does not follow fishing vessels
and tends to avoid nearshore waters. Natural death thus seems to be the major
cause of mortality (Pinedo 1986). Arctocephalus australis preys upon a large
154 C. M. Vooren

variety of organisms, composed of gastropods, shrimp (Pleoticus muelleri,

Artemesia longinaris), cephalopod (Loligo sanpaulensis, Illex argentinus),
and teleost fish, predominantly Engraulididae and Sciaenidae.
The islands South Georgia, Tristan da Cunha, and Gough are reproductive
areas for Arctocephalus tropicalis. Probably owing to a growing population, the
southern Brazilian coast is visited with increasing frequency by mostly sexually
inactive adult males, especially during the winter and spring (Pinedo 1990), as
has been observed for the coast of Chile (Torres et al. 1984). In southern
Brazilian waters, Arctocephalus tropicalis feeds on Anchoa marin ii, Pomatomus
saltatrix, Cynoscion guatucupa, Micropogonias jurnieri, Trichiurus lepturus,
Mugil spp., Odontesthes spp., Ariidae, and on Loligo sanpaulensis. Although
records of fishing interactions are lacking, at least 35% of the 65 dead specimens
found from 1976 to 1994 seem to have died of unnatural causes (Pinedo 1990;
Pinedo, unpubl.).

Marine Turtles

Spawning areas of marine turtles in Brazil are restricted to latitudes north of 19°
38' S (Marcovaldi 1991). Although Dermochelys coriacea, Chelonia mydas,
Eretmochelys imbricata and, more frequently Caretta caretta are seen from the
spring to fall between 29° 20' and 33° 45' S (Pinedo, unpubl.), in the absence of
detailed information, studies on population characteristics, feeding area and
behaviour, displacement patterns, and mortality causes of marine turtles from
this region are urgently needed (Ogren et al. 1989).

6.19
Sea and Shore Birds

C. M. Vooren

The coast of the warm-temperate southwestern Atlantic between 28° 40'

and 34° 34' S (Santa Marta Grande Cape to Chui) and its adjacent waters
sustain 83 species of sea and shore birds (Appendix). Of these, 72 are visi-
tors who breed in distant places and migrate to the area to feed. Two major
environments can be distinguished: the beach and nearshore waters
provide habitats for 50 species of shorebirds and coastal seabirds, including
44 Charadriiformes (gulls, terns, skimmer, waders, and sheathbill), while
the offshore waters of the continental shelf and slope sustain 33 species of
oceanic seabirds, including 25 Procellariiformes (albatrosses and petrels).
Shorebirds find in the swash zone an almost constantly accessible supply of
abundant food, mainly the bivalve molluscs Mesodesma mactroides and
Donax hanleyanus and the crustaceans Emerita brasiliensis and Excirolana
armata, while gulls and hawks feed on stranded debris such as marine
Sea and Shore Birds 155

organisms, wind-blown insects, and discards of the coastal fishery. Terns,

skimmers, herons, and cormorants feed on shrimps and fish which abound
all year round in the coastal waters, and the terns constitute a source of
food for the kleptoparasitic skuas. Zooplankton, small pelagic fishes, and
squid reach peak abundance in offshore waters during the winter and
constitute at that time, together with discards of the fishery, the food
supply for albatrosses, petrels, and penguins (Gianuca 1983; Belton 1984,
1985; Sick 1984; Harrington et al. 1986; Resende 1988; Vooren and Fernan-
des 1989; Vooren and Chiaradia 1990).

Migrations

Of the 11 "resident species", only the oystercatcher Haematopus palliatus

and the plover Charadrius collaris breed on the beach in the spring and are
permanent components of the coastal ecosystem. The terns Sterna trudeaui
and Sterna superciliaris, the gull Larus maculipennis, arid the stilt Himan-
topus himantopus breed in nearby inland habitats, are scarce on the shore
in the spring, and move to the coast in the fall and winter. The herons
Egretta thula and Ardea cocoi and the hawks Milvago chimango and Poly-
horus plancus forage all year round on the beach in small numbers and are
probably non-breeders of populations which nest further inland (Sick 1984;
Vooren and Chiaradia 1990).
Forty-two species are "southern migrants" which breed in the summer in
temperate to antarctic latitudes of the Southern Hemisphere, and most of
these reach peak numbers in southern Brazil during the winter. The cate-
gory includes 27 oceanic seabirds (albatrosses, petrels, and penguins) and
15 coastal seabirds and shorebirds (mainly terns, gulls, and waders; Fig.
6.19.1). Large numbers of dead juveniles of the penguin Spheniscus magel-
lanicus are a typical feature of the beach during the winter, and groups of
live birds are at that time common on the continental shelf. Adults are
scarce, but the shelf waters of southern Brazil appear to be the nursery for
first-year juveniles of Spheniscus magellanicus during the winter, when
food such as the anchovy Engraulis anchoita reaches peak abundance.
Banded juveniles from the penguin rookeries at Valdez Peninsula
(Patagonia, Argentina) have been found, and most of the birds probably
come from this breeding ground. Among the albatrosses, Diomedea
melanophris feeds in flocks on discards of th,e fishery industry on the
continental shelf, while Diomedea exulans is restricted to the edge of the
shelf, where squid such as Illex argentinus abound in winter and constitute
a food supply for the adults of Diomedea exulans which are at that time
rearing chicks on South Georgia. Among petrels, the most common winter
visitors from latitudes between the Malvinas Islands and the Antarctic are
Macronectes sp., Fulmarus glacialoides, Puffinus griseus, Pterodroma incerta,
156 C.M. Vooren

100r-----~==----------------_.
A • OTHERS (9)
o WADERS (13)
80 o TERNS (8)
• GULLS (3)
"E
.:.:
~ 60

>
l-
ii)
Z
W
C

70 B,--

60
- • SOUTHERN MIGRANTS (2)
o NORTHERN MIGRANTS (8)
o PERMANENT SPECIES (3)

"E 50
.:.: ~
,;
:§.
40
>
I-
Ci5
z 30
w
c r--

20

10
- r--
- ~
~
r--
~

I-- Fig. 6.19.1. Monthly mean

rr
f-- density of shore and coastal
'--
-- seabirds (A) and of waders
J F M A M J J A SON D (B) on the Atlantic ocean
MONTHS beach at 32' S

Pachyptila belcheri, Dapiion capense, and Procellaria aequinoctialis

aequinoctialis, the last two species feeding on discards of the fishery
industry. Puffinus gravis, Diomedea chlororhynchus, and Procellaria aequi-
nd'ctialis conspicillata from Tristan da Cunha Island are common all year
round, the former two species being numerous on the continental shelf where
they follow fishing boats, while Procellaria aequinoctialis conspicillata occurs
only at the edge of the shelf. Puffinus gravis is usually said to migrate in the
Sea and Shore Birds 157

winter to the North Atlantic, but it is evident that part of its population
remains in the southern hemisphere at that time. The same is true for
Oceanites oceanicus which breeds in the Antarctic and is seen off southern
Brazil from May to September. Larus dominicanus breeds on the coast of
Uruguay in the spring and is one of the most numerous birds on the coast
of Brazil's extreme south during the remainder of the year. Sterna maxima,
Sterna hirundinacea, and Sterna eurygnatha breed in Argentina, the former
two species wintering on the South Brazilian coast, while Sterna eurygnatha
passes through the area in the spring and fall, with a few birds remaining
during the winter. Flocks of non-breeding Rynchops niger roost during the
summer at coastal lagoons, but it is not known where these birds come
from. Charadrius falklandicus and Charadrius modestus are common on
the beach in the winter and are evidently migrants from Patagonia,
although a small number of the former species breeds in the southern
Brazilian Peixe Lagoon (see Fig. 3.2). In addition, the Andean flamingo
Phoenicopterus chilensis is habitually seen in numbers of up to 500 birds at
the Peixe Lagoon (Escalante 1970; Belton 1984; Yo-oren and Fernandes
1989; Croxall and Prince 1990; Marchant and Higgins 1990; Vooren and
Chiaradia 1990; Prince et al. 1992).
Thirty species are "northern migrants" which breed in temperate to arctic
latitudes of the Northern Hemisphere during the boreal summer and reach
peak numbers in southern Brazil from the austral spring to fall (Fig. 6.19.1).
The category includes 23 waders, 3 skuas, 2 terns, and 2 petrels. Ten species
occur in large numbers and are discussed below. Pluvialis dominica,
Pluvialis squatarola, Calidris alba, Calidris fuscicollis, and Calidris canutus
from Canada are the most common waders. Calidris canutus reaches peak
numbers of up to 150 birds per km of shoreline from March to May,
completing the pre-nuptial moult and then fattening up for the northward
flight to Delaware Bay. The other five species spend the summer in the
area, but populations of Calidris fuscicollis and Calidris alba, which spend
the austral summer in Argentina, use the beach as a staging area in the fall
in the manner of Calidris canutus. Sterna hirundo from North America
roosts on selected places of the coast in groups of up to 14000 birds during
the summer and fall. Stercorarius longicaudus and Stercorarius parasiticus
associate with flocks of terns at sea. Puffinus pUffinus from the British Isles
migrates southward through the area, mostly from August to October, as
evidenced by the number of dead individuals washed ashore during those
months. Puffinus diomedea is common over the edge of the continental
shelf during the summer (Belton 1984; Harrington et al. 1986; Resende
1988; Vooren and Fernandes 1989; Vooren and Chiaradia 1990).
The coast and the waters of the continental shelf and slope of southern
Brazil are of crucial importance for populations of migratory sea and shore
birds which breed in Arctic Canada, the British Isles, oceanic islands off
Spain and Portugal, the Tristan da Cunha Islands, Patagonia, the Malvinas
158 C. M. Va oren: Sea and Shore Birds

(Falkland) Islands, South Georgia and South Orkney Islands, and the
Antarctic Peninsula. Through its role as a feeding and staging area on the
migration routes of these birds, the region occupies a strategic position in a
web of ecological relationships which spans the Americas and the Atlantic
Ocean as a whole. The conservation of the coastal and marine environ-
ments of southern Brazil is of worldwide significance.

Feeding and Resource Partitioning on the Beach

On the lower beach the eight abundant waders Haematopus palliatus,

Pluvialis squatarola, Pluvialis dominica, Charadrius collaris, Charadrius
falklandicus, Calidris canutus, Calidris alba, and Calidris fuscicollis feed on
invertebrates which live buried in the sand of the swash zone and which
only emerge to migrate with the water level as this changes due to wind and
lunar tides. As indicated by the shape and size of the bill and specialized
searching behaviour, Haematopus palliatus preys preferentially on large
individuals of the clam Mesodesma mactroides which remain buried at a
depth of about 5 cm and are therefore not accessible to other birds, except
when the clams emerge to migrate with the tide, and can be taken by the
gull Larus dominicanus. From February to April, small juveniles of the
clams Mesodesma mactroides and Donax hanleyanus and the crustaceans
Emerita brasiliensis and Excirolana armata reach peak numbers in the
swash zone, with up to 7000 to 20 000 individuals per m2 for each of these
species, and the arrival of the migrating sandpipers Calidris canutus,
Calidris juscicollis, and Calidris alba in the area is timed so as to coincide with
the availability of this food supply. Calidris canutus and Calidris juscicollis
probe the slightly submerged or just exposed wet sand of the swash zone
with their straight and fine bills, mostly in the manner known as stitching
(Burton 1974), localizing prey through touch and taste. Calidris canutus
searches in compact flocks which move along as a single unit, while
Calidris fuscicollis does so in loosely scattered groups. The latter species
also searches visually on the dry sand above the swash zone. Calidris alba
has a thicker bill and feeds in dense flocks on the submerged substrate,
running constantly with the advance and retreat of the waves and catching
exposed prey on sight. In contrast to the sandpipers, the five common
plovers of the genera Pluvialis and, Charadrius have powerful running legs,
short thick bills, and very large eyes. These birds search for their prey
exclusively by visual means and in a solitary manner, over an intraspecific
distance of at least 2 to 10 m. The birds feed mostly on the exposed
substrate at the upper margin of the swash zone, and to a lesser extent on
lightly submerged sands. Charadrius falklandicus, Charadrius collaris,
Pluvialis dominica, and Pluvialis squatarola feed in a similar way in the
same places, but while the latter two occur in the summer, Charadrius
Genetic-Biochemical Differentiation among Marine Organisms 159

falklandicus is a winter visitor, and Charadrius collaris lives during the

summer in the dunes and feeds on the lower beach in the winter. Thus, the
seven abundant species of sandpipers and plovers differ in methods of prey
searching and in the distribution in space and time of feeding within the
swash zone, partitioning in this way the space available for feeding and the
prey according to different states of behaviour, i.e. buried or migrating with
the tide. Future research may decide whether through these differences in
feeding the birds partition different species of prey or different population
strata of the same species of abundant prey (Gianuca 1983; Harrington et al.
1986; Vooren and Chiaradia 1990).
Sterna superciliaris fishes mostly at creek mouths, Sterna trudealti does so
in the surf zone, while Rynchops niger fishes in lagoons and in quiet estuarine
and inshore waters. The other terns fish further offshore, Sterna hirundo
associating with schools of skipjack tuna. Thus, resource partitioning occurs
through spatial separation of the fishing areas, as also happens among
piscivorous birds of tropical oceanic islands (Ashmole and Ashmole 1967).
The gulls Larus dominicanus and Larus maculipennis are unspecialized
generalist feeders, taking animal debris on beaches and in coastal waters,
Larus maculipennis also catching insects on the wing and small fishes at
shallow creek mouths, while pellets of Larus dominicanus frequently contain
shells of bivalve molluscs. In order to have access to their food supply, all
these birds need undisturbed and safe places on the beach for roosting within
reach of their feeding areas. Due to the long extent of the beaches and low
level of human disturbance, the birds share bivalves without competitive
.interaction. Gulls and terns roost together on the beaches in multispecific
flocks which are structured as an assemblage of monospecific groups
(Vooren and Chiaradia 1990; Chiaradia 1991; Vooren, unpubl.).

6.20
Genetic-Biochemical Differentiation
among Marine Organisms

J. A. Levy
Genetic-biochemical data have revealed patterns of phylogenetic relation-
ships within the families ofSciaenidae (Cassano 1987) and Scombridae, and
determined the taxonomic status in the genera Squatina (Chondrichthyes;
Sole-Cava et al. 1983; Sole-Cava and Levy 1987), Myliobatis (Chondrichthyes;
Levy and Conceicrao 1989), Menticirrhus (Cassano and Levy 1990),
Cynoscion (Levy and Cassano 1994), Eledone (Cephalopoda; Levy et al.
1988) !lnd Odontesthes (Atherinidae; Prodohl and Levy 1989) from the
warm-temperate southwestern Atlantic.
160 J. A. Levy: Genetic-Biochemical Differentiation among Marine Organisms

Genetic-biochemical data have also provided important information for

fisheries management (FAO/UNEP 1981) because aspects of population
dynamics, speciation, and distribution can be interpreted by the degree of
genetic differentiation within a species, which results from restricted
genetic flow between individuals of the same population. Since populations
of marine species are frequently divided into morphologically and ecologi-
cally different groups with distinct spawning periods and sites, reproduc-
tive isolation enhances the accumulation of genetic differences between
groups, owing to mutation, genetic drift, or selection. Understanding the
genetic variability within and genetic divergence between populations is
paramount for the rational exploitation and conservation of biological
resources (PIa et al. 1990), because decreasing variability in the natural
populations increases the probability of their extinction (Meffe 1986).
Individuals of several economically important populations in the south-
western Atlantic, like Micropogonias furnieri (Maggioni 1992, Matthiensen
et al. 1993; Maggioni et al. 1994), Mesodesma mactroides (Marins et al.
1995), Artemesia longinaris (Weber et al. 1993"),Penaeus paulensis, arid
Odontesthes argentinensis, show normal genetic variability and high homo-
geneity among sampled allelic frequencies, and thus should be considered
as belonging to genetically homogeneous populations. These results sug-
gest that the warm-temperate southwestern Atlantic region might act as
one large marine ecosystem, similar to those of the eastern coast of North
America and the coast of South Africa (Sharp 1988).
7 Relationships and Function of Coastal
and Marine Environments

7.1
The Foredune System

U. Seeliger

The coastal dunes of southern Brazil are remarkably similar because

Holocene sea-level changes have influenced their genesis (pfadenhauer 1980;
Schwarzbold and Schafer 1984), resulting in the sequential formation of
depressions and dune ridges parallel to the shore. The most recent transgres-
sion-regression events moulded the 300-1000 m wide fore dunes, with season-
ally flooded freshwater marshes immediately interior of them. The flat
marine terrace supplies a considerable amount of fine quartz sand to the
beach (pfadenhauer 1980), and the sandy substrate of windblown foredunes
contains a high proportion of shell fragments but little organic matter
(Cordazzo and Seeliger 1987). Since the tidal ranges are low «SO em), the
dominant southeasterly and northeasterly onshore winds in the winter and
summer, respectively, control seawater flooding of the beaches and shore
profiles (pfadenhauer 1980; Costa et al. 1984, 1991; Bernardi et al. 1987).
Different orientations of the coast north and south of 33° S imply variations
in onshore wind approach angles and impact. On the northern coast, NE
onshore winds induce considerable beach-dune sand transport. In contrast,
along the southern coast beach-dune sandflow is reduced because NE winds
blow parallel to the shore, whilst SE winds blow directly onshore and cause
inundation of beaches and landward erosion of backshore and fore dune
sands. Additionally, winter cold front passages lead to severe beach overwash
and differentially affect sand dynamics along the northern and southern
coasts. As a result, the northern beaches are largely intermediate and
occasionally dissipative, whilst the southern beaches with coarser-textured
sand tend to become reflective (Calliari and Klein 1993).

Dune Habitats

In spite of latitudinal disturbance gradients, which are a source of spatial

and temporal environmental heterogeneity (pfadenhauer 1980; Bernardi
and Seeliger 1989; Seeliger 1992b), the entire coastal foredune complex can

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
162 U. Seeliger

be regarded as one functional unit (sensu Doing 1985). Gradients of beach/

foredune sediment transport and edaphic stability of foredunes not only
induce different site-specific environmental conditions (Hesp 1991), but
also influence the composition of vegetation and the abundance of species
(Barbour 1992; Fahrig et al. 1993). Pioneer backshore and foredune plants
are selected in accordance with their physiological tolerance and
morphological and reproductive traits (Barbour et al. 1976, 1987; Hesp
1991). The environmental conditions and plant associations define six typi-
cal habitats on southern Brazilian backshores and foredunes. Plant asso-
ciations, dominanted by Blutaparon portulacoides, Panicum racemosum,
Spartina ciliata, Hydrocotyle bonariensis/Paspalum vaginatum, And-ro-
trichum trigynum, and Andropogon arenarius, occur in response to mean
low water (MLW) height and distance, and along salinity, substrate humid-
ity and stability, and silt/clay contents gradients (Cordazzo and Seeliger
1993; Costa et al. 1995). The different environmental and vegetational
characterstics of each habitat also provide the conditions for a diverse
animal community in coastal foredunes.
Since tidemarks are wind-controlled, they are unstable and often over-
lap, thus leading frequently to life-cycle disruptions of the otherwise
common annual driftline species Cakile maritima, Calycera crassiflora, and
Calystegia soldanella (Barbour et al. 1976; Doing 1985; Moreno-Casasola
and Espejel1986; Fahrig et al. 1993). As a consequence, southern Brazilian
beaches lack driftline habitats because species retreat to more stable and
permanent tidemarks behind the backshore (Cordazzo and Seeliger 1993).
The backshores themselves are inhospitable environments and are sea-
sonally unstable, owing to eolian sandflow and erosive seawater overwash
during winter storms, respectively. The salt-tolerance and rhizome-propa-
gated perennial growth of Blutaparon portulacoides enable this species to
withstand the local conditions of saline stress and sand instability
(Bernardi and Seeliger 1989). The plants capture the windblown sand
(Pfadenhauer 1980) and promote the development of an extensive embryo
dune habitat (pfadenhauer 1978; Bernardi and Seeliger 1989). Although
extreme storms occasionally completely erase this habitat, reestablishment
takes less than 2 years (Bernardi et al. 1987). Only where creeks break the
dunes does the outflowing water destroy the embryo dune habitat; under
increasing freshwater inundation stress, Blutaparon portulacoides is
replaced by a Hydrocotyle/Paspalum association (Cordazzo and Seeliger
1993). Owing to predominantly terrestrial conditions, the fauna of the
embryo dune habitat is primarily composed of insects, although
amphipods of marine origin (Orchestoidea brasiliensis) may also occur
(Gianuca, Sect. 6.5).
The Foredune System 163

Most sand cannot be fixed in the embryo dune habitat but is deposited
in zones beyond seawater advance, where virtually mono specific stands of
Panicum racemosum progressively capture wind-blown sand (Cordazzo
and Seeliger 1993). Accretion rates of up to 70 cm year-I (Costa et al. 1984)
promote the formation of a 4-6 m high frontal dune ridge habitat
(Cordazzo and Seeliger 1993). The continuous deposition of nutrient-rich
sand favours the sprouting of rhizome fragments and vigorous growth of
Panicum racemosum (Costa et al. 1991). On leeward dunes, with lower sand
deposition and fewer nutrients (Costa et al. 1984, 1991), Panicum racemosum
enters a regressive stage, and species like Senecio crassiflorus and
Gamochaeta americana invade (pfadenhauer 1978, 1980; Cordazzo and
Seeliger 1987, 1988a). The relatively dense plant cover and the dry substrate
offer niches for a large number of insects, though frogs, lizards, and small
mammals like the tuco-tuco (Ctenomys flamarioni; Ctenomyidae) are also
important components (Gianuca, Sect. 6.5).
Lower humidity areas beyond the backshore are exposed to disturbance
by occasional seawater overwash. Here, its large number of highly viable
seeds favours the growth of Spartina ciliata (Hueck 1955; Legrand 1959;
Pfadenhauer 1980; Costa and Seeliger 1988a), and the plant's active excre-
tion of salts compensates for sediment salinization (Bastos et al. 1993b).
Since the clumpy growth of Spartina has little effect on sand-binding, a
periodically recycled, typically 1-2 m high, rounded hummock dune habitat
develops (Seeliger 1992a).
Behind the hummock dune and frontal dune ridge habitats, either grad-
ual sand burial of lower freshwater marshes or the formation of blowouts
establishes a mosaic of dry slack habitats and seasonally inundated wet
slack habitats. Owing to greater substrate stability, the diversity of plant
and insect species increases in these habitats (pfadenhauer 1978, 1980;
Costa et al. 1984; Cordazzo and Seeliger 1987, 1988a, 1993). The lower wet
slack habitat is generally inundated during the winter and is dominated by
Androtrichum trigynum and other inundation-tolerant species (Phyla
canensis, Bacopa monnieri, Pluchea sagittalis, Paspalum vaginatum, Hydro-
cotyle bonariensis) and marsh remnants Uuncus acutus, Typha domingensis;
Cordazzo and Seeliger 1987, 1993). The dry slack habitat, with reduced
inundation stress, is characteristically occupied by Andropogon arenarius.
Slacks and adjacent marshes serve as nesting sites for several shore birds
and are inhabited by the skunk Conepathus chinga, the armadillo Dasypus
hybridus, and the fox Dusicyon gymnocercus (Gianuca, Sect. 6.5).
164 U. Seeliger

URUGUAY

20 km

Fig. 7.1.1. Typical coastal foredune physiography changes from frontal dune ridges (A)
over hummock dunes (B) to sand plains '(C) with increasing shore disturbance

Dune Physiography

Ocean-shore disturbance gradients are a consequence of oceanic and

climatic allogenic processes. Their relative significance depends on the
contribution wave and wind energies has on both sand transport (Short
The Foredune System 165

and Hesp 1982) and the selection of beach/foredune plants. Disturbance

not only selects for dominant plant species, but selected plants also modify
the impact of disturbances. As has been observed elsewhere (Barbour et al.
1976, 1987; Godfrey 1977; Wiedemann 1984; Fahrig et al. 1993), shore
disturbance gradients, either acting alone or in concert with the dominant
plant cover, are likely to have influenced the genesis of different fore dune
physiographies along the southern Brazilian coast over time (Costa et al.
1991; Seeliger 1992b; Cordazzo and Seeliger 1993; Fig. 7.1.1).
A change in orientation of the southern Brazilian coastline causes a
higher impact of winds and waves in the south (Calliari and Klein 1993),
and thus disturbances owing to beach overwash and sand transport
patterns increase in a north-south direction. Although disturbances tend to
interact more severely with backshores than with foredunes, their modify-
ing influence is more apparent in foredune areas. On backshores annual
driftline species (Cakile maritima, Calycera crassiflora, Calystegia soldan-
ella) and Blutaparon portulacoides tolerate intense disturbance. Owing to
their reduced sand-binding capacity, the annual growth of driftline species,
and the periodic obliteration of Blutaparon portulacoides popUlations by
sea overwash (Bernardi et al. 1987; Bernardi and Seeliger 1989), their
modifying influence on disturbance is likely to be negligible. However,
different rates of sand flow from the beach and different sand accre-
tion/erosion patterns may have selected for either the Panicum or Spartina
associations in the fore dune areas. Their different strategies of spatial
occupation, growth, and sand-binding capacity (Costa et al. 1984, 1991;
Costa and Seeliger 1988a,b; Cordazzo and Seeliger 1993) are likely to have
ultimately affected the evolution of the fore dune physiography.
At northern, less disturbed shores, above the influence of storm tide
disturbances and saltwater flooding, the clonal spreading and vertical
growth of Panicum racemosum promote efficient sand deposition, which
counteracts sand erosion during periods of prevailing offshore winds
(Pfadenhauer 1978, 1980; Costa et aL 1984, 1991; Cordazzo and Seeliger
1987, 1993). Over time, the continuous sand flow from the beach results in
the formation of a prograding fore dune ridge physiography which protects
dry and wet slack habitats in the lee and prevents sand transgression over
the inland marshes (Costa et al. 1991; Seeliger 1992a).
Further south, Spartina ciliata dominates lower shores characterized by
increased disturbance from periodic wind erosion or occasional sea over-
wash. The clumpy clonal growth of Spartina, with limited sand-binding
capacity, typically forms an extensive hummock dune physiography with
interdispersed dry and wet slacks. Stands of Acacia, Eucalyptus, and Pinus
bind persisting landward sand transport and occasionally form extensive
retention ridges.
166 v. M. T. Garcia and N. M. Gianuca
Along the severely disturbed coast in the extreme south of the region,
the reduced sand flow from the beach and the continuous sand erosion in
fore dune areas by prevailing winds have essentially impeded the establish-
ment of a plant cover (Costa et al. 1991; Costa and Seeliger 1988b, 1990),
and allogenic factors appear to be entirely responsible for the shaping of
the fore dune landscape. As a consequence, the dominant foredune ridge
and hummock dune physiographies of northern coastal regions have been
replaced by a wind-eroded, humid, deflation plain physiography. Beyond
the deflation plains, the continuous landward sandflow formed large trans-
gressive dune fields which covered inland marshes and merged with older
dunes (Gomes et al. 1987).
Since foredunes perform a major function in coastal protection and
wildlife conservation, and human activities often interfere with this func-
tion, management approaches must consider ecological differences within
and between physiographically distinct regions (Godfrey 1977) to preserve
the value of any part of the coast for specific types of land use (van der
Meulen et al. 1985) and to prevent heavy losses or complete destruction cif
pristine dune areas (Westhoff 1985).

7.2
The Beach and Surf Zone

v. M. T. Garcia and N. M. Gianuca

The beach and surf zone of intermediate and dissipative beaches, confined by
the outer limits of circulation cells in the external surf zone and the upper
limits of swash, may act as a single semi-closed and self-sustaining ecosystem,
which is largely independent of food input from the adjacent sea (McLachlan
1980, 1983). Components of the necton and benthic, planktonic, and
microbial communities tend to lend considerable diversity to such a system
(Brown and McLachlan 1990). The extensive beaches of Brazil's southern coast
are largely of the intermediate type (sensu Wright and Short 1984), though
dissipative beaches dominate immediately south of 32° S (Gianuca 1983; Calliari
and Klein 1993). Therefore, the majority of the coast is characterized by wide
sandy beaches with a gentle slope and a well-developed surf zone with
moderate to high wave energy, where sandbars, deposited parallel to the
beach, are cut by rip currents. Sub-aerial and exposed beach profiles are
seasonally modified as a result of wave energy-induced sand accretion during
summer and erosion during the remainder of the year (Calliari and Klein
1993). During most of the year, waves break regularly at a similar distance from
the shore and form the inner surf zone down to approximately 2 meters depth.
Less frequent storm waves, particularly during cold-front passages, generate an
external surf zone between 2 and 10 m depth (Borzone and Gianuca 1990).
The Beach and Surf Zone 167

Organic Matter Sources

Diatom patches are a typical feature of surf zones of intermediate and dissi-
pative beaches and constitute a major source of organic matter (Schaefer and
Lewin 1984; Brown and McLachlan 1990; Talbot et al. 1990). Along the south-
ern Brazilian coast, the phytoplankton of the surf zone is characteristically
dominated by frequent and dense accumulations of the diatom Asterionellopsis
glacialis. Apart from mesoscale cycles (weekly or monthly), patch formation
appears to follow diurnal cycles, with high cell concentrations in the water
column during the day which decline at night. Although high silicate (30-
60 1lM) concentrations, highly variable nitrate (1-30 1lM) levels, and.1ow N:P
ratios (7-8) in surf zone waters suggest a possible nitrogen limitation for
phytoplankton growth, the intensity of mesoscale accumulations appears to
be largely controlled by physical and meteorological factors and the
dynamics of the system (Odebrecht et al. 1995a). Similar to patch formation
during "storm-caIrn-storm" cycles elsewhere (Talbot and Bate 1989), the
development of Asterionellopsis glacialis patches along the southern Brazilian
coast is related to cyclic cold front passages with strong southerly winds,
which force seawater onshore and generate high-energy waves and a wider
surf zone (Gianuca 1983; Odebrecht et al. 1995a). Under these conditions,
high concentrations of Asterionellopsis glacial is cells are resuspended from
nearshore sediments and, owing to the high wave energy, accumulate in the
surf zone where exposure to intense light stimulates further cell growth
(Garcia, unpubl.). Rip currents and undertow return the diatom cells to
nearshore sediments beyond the surf zone where they accumulate under
calm conditions.
Mesoscale cycles of Asterionellopsis glacialis patch formations are accom-
panied by extreme fluctuations in chlorophyll a concentrations (1.2-1600 /-lg rl;
Odebrecht et al. 1995a). Particularly in the summer, centric diatoms
(Skeletonema costatum and large species of the genus Coscinodiscus) become
important components of the phytoplankton community (R6rig, unpubl.)
and contribute to average annual chlorophyll a concentrations of2.7-350 /-lg r 1
in the water column. Finally, occasional nuisance blooms of potentially toxic
dinoflagellates, like Gyrodinium cf. aureolum, Dinophysis acuminata and
Noctiluca scintillans (Machado 1979; Rosa and Buselato 1981; Garcia et al.
1994; Odebrecht et al. 1995b) are associated with massive mortality of clams.
Although information on phytoplankton primary production levels is
still lacking, peak chlorophyll a concentrations in the surf zone (Odebrecht
et al. 1995a) and estimates based on experimental cultures of Asterionellopsis
glacialis isolates with maximum rates of 3.74 mg C mg chlorophyll-l h- 1
(Garcia and R6rig, unpubl.) suggest that primary production rates during
diatom accumulations may vary between 1300 and 6000 mg C m-3 h- 1• These
data are comparable to values reported for surf accumulations of other
diatom species elsewhere (Schaefer and Lewin 1984).
168 v. M. T. Garcia and N. M. Gianuca
Average particulate organic carbon (POC) concentrations in the water
column (460-20 050 f.Lg n and chlorophyll a values provide an estimation
of a mean C:Chl a ratio of 44 and a contribution of phytoplankton biomass
between 20 and 70% to the total POC pool. Furthermore, elevated back-
ground levels (1817 ± 300 f.Lgn of particulate organic detritus in surf zone
waters may contribute between 20 and 95% to the particulate organic
carbon pool (Garcia, unpubl.), and therefore comprise an important food
source for detritivorous organisms and the microbial community.
The intense mucus secretion by Asterionellopsis glacialis cells (Odebrecht
et al. 1995a) releases considerable amounts of dissolved organic matter into
surf zone waters during patch formation. In addition, excretion products
liberated by the abundant benthic intertidal fauna (Gianuca 1983, 1985) as
well as decomposition processes are likely to increase dissolved organic
carbon concentrations further. These compounds are either directly avail-
able to microbial organisms through the microbial loop (Brown and
McLachlan 1990) or may aggregate via bubble formation to make particulate
organic matter (Ogura 1972). As observed for othel'dissipative or intermedi-
ate beaches (Talbot· and Bate 1988; Brown and McLachlan 1990), phyto-
plankton appears to be the principal source of organic matter in the surf zone
along the southern Brazilian coast, although a constant supply of detrital
matter may become important, especially in the absence of diatom patches.

Trophic Relations

Macrobenthic filter, suspension, and detritus feeders are the most important
primary consumers of the beach and surf zone and are the major link
between the abundant phytoplankton and organic detrital matter and higher
level consumers (Gianuca 1985), though zooplanktonic organisms and some
fish species are also important primary consumers.
Polychaetes and the filter-feeding molluscs Amiantis purpuratus, Tivela
ventricosa, and Mactra isabelleana constitute the major macro benthic bio-
mass in the external surf zone, followed by detritus-feeding organisms, which
ingest deposited diatoms and detrital matter. Among the 126 inner surf zone
species (Borzone and Gianuca 1990), dense populations of filter and suspen-
sion feeders, like Donax gemmula (1809 indo m"\ Magelona riojai (7874 indo
m"2), and Phoxocephalopsis zimmeri (2050 indo m\ use phytoplankton and
detrital particles as an important food source. Filter feeders, like Mesodesma
mactroides, Donax hanleyanus, and Emerita brasiliensis, which total 95% of
the secondary production in the lower intertidal zone, feed intensely on the
surf diatom Asterionellopsis glacialis (Gianuca 1983, 1985, 1987). The abun-
dant (91 581 indo m"2) suspension-feeding intertidal polychaete Spio gaucha
can survive under culture conditions on a diet composed exclusively of
Asterionellopsis glacialis (P. Santos, unpubl.). Even beyond the intertidal
The Beach and Surf Zone 169

zone, large depositions of Asterionellopsis glacialis and detrital matter during

storms in supratidal or overwash areas between foredunes serve as food for
dense (2350 indo m-2) populations of beetles (Bledius bonariensis, Bledius
microcephalus, Bledius Jernandezi) and the ghost crab Ocypode quadrata,
which occasionally complements its diet with diatoms (Gianuca 1985). The
high density of supratidal beetles attracts several predators, both insects and
birds, like the collared plover Charadrius collaris and the pipit Anthus
correndera.
The zooplankton community of the surf zone in southern Brazil is domi-
nated by mysids and copepods, although meroplankton, composed of
gastropod, bivalve, cirriped, decapod, and echinoderm larvae, is also_present
(Bersano 1994). Particularly under the influence of longshore currents with
increased hydrodynamic activity, mysids frequently exceed 60% relative
abundance. Metamysidopsis elongata atlantica is the most abundant species
and occurs in 95% of the samples. Copepods are less abundant but more
diverse. The omnivorous Acartia tonsa and Eucalanus pileatus are the most
common species (Bersano 1994). Owing to its high biomass of approximately
320 mg C m-3 (max. 8100 mg C m-3 ), the zooplankton in the surf zone is likely
to exert a strong grazing pressure on the phytoplankton community.
The fish assemblage of the surf zone, mainly represented by Trachinotus
marginatus, Mugil platanus, Mugil curema, Menticirrhus littoralis, Oncopterus
darwini, Odontesthes bonariensis, Atherinella brasiliensis, Lycengraulis
grossidens, Brevoortia pectinata, and Micropogonias furnieri, differs from that
of adjacent deeper (>5 m) coastal waters. Juveniles of many of these species
utilize the surf zone as a nursery and feeding area (Cunha 1981). Juveniles of
Mugil platanus, Mugil curema, and Mugil gaimardianus ingest cyanobacteria,
dinoflagellates, and small crustaceans, though their principal diet is com-
prised of diatoms. The common surf zone diatoms Asterionellopsis glacialis
and Skeletonema costatum may represent 90% of the stomach content of
Mugil platanus and Mugil gaimardianus, respectively (Vieira 1985).
Odontesthes sp. and Atherinella brasiliensis change their diet preference from
diatoms to zooplankton during development (Bemvenuti 1990).
The extreme density (up to 430 000 indo m-') of macrobenthic inner surf
zone invertebrates (Soares 1992) supplies abundant food resources for a variety
of secondary consumers. The diet of juvenile fishes of the species Menticirrhus
littoralis, Oncopterus darwini, Netuma barba, and Trachinotus marginatus is
almost entirely composed of benthic invertebrates (Gianuca 1985; Monteiro-
Neto and Cunha 1990). Large individuals of Pogonias cromis feed on the
bivalves Amiantis purpuratus, Tivela ventricosa, and Mactra isabelleana
(Gianuca 1985). The surf zone gastropods Buccinanops duartei, Olivancillaria
auricularia, Olivancillaria uretai, and the swimming crab Arenaeus cribarius
prey on Mesodesma mactroides, Donax hanleyanus, and Emerita brasiliensis
in the swash zone (Gianuca 1983, 1985). Macrobenthic invertebrates are also
ingested by larger predators, like rays, skates, and small sharks.
170 V. M. T. Garcia and N. M. Gianuca: The Beach and Surf Zone

SEA SURF ZONE AND BEACH LAND

.
SURF DIATOMS I- - - - + / - - - - !
(POC+DOC) / I
[ BIRDS I
EXPORT
[ DETRITUS l---------~[INSECTS I

Fig. 7.2;1. Schematic diagram of trophic relationships in the beach and surf zone eco-
system of the warm-temperate southwestern Atlantic

The abundance of juvenile fish in shallow waters attracts sub-adults and

adults of different ichthyophagus fish as well as gulls, terns, cormorants,
and herons, which are top predators in the surf zone (Vooren and Chiaradia
1990). As the tide retreats, shore birds (plovers, oystercatchers, sandpipers)
become the main predators of intertidal invertebrates. During the summer,
90% of the stomach content of Calidris fuscicollis, Calidris canutus, Calidris
alba, Pluvialis dominica, Larus dominicanus, and Larus maculipennis is
composed of the filter feeders Mesodesma mactroides, Donax hanleyanus,
and Emerita brasiliensis (Gianuca 1983, 1985). The elevated biomass of
benthic invertebrates constitutes an important food supply not only for
resident bird species, but also for many other temporary visitors during
their migrations north and southwards.
In conclusion, the environmental interactions and trophic relations
between different community components suggest that the beach and surf
zone in southern Brazil functions as a semi-closed and largely self-
sustained ecosystem (Fig. 7.2.1), as has been proposed by McLachlan (1980)
for exposed South African beaches.
The Continental Shelf and Slope 171

7.3
The Continental Shelf and Slope

J.P. Castello, M. Haimovici, C. Odebrecht, and C.M. Vooren

The influence of the Subtropical Convergence (STC), with marked seasonal

latitudinal displacement, characterizes the southern Brazilian continental
shelf and slope regions (29°_34° S) as a biogeographic transition zone (Sharp
1988) between the large neritic areas of Patagonia and tropical Brazil. The
composition and abundance of species, the pelagic structure, the spatial
distribution of communities and their trophic interactions, as well as
biological production are largely controlled by the seasonal dominance of
distinct water masses over shelf and slope. Coastal and shelfbreak upwelling
of deep Subtropical Water (STW) (South Atlantic central water) is frequent in
the spring/summer and winter/spring, respectively (Lima and Castello 1995).
The influence of Subantarctic and Tropical Water is greatest in the winter
and summer, respectively, though waters of subantarctic origin may also rise
during the summer along the southernmost shelf break regions (Fig. 7.3.1).
Freshwater runoff from the La Plata River and the Patos Lagoon become
important in the winter and spring. Particularly between Rio Grande and
Chui, oceanographic conditions favour high biological production, which
results in a considerable, still almost unexploited biomass of small pelagic
fishes (Castello and Habiaga 1982; Lima and Castello 1995) and highly
productive, but now overexploited, demersal fishery resources (Haimovici et
al. 1989a; IBAMA 1993). During most of the year, the thermal front on the
western side of the STC provides a suitable habitat for pelagic sharks,
tunafish, and thelike. Together, pulsed sub-surface coastal upwelling,
onshore Ekman transport, and large-scale influence of continental runoff
seem to sustain favourable conditions for fish spawning grounds and larval
survival.

Pelagic Production

Owing to the influence of nutrient-rich Subantartic Water (SAW) and

freshwater runoff from the Patos Lagool1 and La Plata River, chlorophyll a
and primary production rates are highest in the late winter and spring.
Although outflowing freshwater generally has low nitrate concentrations,
owing mainly to in- or nearshore phytoplankton uptake (Ciotti et al. 1995;
Abreu et aL 1995a), nearshore bottom turbulences add nutrients to the
euphotic zone (Odebrecht and Djurfeldt 1996) indicating that recycled
nutrients may efficiently return to the water column. The influence of SAW
on biological production occurs especially over the southern central shelf,
where horizontal and vertical fronts are related to areas of the highest
172 J.P. Castello et al.: The Continental Shelf and Slope

integrated chlorophyll a concentrations (> 100 mg m o2 ; Ciotti et al. 1995). In

the summer, the southward displacement of the oligotrophic Brazil Current
leads to the lowest chlorophyll a and primary production rates. During this
period, sporadically high chlorophyll a values and/or primary production
rates depend on the upwelling intensity of nutrient-rich STW and the mixing
regime, with high chlorophyll a concentrations occurring at sub-surface
(Odebrecht and Djurfeldt 1996) or surface waters (Hubold 1980a). Interannual
differences between phytoplankton chlorophyll a concentrations in shelf
waters are strongly determined by continental runoff, which seems to be a
function of the El Nino-Southern Oscillation cycle (ENSO).The highest
chlorophyll a concentrations off southern Brazil coincide with a large
freshwater outflow after periods of strong El Nino events in the Pacific Ocean
(Ciotti et al. 1995). In addition, changes in the intensity of the Malvinas
Current, probably as a consequence of climatic disturbances related to the
ENSO phenomenum, may affect the variability of primary production from
year to year.
A first evaluation of primary production data suggests moderate to high
mean annual primary production rates (160 g C m o2 i'; Odebrecht and
Garcia, Sect. 6.7) which probably sustain relative high secondary production
(Cushing 1988). The high winter/spring biomass of zooplanktophagous (i.e.
Engraulis anchoita), benthophagous, and predatory organisms, many of
which, owing to the northward shift of the STC and the presence of SAW,
migrate during this period from the south, supports this notion. Among the
over 40 species of the pelagic necton community of the continental shelf, the
fishes Engraulis anchoita, juvenile Cynoscion guatucupa, Trichiurus lepturus,
and the squid Loligo sampaulensis are the most characteristic (Mello et al.
1992). Biomass estimates (in 1000 tons) for Engraulis anchoita range from 35
to 1928 in the summer and winter (Lima and Castello 1995), for Trichiurus
lepturus from 3 to 30 in the winter and spring (Haimovici et al. 1996), and for
Loligo sanpaulensis from 1.2 to 3.5 in the fall and spring (Andrigueto and
Haimovici 1991). Over 50 species of cartilaginous fishes (mostly benthic
feeding sharks, angel sharks, rays, and skates) and over 150 species of bony
fishes (including 11 species of sciaenids) amount to over 80% of the total
biomass, making up the demersal fish community of the shelf. Biomass
estimates based on bottom-trawl surveys range from 43000 to 96 000 tons
for cartilaginous fishes (Vooren, Sect. 6.16) and from 137 000 to 340 000
tons for bony fishes (Haimovici et al. 1996) in the fall and winter/spring,
respectively.

Fig. 7.3.1. Distribution of surface water masses over the shelf and vertical profiles at
different latitudes duringthe winter (A) and summer (B). Coastal water (CW); South
Atlantic Central Water (SACW); Tropical Water (TW); Subantartic Water (SA W);
Antartic Intermediate Water (AIW); mixed Coastal Water with either SAW or TW
(MCW). Mean ranges of chlorophyll a concentrations are given for some water masses
The Continental Shelf and Slope 173

®
20°

SANTA MARTA GRANDE CAPE

29°

m rr---,----,
30°

31°

CHLOROPHYLL a
100 ( mg m- 3 )
150
200
D cw < 0 _5-2.0
250 @ SACW < 0.5-3.0
L--_ _- - ' _ LJ TW < 0.5

~k:]\~0 ~::
1_0-9_0

100 r' , ;."; MCW

150 "',-
200 (';,3'1 ~>.
250 "0,,, ., ':-
50° 49° 48° 47°

®
28°

SANTA MARTA GRANDE CAPE

29°

30°

31°

mD
. ·'·
50
100
150
200 _
. . : ,0:

D cw
CHLOROPHYLL a
(mg m- 3 )
< 0 _5-3 .5
SACW < 0 .5--5 .0
250 (2)
O TW < 0.5

o
m[J
50 _j o SAW
100
150
200
250 ® ';~
50° 49° 48° 47° 46°

Fig. 7.3.1.
174 J.P. Castello et al.: The Continental Shelf and Slope

Biomass maxima of bony fish are associated with cold « 1tc) SAW of the
coastal branch of the Malvinas Current over the central shelf and/or with
the oceanographic front (>1tc) formed at the western side of the STC
(Fig. 7.3.1). In general, demersal teleost diversity and species richness
decrease with depth. The much lower abundance of bony fish over the
upper slope than on the shelf (Haimovoci et al. 1994a) appears to be related
to a poor benthic fauna.
In summer, the biomass oflarger organisms is lower and is mostly com-
posed of juvenile sciaenids, mainly Trichiurus lepturus, Cynoscion guatucupa,
Umbrina canosai, angel sharks, and rays. During this season, the inner shelf
areas form important nursery grounds for young of the year Engraulis
anchoita, juveniles of commercially important bony fishes, and the
neonates of cartilaginous fishes such as Rhinobatos horkelii, Sphyrna
lewini, and myliobatid rays.

Pelagic Trophic Relations

The size structure of the pelagic community and the trophic interactions
are largely determined by the size of primary producer species (Ryther
1969; Pomeroy 1974). Large primary producers (microplankton) give rise
to shorter, less complex food webs with more efficient energy transfer,
whilst small-celled (pico-nanoplankton) based communities display long
and complex food webs with inherently inefficient energy transfer.
Large, single, centric diatoms or long chains of small-celled species are
associated with high chlorophyll a values over the shelf during the late
winter and spring. A considerable bacterial biomass, abundance of proto-
zooplankton, and intense grazing activity by flagellates and ciliates point
towards a complex microbial food web during periods of high phyto-
plankton biomass (Abreu, Sect. 6.6). The high phytoplankton biomass also
supports a rich and abundant pelagic shelf fauna. Copepods are the domi-
nant zooplankters (max. concentration of 2000 org. m-3 ), but cladoceran
and cold water euphausiaceans are also important. The dominance of
planktophagous pelagic fish, like Engraulis anchoita, emphasizes the
importance of short food chains during this period. Early larval stages (4-
12 mm SL) of Engraulis anchoita feed on starch grains probably related to
phytoplankton (Bursa 1968), as well as on nauplii, spores of fungi, and
tintinnids. With increasing larval size (> 12 mm SL), copepods assume a
more important part of the diet (Freire 1995). Calanoid copepods (Calanus
australis, Calanoides carinatus, Temora stylifera, Oncaea sp., Microsetella
sp.), euphausiacean (Euphausia similis) and hyperid amphipods comprise up
to 90% of the diet of adult Engraulis anchoita (Schwingel and Castello 1995).
Paralarvae and juveniles of the Argentinean squid Illex argentinus also feed
on copepods and euphausids (Santos 1992; Vidal 1994b). Other zooplankto-
The Continental Shelf and Slope 175

phagous species include the squid Loligo sanpaulensis, Anchoa marmll,

Trachurus lathami, and juveniles of Cynoscion guatucupa and Trichiurus
lepturus (Haimovici et al. 1996). Large individuals of Trichiurus lepturus,
adult Cynoscion guatucupa (Vieira 1990; Haimovici et al. 1996), Pagrus
pagrus (Capftoli and Haimovici 1993), Galeorhinus galeus, and the skates
Sympterigia acuta and Sympterigia bonapartei prey on small fishes like
Engraulis anchoita, Paralonchurus brasiliensis, and Symphurus jenynsi
(Queiroz 1986). Large icthyophagous pelagic predators include the bluefish
Pomatomus saltatrix (Haimovici and Krug 1992), angel sharks of the genus
Squatina, and Raja castelnaui, which feeds both on pelagic and demersal
fishes and squid. The feeding activity of other large predators, like the
porpoise Pontoporia blainvillei and the sea lion Otaria flavescens, is largely
restricted to coastal waters (Pinedo 1982). Over the outer shelf, pelagic
zooplanktophagous species are represented by Maurolicus muelleri,
myctophids, gonostomatids, juvenile Merluccius hubbsi, Brama spp., Loligo
sanpaulensis, Ornithoteuthis antillarum, and Illex arggntinus, which also
preys on small pelagic and mesopelagic fishes (Santos 1992). Large
ichthyophagous benthic fishes include Galeorhinus galeus, Genypterus
brasiliensis, and Polyprion americanus which feed on young Merluccius
hubbsi and Illex argentinus but also ingest macrocrustaceans. Large
ichthyophagous pelagic predators, like the migratory Xiphias gladius, prey
on Illex argentinus and other ommastrephids (Mello 1992), which also play
an important role in the diet of Thunnus obesus and Thunnus alalunga
(Santos and Haimovici, unpubl.), whilst Thunnus albacares preys on Brama
sp., squids, and hyperid amphipods (Vaske 1992; Fig. 7.3.2).
In contrast, principally small primary producer organisms (pico-
plankton) compose the lower phytoplankton biomass during the summer
(Odebrecht, unpubl.) when phytoplankton and bacterial carbon sources
tend to be of similar magnitude. The pelagic community is more diverse
than during the colder months and gives rise to complex feeding inter-
actions, which is characteristic of oceanic oligotrophic areas where most
zooplankton and all necton depend on secondary food sources (Cushing
1989; Legendre and Le Fevre 1995). Filter feeders, like the Cladocera species
Penilia avirostris and Thaliaceae, are abundant and consume bacteria and
picoplankton. The large biovolume of Thaliaceae is common in subtropical
seas because intensive grazing of large populations on small particles leads
to fast growth. Planktonic molluscs (pteropods) feed on small particles,
phytoplankters, protozoans, and zooplankters which adhere to their exter-
nal mucus produced in the mantle cavity or in the foot (Parsons et al.
1984). The Euphasiaceae present diverse feeding mechanisms, like filter-
feeding Qr "food basket" feeding. Practically all coelenterates are carnivores
and prey on plankters, small fishes, and/or components of the necton.
Zooplanktophagous fishes, like Peprilus paru, young Trichiurus lepturus,
Balistes capriscus, Cynoscion jamaicensis, and the benthophagous
 OUTER SHELF AND UPPER SLOPE
(100.000 m) .

NETlIMA BARBA
POGONIAS CRaMIS ~
\
MICROPOGONJAS FURNIERJ
MUGILPLATANUSD aL.ENGRAULIS ANCHOITA D 'It.TRACHURUS LA/IIATHI
{(
'''F''''.~.~.--~, PENA£US PAULEHSIS D • POMATOMUSSALTATRIX &SCOMBER JAPONICUS XIPHIIISSP.
.01ARIA FLAVESCENS &.,LLEX ARGENTINUS D THUNNUS ALALUNGII
.GALEORHINUS GALEUS • HWCOLENUS SP. SPHYRJo/ASP.
• SAROA SAROA • GEN'YPTERUS BRASILIENSIS THUNUS OBESUS
• SaUA TINA?? .COELHORYHCHUS SP.
• • PAGRUS PAGRUS
~ .MICROPOGONIAS FURNJERJ D. \
-UIIIBRINA CANOSAJ D

&ENGRAULISANCHOITA III .... MYCTOPHIDS

&ANCHOA MARINI &MAUROUCUS MULLER/
& LOL/GO SANPAULENSIS .... GONOSTOMATJDAE
&CYNOSeJON GUATUCUPA II &MERLUCCIUSHUBBSI U
&. MACRODON ANCYLODOH II .... UROPHYCIS M/STACEUS II
• RAJA CASTELNOUJ .... ,LLEK ARGENT/NUS U
.saUATlNA?? &ORNITOTEUTHIS ANTILLARUM
• ARTEMESIA LONGJNARIS ..... TRJCHIURUS LEPTURUS
• MYLIOBATJS SP. .CARCHARINUS SP. 132·sl
• SYMPTERlGJA SPP. .PRIONIlCE GLAUCII
• RAJA?? .AtOP/AS SPP•
• UMBR/NA CANOSAI U eHEPTANCHIAS SP.
• • POL YPRION AMERICANUS

'It.PEPR/LUS PARU
.... ENGRAULISANCHOITA iii
'..... TRJCHIURUS LEPTURUS
.CYNOSCION JAMAICENSIS
• SPHYRNA SP.
• CARCHARINUS SP•
• SARDA SARDA
.RHYNOBATOS SP.
• MYLIOBIITIS SP. - - -
.CTENOSCIAENA GRACJLCRJRHUS j \ \
• M/CROPOGONIAS FURNIERJ U
NETUMA BARBA BAtiSTES CAPRiSCUS
POGON/AS CROM'S
MICROPOGONIAS FURNIER/
MUGIL PLATANUS U III PRE-RECRUITS
PENAEUS PIIULENS/S . . .. LARVAE ISOUTHI
I I JUVENILES
I'lIIIDULTS
\j, C~V~ ... PLANKTOPHAGUS
• ICHTYOPHAGUS
• BENTHOPHAGUS

Fig. 7.3.2.
The Continental Shelf and Slope 177

Ctenosciaenea gracilirrhus arrive from the north, whilst other ichthyo-

phagous and/or benthic feeding species, including Pomatomus saltatrix,
adult Umbrina canosai, Cynoscion guatucupa, Micropogonias jurnieri,
Pagrus pagrus, and Mustelus schmitti migrate southward. Owing to the
southward migration of Galeorhinus, necton abundance decreases in slope
regions, though dense schools of the skipjack tuna (Katsuwonus pelamis),
which feed heavily on macro-zooplankton like Euphausia similis, the
mesopelagic fish Maurolicus muelleri, and squids (Vilela 1990), arrive from
the north (Fig. 7.3.2; Vilela and Castello 1991). Between November and
May, large flocks of sea birds, like Puffinus gravis, Sterna hirundo,
Procellaria aequinoctialis conspicillata, and Diomedea chlororhynchos
(Chiaradia 1991), are associated with schools of Katsuwonus pelamis over
the outer shelf. The rejected small bony fish from bottom-trawl fishing
activities (Haimovici and Palacios 1981; Haimovici and Mendon<;:a 1996)
also provide an important food source for sea birds over the inner shelf
(Vooren and Fernandes 1989).

Pelagic-Benthic Interactions

Pelagic-benthic interactions are either associated with the life-cycle of

species with both planktonic and benthic stages or are related to their
trophic behaviour. Trophic interactions may be bi-directional since either
benthic organisms may benefit from sedimentation of particles from the
water column or demersal and nectonic organisms may feed on the benthic
fauna. Also, diel migrations favour an increase of pelagic-benthic interac-
tions. For example, Illex argentinus feeds during the night at the surface
but rests during the day near the bottom, where it is preyed upon by
Polyprion americanus (Santos 1992) and probably by Galeorhinus galeus.
Benthic food chains appear to be especially important over southern shelf
regions, which coincides with intense bottom-trawl fishing (Haimovici et
al. 1989a). Here, the coastal branch of the Malvinas Current supports high
primary productivity, and areas with sandy and muddy bottoms favour an
abundant invertebrate fauna of polychaetes. Among the benthophagous
species, juvenile and adult Umbrina canosai (Haimovici et al. 1989b),
Micropogonias jurnieri, Myliobatis spp., Sympterigia spp., Netuma spp.,
Mustelus schmitti, Raja spp., and Urophycis brasiliensis are dominant over
the shelf. Depending on the size of predators, the shrimps Artemesia longi-
naris and Pleoticus muelleri, the crabs Portunis spinicarpus and Callinectes
sp., the bivalve mollusc Solen tehuelches, and a variety of polychaetes,

Fig. 7.3.2. Main feeding habits and seasonal movements of principal necton species of
inner and outer shelf and upper slope regions
178 J.P. Castello et al.: The Continental Shelf and Slope

isopods, amphipods, and cumaceans are the principal prey in shallow

coastal waters (Queiroz 1986). Benthic feeding fish species of the outer shelf
include Urophycis mystaceus, Coelorhincus marin ii, and Helicolenus lahillei.
Although many species feed exclusively on the epi- and in faunal organ-
isms, some important species, including adults of Cynoscion guatucupa
(Viera 1990) and Pagrus pagrus (Capitoli and Haimovici 1993), prey on
both benthic and pelagic organisms.
8 Estuarine-Marine Interactions
P.C. Abreu and J.P. Castello

The Patos Lagoon estuary and the adjacent marine system are abiotically
and biologically linked. In general, abiotic estuarine-coast interactions are
exemplified by eutrophication of coastal waters, owing to nutrient and
organic matter runoff which locally enhances primary and secondary
production (Mann and Lazier 1991). Biological interactions are represented
by the transport of neritic invertebrate and fish species which take
advantage of the sheltered conditions and food availability in estuaries as
nursery and feeding grounds (Knox 1986).

Physical Interactions
The topography of the southern Brazilian continental shelf controls the
transport of oceanic water masses, but the dynamics of coastal waters as
well as those of the Patos Lagoon follow short- and long-term
meteorological cycles. The frequent passage of polar fronts augments the
velocity of coastal currents and directs large amounts of water onshore
(Fig. 8.1a). These conditions not only lead to high concentrations of
phytoplankton cells (especially Asterionellopsis glacialis) in the surf zone
which provide an important food source for the intertidal benthic fauna
(Odebrecht et al. 1995a), they occasionally also cause accumulations of
toxic dinoflagellates which result in the death of benthic organisms and fish
(Odebrecht et al. 1995b). Since the tidal amplitudes are small, the exchange
of water between the Patos Lagoon and coastal regions is also influenced by
the passage of polar fronts (Costa et al. 1988a; Moller et al. 1991). SE winds
force saltwater into the lagoon, which may penetrate up to 200 km
upstream during drought periods (Niencheski et al. 1988). The input of
coastal water normally causes sediment resuspension and consequently a
release of inorganic nutrients like phosphate (Kantin and Baumgarten
1982; Fig. 8.1b). In general, intrusion of coastal water increases
phytoplankton diversity in the estuary. If this process persists for a
prolonged period, the residence time of estuarine water increases, and
owing to reduced export of phytoplankton cells, the accumulation of
phytoplankton biomass in the estuary is considerable. A subsequent
change to northerly winds favours the discharge of estuarine waters, and
large amounts of particulate organic carbon are exported as phytoplankton
cells (Abreu 1987). Since much of the exported phytoplankton biomass is

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
180 P.C. Abreu and J.P. Castello

composed of species like Skeletonema costatum, which enter the estuary

during saltwater intrusion, part of the eutrophication process of coastal
waters appears to depend on the addition of extra nitrate to the water
column and on the prior seeding of the estuary with euryhaline neritic cells
during saltwater input (Abreu et aI. 1995a; Fig. 8.1c).
An increase in precipitation in southern Brazil is a consequence of EI
Nino-Southern Oscillation (ENSO) events, and thus long-term phyto-
plankton variability of the region might be related to this large-scale climatic
phenomenon. Chlorophyll a variability in the Patos Lagoon estuary, with
maximum values during years with high precipitation, shows a relationship
between ENSO and the eutrophication process of coastal regions (Abreu et
aI., unpubI.). The freshwater flow from the Patos Lagoon, and especially
from the La Plata River, is the main source of silicate and ammonia to the

A PHYSICAL FORCES B NUTRIENTS

EST~;.;;AR..;.;y.,...>
_______O_CEAN

HY""O'OG~ .~
c PHYTOPLANKTON D DETRITUS

9
ESTUARY

EXPORT ¢::J
OCEAN ESTUARY
-------------------
OCEAN

SEDIMENTATION

EURYHALINE CELL
SEEDING

E ZOOPLANKTON F CRUSTACEANS & FISH

ESTUARY OCEAN ESTUARY OCEAN

ADULTS

Fig. 8.1. Diagrammatic representation of the main estuarine-marine interactions

Estuarine-Marine Interactions 181

coastal euphotic zone, and thus high rainfall in the watershed should cause
an increase of nutrients and primary production. A direct relation between
ENSO events and a rise of chlorophyll a levels over the southern Brazilian
continental shelf has been observed by Ciotti et al. (1995). Additionally, the
large freshwater outflow increases the stability of the water column in
coastal shelf regions, which causes higher spring chlorophyll a concentra-
tions (Ciotti et al. 1995) than in years with reduced outflow. Similarly,
water column stabilization favours the occurrence of large patches of
anchovy Engraulis anchoita larvae, owing to probably both higher food
availability and physical mechanisms which inhibit the dispersal of eggs
and larvae, thus increasing their survival chance (Lima and Castellg 1995).
During years of extreme precipitation, outfiowing estuarine waters form
a plume which may extend 30-50 km along the coast (Hartmann et al.
1980), and high levels of primary and secondary production at the conflu-
ence with coastal water masses are likely to influence the biological
dynamics of the coastal region decisively (Abreu et al. 1995b). Together
with natural and dredge removal of sediments, large amounts of plant
detritus originated from marshes, seagrasses, and algae and animal debris
are exported from the estuary (Castello 1985). These materials may form
dense acoustic scattering layers (>100 mg r l suspended matter) at the
bottom of the estuary inlet (Calliari 1980) and in the mid-water of the shelf.
The sedimentation of detrital particles and of phytoplankton cells (Ciotti et
al. 1995) and the presence of a rich benthic inner shelf community
(Borzone and Griep 1991; Absalao 1991) suggest the significance of deposi-
tion processes in the coastal region (Fig. 8.1d).

Transport of Organisms
The meteorological processes which control water exchange between the
Patos Lagoon and shelf regions are also responsible for the transport of
zooplankton and fish and crustacean larvae into the estuary. Especially
during the summer, the estuarine zooplankton community is dominated by
neritic species which enter with salt water intrusion, whilst limnic species
prevail during years with high precipitation (Montu 1980; Rieger and
D'Incao 1991; Fig. 8.1e). Although fish eggs and larvae occur during the
entire year in the estuary, peaks are associated with the spring and
summer. Probably owing to elevated temperatures in the late spring and
early summer, the spawning activities of most fish species are transferred
to nearshore waters, which coincides with the period of maximum salt
water intrusion into the estuary. The combined effect of these processes
warrants the transport of eggs and larvae into the estuary, where food
availability and sheltered conditions ensure growth (Muelbert and Weiss
1991; Ibagy and Sinque 1995; Fig. S.lf).
182 P.C. Abreu and J.P. Castello: Estuarine-Marine Interactions

The pink shrimp (Peaneus paulensis) is among the most prominent

species which enter the estuary. The larvae of northern stocks (29 0 S) are
transported by coastal currents to shelf regions near the Patos Lagoon inlet.
The species is truly estuarine-dependent and migrates with varying success
into the estuary in the spring and summer, trying to overcome the
freshwater outflow. A negative relationship between the winter/fall
precipitation and shrimp production during the following summer clearly
indicates the impact freshwater runoff has on larval penetration (Castello
and Moller 1978). After migration into the estuary, larvae settle in shallow
embayments where they undergo metamorphosis and develop into adults
(Fig. 8.1f).
Fish migration is perhaps one of the most common and best known
manifestations of estuarine-coastal interactions, where migrations might be
seen as an evolutionary solution to an optimization of population
abundance. The Patos Lagoon estuary provides nursery, protection, and
feeding grounds for several migratory fish species, like the marine catfish
(Netuma barba) and the white croaker (Micropogonias furnieri). After
spawning in coastal water during the late spring and summer, eggs and
larvae of Micropogonias furnieri are transported into estuarine shoals
where they are retained, owing to reduced water circulation and the
presence of seagrass beds (Fig. 8.1f). However, Micropogonias furnieri is
not an estuarine-dependent species as considered by Chao et al. (1985),
since development and growth also occur in more haline adjacent coastal
waters (Vieira and Castello, Sect. 4.13). Migration of fish in and out of the
estuary exemplifies the import or export of biomass, and local fisheries
benefit from this process. A clear understanding of this process by
scientists, resource managers, and fishermen may often be limited by the
unreliable quantifications of the biomasses involved, though issues of
immediate financial profit may also adversely influence the decision-
making policy. These conditions have largely inhibited the development of
an integrated management of estuarine and coastal Micropogonias furnieri
stocks which, together with Netuma barba stocks, eventually collapsed,
leading to severe damage in the estuarine fisheries (Reis 1992).
In conclusion, the eutrophication processes produced by the land and
sea, and the utilization of estuarine nursery and feeding grounds by
organisms reflect two-way interactions which tightly couple the estuary to
adjacent coastal waters and which significantly contribute to the region's
considerable biological production.
9 Fisheries

M. Haimovici, J.P. Castello, and C.M. Vooren

For more than a century the Patos Lagoon estuary (von Ihering 1896) and
the coastal sea have exemplified the fishing potential in the warm-temper-
ate southwestern Atlantic (Yesaki 1973). Today about 6000 artisanal (Reis
1992) and 3000 industrial fishermen are temporarily or permanently
involved in fishing activities. Despite the economic and social importance
fisheries have historically assumed in this region, reliable records on fish-
ing methods and landing statistics are lacking before 1945. Until then, arti-
sanal trammel, gill, and c~annel net fishing with smallwooden sail and row
boats was largely restricted to the Patos Lagoon and estuary. The event of
modern means of storage and transport and the introduction of a large
number of synthetic fibre nets and motor-powered boats (< 10 m, 10-24 HP),
which permitted artisanal trawl fishing, eventually led to the depletion of
estuarine stocks. During the 1980s, artisanal fisheries quickly extended into
shallow coastal waters where larger (12-15 m, 90-120 HP) wooden boats
with up to 20 tons capacity used gill-nets, and occasionally purse seines and
hooks. In the estuary and coastal waters artisanal fisheries follow a clearly
defined seasonal pattern (Reis et al. 1994). The black drum (Pogonias
cromis) and white croacker (Micropogonias jurnieri), which actively feed in
brackish waters, and the catfish (Netuma barba), which uses the estuary for
reproduction, are fished in the spring, the pink shrimp (Penaeus paulensis)
is caught in the summer and fall, the mullet (Mugil platanus) in the fall,
and the bluefish (Pomatomus saltatrix), castanha (Umbrina canosai), and
weakfish (Cynoscion guatucupa) in the winter. The mean annual artisanal
fishery catch totals 21 500 t (Table 9.1; 58% demersal fishes, 15% pelagic
fishes, 14% estuarine shrimp and crab, 12% freshwater fish) but, owing to
often incomplete landing statistics, catch figures may be underestimated by
as much as 25%. Industrial fisheries began in 1947 for demersal species
(Yesaki and Bager 1975), and landings were mostly comprised of sciaenids,
the red porgy Pagrus pagrus (Haimovici et al. 1989), and to a lesser extent
elasmobranchs (Vooren et al. 1990). Pelagic fishery commenced in 1962
(Yesaki and Bager 1975) and intensified after 1977 when foreign vessels
were hired and operated out of Rio Grande port (Zavala-Camin and Antero
da Silva 1991; Antero da Silva 1994).

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
184 M. Haimovici, J.P. Castello, and C.M. Vooren

.-----------------------, • SLOPE AND OCEANIC

a WATERS
...~ 60
0'80

..
~ 40
o SHELF WATERS

0
t:.. 20
1-....-...,.....-.,..........---.---.---.--.-...--..--.--.--..---.--.-.,....-.---......., 0 ESTUARINE AND
COASTAL WATERS

, . . . . - - - - - - - - -- - - - - - - - , . OCEANIC PELAGIC FISHES

Iilll COASTAL PELAGIC FISHES

o CRUSTACEANS

EJ DEMERSALELASMOBRANCHS

..........---,.---.-....,........-...,.-,......-,---,---,--.-...-.,...-,---,.---.-....,....-,--.j D OE M ERSAL TE LEOS TS

......0'80
• HOOKS

Ii]] TRAPS
..
,.... 60
PURSE SEINES
~ 40
o Ii] GILL NETS
t:.. 20
o TRAWL NETS
1976 1979 1982 1985 1988 1991 1994

Fig. 9.1. Landed catches of estuarine and marine fisheries in the southwestern Atlantic
between 29° and 35° S based on depth range (a), type of fishery resource (b), and fishing
methods (c)

Seasonal changes in the composition and abundance of the necton

(Vooren et al. 1990; Haimovici et al. 1994a; Lima and Castello 1995;
Haimovici et al. 1996), and consequently also of fisheries, are induced by
the alternate influence of cold subantarctic water and warm tropical water
over the shelf. Shelf fisheries are more intense in the winter when a variety
of commercially important species (castanha, weakfish, red porgy, bluefish,
and the school shark Galeorhinus galeus immigrate from the south. Off-
shore fishing occurs all year raung; in the winter with long lines for south-
ward migrating tunas and sailfishes and in the summer with live bait fish-
ery for skipjack tuna (Katsuwonus pelamis). Most of the industrial fishery
catches occur over the continental shelf (71 %) and only 4% in oceanic and
slope regions (Fig. 9.1a). About 80% of catches is landed in the fishing port
of Rio Grande. Between 1975 and 1994 average annual catches approxi-
mated 74 000 metric tons but oscillated between a minima of 52 200 t in
1989 and a maxima of91 800 tin 1993 (Haimovici et al. 1989; Vooren et al.
1990; IBAMA 1993, 1995). Demersal teleosts (71%) dominate among the
Fisheries 185

catches, followed by pelagic shelf teleosts (10%), demersal elasmobranchs

(8%), crustaceans (5.3%), and pelagic oceanic fish (4%; Fig. 9.1b). Trawl-net
(61 %) and gill-net (19%) methods account for most of the catch, whilst purse
seines (10%), traps (4.5%), and hooks (40/0) are less important (Fig. 9.1c).

Industrial Fishing Methods

In the early 1980s, when some stocks (Pogonias cromis, Netuma barba, Pagrus
pagrus) collapsed and others (the royal weakfish Macrodon ancylodon,
Umbrina canosai) showed signs of overfishing (Haimovici 1988a, b; Haimovici
et al. 1989; Reis et al. 1994), demersal fishing diversified into double-rigg
trawling, bottom gill-nets, bottom long-lines, and trap methods (Barcellos et al.
1991; Lima and Branco 1991) to exploit new resources.
Pair- and otter-trawling methods were introduced in 1947, but the trawler
fleet first grew rapidly during the 1970s as a result of fiscal incentives (SUDEPE
1974; Yesaki and Bager 1975). Both methods use double sheet codends of 50-
70 mm mesh size between opposite knots, which drastically reduces the chance
of small fish escaping. Fishing vessels, which may employ either method
depending on the season, are 22-35 m long and are powered by 250-650 HP
engines. Since they lack refrigeration, the catch is stocked for up to 15 days in
holds between layers of crushed ice (Haimovici et al. 1989). Although pair- and
otter-trawling methods have changed little over the past few decades, fishing
has become more efficient, owing to satellite navigation and more efficient use
of echo-sounders. Pair-trawling, followed by otter-trawling, are the principal
fishing methods for demersal sciaenids, like Umbrina canosai, Cynoscion
guatucupa, Macrodon ancylodon, and Micropogonias junieri, which represent
about 80% of total mean annual trawl landings. Otter-trawling over the inner
shelf (20-100 m) landed 20 287 t in 1976, but this amount gradually decreased to
2708 t in the early 1990s. However, over the outer shelf (80-200 m) a few larger
trawlers (49-52 m) landed an average of 4632 tin 1993 and 1994. Outer shelf
otter-trawl catches include sharks and rays (31%) and sciaenids (20%), which
are already intensely exploited, and the cutlassfish Trichiurus lepturus (24%),
which is discarded by the inner shelf trawl fishery (Table 9.1).
Double-rigg trawling was first used in 1985 from 20-24 m long wooden
vessels with 250-350 HP engines (Barcellos et al. 1991). Twin nets were
employed for both shrimp fishing (Artemesia longinaris, Pleoticus muelleri) and
demersal teleosts and elasmobranchs, though the length of the ground rope and
the mesh size of the codend differed. Shrimps are fished in shallow waters in the
spring and summer, whilst Paralichthys patagonicus is fished at 20-80 m depth
between the fall and spring and Squatina spp. over the outer shelf at up to
140 m depth. Double-rigg trawlers landed about 30% of the total elasmobranch
catches in 1989-1990, but their contribution decreased to 5% in 1992 (IBAMA
1995). Annual landings of double-rigg trawling totalled 3728 t after 1989
(IBAMA 1995; Haimovici and Mendon<;a, unpubl.).
186 M. Haimovici, J.P. Castello, and C.M. Vooren

Since trawl fishery is a non-selective method, small specimens of

commercially valuable species and species without value are generally
discarded from the catch. The estimated total annual discard ranges from
17 000 to 25 000 t. Onboard rejection by pair and otter-trawling of small
demersal fishes and elasmobranchs (Haimovici and Palacios 1981; Haimovici
and Habiaga 1982) may vary between 26 and 46%, though a larger (90 mm
between opposite knots) mesh size would remedy this situation (Vooren
1983). Onboard discard of small elasmobranchs and teleosts during double-
rigg trawling for fish exceeds 50% of the total catch. Discard during shrimp
fishings may reach 20% (Haimovici and Mendon<;a 1996) which is
significantly lower than the rejection rate reported for subtropical and
tropical regions (Alverson et al. 1994).
Purse seine fishery for pelagic fishes has occurred in shallow shelf waters
ofless than 50 m depth since 1962. Seines (600-800 m length, 70-80 m height)
are operated by vessels of 20-24 m length and 250-450 HP (Krug and
Haimovici 1991). Mullets are caught in the fall and bluefish is the principal
catch in the winter, though sometimes the jack mackerel (Trachurus
lathamis), mackerel (Scomber japonicus), menhaden (Brevoortia pectinata),
and occasionally also demersal fish like white croaker and castanha represent
an important portion of the catch which averages (1990-1994) 4261 t annually
(Table 9.1).
Trap fishing methods at depths greater than 400 m were employed for the
red crab (Chaceon notialis) by hired Japanese vessels in 198411985, and the
catch totalled 1471 t over a 9-month period after which fishing was
abandoned (Lima and Branco 1991). Trawl fishing for Pagrus pagrus dates
back to 1973 (Yesaki and Barcellos 1974), and overfishing made exploitation
uneconomical at the beginning of the 1980s (Haimovici et al. 1989). Between
1987 and 1992 Pagrus pagrus was caught by trap fishing methods, similar to
those used in Argentina (Barcellos et al. 1991), over beachrock outcrops (70-
80 m) and gravel bottoms (110-160 m). The average annual catch was 106 t
and was composed 83% of Pagrus pagrus (Table 9.1).
Hand-line fishery, aimed at the wreckfish (Polyprion americaus), was
developed over hard bottoms of the upper slope in 1973 and was gradually
replaced by vertical long-lines, bottom gill-nets, and bottom long-lines in
recent years. Landing statistics of the commercially valuable wreckfish are
incomplete but amount to several hundred tons annually (Table 9.1). Indi-
vidual fishing grounds are abandoried after intense fishing for short periods
and recover only after several years.
Bottom gill-nets replaced bottom long-lines, which were used for demersal
shark fishing through 1989 (Barcellos et al. 1991). The gill-nets are pulled by
18-30 m long trawlers with 185 to 350 HP engines, and they may exceed
several kilometers in length. Gill-netting is used at all depths (>50 m) of the
continental shelf and has quickly become the main fishing method (IBAMA
1995). Between 1990 and 1994 mean annual catches equalled 5355 t
Fisheries 187

(Table 9.1). The increase of the total elasmobranch landings after 1990 clearly
reflects the growth of the bottom gill-net fishery.
Japanese fishing vessels initiated surface long-line fishery for tunas,
principally Thunnus alalunga, Thunnus obesus, and Thunnus albacares,
and the swordfish Xiphias gladius over the southern shelf in 1961, followed
by Brazilian long-liners in 1969 (Zavala-Camin and Tomas 1990; Zavala-
Camin and Antero da Silva 1991), which dominated this fishery after 1977.
Even so, a number (2-11) oflarge foreign vessels continue to fish for tunas
and swordfish and land their catch in Rio Grande (Mello et al. 1993).
During the 1977-1991 period, the average yield for this fishery was
728 kg/thousand hooks (Antero da Silva 1994). Pole and live bait fishery of
the skipjack tuna (Katsuwonus pelamis) was introduced in 1979 (Castello
and Habiaga 1988) and, owing to high initial yields, rapidly attracted a
growing number (up to 97 in 1982) of 24-28 m long vessels (Matsuura
1982b). Four larger foreign vessels were hired in 1991. Almost 93% of the
catch is composed of Katsuwonus pelamis, and annual landings average
3240 t (Table 9.1).

Table 9.1. Mean annual landed catches (tons) by fishing method in the southwestern
Atlantic between 29° and 34° S from 1975 to 1994

Fishing method 1975-1979 1980-1984 1985-1989 1990-1994

Artisanal fishery 24926 20548 20617 20091
Pair-trawl 32056 33951 32051 29137
Otter-trawl (inner shelf) 15237 9160 10055 2708
Otter-trawl (outer shelf) 1993-1994 4362
Double-rigg trawl 3216
Bottom gill-net Since 1989 435 5355
Fish trap 1988-1992 106
Hand line and vertical 231 82 321 168
long-line
Purse seine 7105 1949 1833 4990
Offshore pelagic long-line Since 1977 1295 1943 2260 3504
Pole and live bait fishery Since 1991 3240

Demersal Teleost Fisheries

Demersal teleost catches (1975-1994) amount to 71% of all landings, and

70% are comprised of the sciaenids Micropogonias jurnieri, Umbrina canosai,
Cynoscion guatucupa, and Macrodon ancylodon. Annual landings do
oscillate, but catch per unit effort (CPUE) for the first three species has
decreased steadily (Fig. 9.2).
188 M. Haimovici, J.P. Castello, and C.M. Vooren

Regional Micropogonias furnieri stocks, represented by a spawning

group near the Patos Lagoon inlet (32 0 S) and another further south (340 S;
Haimovici and Umpierre 1996), have been heavily exploited by artisanal
and industrial gill-net fisheries for more than two decades (Reis 1992),
which has resulted in a drastic decrease in the density of juveniles in coastal
waters between 1980 and 1990 (Ruffino and Castello 1992a). Although
annual catches between 8000 and 17 000 t (mean 14 071 t) appear to be
stable, the declining CPUE (Fig. 9.2) and changes in the age structure
(Schwingel and Castello 1990; Haimovici and Umpierre 1996) indicate that
the spawning stocks are overfished.
Umbrina canosai, a small species of low natural mortality (Haimovici
and Reis 1984; Haimovici 1988b), is heavily fished along the southern
Brazilian coast and to a lesser extent in Uruguay and Argentina (Fig. 9.2).
This species represented the largest (19 000 t) part of landings during the
1970s, but gradually decreased to 10 000 t in recent years. Similarly, the
biomass diminished from 75 000 t in 1976 to about 37 000 in 1983
(Haimovici 1988b). However, in contrast to MicrOpogonias furnieri, the age
structure of Umbrina canosai has not changed in recent years. Therefore, in
spite of overfishing, actual landings should persist if exploitation levels do
not rise.
During the colder months adults of a single Cynoscion guatucupa stock
migrate from fishing grounds in Uruguay and Argentina to more northern
Brazilian shelf regions. Landings in southern Brazil, which represent about
30% of the total catch of the three countries, oscillate between 4000 t (1975
and 1988) and 14 000 t (1986 and 1994). These variations reflect either
interannual differences in recruitment or availability to Brazilian trawlers,
because the CPUE displays similar oscillations (Fig. 9.2).
A single stock of Macrodon ancylodon was found between 29 0 and 340 S
(Yamaguti 1979) and has been exploited since the 1950s, with a maximum
catch of 10 617 t in 1977. Landings gradually decreased to 3000 t in 1992 but
showed an increase over the last 2 years (Fig. 9.2). Although Macrodon
ancylodon may reach a maximum age of 12 years (Yamaguti 1967), since
1977 age classes above 6 years have been absent (Martins Juras 1980;
Haimovici 1988a), which suggests that sustainable annual yields are below
4000 t. The slow growing and long-living Pagrus pagrus was overexploited
by trawl fishing during the 1970s (Yesaki and Barcellos 1974; Haimovici et
al. 1989), and stocks collapsed after a few years without showing signs of
recuperation (Table 9.2). Landings of the flatfishes Paralichthys spp., the
sea robin (Prionotus punctatus), and Urophycis brasiliensis averaged almost
3500 t annually over the last 5 years (Table 9.2). Landings of flatfishes alone
reached a maximum of 2157 tin 1990 but decreased more recently. Since
Pr~ionotus punctatus and Urophycis brasiliensis are not abundant over the
southern shelf (Haimovici et al. 1996), catches of these species probably
cannot be sustained.
 '-r:I
20 MICROPOGONIAS RJRNlERI 3.0 UMBRtNA CANOSAI
24 3.0
(WHITE CROAKER, (CASTANHA,
.,,~2.4 20 2.5 :;;.
If
2.0 '"
1.5 n
)(
'11
C
~ ::~i~ il'~ l~il]llr!l~!i~j'!;i!l~i;!.f:::
U)
Z
1.0 m
-=i
g 0
fI)
z
en
16 CYNOSCION GUATUCUPA 3.0 12 MACRODON ANCYLODON c
2.0 ~
"~o
Z
(WEAKFISH, IROYAL WEAKFISH)
10 L-
:j 12 2.4 1.8 ~
..J 8 en
1.8 1.2 ~
t! I
~ 1.2 0.8

0.1 0.4

1976 1979 1982 1985 1988 1911 1994 1911 11119 1982 1985 1988 1991 1994

o TOTAL LANDED CATCH o PAIR TRAWL • OTTER TRAWL

Fig. 9.2. Total landed catches and catch per unit effort (CPUE) of otter- and pair-trawls of the most abundant
demersal teleosts between 29° and 35° S

.....
00
\C
190 M. Haimovici, J.P. Castello, and C.M. Vooren

2000 1.0

0.8
~ 1500 o
"tI
o c:
m
t:. 0.6 -=l
f/)
(!) oz
z
C 1000
z .. 0.4 e!
~
:3 \
..:]

500
0.2

1983 1985 1987 1989 1991

Fig. 9.3. Landings of guitl!-r fish Rhinobatos horkelii at the PO'rt of Rio Grande (solid line)
and mean annual CPUE of the otter-trawl fleet (dashed line)

The stocks of demersal teleosts, like Pogonias cromis, the catfishes

Netuma barba and Netuma planifrons, and Micropogonias jurnieri, which
use the Patos Lagoon estuary during part of their life-cycle, have been
heavily exploited by the artisanal gill-net fishery during the 1970s and
collapsed after 1980 (Table 9.2; Haimovici et al. 1989; Reis et al. 1994). The
recuperation of Pogonias cromis and Netuma stocks will be difficult, owing
to their high vulnerability to gill-net fishing, slow growth, and late sexual
maturation. As a result, only mullet and pink shrimp actually represent
economically viable fisheries in the estuary.

Demersal Elasmobranch Fisheries

About 10% of the total sea fish landings (1975-1994) in Rio Grande are
comprised of elasmobranchs. Until 1988 almost 80% of all catches were by
bottom-trawl fishing (Vooren et aI. 1990) but in recent years most catches
are being made by bottom gill-nets. Landing statistics are merely sub-
divided into sharks, angel sharks, guitar fish, and rays (Table 9.2).
Fisheries 191

Table 9.2. Mean annual landings (1975-1994) of the main fishery resources fished in the
southwestern Atlantic between 29° and 34° S

Mean annual landings (tons)

1975- 1980- 1985- 1990-
1979 1984 1989 1994
Estuary and shelf pelagic species
White croaker Micropogonias fornieri 14308 14904 12364 14709
Weakfesh C)'noscion guatucupa 6439 7377 9572 8785
Royal weakfish Macrodon anC)'lodon 7941 5865 3659 3966
Castanha Umbrina canosai 16900 14877 11732 9629
Catfish Netumaspp. 3983 1536 452 615
Black drum Pogonias cromis 1044 359 246 63
Hake Merluccius hubbsi 760 101 200 129
Flatfish Paralychthis spp. 424 417 1486 1363
Red porgy Pagurs pagrus 1419 327 294 238
Searobin Prionotus punctatus 12 71 486 988
Abr6tea Uroph)'cis brasiliensis 241 337 665 1186
Cutlassfish Trichiurus lepturus 75 Z9 63 441
Demersal sharks' 1637 2486 3754 3748
Angel shark Squatina spp. 947 1485 2607 2183
Guitar fish Rinobathos horketi 1010 1253 901 460
Rays and skates various species 116 461 718 746
Pink shrimp Peneus paulensis 4143 1830 4354 2627
b
Marine shrimps 1331 1148

Estuary and shelf demersal species

Menhaden Brevoortia pectinata 1549 343 646 1324
Bluefish Pomatomus saltatrix 4290 2290 2456 3521
Mullet Mugilplatanus 2081 1709 1486 1524
Mackerel Scomberjaponicus 969
Jack mackerel Trachurus lathami 1555

Demersal upper slope species

Wreckfish Polyprion americanus 152 68 145 176
Red crab Chaceon notialis

Offshore pelagic species

Bigeye tuna Thunus obesus 312 419 640 500
Albacore Thunus alalunga 269 446 494 1075
Yellowfin tuna Thunus albcarens 212 336 325 684
Swordfish Xiphias gladius 122 285 350 601
Skipjack tuna Katsuwonus pelamis 2402
Pelagic sharks' 182 481 382 547
a Mostly Galeorhinus galeus and Mustelus schmitti.

b Pleoticus muelleri and Artemesia longinaris.

, Isurus oxyrinchus, Prionace glauca, Sphrena spp. among others.

192 M. Haimovici, J.P. Castello, and C.M. Vooren

The division "sharks" consists mostly of the school shark (Galeorhinus

galeus), with the narrownose smooth-hound (Mustelus schmitti) in second
place. The latter species was discarded until about 1985 (Haimovici and
Palacios 1981), but was subsequently landed in increasing quantities. Both
species migrate from the Uruguayan and Argentinian shelf to Brazilian
waters in the winter. Annual landings in Rio Grande rose gradually from
1414 t (1975) to 2482 t (1984), then increased sharply to about 3500 t (1985-
1987), followed by a drop to 1247 t (1989). The decrease after 1987 reflects
the reduced abundance of Galeorhinus galeus and possibly also of Mustelus
schmitti. Owing to increased fishing efforts by bottom gill-netting, total
shark landings increased to 2791 tin 1992. The otter-trawl CPUE showed a
similar pattern, rising to a peak of about 5 t/trip in 1989, then dropping to
1.5 t/trip in 1992. The rise in CPUE until 1989 reflects increased targeting of
shark and a decrease in discarding.
Catches of angel sharks consist mainly of Squatina guggenheim, which
inhabits the inner shelf at depths down to 80 m, and Squatina occulta which
is abundant on the 'outer shelf and the slope doWn to 350 m. Squatina
guggenheim is a resident species, but the migratory status of Squatina
occulta is not clear. Annual landings of angel shark increased from 820 t
(1973) to a peak of 2442 t (1988) and then dropped sharply to 1031 t (1990).
Similarly, otter-trawl CPUE of angel shark reached a maximum of 6.2 t/trip
in 1988, followed by a decline to 0.7 t/trip in 1992, which reflects a major
decrease of the abundance of both species of angel shark, the resident
Squatina guggenheim probably being the most affected. However, while
abundance decreased, landings increased to 1761 t in 1992, owing to
increased fishing efforts by bottom gill-netting.
Adult females of the guitar fish (Rhinobatos horkelii) concentrate
between December and February in shallow inshore waters to breed.
During this period they are particularly vulnerable to intensive pair-trawl
and beach seine fisheries. During the remainder of the year, adults of both
sexes disperse over the inner shelf where catching is done by otter-trawling.
A continuous decline of otter-trawl CPUE from 0.76 t in 1984 to 0.08 t in
1992 provides a measure of decreasing abundance (Fig. 9.3). Similarly,
annual landings of this species decreased from a peak of 1927 t in 1984 to
an all-time low of 178 tin 1992.

Pelagic Teleost Fisheries

The fishery for pelagic teleosts, like Pomatomus saltatrix, the cutlassfish
(Trichiurus lepturus), the anchovy (Engraulis anchoita), and Trachurus
lathami, is little developed and represents only about 10% of total landings.
Pomatomus saltatrix is a fast growing species which migrates from
Argentina to southern Brazilian shelf waters. The species concentrates in
surface waters warmed above 15° C temperature (Krug and Haimovici 1989;
Fisheries 193

Haimovici and Krug 1992). In the winter mainly age classes between 2 and
5 years are fished (Krug and Haimovici 1991). Mean annual landings total
3528 t (1970-1992), with an exceptional peak of 12 126 t in 1971. Since climatic
conditions regulate the abundance and migration cycles of Pomatomus
saltatrix, the fishing potential of this species is highly unpredictable
(Haimovici and Krug 1996). Trichiurus lepturus is fairly abundant, and the
spring biomass may exceed 30 000 t (Haimovici et al. 1996). Since the
commercial value of this species drastically declines when kept between ice
layers, rather than being deep frozen onboard as is done by some hired
Korean trawlers, the stock is underexploited. The small-sized Engraulis
anchoita is the main pelagic species in southern Brazilian shelf waters.
Since the species has no market value, it is unexploited, though the winter
biomass may attain over one million tons. Trachurus lathami and Scomber
japonicus may be caught by purse seines between 30 and 80 m depth in the
winter and therefore may represent a potential resource for midwater
trawling. The irregular occurrence of Scomber japonic~s is probably related
to the intensity of the subantartic water influence. -
Large pelagic fishes are caught along the thermal front of the western
margin of the Subtropical Convergence and landed in Rio Grande (Mello et
al. 1993; Antero da Silva 1994). Landings (1977-1989) contain the yellowfin
tuna (Thunnus albacares; 23%), the bigeye tuna (Thunnus obesus; 21.5%),
the albacore tuna (Thunnus alalunga; 18%), the swordfish (Xiphias gladius;
16%), various pelagic sharks (i.e. Prionace glauca, Sphyrna zygaena,
Sphyrna lewini; 18.5%), and billfishes (i.e. Tetrapturus albidus, Makaira
nigricans; 3%; Mello et al. 1993). Although stock assessments are unreliable
and catches fluctuate, the tuna fishery shows potential for expansion. The
skipjack tuna (Katsuwonus pelamis) is caught during the summer and fall
with pole and live-bait in waters between 80 and 1000 m depth and surface
temperatures between 20 and 26°C (Castello and Habiaga 1988). Since the
exploitation rates of skipjack tuna are moderate (Jablonski and Matsuura
1985; Vilela and Castello 1993), the catch could be increased. The
availability of live bait appears to be the main limiting factor for the
expansion of this fishery (Table 9.2).

Invertebrate Fisheries
Crustacean catches, mostly shrimp, blue crab, and red crab, amount to
approximately 6% of the total landings (1975-1994). Subadults of the pink
shrimp (Penaeus paulensis) migrate from northern coastal waters into the
nursery grounds of the Patos Lagoon estuary and coastal lagoons in
Urugul\Y, where the species is caught in the summer and fall (Valentin et al.
1991). Annual artisanal shrimp fishery landings change considerably as a
consequence of variable recruitment conditions (Castello and Moller 1977;
D'Incao 1991) and growth. Despite predictions that estuarine shrimp fish-
194 M. Haimovici, J.P. Castello, and C.M. Vooren

eries might collapse, owing to excessive catching of juveniles (Valentin et

al. 1991), average annual landings between 1988 and 1994 were close to the
20-year annual mean of 3106 t. This leads to the conclusion that environ-
mental variables rather than fishing pressure are the determinants for
shrimp fishery catches. Juveniles and adults of the shrimps Pleoticus muelleri
and Artemesia longinaris occur in coastal waters (Haran et al. 1992), and
exploitation of Artemesia longinaris stocks appeared to have reached
saturation in 1989 (Ruffino and Castello 1992b). However, the variability of
catches during subsequent years coincided with a reduced catch of largely
migratory demersal species, and thus oscillation in the abundance of Artemesia
longinaris may depend on interannual variations in oceanographic
conditions (Table 9.2).
The crab fishery is hardly developed. Small-scale trawl-net fishery for
the blue crab Callinectes sapidus occurs regularly in the Patos Lagoon estu-
ary. In 1984 the deep water red crab Chaceon notialis was experimentally
fished over the continental slope (34° S; Lima and Branco 1991), but catches
per trap-day decreased sharply in less than 6 mon.ths. Since this species has
a wide low-density distribution, a biomass estimate (Delury method) of
2183 tof the unexploited stock in the fishing area (3000 km2 ) may be
conservative. The squids Loligo sanpaulensis and Illex argentinus
frequently occur over the shelf between 30° and 34° S (Haimovici and Perez
1991a). Despite spring biomass estimates of 3500 t for Loligo sanpaulensis
(Andriguetto and Haimovici 1991), landings of cephalopods are virtually
non -existent.

Fisheries Interactions

Fishing activities interact with both sea birds and marine mammals.
Between November and May flocks of several types of sea birds (Puffinus
gravis, Sterna hirundo, Procellaria aequinoctialis conspicillata, Diomedea
chlororhynchos) are reliable indicators of schools of the skipjack tuna
(Katsuwonus pelamis) over the mid and outer shelf (Chiaradia 1991). A
variety of sea birds (Procellaria a. aequinoctialis, Fulmarus glacialoides,
Diomedea melanophoris) die on the hooks of long-line tuna fisheries along
the western margin of the Subtropical Convergence, and the annual death
toll may exceed 2650 birds (Vaske 1991). Fishery interactions may be the
main cause for the decline of Diomedea exulans populations in the South
Atlantic Ocean (Croxall and Prince 1990). Discard from trawl fisheries is
composed of a large number of either small, floating teleosts or fast-sink-
ing, small sharks and skates, and represents an important additional food
source for sea birds like subantartic albatrosses and petrels (Vooren and
Fernandes 1989) and the sea lion Otaria flavescens. Dead stranded speci-
mens of Otaria flavescens are commonly found on southern Brazilian
beaches. A large number have broken skulls or bullet holes inflicted by
Fisheries 195

fishermen, while others may have died from entrapment in nets (Rosas et
al. 1994). Gill-nets are the principal cause of small cetacean mortality in the
southwestern Atlantic (Pinedo 1994a). The species most affected is the
small dolphin Pontoporias blainvillei, but the bottle-nose dolphin (Tursiops
truncatus) is also occasionally entangled and killed by gill-nets near the
Patos Lagoon inlet (Pinedo 1994b).

Fishery Perspectives

Total annual catches in the southwestern Atlantic (38°-34°S), including a

conservative estimate of 25% discard, range from 70 000 to 120 000 t, inter-
annual differences being partly due to environmental factors. Changes of
landing statistics with time are principally a result of the establishment of
new fisheries (i.e. bottom gill-net fisheries for elasmobranchs and double-
rigg trawling for flatfishes and shrimp) as the older stocks became overex-
ploited. Today, most commercially important estuariQe, coastal, and shelf
stocks are either fully or overexploited, and landings· are expected to
decrease in coming years. Overfished stocks of demersal finfishes include
Netuma spp., Pogonias cromis, and Pagrus pagrus. The abundance of
Rhinobathos horkelii, Galeorhinus galeus, and other shark species have
declined up to 90% in recent years, probably due to recruitment overfish-
ing since these species are low-fecundity viviparous fishes. The assessment
of the status of stocks of less abundant shark species and of many other
species, which are simply grouped as rays and skates in the landing statis-
tics, is at present impossible. Furthermore, about two-thirds of recent
demersal catches have concentrated on migratory stocks (i.e. Micropogonias
furnieri, Umbrina canosai, Cynoscion guatucupa, Mustelus schmitti) which
are shared by Uruguayan and Argentinian fisheries. Estuarine and marine
shrimp, menhaden, bluefish, and mullet catches are highly variable. These
stocks do not clearly confirm overfishing but appear to lack potential for
expansion. In addition, the growth potential of under- or unexploited
stocks seems to be limited. Only Engraulis anchoita and Trachurus lathami
stocks might have some fishery potential, and catches of offshore tunids
may eventually increase.
Demersal teleost and elasmobranch stocks are affected as a whole, and
thus their abundance and diversity are at risk under intensive and uncon-
trolled fishing. For example, if present trends continue, the populations of
Rhinobatos horkelii and Squatina guggenheim will become extinct. The
same may well be true for an unknown number of other species, whose
decline remains unnoticed because they are not listed in the landing statis-
tics. Although proposals to reduce estuarine and all demersal fishing efforts
and to decrease onboard discard (Haimovici and Palacios 1981; Vooren
1983; Haimovici et al. 1989; IBAMA 1993, 1995) have provided an adequate
legal basis, neither fishing efforts nor discard of undersized fish has dimin-
196 M. Haimovici, J.P. Castello, and C.M. Vooren: Fisheries

ished. Unquestionably, future fishery management strategies must not only

involve both the fishermen and the fishing industry in creating effective
legislation, but also encourage international management of shared stocks.
10 Natural and Human Impact
U. Seeliger and C.S.B. Costa

Early accounts of man's interference with the natural resources of coastal

regions in the warm-temperate southwestern Atlantic date from the last
century (von Ihering 1885). In particular, the Patos Lagoon and estuary
have been subject to multiple human impacts during the past few decades.
In the near future though, a gradual degradation of the ozone layer and
increasing sea levels are likely to pose major ecological problems for the
entire region.

Freshwater Flow

The water quality of the Patos Lagoon estuary and coastal nearshore
environments is ultimately related to natural freshwater runoff. Because of
the assumption that runoff is available for expanding agricultural and
industrial activities and domestic needs without any limitation, the
importance of freshwater flow into the estuary is often overlooked. The
annual estimated streamflow through the Patos Lagoon inlet averaged 109 km3
between 1965 and 1975 (Castello and Moller 1978) but varied significantly
within (CV=80%) and between (CV=29%) years. During the drought years
1968-1969 (IPAGRO 1977), the water demand for irrigated rice cultivation
(EPAGRI 1992) and for the population of the watershed approximated 6%
of the expected (75 km3 ) annual runoff. Since then, both the population
and the area of rice cultivation have increased by more than 37% and
120%, respectively (IBGE 1993), and thus today as much as 13% of natural
runoff may be diverted during drought periods (Fig. 10.1). Furthermore,
the construction of locks (1977) in the Sao Gon~alo channel to prevent
salinization of Mirim Lagoon waters used for irrigation has considerably
reduced freshwater flow into the estuary. The growing encroachment on
freshwater discharge into the estuary modifies seasonal variations in flow
rates that may be essential for flushing and maintaining the balance of
salinity and nutrients in the estuary. Many of the commercially important
marine fish and crustaceans in the southwestern Atlantic have similar life-
cycles ~nd visit the estuary periodically to benefit from low salinity waters,
physical protection, and food sources for either growth or reproduction.
Therefore, any further intentions to control or divert freshwater flow must
await the results of studies to establish minimum flow requirements for the
estuary.

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
198 U. Seeliger and C.S.B. Costa

175

e-
~
150

tt
o
125
z _..--- - -- MEAN 109 km'- - - - - -
~
IX: 100

75 -----~--------~--__, 0
4 ~
w
en
8 ~
IX:
12 ~
<t
16 ~

1970 1975 1980 1985 1990 1995

Fig. 10.1. Annual total freshwater runoff between 1965 and 1975 and estimated water use
during drought years

Nutrient Enrichment and Eutrophication

Freshwater runoff not only influences estuarine salinity regimes but also
affects the quality of surface waters through inputs of nutrients and toxic
materials. Most urban centres around the Patos Lagoon (approx.
3.5 million inhabitants) either lack or have inadequate sewage treatment
facilities which tend to overflow during heavy rains and discharge raw
sewage directly into the lagoon or estuary. Nutrients are also added by the
major tributaries from large-scale agricultural activities; phosphate (2-3 fJM)
and nitrogen (>40 11M) concentrations in particular tend to be seasonally
elevated in the receiving waters of the Patos Lagoon (Vilas Boas 1990). In
spite of a substantial reduction during transport, estuarine nutrient levels
are also occasionally elevated (Kantin 1983; Niencheski and Windom 1994),
owing to remobilization of bottom' sediments in the lower estuary (Abreu et
al. 1992), local discharge of industrial and domestic effluents, and
agricultural runoff through the Sao Gon~alo channel. Excess nutrient loads
in the estuary cause eutrophication with the development of blooms and
cbanges in phytoplankton composition. Blooms of the potentially toxic
bluegreen alga Microcystis aeruginosa (Yunes et al. 1994) form in limnic
regions of the lagoon and, during years with high spring tributary
discharge, are transported into the estuary (Odebrecht et al. 1987; Bergesch
Natural and Human Impact 199

1990). During the summer, estuarine embayments with prolonged water

residence time favour blooms of other species (Persich 1993), which
contributes to the decline of the submerged macrophyte biomass in the
estuary, owing to attenuation of light penetration and epiphytic algae
loading on Ruppia leaves (Cafruni 1983; Costa and Seeliger 1989).

Hazardous Materials

Since large extensions of coastline north and south of the inlet are devoid
of industrial discharge, trace metals are added to the estuary either through
freshwater runoff from the Patos Lagoon or through local discharge of
effluents (Seeliger et al. 1988). Average metal concentrations in estuarine
waters with high suspension loads correspond to natural background levels
(Seeliger and Knak 1982a; Niencheski et al. 1994) reported for estuarine
systems elsewhere (Windom 1975), though dissolved copper concentra-
tions are occasionally slightly higher (Seeliger and Knak1982b). Sporadic
increases of suspended copper and lead concentrations in the estuary may
reflect metal input from industrial effluents and mining activities to the
upper limnic part of the Patos Lagoon (Baumgarten 1987; Vilas Boas 1990).
In general, metal concentrations correlate significantly with suspended
matter loads, and estuarine metal gradients are not well-defined (Seeliger
and Cordazzo 1982) because physico-chemical processes add particulate
metals through resuspension of estuarine bottom sediments by inflowing
seawater, wind, and waves (Niencheski et al. 1994). The biological effects of
trace metal pollution have not yet been documented for the Patos Lagoon
estuary, probably because estuarine organisms tend to display elevated
metal tolerance, and the concentrations are close to background levels.
However, in shoals with reduced circulation, prolonged water residence
time, and added domestic and industrial effluents even, low pollution levels
appear to induce changes in the zooplankton community structure and
anatomical anomalies (Montu and Gloeden 1982). Although data on the
effect of agrotoxins on estuarine organisms are still lacking, the excessive
application of pesticides over vast extensions of agricultural lands around
the Patos Lagoon is likely to contribute to the estuary through runoff with
elevated concentrations. Some intertidal estuarine species (i.e. Chasmagnathus
granulata) which decisively influence energy transfer in submersed and
emersed habitats (D'Incao et al. 1992) appear to be highly susceptible to
pesticide concentrations in warmer waters during the summer (Monserrat
and Bianchini 1995). Superficial coastal nearshore waters are especially
prone to pollution by different fractions of hydrocarbons discharged by
vessels during the washing of tanks. These pollutants are likely to interfere
with sea and shore birds and may occasionally cause high mortality among
migrant penguins.
200 U. Seeliger and e.S.B. Costa

Sedimentation, Dredge, and Fill

The introduction of large amounts of suspended sediments from the Patos

Lagoon and the presence of extensive shallow estuarine shoals with Ruppia
beds and fringing marshes provide an ideal setting for natural deposition
processes in the estuary. Over the last two centuries the water area of the
lower estuary has decreased by approximately 11 %, which emphasizes the
depositional character of this environment. Natural erosion commonly
occurs along the southern margins of the estuary and the inlet, with an
estimated annual sediment removal of about 2 500 000 m3 to coastal near-
shore waters (Calliari 1980; Fig. 10.2.). Losses oflower estuarine water areas
not only result from natural sedimentation but also from man-mediated
processes. The expansion of Rio Grande port between 1909 and 1914
generated 8 776 000 m 3 of dredge material which served for the establishment
of islands and was deposited along lower estuary margins. The construction
of jetties in 1917 has probably intensified nat~ral sedimentation at the
northern side of the inlet. Furthermore, significant natural sediment
deposition in the lower estuary has historically interfered with navigation,
and dredging activities have concentrated in the navigation channel and the
inlet since 1833. Modern vessels with increasing draft require deeper
waterways, and today approximately 30 000 m 3 of channel and berth material
needs to be dredged monthly from the main shipping channel. These human
activities may have solved immediate problems but also resulted in often
unexpected and long-term changes by modifying depth, circulation, and
sediment deposition patterns (Calliari 1980). Since the accumulative effects of
frequent dredging are comparable to natural geological processes (Nichols
and Howard-Strobel 1991), the export of fine sediments to the adjacent coast
is likely to have increased (Villwock and Martins 1972; Hartmann et al. 1980;
Calliari and Abreu 1984; Borzone and Griep 1991; Calliari and Fachin 1993).
The filling of intertidal and shallow water flats in the lower estuary for
port, residential, and more recently industrial development has decreased or
destroyed vital seagrass and tidal marsh habitats. Filling along lower
estuarine margins and over and around small islands has destroyed as much
as 10% of the total salt marsh area. Seagrasses are especially susceptible to
filling, and burial by dredged sediments results in their immediate
eradication. Only if plants are lightly covered and the root-rhizome systems
are not damaged may regrowth occur through the sediments. Since dredged
sediments are readily resuspended, they represent a source of excess
suspended matter for long periods of time. The gradually increasing water
turbidity reduces seagrass production and depth distribution. Therefore, the
extension of natural beds and their valuable functions as habitat and shelter
decrease. Although the disposal of dredged material is clearly affecting
estuarine resources, no disposal program has yet been implemented for the
Patos Lagoon estuary.
Natural and Human Impact 201

6'

ATLANTIC OCEAN

10'
r
16' 10'

Fig. 10.2. Areas of the lower Patos Lagoon estuary which have suffered physical changes
since 1776

Burning, Cutting, and Grazing

Several natural and man-induced events interact with salt marshes and
their habitat function. Especially during drought periods, extensive upper
salt marsh areas with large quantities of dead vegetative biomass become
vulnerable to accidental burning. Although fire may cause high mortality of
invertebrates and particularly of nesting birds, it opens the canopy of the
dominant Spartina densiflora and Juncus effusus vegetation to invasion by
secondary species and recycles large amounts of nutrients otherwise locked
up in dead and live plant biomass. Rapid vegetative regrowth is stimulated
by the nutrient enrichment, and the dominant species generally recover
their aerial biomass densities within less than a year. Cutting of Spartina
alterniflora and Scirpus maritimus to protect recently established summer
vegetable crops against desiccation and to supply additional nutrients is a
common practice, which represents only a small-scale disturbance (2-20 m\
In contrast, a large-scale impact of grazing livestock on marginal marshes
has existed since early colonization because the small tidal range, gentle
202 U. Seeliger and C.S.B. Costa

marsh topography, and reduced net of tidal creeks facilitate access by cattle
and horses. Although detailed studies are lacking, differences in plant
diversity and habitat physiography between grazed and ungrazed sites are
apparent, and a total loss of habitat occurs in frequently grazed marshes.

Afforestion

In spite of the virtual absence of human habitations along the ocean shores
between 30° and 34° S, anthropogenic interference is becoming increasingly
apparent. Effective binding of foredune sand by the grass Panicum
racemosum has slowed dune transgression over inland marshes for
centuries. Extensive grazing of Panicum by cattle is now commonplace and
reduces sand accretion on foredunes by as much as 70 cm year- 1 (Costa et
al. 1991), leaving fore dunes susceptible to erosion by onshore winds, with
the formation of blow-outs and massive sand transport over the inland
marshes. Because of their natural stability, these marshlands are preferred
sites of afforestion with Pinus and Eucalyptus in excess of 100 km2• The
seasonally flooded marshes directly control the watertable of foredunes
and associated vegetation. Afforestation has increasingly upset this natural
equilibrium by artificially draining the marshes before planting and
through the high evapotranspiration rates of already established dense
plantations. The combined effects of grazing and hydrodynamic dis-
turbances have reduced the sand stability of foredunes and increased
transgression rates (Seeliger 1992a).

Sea Level Rise and Ozone Layer Depletion

Evidence suggests there is global warming due to the accumulation of

atmospheric CO 2 and other gases (Schneider 1989) with a direct conse-
quence of rising sea levels. Although the absence of conservative data does
not allow us to trace secular trends of sea-level changes along coasts in the
warm-temperate southwestern Atlantic (Schmack 1993), evidence of
erosive processes along the Patos Lagoon margins and the coast (Villwock
1994) implies an estimated rate of sea-level rise of between 2 and 3 mm
yea{l. If these rates are confirmed for the flat and wave-dominated coast
(Costa et al. 1988a; Costa and Seeliger 1988a), storm overwash will more
frequently impact areas further inland than in the past (Bernardi et al.
1987; Calliari and Klein 1993; Costa et al. 1995), coastal marshes and wet-
lands are likely to migrate further inland (Costa and Davy 1992), and
salinization of the upper Patos Lagoon estuary will increase.
The gradual degradation of the ozone layer over Antartic regions is
causing an increase in the transmission of ultraviolet B (UV -B) radiation
and poses a threat to cold and warm-temperate latitudes in the southern
hemisphere. Ozone-poor stratospheric air is spreading to northern
Natural and Human Impact 203

Argentina, Uruguay, and southern Brazil and has already led to a 13-20%
decrease of ozone at 30° S during the spring (Kirchhoff et al. 1994; Santee et
al. 1995). The potentially deleterious effect of increased UV-B exposure on
marine plankton and benthic communities (Haeder and Worrest 1991;
Karentz 1991) is likely to modify their distribution and abundance in the
southwestern Atlantic with concomitant changes in productivity and
trophic dynamics.
The coastal region of the warm-temperate southwestern Atlantic repre-
sents an enormous ecological and economic resource which has satisfied
social and economic interests without management for more than 200 years.
Because natural thresholds are being approached, vital habitats are being
reduced or destroyed and fisheries will continue to decline if modern
management strategies are not applied.
11 Management Considerations
M.L. Asmus and P.R.A. Tagliani

Among coastal regions of the southwestern Atlantic, the Patos Lagoon and
surrounding lands, particularly the estuary and adjacent waters, stand out
because of their ecological (zone of biological production and maintenance
of biodiversity) and socio-economic significance (port, industrial, agricul-
tural, fishing, tourist activities). Historically, socio-economic demands tend
to collide with ecological preservation, and increasing anthropogenic
activities are likely to jeopardize the health of the Patos Lagoon, estuary,
and coast (Seeliger and Cos.ta, Sect. 10). Therefore, the need to understand,
plan, and manage these systems has become an issue of priority. During the
1980s a National Plan for Coastal Management was implemented by the
Brazilian government, aiming to subsidize macrozonation, planning,
management, and monitoring efforts. Thematic mapping (1:100 000) was
expected to offer recommendations for the use of the coastal zone and the
inner continental shelf, considering legal restraints, social needs, and the
heterogeneity and potential of the region.
Initial management efforts were descriptive, but interpretative, simula-
tive, and predictive phases soon followed (Asmus et al. 1991; Assis 1994).
The coastal plain, with Patos Lagoon and estuary, reflects a considerable
environmental heterogeneity, expressed by a mosaic of environmental
units. Environmental units are characterized by small but distinct differ-
ences in, for example, topography, morphology, soil, vegetation, and
climate (Cendrero and Diaz de Teran 1987). They can be defined by
analytical or synthetic methods which correspond to descriptive and inter-
pretative phases (Dyer and Vinogradov 1990), respectively. The analytical
method, represented by the elaboration of thematic maps of the coastal
plain, described geomorphological and hydrophysical aspects, fauna and
flora, land use, and socio-economic attributes and furnished recommenda-
tions for use (Asmus et al. 1991).
The synthetic method, based on the interpretation of satellite imagery
and ground truth verification, offers an integrated view of the different
natural and anthropic components. A total of 33 environmental units have
been identified in both emersed littoral, transitional, and lagoonal systems
around the Patos Lagoon estuary and submersed areas within it. Units of
emersed areas are principally comprised of marshes, lagoonal terraces,
semifixed dunes, and marine deposits. The extensive mono cultures of rice
plantations and large-scale afforestation with Pinus sp. are the main

U. Seeliger et al.
Subtropical Convergence Environments
© Springer-Verlag 1997
 N
o
51°45' 0\

PATOS LAGOON

ATLANTIC OCEAN

s::
i-'
>
~
• AREA OF PRESERVATION §'"
p..
• AREA OF CONSERVATION "d
~
o AREA OF DEVELOPMENT i>
""'l

32° 25'
cap;.
8.

Fig. 11.1. Areas of preservation, conservation, and development of the Patos Lagoon estuary and adjacent coast
Management Considerations 207

causes of their perturbation. These agricultural activities, owing to an

increased equitability of the landscape pattern, induce low spatial hetero-
geneity (Tagliani 1995). Environmental units of submersed estuarine areas
are dominated by protected shoals and channel zones. The ideal use of
different environmental units is determined by their function within the
coastal ecosystem (Tagliani 1995). A matrix of environmental functions can
be constructed, based on the analysis of environmental characteristics, like
biological production, support capacity, resilience, and socio-economic
aspects (Groot 1986). This matrix proved to be an excellent tool to define
and group environmental units into areas of preservation (not to be
exploited), areas of conservation (limited and strictly controlled exploita-
tion), and areas of development (Asmus et al. 1989, 1991; Fig. 11.1). Since
the present use and exploitation by man of the estuary and the land around
it conflict with management recommendations for some areas, mitigations
will be necessary.

I-
~
Il.
~
0::
W
I-

~::I:
(/)
w
0::
LL
ESTUARINE SHOALS DEEPER OPEN WATERS

NUT
DOC
poe

/ COASTAL WATERS

Fig. 11.2. Main modelled environments in the Patos Lagoon estuary. NUT, N nutrients;
poe particulate organic carbon; DOC dissolved organic carbon; L light; T temperature;
Ti tide; W wind; R rain
208 M.L. Asmus and P.R.A. Tagliani: Management Considerations

More recently, ecological modelling, representing simulative and

predictive phases of management, has focused on dynamic aspects of the
estuary and adjacent areas in order to characterize the dominant ecologi-
cal, economic, and social processes of the region. Modelling of estuarine
environments (Fig. 11.2) aims to determine dominant components and
processes, the natural and anthropogenic control of processes, susceptibil-
ity of components/processes to management, and management recom-
mendations (Faccini et al. 1995). All models emphasize the importance of
metereology (wind, precipitation) and hydrodynamics (circulation) on
critical ecological processes in the estuary (Bonilha and Asmus 1994;
Rabelo da Cunha 1994; Teixeira da Silva 1995). Modelling efforts nave
identified gaps in our understanding of the estuary, which is an essential
step for the improved comprehension of processes and indispensable for
present and future management (Bonilha 1996).
The classification of national waters by the Brazilian government, based
on use and quality, provided some legal support to administrate the use·of
waters in the Patos Lagoon estuary. However, environmental laws lack the
desired detail to back urgently required management steps at municipal,
state, and federal levels. Consequently, present and probably future
management efforts of the Patos Lagoon estuary are hampered by technical
and legal problems.
Appendix
Phylogenetic list of the 3193 coastal and marine taxa reported for the warm-temperate
southwestern Atlantic since the beginning of this century.

KINGDOM PROKARYOTAE L. martensiana Menegh. ex Gom.

PHYL UM CYANOBACTERIA L. perelegans Lemm.
(CYANOPHYTA) Microcoleus tenerrimus Gom.
CLASS CYANOPHYCEAE M. chthonoplastes Thuret ex Gom.
Order Chroococcales Oscillaloria amphibia C.Ag. ex Gom.
Family Chroococcaceae O. animalis C.Ag. ex Gom.
Aphanothece castagnei (Breb.) Rab. O. earlei Gardner
A. stagnina (Spreng.) A.Braun O. obscura Bruhl & Biswas
Chroococcus dispersus (Keissl.) Lemm. O. limnetica Lemm.
C. limneticus Lemm. O. limosa C.Ag. ex Gom.
C. membraninus (Menegh.) Nag. O. meslini Fremy
C. minor Klitz. O. nigroviridis Thw. ex Gom.
C. pallidus Nag. O. okeni C.Ag. ex Gom.
C. turgidus (Klitz.) Nag. O. ornata Klitz. ex Gom.
C. turicensis (Nag.) Hansg. O. princeps Vaucher ex Gom.
Gloeocapsa crepidinum Thuret O. proboscidae Gom.
Gleothece confluens Nag. O. subbrevis Schmidle
Gomphosphaeria aponina Klitz. Phormidium cf. arcuatum Skuja
G. lacustris Chodat P. crouani Gom.
Merismopedia convoluta Breb. P.fragile (Menegh.) Gom.
M. elegans A.Braun in Klitz. P. usterii Schmidle
M. glauca (Ehr.) Nag. Schizothrix Jamyi Gom.
M. punctata Meyen S. muelleri Nag. ex Gom.
M. tenuissima Lemm. Spirulina labyrinthiformis (Menegh.)
Microcystis aeruginosa Klitz. Gom.
M. ichthyoblabe Klitz. S. subsalsa Oersted ex Gom.
M. robusta (Clark) Nygard S. subtilissima Klitz.
Trichodesmium erythraeum (Ehr.) ex
Order Pleurocapsales Gom.
Family Hyellaceae
Xenococcus cUuiophorae (Ti.) Setch. & Gard. Family Nostocaceae
Anabaena aphanizomenoides Forti.
Family Tubiellaceae .A. doliolum Bharadw.
A. circinalis Klitz.
Order Nostocales A. fertilissima Rao
Johannesbaptista pellucida (Dickie) A. cf. iyengarii Bharadw.
Taylor & Drouet A. oscillarioides Bory ex Bornet & Flah.
A. variabilis Klitz.
Family Oscillatoriaceae
Lyngbya confervoides C.Ag. ex Gom. Aphanizomenon gracile Lemm.
L. aestuarii Liebm. ex Gom. Cylindrospermopsis raciborskii (Wolos.)
L. epiphytica Hieron. Seen. & Subba Raju
L. lutea (C.Ag.) Gom. Cylindrospermum sp.
210 Appendix

Nodularia harveyana (Thwaites) Thuret Family Bangiaceae

emend. Nordin & Stein Goniotrichum alsiddii (Zanard.) Howe
Nastoc commune Vaucher ex Born. & Erythrocladia subintegra Rosenv.
Flah. Erythrotrichia carnea (Dill.) J.Ag.
N. paludosum Klitz. Bangia atropurpurea (Roth) C.Ag.
Richelia intracellularis Schm. in Ostenf. Porphyra pujalsii Coli & Oliveira F.
&Schm. P.rizzini Coli & Oliveira F.

Family Rivulariaceae CLASS FLORIDEOPHYCIDAE

Calothrix parietina Thur ex Born. & Order Nemalionales
Flah. Family Acrochaetiaceae
Kylinia crassipes (Borg.) Kylin
Order Stigonematales Acrochaetium flexuosum Vickers
Family Nostochopsidaceae A. globosum Borg.
Mastigocoleus testarum Lagerh.
Order Gelidiales
Family Stigonemataceae Family Gelidiaceae
Stigonema mesentericum Geitler Gelidium floridianum Taylor
G. crinale (Turn.) Lamour.
KINGDOM PROTOCTIST A Pteroclaaia-capillacea (Gmelin) Bornet-
PHYLUM RHIZOPODA & Thuret
CLASS LOBOSEA
Order Arcellinida Order Cryptonemiales
Family Arcellidae Family Hildebrandiaceae
Arcella vulgaris Ehr. Hildebrandia prototypus Nardo

Family Centropyxidae Family Corallinaceae

Centropyxis aculeata discoides Penard Amphiroa beauvoisii Lamour.
C. (=Cyclopyxis) arenata (Cushman) Corallina officinalis L.
C. marsupiformis (Wall.) Jania rubens (L.) Lamour.

Family Difflugiidae Family Grateloupiaceae

Difflugia acuminata Ehr. Grateloupia cuneifolia J.Ag.
D. amphora Leidy G. filicina (Wulf.) J.Ag.
D. capreolata Penard
D. corona Wall. Order Gigartinales
D. curvicaulis Penard Family Hypneaceae
D. elegans Penard Hypnea musciformis (Wulf.) Lamour.
D. globularis Wall.
D.lageniformis Wall. Family Phyllophoraceae
D. lebes Penard Gymnogongrus griffithsiae (Turn.)
D. lobostoma Leidy Martius
D. mitriformis Wall.
D. pyriformis Perty Family Gigartinaceae
D. urceolata Carter Gigartina teedii (Roth) Lamour.
Pontigulasia compressa Rhumbler
Order Rhodymeniales
Order Gromida Family Rhodymeniaceae
Family Euglyphidae Rhodymenia pseudopalmata (Lamour.)
Buglypha alveolata Dujard. Silva

PHYLUM RHODOPHYTA Order Ceramiales

CLASS BANGIOPHYCIDAE Family Ceramiaceae
Order Bangiales Ceramium brasiliensis Joly
Appendix 211

C. brevizonatum Pet. & Borg. A. palmata Stein

Callithamnion uruguayense Taylor A. retangulata Kof.
Dinophysis acuminata Clap. & Lach.
Family Delesseriaceae D. argus (Stein) Abe
Cryptopleura ramosa (Hudson) Kylin ex D. caudata Sav.-Kent
Newton D. cuneus (Schutt) Abe
D. diegensis Kof.
Family Rhodomelaceae D. doryphora (Stein) Abe
Polysiphonia subtilissima Mont. D. exigua Kof. & Skogs.
P. tepida Hollenb. D. hastata Stein
Poiysiphonia sp. D. hindmarchii (Murr. & Whit.) Bal.
Bryocladia thyrsigera (J.Ag.) Schmitz D. fortii Pav.
Carradoria virgata (C.Ag.) Kylin D. mitra (SchUtt) Abe
D. odiosa (Pav.) TaL & Skogs.
PHYLUM CROMOPHYTA D. okamurai Kof. & Skogs.
CLASSCRYPTOPHYCEAE D. ovum Schutt
Order Cryptomonadales D. parvula (SchUtt) Bal.
Family Cryptomonadaceae D. paulsenii (Schi.) Bal.
Rhodomonas Karst. D. porodictyum (Stein) Abe
R. salina (Wisl.) Hill & Wether. D. punctata Jorg.
D. pusilla Jorg.
CLASSDYNOPHYCEAE D. rapa (Stein) Abe
Order Ebriales D. recurva Kof. & Skogs.
Family Ebriaceae D. rotundata Clap. & Lach.
Ebria tripartita (Schm.) Lemm. D. schroederi Pav.
Hermesinum adriaticum Zach. D. schuettii Murr. & Whit.
D. tripos Gourr.
Order Oxyrrhinales Histioneis dolon Murr. & Whit.
Family Oxyrrhinaceae H. cymbalaria Stein
Oxyrrhis marina Dujard. H. highleyi Murr. & Whit.
H. hyalina Kof. & Mich.
Order Prorocentrales H. milneri Murr. & Whit.
Family Prorocentraceae H. para Murr. & Whit.
Prorocentrum aporum (Schi.) Dodge Ornithocercus cf. heteroporus Kof.
P. balticum (Lohm.) Loebl. o. quadratus Schutt
P. compressum (Bail.) Abe ex Dodge o. magnificus (Stein) Kof. & Skogs.
P. dentatum Stein O. splendidus SchUtt
P. gracile SchUtt O. steinii Stein
P.lima (Ehr.) Dodge o. thumii (Schm.) Kof. & Skogs.
P. micans Ehr.
P. maximum (Gourr.) Schi. Family Oxyphysaceae
P. minimum (Pav.) Schi. Oxyphysis oxytoxoides Kof.
P. rostra tum Stein
P. scutellum Schrod. Order Actiniscales
Family Actiniscaceae
Order Dinophysiales Achradina pulchra Lohm.
Family Citharistaceae Actiniscus pentasterias (Ehr.) Ehr.
Citharistes apsteinii SchUtt A. acutum Lohm.
A. crassum Lohm.
Family Dinophysiaceae A. oceanicum Lohm.
Amphisolenia bidentata Schrod.
A. globifera Stein Order Dinococcales
A. lemmermanni Kof. Family Thoracosphaeracea
A. palaeotheroides Kof. Thoracosphaera sp.
212 Appendix

Order Pyrocystales e. belone Cl.

Family Pyrocystaceae e. bucephalum (Cl.) Cl.
Dissodinium pseudolunula Swift e. candelabrum (Ehr.) Stein
pyrocystis jUsiformis (Wyv.Thoms.) MUIT. e. carriense (Gourr.)
P. lunula (Schutt) SchUtt e. concilians Jorg.
P. noctiluca Murr. ex Haeckel e. contortum (Gourr.) Cl.
P. obtusa Pav. e. contortum var. longinum (Kars.) Sour.
P. robusta Kof. e. contrarium (Gourr.) Pav.
P. spinifera Lohm. e. declinatum (Kars.) Jorg.
e. declinatum f. brachiatum Jorg.
Order Brachydiniales e. declinatum f. normale Jorg.
Family Brachydiniaceae e. declinatum var. majus Jorg.
Asterodinium sp. e. deflexum (Kof.) Jorg.
Brachidinium sp. e. dens Ostenf. & Schmidt
e. euarctuatum Jorg.
Order Noctilucales e. extensum (Gourr.) Cl.
Family Kofoidiniaceae e. falcatum (Kof.) Jorg.
Cymbodinium sp. C. furca (Ehr.) Clap. & Lach.
e. furca var. eugrammum (Ehr.) Jorg.
Family Noctilucaceae e. furca var, h ircus (Schrod.)
Noctiluca scintillans (Macartney) Kof. & e. fusus (Ehr.) Dujard.
Swezy e. fusus var. seta (Ehr.) Jorg.
e. gibberum Gourr.
Family Protodiniferaceae e. gibberum f. dispar (Pouch.) Jorg.
Pronoctiluca sp. e. gibberum f. subaequale Jorg.
e. hexacanthum Gourr.
Order Gymnodiniales e. hexacanthum f. contortum (Lemm.)
Family Gymnodiniaceae Jorg.
Amphidinium britannicum (C.Herdm.) e. horridum f. denticulatum (Jorg.)
Lebour Bohm
Cochlodinium sp. e. horridum (Cl.) Gran horridum
Gymnodinium catenatum Grah. e. horridum moUe (Kof.) Jorg.
G. exechegloutum Norris e. incisum Kars.
G. splendens Lebour e. inflatum (Kof.) Jorg.
Gyrodinium cf. aureolum Hulb. e. karstenii Pav.
G. fusiforme Kof. & Swezy e. karstenii f. robustum (Kars.) Jorg.
G. spira Ie (Bergh) Kof. & Swezy e. kofoidii Jorg.
Plectodinium nucleovelatum Biech. e. limulus Gourr.
Pseliodinium sp. e. lineatum (Ehr.) Cl.
Torodinium robustum Kof. & Swezy e. longirostrum Gourr.
T. teredo (Pouch.) Kof. Swezy e. macroceros (Ehr.) Vanh.
e. macroceros var. gallicum (Kof.) Jorg.
Family Wamowiaceae e. massiliense (Gourr.) Jorg.
Erythropsidinium sp. e. massiliense f. massiliense Soum.
Warnowia sp. e. massiliense f. armatum (Kars.) Jorg.
e. minutum Jorg.
Family Polykrikaceae e. paradoxides Cl.
Polykrikos schwarzii Buetsch. e. pavillardii Jorg.
e. pentagonum pentagonum Gourr.
Order Peridiniales e. pentagonum var. tenerum Jorg.
Family Ceratiaceae e. praelongum Lemm.
Ceratium arietinum Cl. f. gracilentum e. pulcheUum Schrod.
Jorg. e. ranipes Cl.
e. azoricum Cl. e. setaceum Jorg.
Appendix 213

e. strictum (Okam. & Nish.) Kof. O. sceptrum (Stein) Schrad.

C. symmetricum Pav. O. scolopax Stein
e. symmetricum var. coarctatum (Pav.) O. sphaeroideum Stein
Grah. & Bronik. O. turbo Kof.
e. symmetricum var. symmetricum Schuettiella mitra (Schiitt) Bal.
Grah. & Bronik.
e. tenue (Ostenf. & Schm.)tenue Family Peridiniaceae
e. tenue var. buceros Bal. Diplopelta sp.
e. tenue tenuissimum (Kof.) Oblea sp.
e. teres Kof. Peridinium quinquecorne Abe
e. trichoceros (Ehr.) Kof. Protoperidinium cf. aspidiotum (Bal.)
C. trip os (O.Miill.) Nitz. Bal.
e. tripos breve Ostenf. & Schm. Protoperidinium cf. claudicans (Pauls.)
e. tripos tripodioides Jarg. Bal.
C. tripos tripos (O.Miill.) Nitz. P. conicum (Gran) Bal.
e. vultur Cl. P. depressum (Bail.) Bal.
P. divergens (Ehr.) Bal.
Family Ceratocorythaceae P. excentricum (Pauls.) Bal.
Ceratocorys gourretii Pauls. P. globulus (Stein) Bal.
e. horrida Stein P. granii (Ostenf.) Bal.
P. inclinatum (Bal.) Bal.
Family Cladopyxidaceae P. longipes (Kars.) Bal.
Cladopyxis brathiolata Stein P. obtusum (Kars.) Bal.
Palaeophalacroma sp. P. oceanicum (Vanh.) Bal.
P. pentagonum (Gran) Bal.
Family Goniodomataceae P. punctulatum Pauls.
Goniodoma polyedricum (Pouchet) Jarg. P. pyriforme (Pauls.) Bal.
P. quarnerense (Schrad.) Bal.
Family Gonyaulacaceae P. robustum (Kof.) Bal.
Alexandrium Jraterculus (Bal.) Bal. P. spiniferum (Schi.) Bal.
Alexandrium sp. P. steinii (Jarg.) Bal.
Gonyaulax birostris Stein Scrippsiella trochoidea (Stein) Fine &
G. diegensis Kof. Loebl.III
G. digitalis (Pouch.) Kof.
G. Jragilis (Schiitt) Kof. Family Podolampadaceae
G. grindleyi Rein. Podolampas bipes Stein
G. pacifica Kof. P. elegans Schiitt
G. polyedra Stein P. palmipes Stein
G. polygramma Stein P. reticulata Kof.
G. cf. spinifera (Clap. & Lach.) Dies. P. spinifera Okam.
G. verior Sour.
Spiraulax kofoidii Grah. Family Pyrophacaceae
Pyrophacus horologium Stein .
Family Oxytoxaceae . P. steinii Schi.
Centrodinium sp.
Corythodinium sp. CLASS PRYMNESIOPHYCEAE
e. constrictum (Stein) Tayl. Order Isochrysidales
e. latum (Gaar.) Tayl. Family Gephyrocapsaceae
e. milneri Murr. & Whit. Emiliania huxleyi (Lohm.) Hay &
e. tesselatum (Stein) Mohler
Oxytoxum crassum Schi. Gephyrocapsa oceanica Kampt.
O. gracile Gran
O. longiceps Schi. Family Hymenomonadaceae
O. parvum Schi. Cricosphaera sp.
214 Appendix

Order Coccosphaerales S. pirus Hall. & Mark.

Family Calyptrosphaeraceae S. pulchra Lohm.
Calyptrosphaera depressa Schi. S. spinosa Lohm.
C. oblonga Lohm. S. tenuis Lohm.
Helladosphaera cf. comifera (Schi) Umbellosphaera tenuis (Kampt.) Paas.
Kampt.
Zygosphaera sp. Order Prymnesiales
Family Prymnesiaceae
Family Calciosoleniaceae Chrysochromulina sp.
Anoplosolenia brasiliensis (Lohm.) Deft.
Calciosolenia murrayi Gran Family Phaeocystaceae
Phaeocystis pouchetii (Hariot) Lager.
Family Ceratolithaceae
Ceratolithus cristatus Kampt. CLASS CHRYSOPHYCEAE
Order Ochromonadales
Family Coccolithaceae Family Chromulinaceae
Calcidiscus leptoporus (Murr. & Black.) Chromulina sp.
Loebl.Jr. & Tap.
Coccolith us pelagicus (Wall.) Schi. Family Chrysococcaceae
Oolithotus fragilis (Lohm.) Okada & Chrysocrx;cus cf. omatus Paas.
Melnt.
Umbilicosphaera sibogae (Weber-Van Order Synurales
Bosse) Gaard. Family Mallomonadaceae
Mallomonas sp.
Family Halopappaceae
Halopappus sp. CLASS DICTYOCHOPHYCEAE
H. adriaticus Schi., emend. Manton, Order Dictyochales
Bremer & Oates Family Dictyochaceae
Ophiaster hydroideus (Lohm.) Lohm. Dictyocha fibula Ehr.
O. formosus Gran D. speculum (Haeck.) Ehr.
Mesocena polymorpha Ehr.
Family Pontosphaeraceae Octactis pulchra Schi.
Lohmannosphaera subclausa Schi
Pontosphaera discopora Schi. CLASS BACILLARIOPHYCEAE
P. sessilis Lohm. Order Centrales
P. syracusana Lohm. Family Asterolampraceae
Scyphosphaera apsteinii Lohm. Asterolampra marylandica Ehr.
Asteromphalus arachne (Breb.) Ralfs
Family Rhabdosphaeraceae A. jlabellatus (Breb.) Grev.
Acanthoica quattrospina Lohm. A. heptactis (Breb.) Ralfs ex Pritch.
Algirosphaera quadricomu (Schi) A. hookeri Ehr.
Norris A. robustus Castr.
Discosphaera tubifer (Murr. & Black.) A. sarcophagus Wall.
Ostenf.
Rhabdosphaera claviger Murr. & Black. Family Biddulphiaceae
R. hispida Lohm. Biddulphia biddulphiana J.Sm. (Boy.)
B. tuomeyi (Bail.) Roper
Family Syracosphaeraceae Cerataulina pelagica (Cl.) Hend.
Coronosphaera maxima (Hall. & Mark.) C. daemon (Grev.) Hasle
. Gaard. C. dentata Hasle
Deutschlandia anthos Lohm. Climacodiun biconcavum Cl.
Syracosphaera dentata Lohm. C. frauenfeldianum Gron.
S. heimii Lohm. Eucampia comuta (Cl.) Gron.
S. histrica Kampt. E. zoodiacus Ehr.
Appendix 215

E. zoodiacus cylindrocornis Syvert. c. rostratus Laud.

Eunotogramma dubium Hust. C. saltans Cl.
E. leve Grun. C. seriacanthus Gran
E. marinum (W.Sm.) Perag. e. similis Cl.
E. rostra tum Hust. c. socialis Laud.
Hemiaulus hauckii Grun. ex V.Heurck C. subsecundus (Grun.) Hust.
H. membranaceus Cl. C. subtilis Cl.
H. sinensis Grev. C. subtilis var. abnormis Prosch.-Lavr.
Hydrosera whampoensis (Schw.) Deby C. wighami Brightw.
Isthmia enervis Ehr. e. willei Gran
Terpsinoe americana (Bail.) Ralfs
T. musica Ehr. Family Coscinodiscaceae
Coscinodiscus apiculatus Ehr. "
Family Chaetoceraceae e. apiculatus var. ambigua Grun.
Bacteriastrum biconicum Pay. C. argus Ehr.
B. elegans Pay. C. asteromphalus Ehr.
B. elongatum Cl. e. asteromphalus var. pabelanica Grun.
B. furcatum Shadb. C. brasiliensis M.Melch.
B. hyalinum Laud. C. centralis Ehr.~
B. hyalinum var. princeps (Castr.) Ikari C. concinnus W.Srii~
B. mediterraneum Pay. C. curvatulus Grun. ex Schm.
Chaetoceros affinis Laud. e. gigas Ehr.
e. atlanticus Cl. C. gigas var. praetextus (Jan.) Hust.
c. atlanticus var. neapolitan us (Schoed.) C. granii Gough
Hust. C. janischii Schm.
c. cf. borealis Bail. C. jonesianus (Grev.) Ostenf.
e. brevis SchUtt C.jonesianus (Grev.) Ostenf. var.
e. coarctatus Laud. jonesianus
c. compressus Laud. var. compressus C. jonesianus var. commutata (Grun.)
C. constrictus Gran Hust.
e. costatus Pay. C. kurzii Grun. & Schm.
c. concavicornis Mangin C. marginatus Ehr.
e. curvisetus Cl. C. obscurus A.Schm.
C. dadayi Pay. C. oculus-iridis Ehr.
C. danicus Cl. C. oculus-iridis Ehr. var. loculifera Ratt.
c. debilis Cl. e. oculus-iridis Ehr. var. stelliger Ratt.
C. decipiens Cl. C. pacificus Grun.
C. didymus Ehr. C. perforatus Ehr.
e. didymus var. anglica Gran C. perforatus var. cellulosa Grun.
C. didymus var. protuberans Gran e. perforatus var. pavillardii (Forti)
C. divers us Cl. Hust.
e. eibenii (Grun) Meun. e. radiatus Ehr.
e. laciniosus Schutt C. radiatus var. minor A.Schm.
c. laevis Lend.-Fort. , C. rothii (Ehr.) Grun.
c. lauderi Ralfs e. sublineatus Grun.
e. lorenzianus Grun. e. thorii Pay.
c. messanense Castro e. variabilis Freng.
c. pelagicus Cl. e. vidovichii M.Melch.
c. peruvian us Brightw. e. wailesii Gran & Angst.
e. peruvianus f. gracilis Schrad. Gossleriella tropica Schutt
e. pseudocurvisetus Mangin Palmeria hardmanniana Grev.
e. radians Schutt Stellarima stellaris (Roper) Hasle &
c. radicans Shutt Sims
216 Appendix

Family Psammodiscaceae Family Hemidiscaceae

Psammodiscus nitidus (Greg.) Round & Actinocyclus barkly (Coat.) Grun.
Mann A. brasiliensis M.Melch.
A. cholnokyi Van Land.
Family Cymatosiraceae A. curvatulus Jan.
Campylosira cymbelliformis A. kuetzingii A.Schm.
(A.Schm.)Grun. ex V.Heurck A. normani (Greg.) Hust.
Cymatosira adaroi Azpt. & Moros A. normani (Greg.) Hust. var. normani
C. atlantica Freng. A. octonarius Ehr.
Plagiogrammopsis vanheurckii (Grun.) A. octonarius var. crass us (W.Sm.)
Hasle Hend.
A. octonarius var. ralfsii (W.Sm.) Hend.
Family Eupodiscaceae A. octonarius var. spars~ (Greg.) Hend.
Auliscus pruinosus Bail. A. octonarius var. ten ella (Breb.) Hend.
A. sculptus (W.Sm.) Ralfs ex Pritch. A. platensis M.Melch.
A. cae latus Bail. A. subtilis (Greg.) Ralfs ex Pritch.
A. caelatus var. rhipis (A.Schm.) Perag. Hemidiscus cuneiformis Wall.
Cerataulus smithii Ralfs H. cuneiformisvar. ventricosa (Castr.)
C. turgidus Ehr. Hust.
Eupodiscus antiguus (Cox) Hanna H. ovalis L"ohm.
E. radiatus Bail. Roperia tessellata (Roper) Grun.
Odontella aurita (Lyng.) C.Ag.
O. auritavar. obtusa (Klitz.) Hust. Family Lithodesmiaceae
O. mobiliensis (Bail.) Grun. Bellerochea sp.
O. regia (Schul.) Simon. Ditylum brightwellii (West) Grun. ex
o. rhombus (Ehr.) Klitz. V.Heurck
o. longicruris Grev. Lithodesmium undulatum Ehr.
O. sinensis (Grev.) Grun. Streptotheca thamesis Shrubs.
Pleurosira laevis (Ehr.) Compo
Triceratium alternans Bail. Family Leptocylindraceae
T. antediluviana (Ehr.) V.Heurck Corethron criophilum Castr.
T. antediluviana var. excavata Freng. Leptocylindrus danicus Cl.
T. dubium Brightw. L. minimus Gran
T. favus Ehr.
T. favus var. quadrata (Ehr.) Grun. Family Melosiraceae
T. formosum f. pentagonale A.Schm. Hyalodiscus laevis Ehr.
T. patagonicum A.Schm. H. scoticus (Klitz.) Grun.
T. pentacrinus (Ehr.) Wall. H. subtilis Bail.
T. reticulum (Ehr.) Boy. Melosira juergensii C.Ag.
T. robertsianum f. inermis Grev. M. moniliformis (O.Mlill.) C.Ag.
T. scitulum Brightw. M. nummuloides (Dill.) C.Ag.
T. sculptum Shadb. Paralia sulcata (Ehr.) Cl.
P. sulcata (Ehr.) Klitz. var. sulcata
Family Heliopeltaceae P. sulcata var. biseriata Grun.
Actinoptychus campanulifer A.Schm. P. sulcata f. coronata (Ehr.) Grun.
A. cf. concentricus A.Schm. P. sulcata f. radiata Grun.
A. senarius Ehr. Podosira montagnei Klitz.
A. senarius cf. var. tamanicus Josue P. stelliger (Bail.) Mann
A. splendens (Shadb.) Ralfs ex Pritch. Stephanopyxis palmeriana (Grev.) Grun.
A.vulgaris Schu. S. turris (Grev.) Ralfs ex Pritch.
A. vulgaris f. octonaria Freng.
A. vulgaris f. polymera Freng. Family Rhizosoleniaceae
A. vulgaris f. senaria Freng. Dactyliosolen antarcticus Castro
Aulacodiscus petersii Ehr. D. mediterraneus Perag.
Appendix 217

Guinardia flaccida (Castr.) Perag. T. angulata (Grun.) Jorg.

Proboscia alata (Brightw.) Sund. T. anguste-lineata (A.Schm.) Fryx. &
Proboscia indica (Perag.) Sund Hasle
Proboscia inermis (Castr.) Sund. T. antarctica Comber
Pseudosolenia calcar-avis (Schul.) Sund. T. delicatula (chilensis) Ostenf.
Rhizosolenia acuminata (Perag.) Gran T. eccentrica (Ehr.) Cl.
R. bergonii Perag. T. eccentrica var. micropora (Ehr.)
R. castracanei Perag. Grun.
R. delicatula Cl. T. hendeyi Hasle & Fryx.
R. fragillissima Berg. T. lentiginosa (Jan.) Fryx.
R. hebetata Bail. T. leptopus (Grun.) Hasle & Fryx.
R. hebetata f. hiemalis Gran T. levanderi Goor.
R. hebetata f. semispina (Hens.) Gran T. minuscula Krasske
R. hyalina Ostenf. T. nanolienata (Mann) Fryx. & Hasle
R. imbricata Brightw. T. nodulineata (Hend.) Hasle & Fryx.
R. longiseta Zach. T. oestrupii (Ostenf.) Hasle var.
R. phuketensis Sund. oestrupii
R. pungens Cl.-Eu. T. oestrupii (Ostenf.) Hasle var.
R. robusta Norm. ex Pritch. venrickae
R. setigera Brightw. T. pseudonana Hasle & Heimdal
R. setigera f. pungens (Cl.) Brun. T. punctigera (Castr.) Hasle
R. setigera var. daga (Brightw.) T. rotula Meun.
M.Melch. T. simonsenii Hasle & Fryx.
R. shrubsolei Cl. T. subtilis (Ostenf.) Grun.
R. stolterfothii Perag. T. tenera Prosch.-Lavr.
R. styliformis Brightw. Tryblioptychus cocconeiformis (Cl.)
Hend.
Family Sticodiscaceae
Ethmodiscus gazellae (Jan.) Hust. Order Penn ales
Suborder Araphidineae
Family Thalassiosiraceae Family Fragilariaceae
Aulacoseira ambigua (Grun.) Simon. Asterionellopsis glacialis (Castr.) Round
A. distans (Ehr.) Simon. Climacosphaenia moniligera Ehr.
A. granulata (Ehr.) Simon. Delphineis surirella (Ehr.) G.W.Andr.
A. islandica (O.MUlI.) Simon. Diatoma anceps (Ehr.) Grun.
A. islandica subsp. helvetica (O.Mlill.) Dimeregramma hyalinum Hust.
Simon. D. cf. marinum (Greg.) Ralfs ex Pritch.
A. italica (Ehr.) Simon. D. minor (Greg.) Ralfs in Pritch.
A. nyassensis (O.Mliller) Simon. Fragilaria crotonenensis Kitt.
A. pseudogranulata (Cl.) Simon. F. constricta Ehr.
Cyclotella kuetzingiana Thw. F. construens Grun.
C. meneghiniana Klitz. F. istvanffyi Pant.
C. littoralis Lange & Syvert. F. lapponica Grun.
C. stelligera Cl. & Grun. F. pinnata Ehr.
C. striata (Klitz.) Grun. Glyphodesmis distans (Greg.) Grun.
C. stylorum Brightw. Grammatophora angulosa Ehr.
Cymatotheca weissflogii (Grun.) Hend. G. angulosa Ehr. var. angulosa
Detonula pumila Castro G. hamulifera Klitz.
Lauderia annulata Cl. G. marina (Lyng.) Klitz.
Planktbniella sol (Wall.) Schlitt G. marina (Lyng.) Klitz. var. marina
Skeletonema costa tum (Grev.) Cl. G. maxima Grun.
S. subsalsum (Cl.) Bethge G. oceanica Ehr.
S. cf. tropicum Cl. G. oceanica Ehr. var. oceanica
Thalassiosira allenii Takano G. serpentina Ehr.
218 Appendix

Licmophora abbreviata C.Ag. E. pectinalis (Klitz.) Rab.

L. argentescens C.Ag.
L. jlabellata (Carm.) C.Ag. Family Achanthaceae
L. gracilis (Ehr.) Gron. Achnanthes brevipes C.Ag.
L. lyngbyei (Klitz.) Gron ex V.Heurck A. brevipes angustata (Grev.) Cl.
Opephora marina (Greg.) Petit A. brevipes var. intermedia (Klitz.)
O. martyi Herib. Gron.
Plagiogramma pulchellum Grev. A. brevipes var. parvula (Klitz.) Cl.
P. pulchellum Grev. var. pygmaea A. curvirostrum Bron.
(Grev.) Perag. & Perag. A. delicatula (Klitz.) Gron.
P. simplex Freng. A. exigua var. heterovalvata Krasske
P. staurophorum (Greg.) Herib. A. hauckiana Gron.
Podocystis adriatica (Klitz.) Boyer A. injlata (Klitz.) Gron..
Rhabdonema adriaticum Klitz. A. javanica Gron. var. rhombica Gron
Rhaphoneis amphiceros (Ehr.) Ehr. A. lanceolata (Breb.) Gron.
R. amphiceros var. tetragona Gron. A. longipes C.Ag.
R. castracanei Gron. A. longipes C.Ag. var. longipes
R. disco ides Subrahm. A. promonturii Giffen
R. surirella (Ehr.) Gron. A. saxonica Krasske
Striatella delicatula (Klitz.) Gron. Anorthonezs hyalina Hust.
Synedra acus Klitz. var. angustissima Cocconeis costata Greg.
Grun. e. diminuta Pant.
S. fasciculata (C.Ag.) Klitz. e. dirupta Greg.
S. fasciculata var. truncata (Grev.) Patr. e. disculus (Schum.) Cl.
S. gaillonii (Bory) Ehr. e. dis tans Greg.
S. gaillonii (Bory) Ehr. var. gaillonii e. heteroidea Hantzsch
S. goulardi (Breb.) Gron. e. pediculus Ehr.
S. reinboldii V.Heurck e. pellucida Gron
S. rumpens Klitz. e. placentula Ehr.
S. ulna (Nitz.) Ehr. e. placentula var. euglypta (Ehr.) el.
S. ulna var. contracta Ostr. e. placentula var. lineata (Ehr.) el.
S. undulata Bail. e. pinnata Greg. ex Grev.
Tabellaria sp. e. pseudomarginata Greg.
Thalassionema frauenfeldii (Grun.) e. pseudomarginata Greg. var.
Hallegr. intermedia Gron.
T. nitzschioides Hust. e. quarnerensis (Gron.) A.Schm.
Thalassiothrix delicatula Cupp e. scuttellum Ehr.
T. longissima Cl. & Gron. e. scuttellum var. minutissima Gron.
T. mediterranea Pay. e. scutellum var. ornata Gron.
T. mediterranea var. pacifica Cupp e. scuttelum var. parva Gron. in el.
e. scutellum Ehr. var. scutellum
Suborder Raphidineae e. scutellum var. speciosa (Greg.) el.
Family Eunotiaceae e. stauroneiformis (V.Heurck) Okuno
Eunotia camelus Ehr.
E. camelus var. denticulata Gron. Family Naviculaceae
E. camelus var. didymodon Gron. Amphipleura lindheimeri Gron.
E. diodon Ehr. var. minor Gron. Anomoeoneis serians (Breb.) el.
E. faba (Ehr.) Gron. Caloneis brevis (Greg.) Cl.
E. jlexuosa (Breb.) Klitz. e. permagma (Bail.) el.
E. gracilis (Ehr.) Rab. Diploneis bombus Ehr.
E. indica var. biggiba Freng. D. bombus var. bombiformis (el.) Hust.
E. indica f. gracilis Freng. D. crabro (Ehr.) Ehr.
E. kocheliensis O.Mlill. D. didymus (Ehr.) el.
E. maior (W.Sm.) Rab. D. gruendleri (A.Schm.) el.
Appendix 219

D. incurvata (Greg.) Cl. N. consentanea Hust.

D. littoralis (Donk.) Cl. N. cryptocephala Klitz.
D. littoralis var. clathrata (Ostrup) Cl. N. crucicula (W.Sm.) Donk.
D. oblongella (Nag. ex Klitz.) Ross N. cuspidata (Klitz.) Klitz var. cuspidata
D. ovalis (Hilse) Cl. N. demerara Grun. ex Cl. var. demerara
D. ovalis var. oblongella (Nag.) Cl. N. digito-radiata (Greg.) Ralfs
D. papula (A.Schm.) Cl. N. exigua Greg. ex Grun. var. capitata
D. papula (A.Schm.) Cl. var. papula Patr.
D. puella (Schum.) Cl. N. flagellifera Hust.
D. smithii (Breb.) Cl. N. forcipata Grev.
D. smithii (Breb.) Cl. var. smithii N. gastrum (Ehr.) Klitz.
D. vacilans (A.Schm.) Cl. N. granulata Bail.
D. vetula (A.Schm.) Cl. var. vetula N. gregaria Donk.
D. weissflogii (Schm.) Cl. N. hagelsteinii Hust.
Donkinia carinata (Donk.) Ralfs N. humerosa Breb. in W.Sm.
Frustulia interposta (Lew.) De-Toni N. laevissima Klitz. var. laevissima
F. rhomboides (Ehr.) De Toni N. lorenzii (Grun.) Hust.
Gomphonema constrictum Ehr. N.lyraEhr.
G. gracile Ehr. N. lyroides Hend...
G. gracile Ehr. var. gracile N. marina Ralfs ex Pritch.
G. gracile var.lanceolata Klitz. N. membranaeea (Cl.)
G. parvulum (Klitz.) Klitz. N. mutica Klitz.
G. turris Ehr. N. notha Wall.
Gyrosigma balticum (Ehr.) Rab. N. nummularia Grev.
G. balticum (Ehr.) Cl. var. balticum N. pelagica Cl.
G. eximium (Thw.) Boy. N. pennata A.Schm.
G. exoticum Choln. N. pennata A.Schm. var. pennata
G. littorale (W.Sm.) Griff & Henfr. N. pennata f. maxima Cl.
G. rectum (Donk.) Cl. N. peregrina (Ehr.) Klitz.
G. scalproides (Rab.) Cl. N. pygmaea Klitz.
G. spencerii (W.Sm.) Cl. N. plagiostoma Grun.
Mastogloia apiculata W.Sm. N. porifera Hust.
M. crucicula (Grun) Cl. N. praetexta Ehr.
M. binotata (Grun.) Cl. N. punctulata W.Sm. var. punetulata
M. braunii Grun. N. punctulata var. marina (Ralfs) Cl.
M. decussata Brun. N. pupula Klitz.
M. erythraea Grun. N. radiosa var. tenella (Breb.) Grun.
M. exigua Lew. N. ramosissima (C.Ag.) Cl.
M. exigua Lew. var. exigua N. rostellata Klitz.
M. exilis Hust. N. salinaroides Choln.
M. fimbriata (Brightw.) Cl. N. scopulorum Breb. ex Klitz.
M. jurgensii C.Ag. N. serena Freng.
M. meisteri Hust. N. sovereigne Hust.
M. smithii Thw. N. soodensis Krasske
M. splendica (Greg.) Cl. & O.Mlill. N. spectabilis Greg.
Navicula anglica Ralfs N. transfuga var. plagiostoma (Grun.)
N. bacillum Ehr. var. bacillum Cl.
N. canalis Patrick N. cf. tripuncata (O.Milll.) Bory
N. eancellata Donk. N. tubulosa Brun.
N. capitata Ehr. var. hungarica (Grun.) N. yarrensis Grun. in Schm.
Ross N. zostereti Grun.
N. cariEhr. Neidium affine var. amphyrhynehus
N. cincta (Ehr.) Ralfs var. cineta in (Ehr.) Cl.
Pritch. N. iridis (Ehr.) Cl.
220 Appendix

Pinnularia borealis Ehr. Nitzschia acuminata (W.Sm.) Cl.

P. cruciformis (Dank.) Cl. N. angularis var. affinis Grun.
P. divergens W.Sm. N. apiculata (Greg.) Grun. in Cl. &
P. gibba Ehr. Grun.
P. hemiptera (Klitz.) Cl. N. archibaldii Lange-Bertalot
P. intermedia (Lagers.) Cl. N. arenosa Garc.-Bapt.
P. major (Klitz.) Rab. N. bilobata W.Sm.
P. microstauron (Ehr.) Cl. N. circumsuta (Bail.) Grun. in Cl. &
P. neuquina Freng. Grun.
P. trevelyana (Dank.) Rab. N. coarctata var. oceanica (Grun.)
Pleurosigma angulatum (Quek.) W.Sm. Freng.
P. atlanticum Freng. N. compressa (Bail.) Boy.
P. brasiliana M.Melch. N. debilis (Am.) Grun.
P. diverse-striatum Meister N. dissipata Klitz.
P. elongatum W.Sm. N. epithemioides Grun.
P. hamuliferum Brun. & Temp. N. filiform is (W.Sm.) Schlitt
P. intermedium W.Sm. N. gandersheimiensis Krasske
P. intermedium var. nubecula (W. Sm.) N. granulata Grun. in Cl. & Grun.
Grun. ex Van Heurck N. hantzschiana Rab.
P. lanceolatum Dank. N. insignisooGreg.
P. marinum Dank. N. insignis var. spathulifera Grun.
P. naviculaceum Bn!b. N.lanceolata W.Sm.
P. normanii Ralfs. N. cf. lanceolata var. pygmaea Cl.
P. strigosum W.Sm. N. linearis Wm.Sm. var. linearis
Rhoicosphaenia curvata (Klitz.) Grun. N. longissima (Bn!b. in Klitz.) Ralfs
ex Rab. N. lorenziana Grun.
Stauroneis salina W.Sm N. marginata Hust.
S. phoenicenterum (Nitz.) Ehr. N. obtusa W.Sm.
Trachyneis antillarum Cl. N. pacifica Cupp
T. aspera (Ehr.) Cl. N. palea (Klitz.) W.Sm.
T. asp era var. intermedia (Grun.) Cl. N. panduriformis Greg.
Tropidoneis lepidoptera (Greg.) Cl. N. panduriformis var. minor Grun.
T. lepidoptera Greg. var. minor Cl. N. punctata (W.Sm.) Grun. var.
punctata
Family Epithemiaceae N. scalaris (Ehr.) W.Sm. var. scalaris
Rhopalodia gibberula (Ehr.) O.Mlill. N. sigma (Klitz.) W.Sm.
R. musculus (Klitz.) O.Mlill. N. sigma var. intercedens Grun.
N. sigma var. sigmatella Grun.
Family Nitzschiaceae N. socialis Ralfs
Bacillaria paxillifer (O.Mlill.) Hend. N. spathulata W.Sm
Cylindrotheca closterium (Ehr.) Reim. & N. tryblionella Hantzch
Lew. N. tryblionella var. victoriae Grun.
C. fusiformis var. fusiformis Reim. & Lew. N. ventricosa (Palmer.) Kitt.
C. signata Reim. & Lew. N. vidovichii (Grun.) Grun.
Cymatonitzschia marina (Lew.) Simon. N. vitrea Norm.
Hantzschia amphioxus (Ehr.) Grun. Pseudoeunotia doliolus (Wall.) Grun.
H. amphioxus var. vivax (Hantzsch) Pseudo-nitzschia australis Freng.
H. distinctepunctata (Hust.) Hust. P. cf. cuspidata Hasle
H. longiareolata Garc.-Bapt. P. delicatissima complex
H. jJsammicola Garc. -Bapt. P. pseudodelicatissima (Hasle) Hasle
H. virgata (Roper) Grun. P. pungens Grun. f. pungens
H. virgata var. capitellata Hust. P. seriata complex
H. virgata var. wittii (Grun.) Grun. Tryblionella coarctata (Grun.) Mann
Appendix 221

Family Cymbellaceae Order Chloramoebales

Amphora acutiuscula Klitz. Family Chloramoebaceae
A. angusta Grev. var. angusta Chloromeson parva Carter
A. bigibba Grun.
A. binodis Greg. var. bigibba Grun. Order Vaucheriales
A. bongrainii M.Perag. var. bongrainii Family Vaucheriaceae
A. castellata Giffen Vaucheria longicaulis Hoppaugh
A. coffeaeformis (C.Ag.) Klitz.
A. granulata Greg. CLASSPHAEOPHYCEAE
A. laevis Greg. var.laevissima (Greg.) CL Order Ectocarpales
A. marina (W.Sm.) V.Heurck Family Ectocarpaceae
A. ovalis (Klitz.) Klitz. Ectocarpus siliculosus (Dillwyn)
A. proteoides Hust. Lyngb.
A. sabiniana Reim. Giffordia mitchellae Hamel
A. turgida Greg.
A. ventricosa Greg. Order Punctariales
Cymbella affinis Klitz. Family Punctariaceae
e. amphicephala Nag. Petalonia fascia (MUlL) Kuntze
e. cuspidata Klitz.
C. cymbiformis C.Ag. PHYLUM CHLOROPHYTA
e. tumida (Breb.) V.Heurck CLASS EUGLENOPHYCEAE
e. tumida var. tumida (Breb.) V.Heurck Order Euglenales
C. turgida (Greg.) CL Family Eutreptiaceae
e. ventricosa C.Ag. Eutreptia sp.
Eutreptiella sp.
Family Surirellaceae
Campylodiscus daemelianus Grun. Family Euglenaceae
e. decorus Breb. Euglena viridis Ehr.
e. echineis Ehr. var. echineis
e. ecclesianus Grev. CLASSPRASYNOPHYCEAE
Petrodictyon gemma (Ehr.) Mann Order Chlorodendrales
Surirella bisseriata Breb. Family Halosphaeraceae
S. fastuosa Ehr. Halosphaera viridis Schmitz
S. febigerii Lewis Pterosperma sp.
S. guatimalensis Ehr. Pyramimonas sp.
S. linearis var. constricta (Ehr.) Grun.
S. recendes A.Schm. CLASS CHLOROPHYCEAE
S. rorata Freng. Order Volvocales
S. tenera Greg. Family Chlamydomonadaceae
S. cf. mulleri Hust. Brachiomonas sp.
Stenopterobia intermedia (Lew.) Van Chlamydomonas sp.
Heurck
Order Chlorococcales
Family Entomoneidaceae Family Oocystaceae
Entomoneis alata Ehr. Koliella longiseta Hindak
Lagerheimia citriformis (Snow)
CLASS RAPHIDOPHYCEAE Collins
Order Chattonellales
Family Chattonellacea Family Scenedesmaceae
Chattonella sp. Crucigenia tetrapedia (Kirch.) W. &
Fibrocapsa japonica Toriumi & Takano S.West
Scenedesmus acuminatus (Lager.)
CLASS TRIBOPHYCEAE Chodat
(XANTHOPHYCEAE) S. acutus Meyen
222 Appendix

S. quadricauda (Turp.) Breb. PHYLUM GRANULORETICULOSA

CLASS FORAMINIFEREA
Order Ulotrichales Order Textulariida
Family Ulotrichaceae Family Rzehakinidae
Planctonema lauterbornii Schmidle Miliammina earlandi Loeblich &
Ulothrix flacca (Dillwyn) Thuret Tappan
M. fusca (Brady)
Family Chaetophoraceae Trilocularena patensis Closs
Phaeophila viridis (Reinke) Burrows
Family Hormosinidae
Family Gomontiaceae Protoschista find ens (Parker)
Gomontia lignicola Moorex Reophax arctica Brady
R. nana Rhumbler
Order Ulvales
Family Ulvaceae Family Lituolidae
Enteromorpha bulbosa (Suhr) Klitz. Ammoastuta inepta (Cushman &
E. clathrata (Roth) J.Ag. McCulloch)
E. compressa (L.) Grev. Ammobaculites exiguus Cushman &
E. flexuosa (Wulf) J.Ag. Bronnimann
E. intestinalis (L.) Link Ammosca7aria pseudospiralis
E. linza (L.) J.Ag. (Williamson)
E. micrococca Klitz. Ammotium cassis (Parker)
E. ramulosa (Smith) Hooker A. planissimum (Cushman)
Ulvaria oxysperma (Klitz.) Bliding A. pseudocassis Cushman &
Ulva lactuca L. Bronnimann
U. fasciata Delile A. salsum (Cushman & Bronnimann)
Haplophragmoides wilberti Andersen
Order Cladophorales
Family Cladophoraceae Family Trochamminidae
Chaetophora aerea (Dillwyn) Klitz. Arenoparrella mexicana (Kornfeld)
Cladophora crispata (Roth) J.Ag. emend. Andersen
e. fascicularis (Mert.) Klitz. Trochammina inflata (Montagu)
e. flexuosa (Dill.) Harv. T. macrescens Brady
e. prolifera (Roth) Klitz. T. ochracea (Williamson)
e. scitula (Suhr) Klitz. Trochamminita salsa (Cushman &
Rhizoclonium kerneri Stockm. Bronnimann)
R. ripanum (Roth) Harv.
R. tortuosum Klitz. Family Textulariidae
Textularia candeiana Orbigny
Order Siphonales T. earlandi Parker
Family Bryopsidaceae T. gramen Orbigny
Bryopsis plumosa (Huds.) C.Ag. Spiroplectammina biformis (Parker &
Bryopsis sp. Jones)

Order Zygnematales Order Miliolida

Family Zygnemataceae Family Miliolidae
Mougeotia sp. Massilina secans (Orbigny)
Spirogyra sp. Pyrgo nasuta Cushman
Pyrgo sp.
Family Desmidiaceae Quinqueloculina atlantica Boltovskoy
Cosmarium sp. Q. aff. frigida Parker
Mesotaenium macrococcum var. Q. isabellei Orbigny
micrococcum (Klitz.) West & West Q. lamarckiana Orbigny
Appendix 223

Q. cf. patagonica Orbigny G. hirsuta (Orbigny)

Q. polygona Orbigny G. inflata (Orbigny)
Q. seminulum (Linnaeus) G. menardii (Orbigny)
Q. aff. seminulum (Linnaeus) G. pumilio Parker
Triloculina trigonula (Lamarck) G. scitula (Brady)
G. tumida (Brady)
Order Lagenida G. truncatulinoides (Orbigny)
Family Nodosariidae G. ungulata Bermudez
Nodosaria catesbyi Orbigny Pulleniatina obliquiloculata (Parker &
Jones)
Family Polymorphinidae
Globulina caribaea Orbigny Family Catapsydracidae
Guttulina problema Orbigny Globigerinita clarkei (Rogl & Bolli)
G. glutinata (Egger)
Family Lagenidae G. humilis (Brady)
Lagena laevis (Montagu) G. iota Parker
L. laevis f. perlucida (Montagu)
L. sulcata f.lyellii (Seguenza) Order Rotaliida
Oolina costata Williamson Family Buliminidae
Bolivina striatula Cushman
Order Globigerinida B. pusilla Schwager
Family Globigerinidae Bulimina marginata Orbigny
Globigerina bulloides Orbigny B. marginata f. aculeata Orbigny
G. digitata Brady B. marginata f. echinata Orbigny
G. falconensis Blow B. marginata f. subalata Cushman &
G. rubescens Hofker Parker
Globigerinella aequilateralis (Brady) B. patagonica Orbigny
G. calida (Parker) B. patagonica f. glabra Cushman &
Globigerinoides conglobatus (Brady) Wickenden
G. elongatus (Orbigny) B. pseudoaffinis Kleinpell
G. ruber (Orbigny) Buliminella elegantissima Orbigny
G. ruber f. pyramidalis (van den Broeck) Uvigerina peregrina f. parvula Cushman
G. ruber f. triloba (Reuss) Virgulina cf. pauciloculata H.Brady
G. sacculifer (Brady)
G. trilobus (Reuss) Family Cibicididae
G. trilobus f. sacculifera (Brady) Cibiddes bertheloti (Orbigny)
G. trilobus (Reuss) C. bertheloti f. boueana (Orbigny)
G. tenellus Parker C. umbonatus Phleger & Parker
Orbulina universa Orbigny
Sphaeroidinella dehiscens (Parker & Family Nonionidae
Jones) Noniongrateloupe (Orbigny)
N. tisburyensis Butcher
Family Candeinidae Noniella atlantica Cushman
Candeina nitida Orbigny
Family Rotaliidae
Family Hantkeninidae Bucella frigida (Cushman)
Hastigerina pelagica (Orbigny) B. cf. inusitata Anderson
Hastigerinella digitata (Rhumbler) B. peruviana campsi (Boltovskoy)
Discorbis williamsoni (Chapman & Parr)
Family Globorotaliidae Poroeponides lateralis (Terquem)
Globoquadrina dutertrei (Orbigny) Rolshausenia rolshauseni (Cushman &
G. pachyderma (Ehrenberg) Bermudez)
Globorotalia anfracta Parker Rotalia beccarii ex gr. parkinsoniana
G. crassaformis (Galloway & Wissler) (Orbigny)
224 Appendix

Family Elphidiidae Family Rhabdonellidae

Cancris sagra (Orbigny) Rhabdonellopsis sp.
Elphidium discoidale (Orbigny)
E. aff. discoidale (Orbigny) Family Xystonellidae
E. excavatum (Terquem) Parundella inflata Kof. & Camp.
E. galvestonense Kornfeld Xystonella sp.
E. gunteri Cole
E. incertum (Williamson) KINGDOM PLANTAE
PHYLUM PTERIDOPHYTA
PHYLUM CILIOPHORA Family Lycopodianaceae
CLASS KINETOFRAGMINOPHORA Lycopodium alopecuroides 1.
Order Haptorida
Family Didiniidae Family Equisetaceae
Mesodinium rubrum (Lohm.) Hamb. & Equisetum giganteum L.
Budd.
Family Polypodiaceae
CLASS POLYHYMENOPHORA Acrosticum aureum 1.
Order Oligotriehida
Suborder Oligotriehina Family Sphagnaceae
Family Strombidiidae Sphagnum sp.
Strombidium spp.
PHYLUM ANGIOSPERMOPHYTA
Family Strobilidiidae CLASS MONOCOTYLEDONEAE
Lohmaniella sp. Order Pandanales
Strobilidium sp. Family Typhaceae
Typha domingensis Pers.
Suborder Tintinnina
Family Codonellidae Order Helobiae
Codonaria fimbriata (Meun.) Bal. Family Potamogatonaceae
Codonella sp. Potamogeton striatus Ruiz & Pay.
Leprotintinnus sp. Ruppia maritima 1.
Tintinnidium sp. Zannichellia palustris 1.
Tintinnopsis sp.
T. beroidea Stein Family Juncaginaceae
T. campanula Ehr. Triglochin striata Ruiz & Pay.
T. radix (Imhof) Brandt Triglochin sp.
T. levigata Kof. & Camp.
Order Graminales
Family Codonellopsidae Family Gramineae
Codonellopsis morchella (Cl.) Jarg. Andropogon arenarius Hack.
Stenosemella sp. A. selloanus Kunth.
Axonopus compressus (Sw.) Beauy.
Family Cyttarocylidae Briza erecta Lam.
Favella sp. B. minor 1.
Cenchrus echinatus 1.
Family Tintinnidae Cynodon dactylon (1.) Pers.
Salpingacantha ampla Kof. & Camp. Digitaria aequiglumes (Hack. & Arech.)
Salpingella secata Brandt Parodi
Elyonurus rostratus Nees
Family Dietyocystidae Eragrostis airoides Nees
Dictyocysta lata Kof. & Camp. E. bahiensis (Schrad. ex. Schult.)
D. mitra Haeckel E. cataclasta Nieora. ex Burk.
D. miilleri (Imhof) Jarg. E. tnchocolea Hack. & Arech.
Appendix 225

Erianthus angustifolius Nees Eichhomia crassipes (Mart.) Solms.

Eriochloa punctata (L.)Desv.
Imperata brasiliensis Trin. Family Bromeliaceae
Ischaemum minus Pres!. Bromelia antiacatha Bert.
1. urvilleanum Kunth.
Panicum racemosum (Beauv.) Spreng. Family Xyrydaceae
P. rep ens Berg. Xyris caroliniana Walt.
Paspalum distichum 1.
P. nicorae Parodi Order Liliales
P. vagina tum Sw. Family Juncaceae
Poidium rufum (Pres!.) Matthei funcus acutus L.
Polypogon maritimus Willd. f. capillaceus Lam.
P. monspeliensis (1.) Desf. f. dombeyanus Gay.
Setaria geniculata (Lam.) Beauv. f. effusus 1.
Spartina altemijlora Lois. f. microcephalus H.B.K.
S. ciliata Brongn. funcus spp.
S. densiflora Brongn.
Sporobulus indicus (1.) Brongn. Family Iridaceae
Stenotaphrum secunda tum (Walt.) Sisynnchium sp..
O.Ktze Sisyrinchium graminifolium Lind!.

Family Cyperaceae Order Orchidales

Androtrichum trigynum (Spreng.) Family Orchidaceae
Pfeiffer Habenaria parvijlora Lind!.
Cladium jamaiscensis Crantz.
Cyperus brevifolius (Rottb.) Hassk. CLASS DICOTYLEDONEAE
e. cayennensis (Lam.) Britt. Order Polygonales
e. giganteus Vahl Family Polygonaceae
e. obtusatus (Pres!.) Mattf. & Ktikenth. Polygonum negletum Bess.
e. polystachyus Rottb.
e. reflexus Vahl Order Chenopodiales
e. sesquiflorus (Torr.) Mattf. & Ktikenth. Family Aizoaceae
e. tener (Clarke.) Ostenf. Sesuvium portulacastrum 1.
Eleocharis montevidensis Kunth.
E. nana Kunth. Family Portulacaceae
E. nodulosa (Roth.) Schult. Portulaca grandijlora Hook.
E. obtusa (Willd.) Schult.
E. sellowiana Kunth. Family Caryophyllaceae
Fimbristylis autumnalis (1.) Roem. & Cardionema ramosissima (Weinm.)
Schult. A.Nels. & Macbr.
F. diphylla (Retz.) Vahl Spergularia cf. marina (1.) Griseb.
F. spadicea (1.) Vahl
Scirpus olneyi Gray Family Amaranthaceae
s. americanus Pers. Altemanthera phyloxeroides (Mart.)
S. cemuus Vahl Griseb.
S. maritimus 1. Blutaparon portulacoides (St.-Hi!.)
Scleria hirtella Sw. Mears.

Order Xyridales (Farinosae) Family Chenopodiaceae

Family Eriocaulaceae A triplex cf. hastata 1.
Eriocaulon modestum Kunth. A. cf. patula 1.
Chenopodium album 1.
Family Pontederiaceae e. retusum Juss. ex Moq.
Heteranthera reniformis R. & P. Salicomia ambigua Michx.
226 Appendix

S. gaudichaudiana Moq. Family Primulaceae

Anagallis arvensis 1.
Order Ranales
Family Ceratophyllaceae Order Plumbaginales
Ceratophyllum demersum 1. Family Plumbaginaceae
Limonium brasiliensis (Boiss.) O.Ktze.
Order Papaverales
Family Cruciferae Order Gentianales
Cakile maritima Scop. Family Gentianaceae
Zigostigma australe (Cham. & Schlecht)
Order Sarraceniales Griseb.
Family Droseraceae
Drosera brevifolia Pursh. Family Apocynaceae
Nymphoides indica (1.) O.Ktze.
Order Rosales
Family Rosaceae Family Asc1epiadaceae
Margyricarpus pinnatus (Lam.) O.Ktze. Asclepias mellodora St.-HiI.

Family Leguminoseae Order Tubiflorae

Aeschynomene sensitiva Sw. Family Convolvulaceae
Indigofera sabulicola Benth. Calystegia sepium (1.) R.Br.
Vigna luteola (Jacq.) Benth. e. soldanella (1.) Roem. & Schult.
Cressa truxillensis H.B.K.
Order Geraniales
Family Polygalaceae Family Verbanaceae
Polygala cyparissias St.-Hii. Phyla canescens (H.B.K.) Greene

Order Parietales Family Solanaceae

Family Guttiferae Solanum cf. americanum (Mill.)
Hypencum connatum Lam.
Family Scrophulariaceae
Order Myrtales Agalinis communis (Cham. & Schlecht.)
Family Onagraceae D'Arcy
Oenothera affinis Camb. Bacopa monnieri (1.) Penn.
Lindernia dubia (1.) Penn.
Family Gunneraceae Stemodia hyptoides Cham. & Schlecht.
Gunnera herteri Osten
Family Lentibulariaceae
Family Haloragaceae Utricularia tricolor St.-Hii.
Myriophyllum brasiliense Camb.
Order Plantaginales
Order Umbelliflorae Family Plantaginaceae
Family Umbelliferae Plantago australis Lam.
Apium graveolens 1. P. paralias Dcne.
Centella hirtella Nannf. P. cf. lanceolata 1.
Eryngium nudicaule Lam.
Eryngium sp. Order Campanulales
Hydrocotyle bonarensis Lam. Family Compo sitae
Lilaeopsis attenuata (Hook. & Am.) Achyrocline satureoides (Lam.) DC.
Fern. Aster squamatus (Spreng.) Hieron.
Baccharis patens Baker
Order Primulales B. spicata (Lam.) Baili.
Family Myrsinaceae B. trimera DC.
Rapanea parviflora Mez. Conyza blakei Cabrera
Appendix 227

C. bonariensis (1.) Cronq. Order Epipolasida

C. floribunda H.B.K. Family Jaspidae
C. pampeana (Parodi) Cabrera Jaspis johnstoni (Schmidt, 1868)
Cotula coronopifolia 1.
Eclipta prostrata 1. Order Lithistida
Gamochaeta americana (Mill.) Wedd. Family Corallistidae
G. spicata (Lam.) Cabrera Corallistes typus Schmidt, 1870
Gnaphalium filaginea (DC.) Cabrera
Picrosia longifolia D.Don. Family Pleromidae
Pluchea sagittalis (Lam.) Cabrera Pleroma sp.
Porophyllum cf. americanum DC. Siphonidium sp.
P. obscurum (Spreng.) DC.
Pterocaulon angustifolium DC. Order Hadromerida
P. bakeri Malme Family Tethydae
P. purpurascens Malme Halicometes minuta Sara & Barbosa,
Senecio crassiflorus (Poir.) DC. 1995
S. selloi (Spreng.) DC.
S. tweediei Hook & Am. Family Clionidae
Spilanthes sp. Cliona sp.

Family Calyceraceae Family Suberitidae

Acicarpha spathulata R.Br. Pseudosuberites melanos Laubenfels,
Calycera crassifolia (Miers) Hick. 1934
Suberites caminatus Ridley & Dendy,
Order Campanulales 1886
Family Campanulaceae
Pratia hederacea (Cham.) G.Don. Order Axinellida (= Halicondrida)
Family Axinellidae
KINGDOM ANIMALIA Axinella sp.
PHYLUM PORIFERA Bubaris sp.
CLASS HEXATINELLIDA Phakelia sp.
Order Dyctyonina (= Hexasteropoda)
Family Euretidae Order Poecilosclerida
Dactylocalix pumiceus Stutchubury, Family Microcionidae
1841 Clathria sp.
Margaritella sp. Microciona sp.

CLASS DEMOSPONGIAE Family Raspailiidae

Order Astrophorida Raspailia sp.
Family Ancorinidae
Penares sp. Family Rhabderemiidae
Stelletta sp. Rhabderemia uruguaiensis (Van Soest &
Hooper, 1993)
Family Geodiidae
Cydonium glariosus Sollas, 1886 Family Iophonidae
Erylus oxyaster Lendenfeld, 1910 Iophon sp.
E. formosus Sollas, 1886
Geodia eosaster (Sollas, 1886) Family Tedaniidae
Geodia sp. Tedania spinata (Ridley, 1881)
Tedania sp.
Family Pachastrellidae
Pachastrella monilifera Schmidt, 1870 Family Desmacellidae
Sphinctrella cribrifera (Sollas, 1886) Biemna sp.
228 Appendix

Order Haplosclerida Suborder Limnomedusae

Family Chalinidae Family Olindiidae
Acervochalina sp. Olindias sambaquiensis (Delle Chiaje,
1841)
Family Niphatidae Vallentinia gabriellae Mendes, 1948
Isodictya sp.
Family Proboscidactylidae
Order Dictyoceratida Proboscidactyla ornata (McCrady, 1857)
Family Thorectidae
Ircinia sp. Suborder Anthomedusae
Family Zancleidae
Family Dysideidae Zanclea costata Gegenbaur, 1856
Dysideasp.
Family Cytaeidae
PHYLUM CNIDARIA Cytaeis tetrastyla Eschscholtz, 1829
CLASS HYDROZOA
Order Trachylina Family Bougainvilliidae
Family Geryonidae Bougainvillia niove Mayer, 1894
Geryonia proboscidalis (Forskal, 1775) B. platygaster (Haeckel, 1879)
Liriope tetraphylla (Chamisso &
Eysenhart,1821) Family Clavidae
Cordylophora sp.
Family Halicreidae Turritopsis nutricula McCrady, 1856
Halitrephes maasi Bigelow, 1909
Family Corynidae
Family Rhopalonematidae Dipurena reesi Vannucci, 1956
Aglantha digitale (Muller, 1776) Stauridiosarsia producta (Writh, 1858)
Aglaura hemistoma Peron & LeSueur,
1809 Family Tubulariidae
Persa incolorata McCrady, 1857 Ectopleura dumortieri (Van Beneden,
Rhopalonema vela tum Gegenbaur, 1856 1844)
Sminthea erygaster Gegenbaur, 1856 Euphysorafurcata Kramp, 1948
E. gracilis (Brooks, 1882)
Family Solmariidae
Pegantha clara Bigelow, 1909 Family Pandeidae
P. laevis Bigelow, 1909 Amphinema dinema (Peron & LeSueur,
P. martagon Haeckel, 1879 1809)
P. rubiginosa (Kolliker, 1853) Halitholus intermedius (Browne, 1902)
P. triloba Haeckel, 1879 Pandea conica (Quouy & Gaimard, 1827)
Solmaris corona (Keferstein & Ehlers,
1861) Family Calycopsidae
Calycopsis borchgrevinki (Browne, 1910)
Family Cuninidae Heterotiara anonyma Maas, 1905
Cunina duplicata Maas, 1893
C. frugifera Kramp, 1948 Suborder Leptomedusae
C. octonaria McCrady, 1857 Family Laodiceidae
C. peregrina Bigelow, 1909 Laodicea undulata (Forbes & Goodsir,
1853)
Family Aeginidae
Aegina citrea Eschscholtz, 1829 Family Mitrocomidae
Solmundella bitentaculata Quoy & Cosmetira pilosel/a Forbes, 1848
Gaimard, 1833
Family Eurenidae
Order Hydroida Phialopsis diegensis Torrey, 1909
Appendix 229

Family Phialuciidae S. turgida (Gegenbaur, 1853)

Phialucium carolinae (Mayer, 1900)
Family Abylidae
Family Lovenellidae Abyla trigona Quoy & Gaimard, 1827
Eucheilota ventricularis McCrady, 1857 Abylopsis eschscholtzi (Huxley, 1859)
A. tetragona (Otto, 1823)
Family Campanullaridae Bassia bassensis (Quoy & Gaimard,
Obelia spp. 1834)
Phialidium discoidum (Mayer, 1900) Ceratocymba leuckarti Huxley, 1859
P. ovale (Mayer, 1900) C. sagittata Quoy & Gaimard, 1827
P. simplex Browne, 1902 Enneagonum hyalinum (Quoy &
Gaimard, 1827)
Family Plumulariidae
Kirchenpaueria sp. CLASS SCYPHOZOA
Order Semaeostomeae
Order Siphonophora Family Pelagiidae
Suborder Cystonectae Chrysaora hysoscella (Linnaeus, 1766)
Family Physalidae C. lactea Eschscholtz, 1829
Physalia physalis (Linnaeus, 1759) C. quinquecirrhg (Desor, 1848)

Suborder Physonectae Family Ulmaridae

Family Agalmidae Aurelia aurita (Linnaeus, 1746)
Marrus antarticus Totton, 1954
Stephanomia bijuga (Delle Chiaje, 1841) Family Cyaneidae
Drymonema gorgo F.M tiller, 1883
Suborder Calycophorae
Family Prayidae Order Rhizostomeae
Amphicaryon ernesti Totton, 1954 Family Lychnorhizidae
Lychnorhiza lucerna Haeckel, 1880
Family Hippopodiidae
Hippopodius hippopus (Forskal, 1776) CLASS ANTHOZOA
Vogtiaglabra Bigelow, 1918 SUBCLASS OCTOCORALLIA
Order Gorgonacea (=Alcyonaria)
Family Diphyidae Family Anthothelidae
Chelophyes appendiculata (Eschscholtz, Tripalea clavaria (Studer, 1878)
1829)
C. contorta (Lens & van Riemsdijk, Family Plexauridae
1908) Muricea midas (Bager, 1959)
Dimophyes arctica (Chun, 1897) Muricea sp.
Diphyes bojani (Eschscholtz, 1829) Muriceides hirtus (Pourtales, 1867)
D. dispar Chamisso & Eisenhardt, 1821 Muriceides sp.
Diphyopsis mitra (Huxley, 1859)
Eudoxoides spiralis (Bigelow, 1911) Family Elliselidae
Lensia campanella (Moser, 1925) Elisella barbadensis (Duchassaing &
L. conoidea (Keferstein & Ehlers, 1860) Michelotti,1864)
L. cossack Totton, 1941 E. elongata (Pallas, 1766)
L. grimaldi (Leloup, 1933)
L. hardy Totton, 1941 Family Primnoidae
L. metegri (Leloup, 1934) Primnoella biserialis Wright & Studer,
L. subtilis (Chun, 1886) 1889
Muggiaea kochi (Wild, 1844) P. compressa Kukenthal, 1908
Sulculeolaria biloba (Sars, 1846) P. distans Studer, 1878
S. chuni (Lens & van Riemsdijk, 1908) P. magelhaenica Studer, 1878
S. quadrivalvis Blainville, 1834 P. philippii Aurivillius, 1931
230 Appendix

Family Acanthogorgidae Deltocyathus conicus Zibrowius, 1980

Acanthogorgia scharammi(Duchassaing Lophelia pertusa (Linnaeus, 1758)
& Michelotti, 1864)
Family FlabelIidae
Family Paramuriceidae Flabellum brasiliensis
Bebryce cinerea (Deichamann, 1936) Flabellum spp.
Javania cailleti (Duchassaing &
Family Gorgoniidae Michelotti, 1846)
Leptogorgia setacea (Pallas, 1766)
Family Dendrophylliidae
Family Isididae Rhizopsammia manuelensis Chevalier,
Acanella eburnea (Pourtales, 1868) 1966

Order Pennatulacea Order Antipatharia

Family Renillidae Family Antipathidae
Renilla kollikeri Antipathes sp.
R. mulleri Kolliker, 1872
R. musaica PHYLUM NEMERTINA
R. reniformis (Pallas, 1766) Family Ototyphlonemertidae
R. tentaculata Ototyphlonemertes sp.

SUBCLASS HEXACORALLIA (= PHYLUM PLATYHELMINTHES

ZOANTHARIA) CLASS TURBELLARIA
Order Zoanthidea Order Polycladida
Family Zoanthidae Family Stylochidae
Epizoanthus sp. Stylochoplana sp.
Stylochus sp.
Order Actiniaria
Family Hormathiidae PHYLUM SIPUNCULIDA
Actinauge longicornis Verril, 1882 Family Sipunculidae
Phelliactis lophohelia Riemann- Sipunculus nudus
Zlirneck, 1973
P. pelophyla Riemann-Zlirneck, 1973 Family Golfingiidae
Golfingia sp.
Family Actiniidae Nephasomma sp.
Bunodosoma caicarum Themistes sp.
B. cangicum
Phlyctenanthus australis Carlgren, 1949 PHYLUM ECHIURIDA
Family Echiuridae
Family Edwardsiidae Echiurus sp.
Edwardsia sp.
Family Bonelliidae
Order Scleractinia Bonellia sp.
Family Faviidae
Cladocora debilis Milne-Edwards & PHYLUM PRIAPULIDA
Haime, 1849 Priapulus sp.

Family Rhizangiidae PHYLUM ANNELIDA

Astrangia rathbuni Vauhan, 1906 CLASS POLYCHAETA
Order Orbiniida
Family Caryophylliidae Family Orbiniidae
Caryophyllia smithii Stokes & Broderip, Phylo felix Kinberg, 1866
1828 Scoloplos (= Leodomas) ohlini (Ehlers,
Dasmosmilia lymani Pourtales, 1880 1901)
Appendix 231

Family Paraonidae Order Phyllodocida

Aedicira belgicae (Fauvel, 1936) Family Phyllodocidae
Anaitides mucosa (Oersted)
Order Cossurida Anaitides sp.
Family Cossuridae Dodecaceria spp.
Cossura candida Hartman Eulalia spp.
Phyllodoce mucosa
Order Spionida Polynoe sp.
Family Spionidae
Apoprionospio pygmaea (Hartman, Family Alciopidae
1961) Alciopa reynaudii Audouin & Milne-
Boccardia hamata (Webster, 1879) Edwards, 1833
Dispio sp. Alciopina parassitica ClaparMe &
Laonice cirrata (Sars, 1851) Panceri, 1867
Paraprionospio pinnata (Ehlers, 1901) Plotohelmis capitata (Greeff, 1876)
Polydora ligni Webster, 1879 Rhynchonereella gracilis Costa, 1862
Prionospio steenstrupi Malmgren, 1867 R. petersii (Langerhans, 1880)
Spio gaucha Orensanz & Gianuca, 1974 Vanadis crystallina Greeff, 1876
Spiophanes bombyx (Claparede, 1870) v. formosa Claparede, 1870
S. kroeyeri (Grube, 1860) V. longissima Levifisen, 1885
S. soederstroemi Hartman, 1953 V. minuta Treadwell, 1906
V. studeri Apstein, 1893
Family Magelonidae
Magelona riojai Jones, 1963 Family Aphroditidae
Aphrodita longicornis Kinberg, 1855
Family Poecilochaetidae Aphroditella alta (Kinberg, 1855)
Poecilochaetus australis Nonato, 1963 Laetmonice parva Amaral & Nonato,
1982
Family Chaetopteridae Lamelleulepethus orensanzil Pettibone,
Phyllochaetopterus socialis 1986
Spiochaetopterus spp.
Family Polynoidae
Family Cirratulidae Euphionella besnardi Amaral & Nonato,
Dodecaceria sp. 1982
Halosydnella australis (Kinberg, 1855)
Order Capitellida H. brasiliensis (Kinberg, 1858)
Family Capitellidae Lagisca lepida Amaral & Nonato, 1982
Heteromastus similis Southern, 1921 Lepidastenia esbelta Amaral & Nonato,
Notomastus lobatus Hartman, 1947 1982
Lepidastenia sp.
Family Maldanidae
Asychis amoena Kinberg Family Polyodontidae
A. atlanticus Kinberg Eupanthalis kinbergi McIntosh, 1876
A. brasiliensis Kinberg E. rudipalpa Amaral & Nonato, 1984
Euclymene sp. Panthalis bicolor Grube, 1877
Nicomache sp. Polyodontes oculea (Treadwell, 1901)
Rhodine sp. P. oerstedi Kinberg, 1855

Order Opheliida Family Pholoididae

Family Opheliidae Pholoe minuta (Fabricius, 1780)
Ammotrypane trigintae Hartman, 1953
Ammotrypane sp. Family Eulepethidae
Euzonus furciferus (Ehlers, 1897) Grubeulepis bracteata Nonato
Travisia forbesii Johnston, 1840
232 Appendix

Pareulepis multibranchiata Amaral & Nephtys acrochaeta Hartman, 1950

Nonato, 1984 N.fluviatilis Monro, 1937
N. simoni Perkins, 1980
Family Sigalionidae N. squamosa Ehlers
Ehlersileanira incisa (Grube, 1877)
Psammolyce flava Kinberg, 1855 Family Tomopteridae
Sigalion cirriferum Orensanz & Gianuca, Tomopteris elegans Chun, 1888
1974 T. kefersteini Greeff, 1879
Sthenelais limicola (Ehlers, 1901) T. planktonis Apstein, 1900
Sthenolepis oculata (Hartman, 1942) T. rolasi Greeff, 1885
Thalenessa edwardsi (Kinberg, 1855) T. septentrionalis Quatrefages, 1865

Family Chrysopetalidae Family Typhloscolecid~e

Dysponetus sp. Sagitella kowalevskii N.Wagner, 1872
Paleo notus heteroseta Hartman, 1945 Travisiopsis dubia Stop-Bowitz, 1948
T. levinseni Southern, 1910
Family Pilargiidae T. lobifera Levinsen, 1885
Parandalia americana (Hartman, 1947) Typhloscolex muelleri Busch, 1851
Sigambra grubii Fritz Muller, 1858
Order Amphinomida
Family Syllidae Family Amphinomidae
Odontosyllis hetero1alcata Temperine, Amphinome rostrata (Pallas, 1766)
1981 Hipponoa gaudichaudi agulhana Day,
Syllis prolixa Ehlers, 1901 1967
Typosyllis ehlersioides Merenzeller, 1890
Order Eunicida
Family Nereidae Family Onuphidae
Eunereis patagonica (Mcintosh, 1885) Americonuphis casamiquelorum
Laeonereis acuta (Treadwell, 1923) Orensanz, 1974
Lycastopsis pontica (Bobretzky, 1872) Diopatra cup rea (Bose, 1802)
Neanthes bruaca Lana & Sovierzoski, D. tridentata Hartman, 1944
1987 D. viridis Kinberg, 1865
N. succinea (Frey & Leuckart, 1847) Hyalonecia juvenalis Moore, 1911
Nereis falsa Quatrefages, 1865 H. tubicola (Muller, 1788)
Kinbergonuphis difficilis (Fauchauld,
Family Glyceridae 1982)
Glycera americana Leidy, 1855 Nothria benthophyla Temperini, 1981
Hemipodus olivieri Orensanz & Onuphis eremita Audouin & Milne-
Gianuca,1974 Edwards, 1833
O. eremita oculata Hartman, 1951
Family Goniadidae O. setosa Orensanz, 1974
Glycinde multidens Fritz Muller, 1858 O. tenuis Hansen, 1881
Goniada littorea Hartman, 1950 Ramphobrachium verngreni (Kinberg,
G. maculata Oersted 1865)
Ophioglycera eximia (Ehlers, 1900)
Family Eunicidae
Family Iospilidae Eunice argentinensis (Treadwell, 1929)
Iospilus affinis (Viguier, 1911) E. frauenfeldi Grube, 1868
Phalacrophorus pictus Greeff, 1879 E. rubra Grube, 1856
P: uniformis Reibisch, 1893 E. vittata (Delle Chiaje, 1828)
Marphysa conferta Moore
Family Nephtyidae M. disjuncta Hartman
Aglaophamus praetiosus (Kinberg, 1866) M. kinbergi Mcintosh, 1910
A. uruguayi Hartman, 1953 M. sanguinea (Montagu, 1815)
Appendix 233

Family Lumbrineridae Family Ampharetidae

Aglaurides fulgida (Savigny) Amphicteis latibranchiata Nonato
Arabellonereis janeiroensis (Augener, Melinna uruguayi Hessle, 1917
1934)
Lumbrinerides gesae Orensanz, 1973 Family Terebellidae
Lumbrineriopsis nucronata (Ehlers, Artacama benedeni Kinberg, 1867
1908) Telephus setosus (Quatrefages, 1865)
Lumbrineris atlantica Kinberg, 1865
L. cingulata (Ehlers, 1897) Family Trichobranchidae
L. latreilli Audouin & Milne-Edwards, Terebellides stroemi Sars
1834 Tricobranchus lobiungens Hessle, 1917
L. magalhaensis Kinberg, 1865
L. tetraura (Schmarda, 1861) Order Sabellida
Ninoe brasiliensis Kinberg, 1865 Family Sabellidae
N. gayheadia Hartman, 1965 Chone infundibiliformis Kroyer, 1856
Ninoe sp. Megalomma sp.
Paraninoe antarctica (Monro, 1930) Sabellaria bella Grube, 1870

Family Arabellidae Family Serpulida,~

Arabella iricolor (Montagu, 1804) Filograna implexa Berkeley, 1827
Drilonereis filum (Claparede, 1868) Hydro ides parvus Treadwell, 1901
D. simplex Orensanz, 1974 Omphalopoma sp.
D. tenuis (Ehlers, 1900) Protula sp.
Notocirrus lorum Ehlers, 1897
Family Protodrilidae
Family Lysaretidae Protodrilus sp.
Lysarete brasiliensis Kinberg, 1865
PHYLUM MOLLUSCA
Family Dorvilleidae CLASSPOLYPLACOPHORA
Pettiboneia sanmatiensis Orensaz, 1973 Order Neoloricata
Family Ischnochitonidae
Order Sternaspida Chaetopleura angulata (Splenger, 1797)
Family Sternaspidae C. isabellei (Orbigny, 1841)
Sternaspis sp.
CLASS GASTROPODA
Order Oweniida SUBCLASS PROSOBRANCHIA
Family Oweniidae Order Archeogastropoda
Owenia fusiformis Delle Chiaje, 1844 Family Pleurotomariidae
Owenia spp. Mikadotrochus notialis Leme & Penna,
1969
Order Flabelligerida Perotrochus amabilis Bayer, 1963
Family Flabelligeridae P. atlanticus Rios & Matthews, 1968
Pherusa capitata N onato, 1966
P. scutigera Ehlers Family Scissurellidae
Anatoma aedonia (Watson, 1886)
Order Terebellida
Family Sabellariidae Family Fissurellidae
Phragmatopoma sp. Diodora patagonica (Orbigny, 1847)
Sabellaria bellis Hansen, 1882 Emarginula tuberculosa Libassi, 1859
Fissurellidea megatrema (Orbigny,
Family Pectinariidae 1841)
Cystenides sp. Lucapinella henseli (Martens, 1900)
Pectinaria laelia Nonato, 1981 L. limatula (Reeve, 1850)
Puncturella granulata Seguenza, 1863
234 Appendix

Family Acmaeidae Crepidula aculeata (GmeIin, 1791)

Collisella subrugosa (Orbigny, 1846) C. pro tea Orbigny, 1835

Family Trochidae Family Xenophoridae

Calliostoma axelolssoni Quinn, 1992 Xenophora caribaea Petit, 1856
C. carcellesi Clench & Aguayo, 1939
C. coppingeri (E.A.Smith, 1880) Family Ovulidae
C.jucundum (Gould, 1849) Primovula vanhyningi (M.Smith, 1940)
C. militaris (Ihering, 1907)
Calliotropis actinophora (DaII, 1890) Family LameIIariidae
C. calatha (DaIl, 1927) Lamellaria patagonica E.A.Smith, 1881
Halistylus columna DaIl, 1890
Mirachelus clinocnemus Quinn, 1979 Family N aticidae
Solariella lubrica DaII, 1881 Natica limbata Orbigny, 1840
S. patriae CarceIIes, 1953 Polinices hepaticus (Roding, 1798)
Tegula patagonica (Orbigny, 1840) Polin ice sp.

Family Turbinidae Family Tonnidae

Arene microforis (DaII, 1889) Tonna galea Brunnich, 1772
A. variabilis (DaII, 1889)
Family Cassidae
Family Seguenziidae Phalium granulatum (Born, 1778)
Ancistrobasis costulata (Watson, 1879) P. labiatum iheringi CarceIIes, 1953

Order Mesogastropoda Family RaneIlidae

Family Hidrobiidae Cabestana felipponei (Ihering, 1907)
Heleobia australis (Marcus & Marcus, Cymatium parthenopeum (von Salis,
1963) 1793)
H. parchapei (Orbigny, 1835) Fusitriton magellanicus (Roding, 1798)

Family Littorinidae Family Triphoridae

Littorina ziczac (Gmelin, 1791) Triphora pulchella (C.B.Adams, 1850)

Family Barleeidae Family Epitoniidae

Barleeia rubrooperculata (CasteIIanos, Cylindriscala andrewsii (Verril, 1882)
1927) C. watsoni (de Boury, 1911)
Epitonium albidum (Orbigny, 1842)
Family Caecidae E. angulatum (Say, 1830)
Caecum achironum (Folin, 1867) E. candeanum (Orbigny, 1842)
C. antillarum Carpenter, 1857 E. celesti (Aradas, 1854)
Mioceras nitidum (Stimpson, 1851) E. dallianum Verril & Smith, 1880
E. georgettina (Kiener, 1839)
Family VitrineIlidae E. humpheysii (Kiener, 1838)
Solariorbis shimeri (Clapp, 1914) E. magellanicum (Philippi, 1845)
E. pourtalesii Verril & Smith, 1880
Family Tornidae E. tenuistriatum (Orbigny, 1840)
Macromphalina argentina Castellanos, E. unifasciatum (Sowerby, 1844)
1975
Family Eulimidae
Family Fossaridae Eulima auricincta Abbot, 1959
Fossarus perplexa (DaII, 1927) E. bifasciata (Orbigny, 1842)
Hemiaclis aff. georgiana (DaII, 1927)
Family Calyptraeidae Melanella arcuata (C.B.Adams, 1850)
Calyptraea centralis (Conrad, 1841) M. conoidea (Kurtz & Simpson, 1851)
Appendix 235

Niso aeglees Bush, 1895 O. deshayesiana (Dudos, 1857)

o. urceus (Roding, 1798)
Order Neogastropoda O.uretaiKlappenbach,1965
Family Muricidae Olivella formicacorsii Klappenbach,
Chicoreus beauii (Fischer & Bernardi, 1962
1857) O. orejasmirandai Klappenbach, 1986
Murexiella hidalgoi (Crosse, 1869) o. plata (Ihering, 1909)
Muricopsis necocheana (Pilsbry, 1900) O. puelcha (Dudos, 1840)
Oraniafusulus (Brocchi, 1814) O. riosi Klappenbach, 1991
Trophon acanthodes Watson, 1882 O. tehuelcha (Dudos, 1840)
T. pelseneeri E.A.Smith, 1915
T. plicatus (Lightfoot, 1786) Family Marginellidae
Typhis cleryi (Petit, 1842) Granulina ovuliformis Orbigny, 1841
T. riosi Bertsch & d'Attilio, 1980 Prunum martini (Petit, 1853)
Urosalpix cala (Pilsbry, 1897) P. rubens (Martens, 1881)
U. haneti (Petit, 1856)
Family Mitridae
Family Thaididae Mitra larrangai (Carcelles, 1947)
Thais haemastoma (Linnaeus, 1767)
Family Conidae
Family Coralliophilidae Conus clerii Reeve, 1844
Coralliophila aedonius (Watson, 1886) C. mazei Deshayes, 1874
C. tostesi Petuch, 1986
Familiy Columbellidae C. villepinii Fisher & Bernardi, 1867
Aesopus metcalfei (Reeve, 1858)
Amphissa acuminata (E.A.Smith, 1915) Family Volutidae
A. cancellata (Castellanos, 1982) Adelomelon ancilla (Lighfoot, 1786)
Anachis isabellei (Orbigny, 1841) A. beckii (Broderip, 1836)
A. sertulariarum (Orbigny, 1841) A. brasiliana (Lamarck, 1811)
A. riosi Clench & Turner, 1964
Family Nassariidae Nanomelon vossi Leal & Rios, 1990
Buccinanops deformis (King, 1831) Odontocymbiola americana (Reeve,
B. duartei Klappenbach, 1961 1856)
B. gradatum (Deshayes, 1844) O. corderoi (Carcelles, 1953)
B.lamarckii (Kiener, 1834) Zidona dufresnei (Donovan, 1823)
B. uruguayensis (Pilsbry, 1897)
Dorsanum moniliferum (Valenciennes, Family Turridae
1834) Benthomangelia macra (Watson, 1881)
Nassarius scissuratus (Dall, 1889) Brachytoma rioensis (E.A.Smith, 1915)
Carinodrillia braziliensis (E.A.Smith,
Family Fasciolariidae 1915)
Fusinus frenguelli (Carcelles, 1953) Cochlespira radiata (Dall, 1889)
Metula anfractura Matthews & Rios, Cryoturris adamsi (E.A.Smith, 1884)
1968 Fusiturricula lavinoides limonensis
(Olsson, 1922)
Family Vasidae F. taurina (Olsson, 1922)
Fulgurofusus coronatum (Penna-Neme Kurtziella dorvillae (Reeve, 1845)
& Leme, 1978) K. padolina (Fargo, 1953)
Leucosyrinx dalli (Busch, 1893)
Family Ofividae L. hemimeres(Watson, 1881)
Ancilla dimidiata (Sowerby, 1850) Oenopota magellanica (Martens, 1881)
Olivancillaria auricularia (Lamarck, Pleurotomella aguayoi (Carcelles, 1953)
1810) P. cala (Watson, 1886)
O. carcellesi Klappenbach, 1965 P. ipara (Dall, 1881)
236 Appendix

P. meriacum (Locard, 1896) Philine falklandica Powell, 1951

Polystira formosissima (E.A.Smith,
1915) Family Bullidae
Spirotropis lithocolleta (Watson, 1881) Bulla striata Bruguiere, 1792
S. patagonica (Orbigny, 1841)
S. stirophora (Watson, 1881) Family Retusidae
Splendrillia carolinae (Bartsch, 1934) Pyrunculus caelatus (Bush, 1885)
Veprecula morra (DaIl, 1881) Volvulella paupercula (Watson, 1883)
V. persimilis (Morch, 1875)
Family Terebridae
Terebra doello-juradoi Carcelles, 1953 Order Thecosomata
T. gemmulata Kiener, 1839 Family Limacinidae
Limacina bulimoides (Orbigny, 1836)
SUBCLASS HETEROBRANCHIA L. inflata (Orbigny, 1836)
Order Heterostropha L. lesueuri (Orbigny, 1836)
Family Architectonicidae L. retroversa f. retroversa (Fleming,
Heliacus bisulcatus (Orbigny, 1845) 1823)
Philippia uruguaya (CarceIles, 1953) L. trochiformis (Orbigny, 1836)
Pseudomalaxis nobilis Verril, 1885
Family Cavoliniidae
Family Pyramidellidae Cavolinia gibbosa f. flava (Orbigny,
Eulimastoma engonium surinamense 1836)
Altena, 1975 C. inflexa(LeSueur,1813)
Odostomia canaliculata C.B.Adams, C. inflexa £ inflexa (LeSueur, 1813)
1850 C. inflexa £ imitans (Pfeffer, 1880)
O. jadisi Olsson & McGinty, 1958 C. inflexa f.labiata (Orbigny, 1836)
O. seminuda (C.B.Adams, 1837) C. longirostris f. longirostris (Blainville,
Turbonilla americana (Orbigny, 1840) 1821)
T. atypha Bush, 1900 C. longirostris £ limbata (Orbigny, 1836)
T. dispar Pilsbry, 1897 C. longirostris f. strangulata (Deshayes,
T.fasciata (Orbigny, 1840) 1823)
T. interrupta (Totten, 1835) C. tridentata (Niebuhr, 1775)
T. rushii Bush, 1899 C. uncinata (Rang, 1829)
T. uruguayensis Pilsbry, 1897 C. uncinata f. uncinata (Rang, 1829)
Clio cuspidata (Bose, 1802)
Family Amathinidae C. pyramidata f. antartica (Dall, 1908)
Iselica anomala (C.B.Adams, 1850) C. pyramidata f. lanceolata (LeSueur,
1813)
SUBCLASS OPISTHOBRANCHIA Creseis. acicula Rang, 1828
Order Cephalaspidea C. acicula f. acicula (Rang, 1828)
Family Acteonidae C. acicula f. clava (Rang, 1828)
Acteon pelecais Marcus, 1981 C. virgula f. conica Eschscholtz, 1929
Mysouffa cumingii (A.Adams, 1854) C. virgula f. virgula (Rang, 1828)
Cuvierina columnella £ atlantica Van
Family Cylichnidae der Spoel, 1970
Acteocina bidentata (Orbigny, 1841) Diacria rampali Dupont, 1979
A. candei (Orbigny, 1842) D. trispinosa f. trispinosa (Blainville,
A. inconspicua Olsson & McGinty, 1958 1821)
Cyliclma aff. crispula Watson, 1883 Hyalocylis striata (Rang, 1828)
C. discus Watson, 1883 Styliola subula (Quoy & Gaimard, 1827)
C. verrillii Dall, 1889
Scafander watsoni Dall, 1881 Order Nudibranchia
Family Tritoniidae
Family Philinidae Marionia cucullata (Gould, 1852)
Appendix 237

Tritonia eriosi Ev.Marcus, 1983 Brachidontes exustus (Linnaeus, 1758)

B. rodriguezi (Orbigny, 1846)
Order Basommatophora B. solisianus (Orbigny, 1846)
Family Siphonariidae Crenella divaricata (Orbigny, 1846)
Siphonaria lessoni (Blainville, 1824) Lithophaga patagonica (Orbigny, 1846)
Modiolus carvalhoi Klappenbach, 1966
CLASS PELECYPODA Musculus viator (Orbigny, 1846)
SUBCLASSPROTOBRANCHIA Mytilus edulis platensis (Orbigny, 1846)
Order Solemyoida Perna perna (Linnaeus, 1758)
Family Solemyidae
Solemya patagonica E.A.Smith, 1885 Order Pterioida
Family Pteriidae
Order Nuculoidea Pteria hirundo (Linnaeus, 1758)
Family Nuculidae
Nucula puelcha Orbigny, 1846 Family Pinnidae
N. semiornata Orbigny, 1846 Atrina seminuda (Orbigny, 1846)

Family Tindariidae Order Limoida

Tindaria striata (King, 1831) Family Limidae
Limatula pigmaea (Philippi, 1845)
Family Nuculanidae
Adrana electa (A.Adams, 1846) Order Ostreoida
A. patagonica (Orbigny, 1846) Family Ostreidae
Nuculana fortiana Esteves, 1984 Ostrea equestris Say, 1834
N. larranagai Klappenbach & Scarabino, O. puelchana (Orbigny, 1841)
1968
Family Pectinidae
Family Yoldiidae Chlamys felipponei (Dall, 1922)
Yoldia riograndensis Esteves, 1984 C. patagonica (King, 1832)
C. tehuelchus (Orbigny, 1846)
Family Malletiidae Cyclopecten leptaleus (Verrill, 1884)
Malletia cumingii (Hanley, 1860) Leptopecten bavayi (Dautzenberg, 1900)

SUBCLASS PTERIOMORPHA Family Propeamussiidae

Order Arcoida Propeamussium pourtalesianum (Dall,
Family Arcidae 1886)
Anadara chemnitzi (Philippi, 1851)
A. ovalis (Bruguiere, 1789) Family Anomiidae
Anomia ephippium Linnaeus, 1758
Family Noetiidae Pododesmus leloiri Carcelles, 1941
Noetia bisulcata (Lamarck, 1819) P. rudis (Broderip, 1834)

Family Limopsidae SUBCLASS HETERODONTA

Limopsis davinae Esteves, 1984 Order Veneroida
L. janeiroensis E.A.Smith, 1915 Family Lucinidae
L. minuta (Philippi, 1836) Myrtea lens (Verrill & Smith, 1880)

Family Glycymerididae Family Thyasiridae

Glycymeris longior (Sowerby, 1833) Thyasira croulinensis Jeffreys, 1874

Order Mytiloida Family Ungulinidae

Family Mytilidae Diplodonta patagonica (Orbigny, 1842)
Amygdalum dendriticum Muhlfels, 1811 Felaniella vilardeboana (Orbigny, 1846)
A. sagitta tum Rehder, 1934
238 Appendix

Phlyctiderma semiaspera (Philippi, Family Psammobiidae

1836) Tagelus plebeius (Lightfoot, 1786)

Family Carditidae Family Donacidae

Carditamera floridana Conrad, 1838 Donax gemmula Morrison, 1971
C. plata (Ihering, 1907) D. hanleyanus Philippi, 1842
Cyclocardia moniliata (Dan, 1902)
Pleuromeris sanmartini Klappenbach, Family Veneridae
1971) Amiantis purpuratus (Lamarck, 1818)
Callista eucymata (Dall, 1889)
Family Condylocardiidae Chione paphia (Linnaeus, 1767)
Americuna besnardi Klappenbach, 1962 Ciausinellagayi (Hupe, 1854)
Carditopsis smithii (Dan, 1896) Eurhomalea exalbida (Dill. 1817)
Pitar rostratus (Koch, 1844)
Family Chamidae Protothaca antiqua (King & Broderip,
Arcinella brasiliana (Nicol, 1953) 1835)
Tivela isabelleana (Orbigny, 1846)
Family Crassatellidae T. ventricosa (Gray, 1838)
Crassatella riograndensis Vokes, 1973 Transenpitar americana (Doello-Jurado,
Crassinella lunulata (Conrad, 1834) 1951)
C. marplatensis Castellanos, 1970
Family Ptricolidae
Family Cardiidae Petricola lapicida (Gmelin, 1791)
Trachycardium muricatum (Linnaeus, P. stellae Narchi, 1975
1758)
Order Myoida
Family Mactridae Family Corbulidae
Mactra isabelleana Orbigny, 1846 Corbula caribaea Orbigny, 1842
M. janeiroensis Smith, 1915 C. lyoni Pilsbry, 1897
M. marplatensis Doeno-Jurado, 1918 C. patagonica Orbigny, 1846
M. patagonica Orbigny, 1846 C. tryoni E.A.Smith, 1880
M. petiti Orbigny, 1846
Mulinia aff. branneri Dall, 1901 Family Erodonidae
Raeta plicatella (Lamarck, 1818) Erodona mactroides Bose, 1802

Family Mesodesmatidae Family Hiatellidae

Mesodesma mactroides Deshayes, 1854 Panopea abbreviata Valenciennes, 1839

Family Solenidae SUBCLASS ANOMALODESMATA

Solen tehuelchus Orbigny, 1843 Order Pholadina
Family Pholadidae
Family Teninidae Barnea lamellosa (Orbigny, 1846)
Macoma uruguayensis (E.A.Smith, 1885) Cyrtopleura lanceolata (Orbigny, 1846)
Strigilla carnaria (Linnaeus, 1758) Pholas campechiensis Gmelin, 1791
Tellina alerta Boss, 1964
T. gibber Ihering, 1907 Order Pholadomyoida
T. petitiana Orbigny, 1846 Family Thraciidae
T. sandix Boss, 1968 Asthenothaerus rushii (Pilsbry, 1987)
Thracia distorta (Montagu, 1808)
Family Semelidae T. similis Couthony, 1839
Abra lioica (Dall, 1881)
A. uruguayensis (Pilsbry, 1897) Family Periplomatidae
Semele casali Doeno-Jurado, 1949 Periploma compressa Orbigny, 1846
S. proficua (Pulteney, 1799) P. ovata Orbigny, 1846
Appendix 239

Family Lyonsiidae Ornithoteuthis antillarum Adam, 1957

Entodesma alvarezi Orbigny, 1846 O. volatilis Sasaki, 1915
E. patagonica (Orbigny, 1846) Ommastrephes bartramii (LeSueur,
1821)
Family Pandoridae Todarodes angolensis Adam, 1962
Pandora braziliensis Sowerby, 1874
P. bushiana Dall, 1886 Family Thysanoteuthidae
Thysanoteuthis rhombus Troschel, 1857
Family Cuspidariidae
Cardiomya cleryana (Orbigny, 1846) Family Onychoteuthidae
Cuspidaria platensis E.A.Smith, 1915 Moroteuthis ingens (Smith, 1881)
M. robsoni Adam, 1962
Family Verticordiidae Onychia caribaea LeSueur, 1821
Euciroa grandis (E.A.Smith, 1885) Onychoteuthis banksi (Leach, 1817)
Verticordia acuticostata (Philippi, 1844)
V. fisheriana Dall, 1881 Family Enoploteuthidae
V. ornata (Orbigny, 1842) Abralia redfieldi (Voss, 1955)
A. veranyi (Ruppel, 1844)
CLASS SCAPHOPODA Abraliopsis pfefferiJoubin, 1896
Order Dentaliida Ancistrocheirus leusueri (Vurany, 1851)
Family Dentaliidae Pterygioteuthis giardi Fischer, 1896
Antalis cera tum (Dall, 1881)
A. infractum (Odhner, 1931) Family Histioteuthidae
Fissidentalium amphialum (Watson, Histioteuthis atlantica (Hoyle, 1885)
1879) H. dofleini (Pfeffer, 1912)
F. carduum (Dall, 1889)
Family Cranchiidae
Order Gadilida Liocranchia reinhardti (Steenstrup,
Family Entalinidae 1856)
Pertusiconcha callithrix (Dall, 1889) Megalocranchia sp.

Family Gadilidae Family Chiroteuthidae

Cadulus braziliensis Henderson, 1920 Chiroteuthis veranyi (Furussac, 1835)
Polyschides tetraschistus (Watson, 1879)
Family Lycoteuthidae
CLASS CEPHALOPODA Lycoteuthis diadema (Chun, 1900)
Order Sepioidea
Family Spirulidae Order Octopoda
Spirula spirula Lamarck, 1801 Family Octopodidae
Eledone gaucha Haimovici, 1988
Family Sepiolidae E. massyae Voss, 1964
Heteroteuthis atlantis Voss, 1955 Octopus tehuelchus Orbigny, 1834
Semirossia ten era (Steenstrup, 1887) O. vulgaris (Cuvier, 1797)
Pareledone charcoti (Joubin, 1905)
Order Teuthoidea P. turqueti (Joubin, 1905)
Family Loliginidae Pteroctopus tetracirrhus Delle Chiaje,
Loligo plei (Blainville, 1823) 1830
L. sanpaulensi5 Brakoniecki, 1984 Scaeurgus unicirrhus (Orbigny, 1840)
Vosseledone charrua Palacio, 1978
Family Ctenopterygidae
Ctenopteryx siculus Family Bolitaenidae
Eledonella pigmaea Verril, 1884
Family Ommastrephidae Japetella diaphana Hoyle, 1885
Illex argentinus (Castellanos, 1960)
240 Appendix

Family Allopsidae Family Daphniidae

Allopsus mollis Verril, 1880 Ceriodaphnia cornuta cornuta (Sars,
1886)
Family Argonautidae C. comuta rigaudi (Richard, 1894)
Argonauta argo Linnaeus, 1758 C. dubia Richard, 1894
A. hians Solander, 1786 C. reticulata (Jurine, 1820)
A. nodosa Lightfoot, 1786 Daphnia ambigua Scourfield, 1946
Scapholeberis spinifera (Nicolet, 1879)
Family Tremoctopodidae Simocephalus exspinosus (Koch, 1841)
Tremoctopus violaceous Della Chiaje, S. serrulatus Schoedler, 1858
1830 S. vetulus (O.F.MUller, 1776)

Family Ocythoidae Family Bosminidae

Ocythoe tuberculata Rafinesque, 1814 Bosminopsis deitersi Richard, 1895
Eubosmina tubicen (Brehm, 1953)
PHYLUM ARTHROPODA
SUBPHYLUM CHELICERATA Family Macrothricidae
CLASS ARACHNIDA Echinisca triserialis Brady, 1886
Order Araneae Ilyocryptys spinifer Herrick, 1884
Family Licosidae 1. verrucosus Daday, 1905
Glieschiella (==Moenkhausiana) Macrothrix laticomis (Jurine, 1820)
halophila (Mello-Leitao, 1932)
Family Chydoridae
CLASS PYCNOGONIDA Acroperus harpae (Baird, 1834)
Family Tanystylidae Alana costata Sars, 1862
Tanystylum orbiculare Wilson, 1878 A. davidi davidi Richard, 1895
A. monacantha monacantha Sars, 1901
Family Colossendeidae Alonella dentifera Sars, 1901
Colossendeis geoffreyi Mane-Garzon, Biapertura affinis affinis (Leydig, 1860)
1944 B. karua (King, 1852)
B. pseudoverrucosa verrucosa Sars, 1901
SUBPHYLUM CRUSTACEA Camptocercus australis Sars, 1896
CLASSBRANCHIOPODA Chydorus ciliatus Poggenpol, 1874
Order Cladocera C. eurynotus eurynotus Sars, 1901
Family Polyphemidae C. faviformis Birge, 1893
Evadne nordmanni Loven, 1835 C. sphaericus sphaericus (O.F.MUller,
E. spinifera MUller, 1868 1785)
Pleopis polyphemoides (Leuckart, 1859) Dunhevedia odontoplax Sars, 1901
Podon intermedius Lilljeborg, 1859 Euryalona orientalis Daday, 1898
P. schmackeri (Poppe) Eurycercus lamellatus (O.F.MUller,
Pseudoevadne tergestina (Claus, 1877) 1785)
Graptoleberis testudinaria (Fisher, 1851)
Family Sididae Kurzia latissima (Kurz, 1875)
Diaphanosoma brachyurum (Lievin, Pleuroxus denticulatus Birge
1848) P. similis Vavra, 1900
D. brevireme Sars, 1901 P. striatus Schoedler, 1858
D. fluviatile Hansen, 1899 P. truncatus truncatus (O.F.MUller,
D. sarsi Richard, 1895 1785)
~enilia avirostris Dana, 1852
Pseudosida bidentata Herrick, 1884 CLASS OSTRACODA
Family Cypridinidae
Family Moinidae Gigantocypris muelleri Skogsberg, 1920
Moina micrura Kurz, 1874 Macrocypridina castanea (Brady, 1897)
Appendix 241

Family Asteropidae C. nasotuberculata MUller, 1906

Leuroleberis poulseni (Moguilevsky & C. oblonga (Claus, 1890)
Ramirez, 1970) C. obtusata Sars, 1866
C. parthenoda Muller, 1906
Family Archiconchoecidae C. parvidentata Muller, 1906
Archiconchoecia cucullata (Brady, 1902) C. plactolycos Muller, 1906
A. striata Muller, 1894 C. plinthina Muller, 1906
C. porrecta Claus, 1890
Family Euconchoecidae C. procera Muller, 1890
Euconchoecia chierchiae Muller, 1891 C. pseudoparthenoda Angel, 1972
C. pusilla Muller, 1906
Family Halocyprididae C. reticulata Muller, 1906
Halocypris injlata (Dana, 1852) C. rhynchena Muller, 1906
H. globosa Claus, 1874 C. secernenda Vavra 1906
Fellia cornuta (MUller, 1906) C. serrulata serrulata Claus, 1874
C. spinifera (Claus, 1890)
Family Conchoecidae C. spinirostris Claus, 1874
Conchoecia acuminata (Claus, 1890) C. stigmatica Muller, 1906
C. acuticosta Muller, 1906 C. subarcuata Claus, 1890
C. aequiseta Muller, 1906 C. symmetrica MuHer, 1906
C. allotherium Muller, 1906 C. teretivalvata lIes, 1953
C. amblypostha Muller, 1906 C. valdiviae Miiller, 1906
C. ametra Muller, 1906
c. antipoda Muller, 1906 CLASS MAXILLOPODA
C. atlantica (Lubbock, 1856) SUBCLASS COPEPODA
C. bispinosa Claus, 1890 Order Calanoida
C. brachyaskos Muller, 1906 Family Calanidae
C. chuni Muller, 1906 Nannocalanus minor (Claus, 1863)
C. ctenophora Muller, 1906 Calanus tenuicornis (Dana, 1849)
C. curta Lubbock, 1860 Neocalanus gracilis (Dana, 1849)
C. daphnoides (Claus, 1890) Calanus australis Brodskii, 1959
C. dentata MUller, 1906 Undinula vulgaris (Dana, 1849)
C. dichotoma Muller, 1906 Calanoides carinatus Kr0yer, 1848
C. discophora Muller, 1906
C. echinata Muller, 1906 Family Eucalanidae
C. edentata Muller, 1906 Eucalanus elongatus (Dana, 1849)
C. elegans Sars, 1866 E. longiceps Matthews, 1925
C. gaussae MUller, 1908 E. crassus Giesbrecht, 1888
C. glandulosa Muller, 1906 E. monachus Giesbrecht, 1888
C. hadoni Brady & Norman, 1896 E. subtenuis Giesbrecht, 1888
C. hettacra Muller, 1906 E. pileatus Giesbrecht, 1888
C. hirsuta MUller, 1906 Rhincalanus cornutus (Dana, 1849)
C. hyalophyllum Claus, 1890
C. imbricata (Brady, 1880) Family Paracalanidae
C. incisa Muller, 1906 Paracalanus aculeatus Giesbrecht, 1888
C. isocheira Muller, 1906 P. campaneri Bjornberg, 1979
C. kyrtophora Muller, 1906 P. crassirostris F.Dahl, 1894
C.lophura Muller, 1906 P. parvus (Claus, 1863)
C. loricata Claus, 1894 P. quasimodo Bowman, 1971
C. macrcJcheira Muller, 1906 P. indicus Wolfenden, 1905
C. macromma Muller, 1906 Delius sewelli Bjornberg, 1979
C. magna Claus, 1874
C. major MUller, 1906 Family Calocalanidae
C. mamillata Muller, 1906 Calocalanus pavo (Dana, 1849)
242 Appendix

C. pavoninus Farran, 1936 Family Metridinidae

C. styliremis Giesbrecht, 1888 Pleuromamma xiphias Giesbrecht, 1889
Ischnocalanus plumulosus Claus, 1963 P. piseki Farran, 1929
Dolichocerea tenuis (Farran, 1926) P. gracilis Claus, 1863

Family Mecynoceridae Family Centropagidae

Mecynocera clausi J.C.Thompson, 1888 Centropages velificatus (deOliveira,
1947)
Family Clausocalanidae C. brachiatus (Dana, 1849)
Clausocalanus Jurcatus (Brady, 1883) C. violaceus (Claus, 1863)
C. pergens Farran, 1926 C. bradyi Wheeler, 1899
C. parapergens Frost & Fleminger, 1968
C. brevipes Frost & Fleminger, 1968 Family Lucicutiidae
Ctenocalanus vanus Giesbrecht, 1888 Lucicutia wolfendeni Sewell, 1932
C. citer Heron & Bowman, 1971 L. ovalis (Giesbrecht, 1889)
L. gaussae Grice, 1963
Family Aetideidae L. longicornis (Giesbrecht, 1889)
Aetideus armatus (Boeck, 1872) L.flavicornis (Claus, 1863)
Euaetideus bradyi (A.Scott, 1909)
E. giesbrechti (Cleve, 1904) Family Heterorhabdidae
E. acutus (Farran, 1929) Heterorhabdus spinifrons (Claus, 1863)
Paracomantena magalyae Campaner,
1978 Family Augaptilidae
Euaugaptilus hecticus (Giesbrecht, 1889)
Family Euchaetidae E. filiger (Claus, 1863)
Euchaeta spinosa Giesbrecht, 1892
E. acuta Giesbrecht, 1888 Family Diaptomidae
Paraeuchaeta barbata Brady, 1883 Notodiaptomus incompositus (Brian,
P. scotti Farran, 1909 1925)
N. amazonicus (Wright, 1935)
Family Phaennidae N. carteri (Lowndes, 1934)
Phaenna spinifera Claus, 1863 Argyrodiaptomus denticulatus Pesta,
Xanthocalanus marlyae Campaner, 1978 1927
X. minor Giesbrecht, 1892
X. agilis Giesbrecht, 1892 Family Pseudodiaptomidae
Brachycalanus bjornbergae Campaner, Pseudodiaptomus acutus (F.Dahl, 1894)
1978 P. richardi F.Dahl, 1894

Family Scolecithricidae Family Candaciidae

Scottocalanus securifrons (J.Scott, 1894) Paracandacia simplex (Giesbrecht,
Scaphocalanus magnus (T.Scott, 1894) 1892)
S. curtus (Farran, 1926) Candacia longimana Claus, 1863
S. echinatus (Farran, 1905) C. bipinnata (Giesbrecht, 1888)
S. impar (Wolfenden, 1911) C. pachydactyla (Dana, 1848)
Lophotrix latipes (T.Scott, 1893) C. curta (Dana, 1849)
Scolecithrix danae (Lubbock, 1856) C. armata Boeck, 1872
Scolecithricella bradyi (Giesbrecht,
1888) Family Pontellidae
S. dentata (Giesbrecht, 1892) Calanopia americana F.Dahl, 1894
Labidocera fluviatilis F.Dahl, 1894
Family Temoridae Pontellopsis brevis (Giesbrecht, 1889)
Temora stylifera (Dana, 1848) P. villosa Brady, 1883
T. turbinata (Dana, 1849) Pontella marplatensis Ramirez, 1966
Temeropia mayumbaensis T.Scott, 1893 Pontellina plumata (Dana, 1849)
Appendix 243

Family Acartiidae Family Corycaeidae

Acartia longiremis (Lilljeborg, 1853) Corycaeus typicus (Kroyer, 1849)
A. lilljeborgi Giesbrecht, 1892 C. speciosus Dana, 1849
A. danae Giesbrecht, 1889 C. furcifer Claus, 1863
A. negligens (Dana, 1848) C. limbatus Brady, 1888
A. tonsa Dana, 1848 C. flaccus Giesbrecht, 1891
C. amazonicus F.Dahl, 1894
Order Cyclopoida C. giesbrechti F.Dahl, 1894
Family Oithonidae C. latus (Dana, 1849)
Oithona plumifera Baird, 1843 C. lautus Dana, 1852
O. similis Claus, 1863 C. ovalis Claus, 1863
O. setigera (Dana, 1849) Farranulla rostrata (Claus, 1863)
O. oculata Farran, 1913 F. gracilis (Dana, 1853)
O. nana Giesbrecht, 1892
O. hebes Santos, 1973 Family Ergasilidae
O. oswaldocruzi Oliveira, 1945 Ergasilus euripedesi Montu, 1980
O. atlantica Farran, 1908
Order Harpacticoida
Family Cyclopidae Family Miraciidae
Macrocyclops albidus (Jurine, 1820) Macrosetella gracills (Dana, 1848)
M. ater (Herrick, 1882) Miracia efferata (Dana, 1852)
Mesocyclops annulatus (Wierzejski,
1892) Family Ectinosomatidae
M. meridianus (Kiefer, 1926) Microsetella rosea (Dana, 1848)
M. longisetus (Thiebaud, 1914) M. norvegica (Boeck, 1864)
Microcyclops mendocinus (Wierzejski,
1902) Family Pseudopeltiidae
Eucyclops ensifer Kiefer, 1936 Clytemnestra rostrata Brady, 1883
C. scutellata Dana, 1848
Family Clausidiidae
Hemicyclops subadhaerens Gooding, Family Tachydiidae
1960 Euterpina acutifrons (Dana, 1852)

Order Poecilostomatoida SUBCLASS CIRRIPEDIA

Family Oncaeidae Order Thoracica
Oncaea conifera Giesbrecht, 1891 Suborder Lepadomorpha
O. venusta Philippi, 1843 Family Scalpellidae
O. venusta var.venella Farran, 1929 Arcoscalpellum boubaloceros Young,
o. media Giesbrecht, 1891 1992
O. mediterranea (Claus, 1863) A. portoricanum (Pilsbry, 1907)
O. curta G.O.Sars, 1916 A. triangulare Hock, 1883
O. minuta Giesbrecht, 1892 Hamatoscalpellum rathbunae (Pilsbry,
O. dentipes Giesbrecht, 1891 1907)
Pachos punctatum (Claus, 1863) Litoscalpellum henriquecostai (Weber,
Lubbockia squillimana Claus, 1863 1960)
L. aculeata Giesbrecht, 1892 L. regina (Pilsbry, 1907)
Conea rapax Giesbrecht, 1891 Heterolepas cornuta (Darwin, 1851)
Ornatoscalpellum gibberum (Aurivillius,
Family Sapphirinidae 1892)
Sapphirina angusta Dana, 1849
S. auronitens-sinuicauda Brady, 1883 Suborder Verrucomorpha
S. nigromaculata Claus, 1863 Family Verrucidae
S. ovatolanceolata Dana, 1849 Altiverruca sp.
Copilia mirabilis Dana, 1849 Rostratoverruca nexa (Darwin)
244 Appendix

Verruca caribea Pilsbry, 1916 Order Isopoda

Family Idoteidae
Suborder Balanomorpha Cleantoides sp.
Family Balanidae Edotea sp.
Balanus amphitrite Darwin, 1854 Idotea balthica (Pallas, 1772)
B. improvisus (Darwin, 1854) Macrochiridotea giambiagiae Torti &
B. venustus Darwin, 1854 Bastida, 1969
Megabalanus coccopoma (Darwin, 1854) M. lilianae Moreira, 1973
M. robusta Bastida & Torti, 1967
Family Chthamalidae Synidotea marplatensis Giambiagi, 1922
Chthamalus bisinuatus Pilsbry, 1916
Family Munnidae
Family Tetraclitidae Munna peterseni (Vanip., 1985)
Tetraclita stalactifera (Lamarck, 1818)
Family Sphaeromatidae
CLASS MALACOSTRACA Ancinus gaucho Pires, 1987
Superorder Hoplocarida Dies fluminensis (Mafie-Garz6n, 1944)
Order Stomapoda Pseudosphaeroma mourei Loyola, 1960
Family Squillidae
Squilla brasiliensis Claman, 1917 Family Serolidae
Serolis bonaerensis Bastida & Torti,
Family Hemisquillidae 1967
Hemisquilla brasiliensis (Moreira, 1903) S. elliptica Sheppard, 1933
S. exigua Nordenstam, 1933
Superorder Peracarida S. foresti Bastida & Torti, 1970
Order Mysidacea S. marplatensis Bastida & Torti, 1970
Family Mysidae S. polaris Richardson, 1911
Bowmaniella (=Coifmanniella) S. uaperta Moreira, 1971
brasiliensis Bacescu, 1968 Seralis sp.
Bowmaniella sp.
Mysidopsis caelhoi Bacescu, 1968 Family Aegidae
M. tortonesi Bacescu, 1968 Aega sp.
Metamysidopsis elongata Holmes, 1900
Neomysis americana (S.I.Smith, 1873) Family Cyrnothoidae
Promysis atlantica Tattersall, 1923 Anilocra sp.
Brasilomysis castroi Bacescu, 1968
Family Cirolanidae
Order Tanaidacea Bathynomus giganteus (Milne-Edwards)
Family Tanaidae Cirolana spp.
Tanais stanfordi (Richardson, 1901) Excirolana armata (Dana, 1852)
E. braziliensis Richardson, 1912
Family Kalliapseudidae
Hemikalliapseudes cavooreni Bacescu & Order Amphipoda
Absalao, 1985 Family Ampeliscidae
Kalliapseudes gianucai Bacescu, 1979 Ampelisca spp.
K. schubarti Mafie-Garz6n, 1949 Byblis sp.
Haploops sp.
Order Cumacea
Family Pseudocumatidae Family Amphithoidae
Leptocuma kinbergii Sars, 1873 Amphitoe ramondi Audouin, 1826

Family Diastylidae Family Atylidae

Diastylis sympterigiae Bacescu & Atylus minikoi (Walker, 1916)
Queiroz, 1985
Appendix 245

Family Cheidae Euphausia americana Hansen, 1911

Cheus sp. E. tenera Hansen, 1905
E. brevis Hansen, 1905
Family Corophiidae E. gibboides Ortmann, 1893
Corophium acherusicum Costa, 1857 E. hemigibba Hansen, 1910
Leptocheirus sp. E. longirostris Hansen, 1908
Paracorophium sp. E. lucens Hansen, 1905
E. recurva Hansen, 1905
Family Gammaridae E. similis G.O.Sars, 1883
Ceradocus sp. E. spinifera G.O.Sars, 1883
Maera spp. E. vallentini Stebbing, 1900
Melita appendiculata Nematobrachion jlexipes (Ortmann,
Melita mangrovi Oliveira, 1953 1893)
Melita spp. Nematoscelis mega lops G.O.Sars, 1883
Netamelita sp. N. microps G.O.Sars, 1883
N. ten ella G.O.Sars, 1883
Family Isaeidae Stylocheiron abbreviatum G.O.Sars,
Cheiriphotis sp. 1883
Photis sp. S. affine Hansen, 1910
S. carina tum G.O.Sars, 1883
Family Lysianassidae S. elongatum G.O.Sars, 1883
Wecomedon sp. S. longicorne G.O.Sars, 1883
S. maximum Hansen, 1908
Family Oedicerotidae S. suhmii G.O.Sars, 1883
Bathyporeiapus bisetosus Escofet, 1970 Thysanoessa gregaria G.O.Sars, 1883
B. ruffoi Escofet, 1971 Thysanopoda aequalis Hansen, 1905
T. monacantha Ortmann, 1893
Family Phoxocephalidae T. obtusifrons G.O.Sars,1883
Heterophoxus sp. T. orientalis Hansen, 1910
Metharpinia sp. T. acutifrons Holt & Tattersall, 1905
Paraphoxus sp. T. tricuspidata Milne-Edwards, 1830
Phoxocephalopsis sp.
P. zimmeri Schellenberg, 1931 Order Decapoda
Pseudoharpinia sp. Suborder Dendrobranchiata
Puelche orensanzi Barnard & Clark, Infraorder Penaeidea
1982 Family Aristeidae
Urothoe sp. Aristeomorpha foliacea (Risso, 1827)

Family Platyischnopidae Family Penaeidae

Platyischnopus sp. Artemesia longinaris Bate, 1888
Funchalia villosa (Bouvier, 1905)
Family Talitridae Parapenaeus americanus Rathbun, 1901
Orchestia platensis Kroyer, 1845 Penaeopsis serrata Bate, 1881
Orchestoidea brasiliensis Dana, 1853 Penaeus brasiliensis Latreille, 1817
P. paulensis Perez-Farfante, 1967
Family Urohaustoriidae P. schmitti Burkenroad, 1936
Huarpe escofeti Barnard & Clark, 1982 Xiphopenaeus kroyeri (Heller, 1862)

Family Caprellidae Family Sicyoniidae

Caprella dilatata Kroyer, 1842 Sicyonia laevigata Stimpson, 1871
S. typica (Boeck, 1864)
Superorder Eucarida
Order Euphausiacea Family Solenoceridae
Family Euphausiidae Mesopenaeus tropicalis (Bouvier, 1905)
246 Appendix

Pleoticus muelleri (Bate, 1888) P. exilis Benedict, 1892

Solenocera necopina Burkenroad, 1939 P. trichocerus Forest & Laurent, 1967
Rodochirus rosaceus (Milne-Edwards &
Family Sergestidae Bouvier, 1893)
Lucifer faxoni Borradaile, 1915
Lucifersp. Family Galatheidae
Munida flinti Benedict, 1819
Suborder Pleocyemata M. forceps Milne-Edwards, 1880
Infraorder Caridea M. irrasa Milne-Edwards, 1880
Family Pasiphaeidae M.longipes Milne-Edwards, 1880
Leptochela serratorbita Bate, 1888 M. valida Smith, 1883

Family Palaemonidae Family Porcellanidae .

Palaemonetes argentinus Nobili, 1901 Pachycheles chubutiensis Boschi, 1963
P. haigae Stimpson, 1858
Family Hippolytidae Porcellana sayana Leach, 1820
Latreutes sp.
Family Hippidae
Family Pandalidae Emerita brasiliensis Schmitt, 1935
Dichelopandalus sp.
Hexhippolysmata oplophoroides Infraorder Brachyura
(Holthius, 1948) Family Homolidae
Parapandalus sp. Homola barbata Fabricius, 1793
Pleistonika sp.
Pontocaris boschi Christoffersen, 1988 Family Latreilliidae
Latreillia sp.
Infraorder Astracidea
Family Nephropidae Family Calappidae
Metanephrops rubellus Moreira, 1903 Acanthocarpus alexandri Stimpson,
1871
Infraorder Thalassinidea Hepatus pudibundus (Herbst, 1785)
Family Axiidae
Meticonaxius sp. Family Leucosiidae
Myropsis quinquespinosa Stimpson,
Family Callianassidae 1871
Callichirus mirim (Rodrigues, 1971) Persephona punctata Stimpson, 1859

Infraorder Palinura Family Majidae

Family Polychelidae Collodes rostratus Milne-Edwards, 1878
Stereomastis sp. Leurocyclus tuberculosus (Milne-
Edwards & Lucas, 1842)
Family Scyllaridae Libinia spinosa Milne-Edwards, 1834
Scyllarides deceptor Holthius, 1963 Pelia rotunda Milne-Edwards, 1875
Persephona punctata
Infraorder Anomura Podochela sp.
Family Diogenidae Rochinia gracilipes Milne-Edwards, 1875
Dardanus arrosor insignis (De Saussure, Stenocionops sp.
1858)
Loxopagurus loxochelis (Moreira, 1901) Family Parthenopidae
Paguristes robustus Forest & Saint Parthenope pourtalesi (Stimpson, 1871)
Laurent, 1967
Family Atelecyclidae
Family Paguridae Corystoides chilensis Milne-Edwards &
Pagurus criniticornis (Dana, 1852) Lucas, 1844
Appendix 247

Family Geryonidae SUBPHYLUM UNIRAMIA

Chaceon notialis Manning & Holtius, CLASS INSECTA
1989 SUBCLASSPTERYGOTA
Order Orthoptera
Family Portunidae Family Gryllotalpidae
Arenaeus cribarius (Lamarck, 1818) Scapteriscus riograndensis Corseuil,
Callinectes danae Smith, 1869 1992
C. ornatus Ordway
C. sapidus Rathbun, 1895 Family Tridactylidae
Ovalipes trimaculatus (de Haan, 1833) Neotridactylus carbonelli Gunther, 1972
Portunus spinicarpus (Stimpson, 1815)
P. spinimanus Latreille, 1919 Order Dermaptera
Family Labiduridae
Family Goneplacidae Labidura riparia Pallas, 1773
Batyplax typhla Milne-Edwards, 1880
Chasmocarcinus rathbuni Bouvier, 1923 Order Homoptera
C. typicus Rathbun, 1898 Family Cicadidae
Cyrtoplax spinidentata (Benedict) Proarna uruguayensis Berg, 1882
Frevillea hirsuta (Borradaile, 1916) Proarna sp.
Speocarcinus meloi D'Incao & Silva,
1991 Family Delphacidae
Delphacodes kuscheli Fennah, 1955
Family Xanthidae
Allactaea lithostrata Williams, 1974 Order Coleoptera
Hexapanopeus paulensis Rathbun, 1930 Family Cicindelidae
Panopeus sp. Cicindela nivea conspersa Dejean, 1825
Pilumnoides hassleri Milne-Edwards, C. patagonica bergiana W. Horn, 1895
1880 Megacephala (=Tetracha) brasiliensis
Pilumnus reticulatus Stimpson, 1860 Kirby, 1912
Platyxanthus patagonicus Milne-
Edwards, 1879 Family Carabidae
Rhithropanopeus harrissii Gould, 1841 Bembidion (=Notaphus) sp.
Tetraxanthus rathbunae Chace, 1939 Scarites spp.
Schizogenius costiceps Steinheil, 1869
Family Grapsidae Tetragonoderus undatus Dejean, 1829
Chasmagnathus granulata Dana, 1851 T. variegatus Dejean, 1829
Cyrtograpsus affinis (Dana, 1851)
C. altimanus Rathbun, 1914 Family Hydrophilidae
C. angulatus Dana, 1851 Paracimus rufocinctus Bruch, 1915
Hepatus pudibundus (Herbst, 1785)
Metasesarma rubripes (Rathbun, 1897) Family Staphylinidae
Bledius bonariensis Bernhauer, 1912
Family Pinnotheridae B. fernandezi Bernhauer, 1939
Dissodactylus crinitichelis Moreira, 1901 B. microcephalus Fauvel, 1901
Favia emiliae (de Melo, 1971)
Pinnixa brevipolex Rathbun, 1898 Family Heteroceridae
P. chaetopterana Stimpson Efflagitatus freudei Pacheco, 1973
P. patagoniensis Rathbun, 1918
P. rapax.Bouvier Family Anthicidae
Pinnotheres maculatus Say Lagrioida nortoni Costa & Van in, 1995

Family Ocypodidae Family Oedemeridae

Ocypode quadrata (Fabricius, 1787) Hypasclera dorsalis (Melsheimer, 1846)
Uca uruguayensis Nobili, 1901
248 Appendix

Family Tenebrionidae Family Pompilidae

Phaleria testacea Say, 1824 Anoplius (=Arachnophroctonus)
bilunulatus Saussure, 1867
Family Scarabaeidae
Plectris bonariensis Frey, 1966 Family Scoliidae
Psammodius sp. Campsomeris (Pygodasis) cineraria
(Sichel, 1864)
Family Dinastidae
Ligyrus gianucai Dechambre & Lumaret, PHYLUM PHORONIDA
1985 Phoronis psammophila Cori
Thronistes rouxi Burmeister, 1847
PHYLUM ECHINODERMATA
Family Geotrupidae CLASS CRINOIDEA
Athyreus chalibeatus Fairmaire, 1892 Order Isocrinida
Family Pentacrinidae
Family Curculionidae Diplocrinus maclearanus (W.Thomson,
Listroderes uruguayensis Kuschel, 1952 1877)

Order Lepidoptera Order Millecrinida

Family Arctiidae Family Bourgueticrinidae
Ecpanteria indecisa Walker, 1855 Rhizocrinus lofotensis Sars, 1864

Order Diptera Order Comatulida

Family Agromyzidae Family Charitometridae
Aplopeudes vogti Steysical, 1980 Crinometra brevispina
brasiliensis{Pourtaies, 1868)
Family Asilidae
Eccritosia rubriventris (Macquart, 1850) Family Antedonidae
Isometra angustipinna (P.H.Carpenter,
Family Tabanidae 1888)
Scepsis nivalis Walker, 1850 I. challengeri (A.H.Clark, 1907)
I. vivipara Mortensen, 1918
Order Hymenoptera Phrixometra longipinna (P.H.Carpenter,
Family Cynipidae 1888)
Eucoila sp.
CLASSSTELLEROIDEA
Family Formicidae SUBCLASS OPHIURIDEA
Subfamily Dolichoderinae Order Phrynophiurida
Conomyrma spp. Family Gorgonocephalidae
Astrocyclus caecilia L0.tken, 1856
Subfamily Formicinae Gorgonocephalus chilensis Philippi, 1858
Brachymyrmex sp.
Camponotus (=Tanaemyrmex) Order Ophiurida
punctulatus Mayer, 1868 Family Ophiuridae
Ophiomastus satellitae Tommasi &
Subfamily Myrmicinae Abreu, 1976
Monomorium sp. Ophioplocus januarii (L0.tken, 1856)
Mycetophylax simplex Emery, 1887 Ophiura sp.
Pheidole spininodis Emery, 1887
Solenopsis geminata Fabricius, 1804 Family Ophiacantidae
Ophiacantha vivipara Ljungman, 1870
Family Larridae
Tachytes ornatipes Cameron, 1889 Family Amphiuridae
Amphilimna olivacea (Lyman, 1869)
Appendix 249

Amphiodia pulchella (Lyman, 1869) Order Platyasterida

Amphioplus lucyae Tommasi, 1971 Family Luidiidae
Amphiura complanata Ljungman, 1867 Luidia clathrata (Say, 1825)
A. crassipes Ljungman, 1866 L. quequenensis Bernasconi, 1943
A. flexuosa Ljungman, 1867 L. scotti (Bell, 1917)
A. joubini Koehler, 1912
A. kinberi Ljungman, 1871 Order Valvatida
A. latispina Ljungman, 1867 Family Ophidiasteridae
A. stimpsonii Lutken, 1859 Calliophidiaster psicodelica Tommasi,
Axiognathus squamatus (Delle Chiaje, 1970
1829)
Diamphiodia planispina (Martens, Order Spinulosida
1873) Family Asterinidae
Hemipholis elongata (Say, 1825) Enoplopatiria stellifera (Mobius, 1859)
Micropholis subtilis Ljungman, 1871
Nudamphiura carvalhoi Tommasi, 1965 Family Echinasteridae
Ophiocnida sp. Allostichaster hartii (Rathbun, 1879)
Ophiurolepis sp. Echinaster brasiliensis Muller &
Troschel, 1M2-
Family Ophiactidae
Ophiactis lymani Ljungman, 1871 CLASS ECHINOIDEA
Order Cidaroida
Family Ophiotrichidae Family Cidaridae
Ophiotrix angulata (Say, 1825) Australocidaris canaliculata (A.Agassiz,
O. rathbuni Ludwig, 1882 1836)
Stylocidaris affinis (Philippi, 1845)
Family Ophiodermatidae
Ophioderma januarii Lutken, 1856 Order Spatangoida
Family Brissidae
Family Ophiochitonidae Brissopsis atlantica Mortensen, 1907
Ophioplax clarimundae Tommasi, 1970
Order Clypeasteoidea
Family Ophiolepididae Family Scutellidae
Ophioceramis januarii (Lutken, 1856) Mellita quinquiesperforata (Leske, 1778)
Ophiomisidium pulchellum (Wyville & Encope emarginata (Leske, 1778)
Thompson, 1877)
CLASS HOLOTUROIDEA
SUBCLASS ASTEROIDEA Order Dendrochirotida
Order Paxillosida Family Psolidae
Family Astropectinidae Psolus patagonicus Ekman, 1925
Astropecten armatus brasiliensis (Mlill &
Troschel, 1842) Order Molpadiida
A. articulatus (Say, 1825) Family Molpadiidae
A. cingulatus Sladen, 1883 Molpadia sp.
A. marginatus Gray, 1840
Bathybiaster herwigi Bernasconi, 1971 PHYLUM CHAETOGNATHA
B. loripes Sladen, 1889 Eukrohnia hamata (Mobius, 1875)
E. bathypelagica Alvarifio, 1962
FamilyGoniasteridae Krohnitta pacifica (Aida, 1897)
Anthenoides brasiliensis Bernasconi, K. subtilis (Grassi, 1881)
1956 Pterosagitta draco (Krohn, 1853)
Hippasteria argentinensis Bernasconi, Sagitta bipunctata QUoy & Gaimard,
1961 1827
Mediaster pedicellaris (Perrier, 1881) S. enflata Grassi, 1881
250 Appendix

S. tenuis Conant, 1896 F. venusta Lohmann, 1896

S. helenae Ritter-Zah6ny, 1910
S. hexaptera Orbigny, 1843 Family Oikopleuridae
S. hispida Conant, 1895 Folia gracilis Lohmann, 1892
S. lyra Krohn, 1853 F. mediterranea (Lohmann, 1899)
S. macrocephala Fowler, 1905 Oikopleura albicans (Leuckart, 1854)
S. maxima (Conant, 1896) O. cophocerca (Gegenbaur, 1855)
S. minima Grassi, 1881 O. drygalskii Lohmann, 1926
S. planctonis Steinhaus, 1896 O. fusiformis Fol, 1872
S. serratodentata Krohn, 1853 O. gracilis Lohmann, 1896
S. tasmanica Thomson, 1947 O. longicauda (Vogt, 1854)
S. decipiens Fowler, 1905 O. parva Lohmann, 1896
S. gazellae Ritter-Zahony, 1909 O. rufescens Fol, 1872
Stegosoma magnum (Langerhans, 1880)
PHYLUM HEMICHORDATA
CLASSPTEROBRANCHIATA SUBPHYLUM GNASTHOTOMATA
Order Cephalodiscida SUPER CLASS PISCES
Cephalodiscus sp. CLASS CHONDRICHTHYES
Order Squatiniformes
PHYLUM CHORDATA Family Squatinidae
SUBPHYLUM UROCHORDATA Squatina argentina (Marini, 1930)
CLASS THALIACEA S. guggenheim Marini, 1936
Family Salpidae S. occuita Vooren & Silva, 1991
Cyclosalpa bakeri Ritter, 1905
C. polae Sigl, 1912 Order Carcharhiniformes
Brooksia rostrata (Traustedt, 1893) Family Triakidae
Ihlea magalhanica (Apstein, 1894) Galeorhinus galeus (Linnaeus, 1758)
I. punctata (Forskal, 1775) Mustelus canis (Mitchill, 1815)
Ritteriella retracta (Ritter, 1906) M.fasciatus Garman, 1913
Salpa aspera Chamisso, 1819 M. schmitti (Springer, 1940)
S. fusiform is Cuvier, 1804
S. thompsoni Foxton, 1961 Family Sphyrnidae
S. maxima Forskal, 1775 Sphyrna lewini (Griffith & Smith, 1834)
Thalia cicarvan Soest, 1973 S. zygaena (Linnaeus, 1758)
T. democratica (ForskaI, 1775)
T. orientalis Tokioka, 1937 Family Carcharhinidae
T.longicauda (Quoy & Gaimard, 1824) Carcharhinus acronotus (Poey, 1860)
Weelia cylindrica (Cuvier, 1804) C. brachyurus (GUnther, 1870)
Traustedtia multitentaculata (Quoy & C. brevipinna (MUller & Henle, 1839)
Gaimard, 1824) C.obscurus(LeSueur,1818)
C. plumbeus (Nardo, 1827)
CLASS APPENDICULARIA C. signatus (Poey, 1868)
Family Fritillaridae Galeocerdo cuvier (Peron & LeSueur,
Appendicularia sicula Fol, 1874 1822)
Fritillaria aberrans Lohmann, 1896 Rhizoprionodon lalandii (Valenciennes,
F. aequatorialis Lohmann, 1896 1839)
F. borealis Lohmann, 1896
F. formica Fol, 1872 Family Scyliorhinidae
F. fraudax Lohmann, 1896 Schroederichthys bivius (Smith, 1838)
F. haplostoma Fol, 1872 Scyliorhinus besnardi Springer &
F. megachile Fol, 1872 Sadowsky, 1970
F. messanensis Lohmann, 1899
F. pellucida (Busch, 1851) Order Lamniformes
F. tenella Lohmann, 1896 Family Odontaspididae
Appendix 251

Eugomphodus taurus (Rafinesque, 1810) M. goodei Garman, 1885

M. NT (Vooren, Sect. 6.16)
Family Alopiidae M. BT (Vooren, Sect. 6.16)
Alopias vulpinus (Bonnaterre, 1788)
Family Dasyatidae
Order Squaliformes Dasyatis centroura (Mitchill, 1815)
Family Saqualidae D. say (LeSueur, 1817)
Centroscymnus cryptacanthus Regan,
1906 Family Gymnuridae
Etmopterus hillianus (Poey, 1861) Gymnura altavela (Linnaeus, 1758)
E. pusillus (Lowe, 1839)
Squaliolus laticaudus Smith & Radcliffe, Family Rhinopteridae
1912 Rhinoptera bonasus (Mitchill, 1815)
Squalus acanthias Smith & Radcliffe,
1912 Order Torpediniformes
S. cubensis Howell-Rivero, 1936 Family Narcinidae
S. mega lops (Macleay, 1881) Benthobatis sp.
S. mitsukurii Jordan & Snyder, 1903 Discopyge tschudii (Heckel, 1845)
Narcine brasiliensis (Olfers, 1831)
Family Echinorhinidae
Echinorhinus brucus (Bonnaterre, 1788) Family Torpedinidae
Torpedo puelcha Lahille, 1928
Order Hexanchiformes
Family Hexanchidae Family Callorhynchidae
Heptranchias perlo (Bonnaterre, 1788) Callorhynchus callorhynchus (Linnaeus,
Hexanchus griseus (Bonnaterre, 1788) 1758)
Notorhynchus cepedianus (Peron, 1807)
CLASS OSTEICHTHYES
Order Rhinobatiformes Family Megalopidae
Family Rhinobatidae Tarpon atlanticus (Valenciennes, 1846)
Rhinobatos horkelii (Muller & Henle,
1841) Family Xenocongridae
Zapteryx brevirostris (Muller & Henle, Chlopsis bicolor Rafinesque, 1810
1841)
Family Muraenidae
Order Rajiformes Gymnothorax conspersus Poey, 1867
Family Rajidae G. ocellatus Agassiz, 1831
Gurgesiella dorsalifera McEachran &
Compagno, 1980 Family Muraenesocidae
Psammobatis bergi Marini, 1932 Hoplunnis tenuis Ginsburg, 1951
P. glansdissimilis McEachran, 1983
P. lentiginosa McEachran, 1983 Family Congridae
P. rutrum Jordan, 1890 Ariosoma opisthophthalma (Ranzani,
Raja agassizi (Muller & Henle, 1841) 1838)
R. castelnaui Ribeiro, 1907 Conger orbignyanus Valenciennes, 1847
R. cyclophora Regan, 1903 Rechias dubius (Breder, 1927)
R. leptocauda Krefft & Stehmann, 1975
R. platana Gunther, 1880 Family Ophichthidae
Sympterigia acuta Garman, 1877 Myrophis frio Jordan & Davis, 1892
S. bonapartei Muller & Henle, 1841 Ophichthus gomesii (Castelnau, 1855)

Order Myliobatiformes Family Clupeidae

Family Myliobatidae Brevoortia pectinata (Jenyns, 1842)
Myliobatis freminvillei LeSueur, 1824
252 Appendix

Chirocentrodon bleekerianus (Poey, Gymnoscopelus piabilis (Whitley, 1931)

1867) Lepidophanes guentheri (Goode & Bean,
Opisthonema oglinum (LeSueur, 1818) 1896)
Pellona harroweri (Fowler, 1917) Symbolophorus barnardi (Taning, 1932)
Ramnogaster arcuata (Jenyns, 1842) S. veranyi (Moreau, 1888)
Sardinella brasiliensis (Steindachner,
1789) Family Polymixiidae
Polymixia lowei Gunther, 1859
Family Engraulididae
Anchoa marinii Hildebrand, 1943 Family Bregmacerotidae
A. tricolor (Agassiz, 1829) Bregmaceros atlanticus Goode & Bean,
Engraulis anchoita Hubbs & Marini, 1886
1935
Lycengraulis grossidens (Agassiz, 1829) Family Gadidae
Urophycis brasiliensis (Kaup, 1858)
Family Gonostomatidae U. mystaceus Ribeiro, 1903
Maurolicus muelleri (Gmelin, 1788)
Vicinguerria sp. Family Merlucciidae
Macruronus magellanicus Lonnberg,
Family Argentinidae 1907
Argentina striata Goode & Bean, 1896 Merluccius hubbsi Marini, 1933

Family Ariidae Family Ophidiidae

Genidens genidens (Valenciennes, 1839) Benthocometes robustus (Goode & Bean,
Netuma barba (Lacepede, 1803) 1886)
N. planifrons Higuchi Reis & Araujo, Genypterus brasiliensis Regan, 1903
1982 Raneya jluminensis (Ribeiro, 1903)

Family Pimelodidae Family Macrouridae

Parapimelodus nigribarbis (Boulenger, Coelorhynchus coelorhynchus (Risso,
1889) 1810)
Pimelodus maculatus (Lacepede, 1803) C. marinii Hubbs, 1934
Malacocephalus laevis (Lowe, 1843)
Family Synodontidae M. occidentalis (Goode & Bean, 1885)
Saurida brasiliensis Norman, 1935 Ventrifossa ori (Smith, 1968)
S. caribbaea Breder, 1927
Synodus foetens (Linnaeus, 1766) Family Batrachoididae
Porichthys porosissimus (Valenciennes,
Family Chlorophthalmidae 1837)
Chlorophthalmus agassizi Bonaparte, Thalassophryne montevidensis Berg,
1840 1893
Parasudis truculenta (Goode & Bean,
1895) Family Lophiidae
Lophius gastrophysus Ribeiro, 1915
Family Paralepididae
Lestidium atlanticum Boradin, 1928 Family Ogcocephalidae
Ogcocephalus vespertilio (Linnaeus,
Family Alepisauridae 1758)
Alepisaurus brevirostris Gibbs, 1960
Family Gobiesocidae
Family Myctophidae Gobiesox strumosus Cope, 1870
Diaphus brachycephalus Taning, 1928
D. dumerilii (Bleeker, 1856) Family Antennariidae
D. effulgens (Goode & Bean, 1896) Phrynelox scaber Cuvier, 1817
Appendix 253

Family Exocoetidae Family Fistulariidae

Hiporhamphus unifasciatus (Ranzani, Fistularia petimba Lacepede, 1803
1842)
Cypselurus sp. Family Macrorhamphosidae
Macrorhamphosus scolopax (Linnaeus,
Family Anablepidae 1758)
Jenynsia lineata (Jenyns, 1842) Notopogon fernadezianus (Delfin, 1899)

Family Poeciliidae Family Syngnathidae

Cnesterodon decemmaculatus (Jenyns, Hippocampus erectus Perry, 1810
1842) H. reidi Ginsburg, 1933
Phalloceros caudimaculatus (Hensel, 1868) Sygnathus elucens Poey, 1867.
Phalloptychus januaris (Hensel, 1868) S. folletti Herald, 1942
Poecilia vivipara Schneider, 1841 S. pelagicus Linnaeus, 1758

Family Belonidae Family Scorpaenidae

Strongylura marina (Walbaum, 1792) Helicolenus lahillei Norman, 1937
S. timucu (Walbaum, 1792) Pontinus rathbuni Goode & Bean, 1896
Setarches guentheri Johnson, 1862
Family Scomberesocidae
Scomberesox saurus (Walbaum, 1792) Family Peristediidae
Peristedion altipinne Regan, 1903
Family Atherinidae Peristedion sp.
Austroatherina incisa (Jenyns, 1842)
Odontesthes bonariensis (Valenciennes, Family Triglidae
1835) Bellator brachychir (Regan, 1914)
O. argentinensis (Valenciennes, 1835) Prionotus nudigula Ginsburg, 1950
Atherinella brasiliensis (Chernoff, 1986) P. punctatus (Bloch, 1797)

Family Lamprididae Family Congiopodidae

Lampris guttatus (Brunnich, 1788) Congiopodus peruvian us (Cuvier, 1829)

Family Trachichthyidae Family Dactylopteridae

Hoplostethus occidentalis Woods, 1973 Dactylopterus volitans (Linnaeus, 1758)
Paratrachichthys atlanticus Menezes, 1971
Family Centropomidae
Family Berycidae Centropomus ensiferus Poey, 1860
Beryx splendens Lowe, 1834 C. parallelus Poey, 1860
C. undecimalis (Bloch, 1792)
Family Anoplogasteridae
Anoplogaster cornuta (Valenciennes, Family Serranidae
1833) Acanthistius brasilianus (Cuvier, 1828)
A. patachonicus (Jenyns, 1840)
Family Zeidae Anthias menezesi Anderson &
Zenopsis conchifer (Lowe, 1850) Heemstra, 1980
Diplectrum formosum (Linnaeus, 1766)
Family Grammicolepididae D. radiale (Quoy & Gaimard, 1824)
Xenolepidichthys dalgleishi Gilchrist, Dules auriga Cuvier, 1829
1922 Epinephelus Jlavolimbatus Poey, 1865
E. itajara (Lichtenstein, 1822)
Family Caproidae E. marginatus (Lowe, 1834)
Antigonia capros Lowe, 1843 E. morio (Valenciennes, 1828)
254 Appendix

E. nigritus (Holbrook, 1855) S. vomer (Linnaeus, 1758)

E. niveatus (Valenciennes, 1828) Seriola lalandi Valenciennes, 1833
Hemanthias vivanus (Jordan & Swain, S. rivoliana Valenciennes, 1833
1884) Trachinotus carolinus (Linnaeus, 1766)
Holanthias martinicensis (Guichenot, T. fa lcatus (Linnaeus, 1758)
1868) T. goodei Jordan & Evermann, 1896
Pikea rosea (Gunther, 1880) T. marginatus Cuvier, 1832
Serranus atrobranchus (Cuvier, 1829) Trachurus lathami Nichols, 1920
S. flaviventris (Cuvier, 1829)
Family Caristiidae
Family Polyprionidae Caristius sp.
Polyprion americanus (Schneider, 1801)
Family Pomadasyidae
Family Priacanthidae Boridia grossidens (Cuvier, 1830)
Cookeolus japonicus (Bloch & Orthopristis ruber (Cuvier, 1830)
Schneider, 1801) Pomadasys corvinaeformis
Priacanthus arenatus Cuvier, 1829 (Steindachner)

Family Apogonidae Family CQryphaenidae

Apogon pseudomaculatus Longley, 1932 Coryphaenaequiselis Linnaeus, 1758
Synagrops bella (Goode & Bean, 1895) C. hippurus Linnaeus, 1758)
S. spinosus Schultz, 1940
Family Bramidae
Family Branchiostegidae Brama brama (Bonnaterre, 1788)
Caulolatilus chrysops (Valenciennes, Pteraclis aesticola (Jordan & Snyder,
1833) 1901)
Lopholatilus villarii Ribeiro, 1915
Family Lutjanidae
Family Pomatomidae Pristipomoides freemani Anderson, 1966
Pomatomus saltatrix (Linnaeus, 1766)
Family Lobotidae
Family Rachycentridae Lobotes surinamensis (Bloch, 1790)
Rachycentron canadus (Linnaeus, 1766)
Family Gerreidae
Family Echeneididae Diapterus rhombeus (Cuvier, 1829)
Echeneis naucrates Linnaeus, 1758 Eucinostomus gula (Cuvier, 1830)
Remora brachyptera (Lowe, 1839) E. melanopterus (Blecker, 1863)
R. osteochir (Cuvier, 1829) Eugerres brasilianus (Cuvier, 1830)
Ulaema lefroyi (Goode, 1874)
Family Carangidae
Caranx chrysos (MitchilI, 1815) Family Sparidae
C. hippos (Linnaeus, 1766) Diplodus argenteus (Valenciennes, 1830)
C. latus Agassiz, 1831 Pagrus pagrus (Linnaeus, 1758)
Chloroscombrus chrysurus (Linnaeus,
1766) Family Sciaenidae
Decapterus tabl Berry, 1968 Ctenosciaena gracilicirrhus (Metzelaar,
Naucrates ductor (Linnaeus, 1758) 1919)
Oligoplites sa liens (Bloch, 1793) Cynoscion guatucupa (Cuvier, 1829)
O. saurus (Schneider, 1801) C. jamaicensis (Vaillant & Bocourt,
Parona signata (Jenyns, 1842) 1883)
Pseudocaranx dentex (Bloch & Macrodon ancylodon (Schneider, 1801)
Schneider, 1801) Menticirrhus americanus (Linnaeus,
Selar crumenophthalmus (Bloch, 1793) 1758)
Selene setapinnis (MitchilI, 1815) M. littoralis (Holbrook, 1860)
Appendix 255

Micropogonias furnieri (Desmarest, Family Percophidae

1823) Bembrops heterurus (Ribeiro, 1903)
Paralonchurus brasiliensis Percophis brasiliensis Quoy & Gaimard,
(Steindachner, 1875) 1824
Pogonias cromis (Linnaeus, 1766)
Stellifer brasiliensis (Schultz, 1945) Family Pomacentridae
S. rastrifer (Jordan, 1889) Abudefdufsaxatilis (Linnaeus, 1758)
Umbrina canosai Berg, 1895
Family Pinguipedidae
Family Ephippidae Pseudopercis numida Ribeiro, 1903
Chaetodipterus faber (Broussonet, 1782) P. semifasciata (Cuvier, 1829)

Family Cichlidae Family Blenniidae

Cichlaurus facetus (Jenyns) Hypleurochilus fissicornis (Quoy &
Geophagus brasiliensis (Quoy & Gaimard, 1824)
Gaimard)
Family Eleotrididae
Family Mullidae Guavina guavina (Valenciennes, 1837)
Mullus argentinae Hubbs & Marini, 1935
Family Gobiidae
Family Kyphosidae Gobioides broussonnetii Lacepede, 1800
Kyphosus incisor (Cuvier, 1831)· Gobionellus oceanicus (Pallas, 1770)
G. shufeldti (Jordan & Eigenmann, 1886)
Family Chaetodontidae G. smaragdus (Valenciennes)
Chaetodon guyanensis Durand, 1960
Family Gempylidae
Family Cheilodactylidae Promethichthys prometeus (Cuvier,
Sciaenoides bergi (Norman, 1937) 1832)
Thyrsitops lepidopoides (Cuvier, 1831)
Family Mugilidae
Mugil curema Valenciennes, 1836 Family Trichiuridae
M. gaimardianus Desmarest, 1831 Benthodesmus elongatus (Clarke, 1879)
M. platanus Gunther, 1880 Evoxymetopon taeniatus Poey, 1863
Lepidoupus sp.
Family Sphyraenidae Trichiurus lepturus Linnaeus, 1758
Sphyraena barracuda (Walbaum, 1792)
S. guachancho Cuvier, 1829 Family Scombridae
S. tome Fowler, 1903 Auxis thazard (Lacepede, 1803)
Euthynnus alletteratus (Rafinesque,
Family Polynemidae 1810)
Polydactylus oligodon (Gunther, 1860) Katsuwonus pelamis (Linnaeus, 1758)
P. virginicus (Linnaeus, 1758) Sarda sarda (Bloch, 1793)
Scomber japonicus Houttuyn, 1780
Family Labridae Thunnus alalunga (Bonaterre, 1788)
Decodon puellaris (Poey, 1860) T. albacares (Bonnaterre, 1788)
T. atlanticus (Lesson, 1830)
Family Opistognathidae T. obesus (Lowe, 1839)
Lonchopisthus meadi Menezes & T. thynnus (Linnaeus, 1758)
Figttdredo,1971
Family Xiphiidae
Family Uranoscopidae Xiphias gladius Linnaeus, 1758
Astroscopus sexspinosus (Steindachner,
1877)
256 Appendix

Family Istiophoridae Family Tetraodontidae

Istiophorus albicans (Latreille, 1804) Lagocephalus laevigatus (Linnaeus,
Makaira indica (Cuvier, 1831) 1766)
M. nigricans Lacepede, 1803
Tetrapturus albidus Poey, 1860 Family Diodontidae
Chilomycterus spinosus (Linnaeus, 1758)
Family Centrolophidae
Centrolophus niger (Gmelin, 1788) Family Molidae
Mola mola (Linnaeus, 1758)
Family Nomeidae
Cubiceps athenae Haedrich, 1965 SUPERCLASSTETRAPODA
CLASS AMPHIBIA
Family Ariommidae SUBCLASS ANURA
Ariomma bondi Fowler, 1930 Family Bufonidae
Bufo arenarum arenarum Hensel, 1867
Family Stromateidae
Peprilus paru (Linnaeus, 1758) Family Leptodactylidae
Stromateus brasiliensis Fowler, 1906 Odontophrynus sp.

Family Gobiesocidae CLASS REPTILIA

Gobiesox strumosus Cope, 1870 Order Squamata
Family Iguanidae
Family Bothidae Liolaemus occipitalis Boulenger, 1885
Citharichthys comutus (Gunther, 1880)
C. spilopterus (Valenciennes) Family Colubridae
Etropus longimanus Norman, 1933 Lystrophis sp.
Mossolene antillarum Norman, 1933
Paralichthys brasiliensis (Ranzani, 1840) Order Testudines (Chelonia)
P. isoceles Jordan, 1890 Suborder Cryptodira
P. orbignyanus (Valenciennes, 1839) Family Chelonidae
P.patagonicusJordan,1886 Caretta carreta (Linnaeus, 1758)
Xistreuris rasile (Jordan, 1890) Chelonia mydas (Linnaeus, 1758)
Erectmochelys imbricata (Linnaeus,
Family Pleuronectidae 1766)
Oncopterus darwini Steindachmer, 1875
Family Dermochelyidae
Family Soleidae Dermochelys coriacea (Vandelli, 1761)
Achirus garmani (Jordan & Goss, 1889)
A. lineatus (Linnaeus, 1758) CLASS AVES
Order Sphenisciformes
Family Cynoglossidae Family Spheniscidae
Symphurus ginsburgi Menezes & Eudyptes chrysolophus (Brandt, 1837)
Benvegnu,1976 E. crestatus (Forster, 1781)
S. jenynsi Evermann & Kendall, 1907 Spheniscus magellanicus (Forster, 1781)
S. trewavasae Chabanaud, 1948
Order Podicipediformes
Family Balistidae Family Podicipedidae
Balistes capriscus Gmelin, 1788 Podiceps major (Boddaert, 1783)
B. vetula Linnaeus, 1758
Order Procellariiformes
Family Monacanthidae Family Diomedeidae
Monacanthus ciliatus (Mitchill, 1818) Diomedea chlororhynchos Gmelin, 1789
Stephanolepis setifer (Bennett) D. chrysostoma Forster, 1758
D. epomophora (Lesson, 1825)
Appendix 257

D. exulans (Linnaeus, 1758) Order Falconiformes

D. melanophrys Temminck, 1828 Family Falconidae
Phoebetria palpebrata (Forster, 1758) Milvago chimango (Vieillot, 1816)
Polyborus plancus (Miller, 1777)
Family Procellariidae
Calonectris diomedea (Scopoli, 1769) Order Gruiformes
Daption capense (Linnaeus, 1758) Family Rallidae
Fulmarus glacialoides (Smith, 1840) Fulica arm illata Vieillot, 1817
Macronectes sp.
Pachyptila belcheri Mathews, 1912 Order Charadriiformes
P. desolata (Gmelin, 1789) Family Haematopodidae
P. vittata (Forster, 1777) Haematopus palliatus Temminck, 1820
Procellaria aequinioctialis
aequinioctialis Linnaeus, 1758 Family Charadriidae
P. aequinioctialis conspicillata Gould, Charadrius collaris Vieillot, 1818
1844 C. falklandicus Latham, 1790
P. cinerea Gmelin, 1789 C. modestus Lichtenstein, 1823
Pterodroma brevirostris (Lesson, 1831) C. semipalmatus Bonaparte, 1825
P. incerta (Schlegel, 1863) Oreopholus ruficQllis (Wagler, 1929)
P. lessoni (Garnot, 1826) Pluvialis dominica (Muller, 1766)
P. mollis (Gould, 1844) P. squatarola (Linnaeus, 1758)
Puffinus gravis (O'Reilly, 1818)
P. griseus (Gmelin, 1789) Family Scolopacidae
P. puffinus (Brunnich, 1764) Actitis macularia (Linnaeus, 1766)
Arenaria interpres (Linnaeus, 1758)
Family Hydrobatidae Bartramia longicauda (Bechstein, 1812)
Oceanites oceanicus (Kuhl, 1820) Calidris alba (Pallas, 1764)
C. bairdii (Coues, 1861)
Family Pelecanoididae C. canutus rufa (Wilson, 1813)
Pelecanoides magellani (Mathews, 1912) C. fuscicollis (Vieillot, 1819)
C. melanotos (Vieillot, 1819)
Order Pelecaniformes C. pusilla (Linnaeus, 1766)
Family Phalacrocoracidae Catatrophorus semipalmatus (Gmelin,
Phalacrocorax olivaceus (Humboldt, 1789)
1805) Limosa haemastica (Linaeus, 1758)
Micropalama himantopus (Bonaparte,
Family Fregatidae 1826)
Fregata magnificens Mathews, 1914 Numenius phaeopus (Linnaeus, 1758)
Tringaflavipes (Gmelin, 1789)
Order Ciconiiformes T. melanoleuca (Gmelin, 1789)
Family Ardeidae T. solitaria Wilson, 1813
Ardea cocoi Linnaeus, 1766 Tryngites subruficollis (Vieillot, 1819)
Butorides striatus (Linnaeus, 1785)
Egretta thula (Molina, 1782) 'Family Recurvirostridae
Himantopus himantopus melanurus
Order Phoenicopteriformes Vieillot, 1817
Family Phoenicopteridae
Phoenicopterus chilensis Molina, 1782 Family Phalaropidae
Steganopus tricolor Vieillot, 1819
Order Anseriformes
Family Anatidae Family Chionidae
Cygnus melancorhyphus (Molina, 1782) Chionis alba (Gmelin, 1789)
258 Appendix

Family Stercorariidae Suborder Odontoceti

Catharacta skua antarctica (Lesson, Family Physeteridae
1831) Physeter macrocephalus Linnaeus, 1758
C. skua chilensis (Bonaparte, 1857)
Stercorarius longicaudus Vieillot, 1819 Family Kogiidae
S. parasiticus (Linnaeus, 1758) Kogia breviceps (Blainville, 1838)
S. pomarinus (Temminck, 1815) K. simus Owen, 1866

Family Laridae Family Ziphiidae

Gelochiledon nilotica Mathews, 1912 Hyperoodon planifrons (Flower, 1882)
Larus belcheri atlanticus Olrog, 1958 Mesoplodon densirostris (Blainville,
L. cirrocephalus Vieillot, 1818 1817)
L. dominicanus Lichtenstein, 1823 Ziphius cavirostris (G. Cuvier, 1823)
L. maculipennis Lichenstein, 1823
Phaetusa simplex (Gmelin, 1789) Family Delphinidae
Sterna eurygnatha Saunders, 1876 Delphinus delphis Linnaeus, 1758
S. hirundinacea Lesson, 1831 Globicephala melas (Traill, 1809)
S. hirundo Linnaeus, 1758 Grampus griseus (G. Cuvier, 1812)
S. maxima Boddaert, 1783 Orcin us orca (Linnaeus, 1758)
S. paradisea Pontoppidan, 1763 Pseudorca"crassidens (Owen, 1846)
S. superciliaris Vieillot, 1819 Sotalia fluviatilis (Gervais, 1853)
S. trudeaui Audubon, 1838 Stenella attenuata
S. vittata Gmelin, 1789 S. clymene (Gray 1846)
S. coeruleoalba (Meyen, 1833)
Family Rynchopidae S. frontalis (G. Cuvier, 1829)
Rynchops nigra Linnaeus, 1758 S. longirostris (Gray, 1828)
Steno bredanensis (Lesson, 1828)
Order Passeriformes Tursiops truncatus (Montagu, 1821)
Family Motacillidae
Anthus correndera Vieillot, 1818 Family Pontoporiidae
Pontoporia blainvillei (Gervais &
CLASS MAMMALIA Orbigny, 1844)
Order Rodentia
Family Cricetidae Family Phocoenidae
Calomys laucha (Olfers, 1818) Australophocaena dioptrica (Lahille,
1912)
Family Ctenomyidae Phocoena spinipinnis Burmeister, 1865
Ctenomys flamarioni Travi, 1981
Order Carnivora
Order Cetacea Family Otariidae
Suborder Mysticeti Arctocephalus australis (Zimmerman,
Family Balaenopteridae 1783)
Balaenoptera acutorostrata Lacepede, A. gazella (Peters, 1875)
1804 A. tropicalis (Gray, 1872)
B. edeni Anderson, 1878 Otaria fla vescens (Shaw, 1800)
B. musculus (Linnaeus, 1758)
Megaptera novaeangliae (Borowski, Family Phocidae
1781) Lobodon carcinophagus (Hombron &
Jacquinot, 1842)
Family Balaenidae Hydrurga leptonyx (Blainville, 1820)
Eubalaena australis Desmoulins, 1822 Mirounga leonina Linnaeus, 1728
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Zavala-Camin LA (1978) Algunos aspectos sobre la estructura populacional del rabil
(Thunnus albacares) en el sudeste y sur de Brasil (1969-1977) con presentaci6n de la
hip6tesis de la migraci6n semestral. Bolm Inst Pesca (S Paulo) 5(11):1-25
Zavala-Camin LA (1981) Habitos alimentares e distribui\=iio dos atuns e afins
(Osteichthyes: Teleostei) e suas rela\=oes ecol6gicas com outras especies peIagicas das
regioes sudeste e suI do Brasil. PhD Thesis, Univ S Paulo, Brazil
Zavala-Camin LA, Antero da Silva (1991) Hist6rico da pesquisa e pesca de atuns coin
espinhel no Brasil. Atlantica (Rio Grande) 13(1):107-11
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67(3):269-277
Zenkovitch VP (1967) Processes of coastal development. Oliver and Boyd, London
Subject Index

Abra 117 Anablepidae (=Jenynsiidae) 58, 59, 78

Abralia 148, 150 Anatidae 62
Abraliopsis 148 Anchoa 51, 52, 53, 54, 55, 57, 59, 60, 61, 120,
Abylopsis 110 122,123, 124, 127, 128, 134, 154, 175, 176
Acacia 165 Ancinus 115, 116, 117
Acanella 120 Ancistrocheiros 136, 148, 150
Acanthogorgia 120 Andropogon 100, 101, 102, 162, 163
Acartia 40, 42, 43, 111, 169 Androtrichum 100,101, 102, 162, 163
Achirus 51, 52, 53, 54, 55, 56, 57 Anguiliform 57, 121, 131
Achrochaetium 83 Annelid 77
AchyroC/ine 99 Anoplius 103
Acroscalpellum 119 Ant 102
Acrosticum 24 Antarctic Bottom Water (ABW) 95
Acteon 117 Antarctic Intermediate Water (AIW) 95,
Actiniaria, actinia 115, 120 172,173
Adelomelon 119 Anthenoides 120
Adrana 118, 119 Anthus 115, 169
Aegina 110 Anticyclone 1, 5, 96
Afforestation 202, 205 Antigonias 122, 130, 131
Aglaura 110 Antipathes 120
Agriculture 12, 23, 197, 198, 199, 205, 207 Apium 26
Ajaia 62 Apoprionospio 117
Albatross 154, 155, 194 Appendicularia 112
Allopsus 149 Araunus 62
Alona 40 Arctocephalus australis 150, 151, 153, 154
Alopias 141, 143, 176 Ardea 155
Alternanthera 24 Arenaeus 115, 116, 169
Amiantis 168, 169 Argentina 130
Ammonium 23, 50, 96, 97, 98, 180 Argonauta 147, 149
Amphicteis 117 Ariid 58, 59, 154
Amphioplus 117 Ariomma 130, 131
Amphipod 44, 45, 72, 77, 78, 84, 86, 88, Ariommatidae 131
119, 125,133,134,135,147,162,174,175, Armadillo 104, 163
178 Artemesia 119, 132, 134, 152, 154, 160, 176,
Amphitoe 78, 86 177,185, 191, 194
Amphiura 118 Artemia 141
Amphora 35, 82 Ascidian 135
Anabaena 108 Asterionellopsis 108, 167, 168, 169, 179
296 Subject Index

Astrangia 119 114, 115, 116, 119, 120, 125, 129, 131, 142,
Astropecten 118, 119 147,168,169,170,171,172,174,175,
Asychis 119 179,182,188,193,194,199,201
Athene104 Bioturbation 46
Atherinella (=Xenomelaniris) 57, 60, 61, Bird 62, 77, 115, 116, 154, 155, 156, 157, 158,
77,84,136,169 159,163,169,170,177,194,199,201
Atherinidae 51, 52, 55, 56, 58, 59, 78, 85, 121, Bivalve 115, 116, 134, 154, 159, 169, 177
123, 136, 138, 159 Bledius 103, 114, 115, 169
Atherinops 137 Blenniidae 51, 52, 55, 56, 57
Athyreus 103 Bloom 34,108, 167, 198, 199
Atrina 119 Blutaparon 99, 100, 103, 162, 165
Atriplex 24,26 Boccardia 86
Australophocaena 150, 151 Bothidae 78, 121, 131
A uxis 121, 123, 127 Bougainvillia 110, 113
Brachydontes 88
Bacillaria 34 Brachymyrmex 102
Bacopa 24, 102, 163 Brachyuran~120, 135
Bacteria 37,39,59,84,104; 105, 174, 175 Brama 127, 175
Balaeonoptera 151 Brazil Current (BC) 1, 94, 95, 96, 105, 106,
Balanus 40, 42, 73, 79, 81, 86, 88, 133 109, 110, 111, 122, 123, 124, 129, 146, 172
Balistes 124, 127, 128, 130, 131, 176, 177 Bregmaceros 121, 122, 135
Balistidae 57, 121 Bregmacerotidae 121, 123
Balloniscus 77 Brevoortia 51, 52, 53, 54, 55, 56, 60, 61, 120,
Bangia 88 121,123,124, 169,186,191
Barnacle 40, 120 Brissopsis 120
Barrier 13, 14, 18, 89, 91 Brooksia 113
Basement 13, 91 Bryocladia 88
Basin 1, 9, 13, 14, 16, 18, 91 Buccinanops 115, 116, 117, 118, 119, 133,169
Bassia 110 Bufo 104
Bathynomus 120 Burning 201
Bathyporeiapus 44, 77 Butorides 62
Batrachoid 58
Batyplax 120 Cadulus 117
Beach 9, 13, 14, 15, 58, 59, 61, 62, 63, 85, 91, Cakile 100, 162, 165
92, 94, 100, 109, 114, 115, 116, 138, 143, Calanoides 111
152, 154, 155, 156, 157, 158, 159, 161, 162, Calanus 174
165,166,167,168,170,192,194 Calidris 157, 158, 170
Bebryce 120 Callinectes 43, 47, 72, 73, 78, 79, 84, 85,
Beetle 103, 114, 115, 169 133, 177, 194
Bembidion 103 Calocalanus 111
Bembrops 130 Calomys 104
Benthobatis 143 Calycera 100, 162, 165
Benthodesmus 130 Calystegia 100, 162, 165
Biapertura 40 Camponotus 102
Biodiversity 3,205 Campsomeris 103
Biomass 27, 29,30,32,33,34,36,37,39, Camptocercus 40
42,44,45, 66, 67, 68, 69,70,73,77,81, Caproidae 121
84, 100, 101, 102, 104, 105, 107, 112, 113, Carabid 103, 114
Subject Index 297

Carangidae 57, 121, 123, 131 Circulation 1, 18, 19, 20, 27, 30, 40, 42, 43,
Carbon 18, 36, 37, 39, 67, 68, 69, 70, 77, 94,95,166,182,199,200,208
104, 113, 168, 175, 179 Cirolana 120
Carbonate 18, 92 Cirriped 18, 40, 81, 86, 88, 169
Carcharhinidae 141 Cladocera 40, 42, 110, 174, 175
Carcharhiniforme 145 Cladocora 120
Carcharhinus 141, 142, 143, 144, 176 Cladophora 88
Caretta 151, 154 Clausocalanus 111
Cavolinia 110, 111 Climate 1, 2, 5, 6, 20, 66, 205
Centengraulis 123 Clio 110
Centropages 42, 111 Clupeidae 51, 61, 73, 120, 121, 124,,131
Centroscymnus 143 Clytemnestra 111
Cephalodiscus 120 Coastal Water (CW) 20, 30, 34, 37, 39, 40,
Cephalopod 127, 132, 134, 135, 146, 147, 47,5°,51,52,76,85,95,96,97, 104, 105,
148, 149, 150, 152, 154, 159, 194 106,107, 108, 109, 111, 112, 113, 122, 123,
Cerataulina 34, 35 124, 125, 129, 131, 132, 133, 138, 143, 152,
Ceratium 105, 106, 108 155, 159, 169, 172;'173, 175, 178, 179, 180,
Ceratophyllum 27 181, 182, 183, 188, 193, 194, 199, 200
Ceriodaphnia 42 Coccolithophorid 105
Cetacean 150, 151, 195 Cocconeis 35, 82
Chaceon 136, 191, 194 Coelorhynchus 121, 130, 176, 178
Chaetoceros 34, 35, 36, 105, 106 Coleoptera 103, 114
Chaetognat 42, 110, 112, 113 Collis ella 88
Channel 9, 14, 15, 16, 18, 19, 20, 21, 22, 23, Conea 111
36, 40, 45, 59, 63, 73, 78, 80, 81, 82, 85, Conepathus 104, 163
86,88,91,183,197,198,200, 207 Conger 130
Charachiform 57 Conomyrma 102
Charadriiformes 154 Conservation 63, 158, 160, 166, 206, 207
Charadrius 63, 115, 155, 157, 158, 169 Consumer 37, 65, 70, 71, 72, 84, 109, 124,
Chasmagnathus 43, 44, 47, 48, 49, 50, 72, 168, 169, 207
76, 77, 79, 199 Copepod 40, 42, 43, 71, 72, 110, 111, 114,
Chasmocarcinus 119 125, 133, 134, 169, 174, 175
Chelonia 151, 154 Coral 120, 131, 147
Chelophyes 110 Corbula 117, 118, 119, 133
Chenopodium 26 Cormorant 62, 155, 170
Chiclid 57 Corycaeus 111
Chiroteuthis 149 Coryphaena 128
Chlamys 119 Coscinodiscus 34,105, 108, 167
Chlorophyceae, Chlorophyta 30, 32, 34 CPUE 131, 187, 188, 189, 190, 192
Chlorophyll 34, 36, 66, 98, 104, 106, 107, Crab 4°,43,44,46,47,49,72,76,78,84,
108,109,122,167,168,171,172,173,174, 85,116,136,147,169,177,183,186,193,
180,181 194
Chloroscombrus123 Crassinella 118
Chthamalus 88 Crepidula 119
Chydorus40 Creseis 110, 111
Cicindella 114, 115 Crinometra 120
Ciliate 34, 37, 39, 106, 109, 174 Crustacean 40, 44, 45, 47, 50, 71, 78, 109,
115,116, 124, 126, 127, 132, 133, 134, 135,
298 Subject Index

Crustacean (cont.) 147,150,152,154,158, Dictyocha 106

169, 175, 180, 181, 184, 185, 193, 197 Dies 78
Cryptophyceae 34 Dinoflagellate 34, 36, 39, 105, 106, 108,
Ctenocalanus 111 109, 167, 169, 179
Ctenomys 104, 163 Dinophysis 105, 106, 108, 167
Ctenophore 42 Diogenichthys 121, 128
Ctenosciaena 57, 59, 60, 130, 131, 177 Diomedea 155, 156, 177, 194
Cum ace an 44, 116, 117, 119, 135, 178 Diopatra 117, 119
Cunina 110, 113 Diphyes 110
Current 19, 20, 94, 95, 96, 116, 125, 166, Diphyopsis 110
167, 169, 179, 182 Diplophos 121
Cyanobacteria (=Cyanophyta) 30, 32, 34, Diplospinus 121, 123
66,67,69,70,88,108,169 Discard 155, 156, 185, 186, 192, 194, 195
Cyclosalpa 113 Discopyge 143
Cyclothone 121, 122 Diversity 3, 18,30,40,42,43, 44, 56, 57,
Cygnus 62, 84 72,75,78, 81, 86, 88, 102, 105, 110, 114,
Cylindrotheca 34, 35 117, 125, 134,-141, 146, 163, 166, 174, 179,
Cynoglossidae 121, 123 195,202
Cynoscion 51, 52, 53, 55, 57, 58, 60, 127, 128, Dolphin 63, 64, 195
129, 130, 131, 133, 134 136, 152, 153, 154, Donax 47,48,49,109, 115, 116, 117, 124,
159,172,174,175,176,177,178,183,185, 154, 158, 168, 169, 170
187,188, 189, 191, 195 Dorsanum n6, 119
Cyperus 24 Dune 13, 14, 16, 24, 94, 98, 99, 100, 101,
Cyrtograpsus 43, 72, 78, 79, 84 102, 103, 104, 114, 115, 159, 161, 162, 163,
Cytaeis 110 164,165,166,169,202,205
Dusicyon 104, 163
Daption 156 Dyastilis 44
Dardanus 119
Dasyatis 143, 144 Eccritosia 103
Dasypus 104, 163 Echinoderm 40, 117, u8, 120, 169
Decapod 43, 44, 46, 47, 72, 73, 78, 80, 81, Echinorhinus 143
82, 88, 134, 169 Ecpantheria 103
Decapterus 127 Efflagitatus 103, 115
Deep North Atlantic Water (DNAW) 95 Effluent 22, 43, 198, 199
Delphacodes 103 Egretta 62, 155
Delphinus 151, 153 Ekman transport 124, 171
Dentalium 118 El Niiio 6, 19, 66, 67, 106, 172, 180, 181
Deposition 9, 14, 16, 18, 75, 81, 91, 92, 100, Elasmobranch 57, 141, 142, 144, 145,
117,163,165,169,181,200 146, 153, 183, 184, 185, 186, 187, 190,
Dermochelys 151, 154 195
Deterioration 3, 12, 43 Eledone 146, 147, 149, 159
Detritus 16, 39, 45, 65, 70, 71, 77, 82, 84, Eledonella 149
85, 115, 168, 170, 180, 181 Eleocharis 24
Diaphanosoma 40, 42 Elisella 120
Diaphus 121, 122, 128, 130 Emerita 109, 115, 116, 124, 154, 158, 168,
Diastylis 115, 116, 117 169,17°
Diatom 34, 35, 36, 59, 67, 72, 82, 83, 88, Encope 118, 119
105,106,1°7,108,167,168,169,170,174
Subject Index 299

Energy 9, 18, 19, 37, 47, 48, 49, 50, 65, 70, Evolution 13, 14, 15, 18, 91, 145, 146, 165,
71,72,82,85, 113, 139, 145, 164, 166, 167, 182
170,174,199 Excirolana 47, 48, 49, 50, 115, 116, 154, 158
Engraulididae 51, 58, 61, 73, 120, 124, 154 Exocoetidae 121, 123
Engraulis 52, 54, 55, 61, 120, 121, 122, 123, Exploitation 2, 3, 132, 134, 151, 153, 160,
124, 125, 127, 128, 134, 135, 136, 155, 172, 185,186,188, 193, 194, 195, 207
174,175, 176, 181, 192, 193, 195
Enneagonum 110 Feeder, feeding 1, 39, 44, 45, 50, 57, 59, 61,
Enteromorpha 32, 33, 67, 83, 86, 88 62, 63, 64, 71, 72, 73, 77, 82, 84, 88, 103,
Epibenthic 58, 65, 70, 71, 72, 78, 133 104,115,116,121,124,125,126, 127, 132,
Epifauna 43, 45, 46, 72, 77, 78, 80, 82, 84, 133,134, 135, 138, 139, 147, 150, ',52, 153,
178 154,155, 156, 158, 159, 168, 169, 170,172,
Epinephelus 51 174,175,177,178,179,182,183
Epiphyte 28, 35, 45, 67, 69,70,71,72,77, Fish 10, 12, 45, 47, 50, 51, 52, 53, 54, 56, 57,
82, 83, 84, 199 58,59,62,64,71,72,73,77, 78, 80, 81,
Epipsamic 77 82, 84, 85, 88, 116, 119, 120, 121, 122, 123,
Epitonium 117 126,127, 128, 129, 1~1, 132, 133, 134, 135,
Eretmochelys 151, 154 136,141,145, 147, 150, 152, 153, 154, 155,
Eriochloa 26 159, 168, 169, 170, 171, 172, 174, 175, 177,
Erodona 43, 44, 45, 73, 79, 81, 84, 133 178, 179, 180, 181, 182, 183, 184, 185, 186,
Erosion 14, 16, 21, 75, 94, 100, 101, 161, 165, 187,190,192,193,195, 197
166, 200, 202 Fisherman 64, 153, 182, 183, 195, 196
Etmopterus 141, 143 Fishery 1, 3, 12, 64, 85, 120, 121, 126, 128,
Eubalaena 151, 152, 153 129,141, 142, 143, 146, 152, 155, 156, 160,
Eubosmina 40 171,182,183,184,185,186,187, 188, 190,
Eucalanus 111, 169 191, 192, 193, 194, 195, 196, 203
Eucalyptus 165, 202 Flabellum 120
Eucheilota 110 Flagellate 34, 36, 37, 39, 66, 105, 106, 108,
Eucinostomus 60 109,167,169,174,179
Eucoila 103 Flamingo 157
Eudoxoides 110 Flatfish 50, 51, 136, 188, 195
Eugomphodus 143, 144 Flora 5, 16, 24, 30, 32, 67, 88, 98, 205
Eukrohnia 112, 113 Foodweb 35, 37, 39
Eunice 120 Foraminifera 110
Eupanthalis 119 Freshwater 1, 6, 9, 18, 19, 20, 21, 23,37, 40,
Euphausia 111, 127, 134, 135, 174, 177 42,43,51,52,57, 88, 95, 96, 105, 114, 117,
Euphausiid 110, 111, 114, 125, 133, 134, 174, 124, 132, 136, 137, 138, 161, 162, 163, 171,
175 172,180, 181, 182, 183, 197, 198, 199, 207
Euphotic 98, 171, 181 Frittilaria 113
Euphysora 113 Frog 104, 163
Eurycercus 40 Front 5, 94, 95, 96, 108, 124, 161, 166, 167,
Euterpina 40, 42, 111 171, 174, 179, 193
Euthynnus 121, 123, 128 Fulica 62, 63, 84
Eutintinnus 109 Fulmarus 155, 194
Eutrophication 179, 180, 182, 198
Euzonus 15 Gadidae, gadiformes 121
Evadne 111 Galeocerdo 143
Evaporation 5
300 Subject Index

Galeorhinus 128, 142, 143, 144, 145, 175, Hawk 154, 155
176,177,184,191,192,195 Heleobia 43, 44, 45, 72, 77, 78, 79, 80, 84,
Gamochaeta 99, 100, 163 133
Gastropod 43, 44, 77, 78, 80, 84, 116, 133, Helicolenus 130, 131, 136, 176, 178
134, 154, 169 Hemicyclops 115
Gelidium 88 Hemipodus 44, 79
Gempylidae 121, 123 Hepatus 119, 133
Genidens 57, 58, 59, 60, 73 Heptranchias 143,176
Genypterus 130, 175, 176 Heron 62, 63, 155, 170
Germination 27, 99, 100, 101 Heteromastus 43, 45, 71, 72, 77, 79, 80
Gerreidae 57 Heteroteuthis 148
Gill-net 64, 152, 183, 184, 185, 186, 187, 188, Hexanchidae, hexanchiforme 141, 145
190, 192, 195 Hexanchus 143
Globicephala 151, 152 Himantopus 63, 155
Globigerinoides 110 Hippopodius 110
Globoquadrina 110 Histioteuthis 148
Globorotalia 110 Holothurian 12.0
Glycymeris 118, 133 Hook 152, 183, 184, 185, 187, 194
Gobiesocidae 51 Hyalocylis 110
Gobiesox 53, 55 Hydrobiidae 78, 80
Gobiidae 51, 52, 57, 121 Hydrocotyle 24, 99, 100, 101, 102, 103, 162,
Gobionellus 55, 56, 60 163
Gobiosoma 55, 56 Hydruga 150
Gonostoma 121 Hygophum 121
Gonostomatidae 120, 122, 175, 176 Hyperoodon 150, 151
Grampus 151 Hypleurochilus 88
Grateloupia 88 Hyporhamphus 55, 56
Grazing 39,66,70,73,169,174,175,201,
202 Ichthyoplankton 51, 52, 55, 56, 120, 121,
Growth 124, 132, 138, 139, 141, 145, 150, 163, 122, 123
165,167,175,182,190,193,195,197 Idiacanthidae 120
Grubeulepis 115 Idiacanthus 120
Gull 62, 63, 154, 155, 156, 158, 159, 170 Ihlea 113
Gurgesiella 143 Illex 128, 134, 135, 136, 147, 148, 150, 154,
Gymnodinium 35 174,175,176,177,194
Gymnogongrus 88 Imperata 101
Gymnura 142, 144 Indian 10
Gyrodinium 108, 109, 167 Industry, industrial 2, 12, 22, 23, 43, 59,
146, 155, 156, 183, 184, 185, 188, 196, 197,
Habitat 9, 44, 45, 46, 47, 51, 62, 65, 70, 76, 198, 199, 200, 205
78, 79, 81, 82, 85, 86, 88, 104, 137, 146, Infauna 43, 44, 46, 71, 72, 73, 77, 80, 82,
149,151,154,155,161, 162, 163, 165, 171, 84,178
199,200,201,202,203 Inlet 9, 16, 18, 19, 20, 21, 23, 30, 40, 44, 52,
Haematopus 155, 158 62,63,78, 86, 88, 92, 133, 153, 181, 182,
Halistylus 118 188,195,197,199,200
Hapopleudes 103 Insect 44, 77, 102, 103, 104, 114, 115, 155,
Haptophyceae 34 159,162,163,169,170
Harengula 120 Interstitial 73
Subject Index 301

Intertidal 24, 27, 40, 44, 47, 78, 80, 81, 82, Ligyrus 103
86, 88, 109, 114, 115, 116, 168, 170, 179, Limacina 110, 111, 113
199,200 Limonium 24
Invertebrate 43, 47, 70, 71, 81, 84, 85, 114, Limosa 63
115,116, 118, 120, 129, 158, 169, 170, 177, Liocranchia 149
179, 193, 201 Liolaemus 104
Irradiance 28, 36, 66 Liriope 110
Irrigation 197 Listroderes 103
Ischaemum 24 Littorina 88
Isopod 44, 45, 47, 72, 77, 78, 80, 116, 117, Lizard 104, 163
119,120,147, 178 Lobodon 150
Isthiophorus 128 Loligo 127, 128, 131, 134, 135, 136, 146, 147,
148, 152, 154, 172, 175, 176, 194
fapetella 149 Lolliguncula 146
Jellyfish 135 Lophius 130
fenynsia 57, 60, 61 Lopholatilus 130
Jetty 30,86,88,200 Loxopagurus 119, 133
funcus 24, 25, 102, 163, 201 Lucicutia 111
Lucifer 134
Kalliapseudes 43, 44, 45, 71, 72, 79, 81, 84 Lycengraulis 51, 52, 53, 54, 55, 56, 57, 60,
Katsuwonus 121, 123, 124, 126, 127, 128, 176, 61, 120, 122, 123, 124, 169
177,184,187,191,193,194 Lycoteuthis 136, 149, 150
Kinbergonuphis 116, 117 Lyngbya 32, 33, 67, 86
Kogia 150, 151, 152 Lystrophis 104
Krohnitta 112, 113
Kurzia 40 Macroalgae 26, 30, 31, 32, 33, 39, 45, 65,
67,69,70,72,78,80,82,83,181
La Plata River 1, 92, 95, 96, 97, 105, 113, Macrochiridothea 115, 116, 117
121, 124, 171, 180 Macrodon 51, 52, 53, 55, 56, 57, 58, 60, 121,
Labidura 103 128,130, 131, 132, 134, 153, 176, 185, 187,
Laeonereis 43, 44, 45, 71, 77, 79, 80 188, 189, 191
Lagocephalus 57, 127 Macrofauna 43,44,45,114,116,117,118,
Lagrioida 103 119, 168, 169
Lamniform 141 Macronectes 155
Lampanyctinae 128 Macrophyte 23,39,65,70,72,77,84,199
Landing statistics 183, 185, 186, 187, 188, Macrothrix 40
190 ,192,193,194,195 Macrouridae 121, 131
Landscape 166, 207 Mactra 115, 116, 117, 119, 133, 168, 169
Larus 62, 63, 155, 157, 158, 159, 170 Magarethia 121
Lensia 110, 113 Magelona 44, 115, 116, 168
Leocrates 115 Makaira 128, 193
Lepidopus 121, 123 Malacocephalus 130
Leprotintinnus 39 Malvinas Current (MC) 1,94,95,96,105,
Leptocheirus 78 122, 124, 129, 132, 146, 150, 172, 174, 177
Leptochela 135 Mammal 63, 150, 151, 163, 194
Libina 133 Management 3, 12, 160, 166, 182, 196, 203,
Light 25, 27, 28, 29, 30, 32, 34, 36, 66, 67, 205,207,208
99, 107, 108, 138, 167, 199, 207
302 Subject Index

Marsh 9, 16, 18, 24. 25, 26, 27, 43, 44. 62, Moenkhausiana 103
63,67,68,69,70,71,72,73,74,75,76, Moina 42
77, 78, 102, 114, 161, 163, 165, 166, 181, Mollusc 18, 40, 47, 84, 109, 110, 116, 117,
200, 201, 202, 205 119, 124, 132, 133, 154, 159, 168, 175, 177
Mastogloia 35 Molpadia 120
Maurolicus 121, 122, 127, 128, 175, 176, 177 Monomorium 102
Mecynocera III Moroteuthis 148
Medusae 42, 110, 113 Mortality 28, 49, 50, 80, 81, 102, 109, 125,
Megabalanus 88 133, 138, 139, 141, 145, 152, 153, 154, 167,
Megalocranchia 149 188, 195, 199, 201
Megaptera 151 Mudflat 25, 43, 44
Meiobenthos 70, 71 Muggiaea 110
Melita 72, 78, 79 Mugil50, 52, 55, 56, 57, 59, 60, 72, 77, 84,
Mellita 115, 116, 117 124,125,154,169,176,183,191
Melosira 35, 86 Mugilidae 58, 59, 85, 121, 123
Menticirrhus 51, 52, 55, 57, 58, 60, 61, 130, Mullus130
159,169 Munida 135
Merluccidae 121 Munna 78, 79
Merluccius 121,122,127,128,130,131,135, Muricea 120
136, 175, 176, 191 Muriceides 120
Meroplankton 40, 169 Mustelus 142, 143, 144, 145, 177, 191, 192,
Mesocyclops 40 195
Mesodesma 47, 48, 109, 115, 116, 124, 154, Mycetophylax 102
158, 168, 169, 170 Myctophidae 121, 122, 128, 131, 175, 176
Mesodinium 34, 35, 36, 106 Myctophum 121, 128
Mesoplodon 150, 151 Myliobatidae, myliobatiforme 141, 145,
Metal 12, 18, 23, 47, 199 174
Metamysidopsis 42, lll, 169 Myliobatis 141, 142, 143, 144, 159, 176, 177
Metasesarma 43, 44, 72, 76, 79 Myriophyllum 27
Meteorology 34, 36,66,106,167,179,181, Myriopsis 120
208 Mysidopsis 42
Microalgae 34, 35, 65, 67, 69, 70, 71, 72, 77, Mysid 40, 42, 110, Ill, 113, 133, 134, 135,
78,82,103 169
Microbe, microbial 37, 39, 70, 71, 72, 77,
109,166,168,170,174 N:P ratio 97, 167
Microcoleus 32, 33, 67 Nanoflagellate 105, 106
Microcystis 34,198 Nanoplankton 36, 174
Micropogonias 51, 52, 53, 54, 55, 56, 57, 58, Narcine 143
59, 60, 64. 73, 77, 121, 129, 130, 131, 132, Natica 117, 118
152,154,160,176, 177,182,183,185,187, Navicula 35
188, 189, 190, 191, 195 Neanthes 117
Microsetella 174 Nematobrachion III
Migration 5, 15, 42, 45, 57, 82, 95, 113, 115, Nematod 71, 72, 77
125, 126, 127, 128, 131, 132, 133, 134, 142, Nematoscelis 111
143, 153, 155, 156, 157, 158, 159, 170, 177, Neocallichirus 115, 116
182, 192, 193 Neolatus 121, 123
Milvago 155 Neomysis 42, III
Modelling 20, 208 Neotridactylus 103, 115
Subject Index 303

Nephasomma 119 Orcinus 152

Nephtys 43, 45, 46, 72, 77, 79, 80, 116 Organic matter 37, 39, 65, 71, 72, 77, 84,
Netuma 57, 58, 59, 60, 73, 131, 130, 133, 169, 98,102, 103, 104, 161, 167, 168, 179
176, 177,182, 183, 185, 190, 191, 195 Ornithotheuthis 127, 148, 150, 175, 176
Nitrate 23, 96, 97, 98, 100, 167, 171, 180 Orthoptera 103, 115
Nitrite 50, 96, 97, 98 Osmoregulation 47, 48, 49, 50, 80
Nitrogen 22, 36, 47, 50, 67, 97, 167, 198 Ostracod 71, 72, 110
Nitzschia 35, 82, 105, 108 Ostrea 88
Noctiluca 105, 108, 109, 167 Otaria 63, 153, 175, 176, 194
Notodiaptomus 40, 42 Overfishing 185, 186, 188, 195
Notorhynchus 143 Owenia 116, 118, 119
Nucula 118 Oxygen 21, 49, 50, 97, 98, 127, 138, 139
Nursery 1,51,52,56,57,59,73,78,82,85, Oxytoxum 105
121, 122, 125, 132, 155, 169, 174, 179, 182, Ozone 197, 202, 203
193
Nutrient 1, 12, 22, 25, 34, 37, 66, 67, 96, 97, Pachyptila 156
98, 100, 101, 102, 106, 107, 109, 163, 171, Pagrus 130, 131, 135, 175, 176, 177, 178, 183,
172, 179, 181, 197, 198,201, 207 186,188, 191, 195
Palaemonetes 43
Obelia 86, 110 Palmeria 106
Oceanites 157 Panicum 24, 100, 103, 162, 163, 165, 202
Octopodidae 147 Panthalis 119
Octopus 146, 147, 149 Paracalanus 40, 111
Ocypode 114, 115, 169 Paracymus 103, 115
Ocythoe 149 Paralichthys 50, 51, 55, 56, 57, 130, 136, 185,
Odontella 34 188, 191
Odontesthes 57,60, 61,77,84, 136, 137, 138, Paralonchurus 51, 52, 53, 55, 56, 57, 58, 60,
139, 140, 141, 154, 159, 160, 169 121,130, 131, 132, 134, 153, 175
Odonthosy/lis 116 Parandalia 116, 117
Oenothera 99 Parapimelodus 51, 52, 53, 55, 56, 57
Oikopleura 112, 113 Paraprionospio 117
Oithona 111 Pareledone 147, 149
Oligotrichid 39 Parana 52, 53, 55, 123, 127, 128
Olivancillaria 115, 116, 117, 118, 119, 169 Particulate organic carbon 37, 168, 179,
Olivella 117 207
Ommastrephes 148, 150 Parvanachis 116, 117
Ommastrephidae 127, 175 Paspalum 24, 101, 102, 162, 163
Oncaea 40, 111, 174 Pelecipod 43, 80, 81
Oncopterus 169 Penaeus 44, 45, 47, 48, 49, 50, 72, 78, 79,
Onuphis 44, 79, 119 84, 85, 152, 160, 176, 182, 183, 191, 193
Onychia 148 Penguin 155, 199
Onychoteuthis 148 Pen ilia 42, 111, 175
Ophiomisidium 120 Peprilus 51, 52, 55, 56, 57, 61, 124, 127, 128,
Ophiuroid 118, 119, 120, 132, 134 130, 131, 135, 175, 176
Ophiurolepis 120 Peracarid 44, 45,72,78,80, 81
Ophyctus 51 Peridinium 34
Orchestia 44, 77, 79 Perna 88
Orchestoidea 114, 115, 162 Persephona 119
304 Subject Index

Pesticide 47, 50, 199 Pollichthys 120, 122

Petrel 154, 155, 157, 194 Pollutant, pollution 43, 50, 199
pH 48, 50, 97 Polyborus 155
Phaeophyta 30 Polychaete 40, 43, 44, 46, 72, 80, 81, 84,
Phalacrocorax 62 86, 110, 115, 116, 117, 118, 119, 120, 132, 133,
Phaleria 114 134, 135, 147, 168, 177
Pheidole 102 Polykrikos 109
Phocoena 150, 151, 152 Polymixia 130
Phoenicopterus 157 Polypogon 24
Phosphate 22, 23, 92, 96, 97, 98, 100, 179, Polyprion 130, 131, 136, 150, 175, 176, 177,
19 8 186,191
Photichthtidae 120 Polyprionidae 131
Photosynthetic, photosynthesis 36, 67 Polysiphonia 86
Phoxocephalopsis 115, 116, 168 Pomacentridae 57
Phyla 102, 163 Pomatomidae 61
Phyllochaetopterus 119 Pomatomus 57, 124, 125, 126, 127, 128, 154,
Physeter 151, 152 176, 177, 183,191, 192, 193
Physiology 47, 48, 50 Pontella 111
Phytoplankton 34, 35, 36, 37, 39, 61, 66, Pontoporia 63, 150, 151, 152, 175, 195
67, 69, 70, 71, 73, 104, 105, 106, 107, 108, Porichthys 51, 52, 55, 57, 58, 60, 130, 131,
109, 125, 167, 168, 169, 171, 172, 174, 175, 135
179, 180, 181, 198 Porifera 120
Pickfordiateuthis 146 Port 10, 12, 23, 86, 183, 184, 200
Picoplankton 36, 104, 174, 175 Portunus 119, 135, 177
Pimelodus 57 Potamogeton 27
Pinniped 150, 153 Prasinophyceae 34
Pinnixa 115, 116 Precipitation 5, 6, 9, 16, 21,34,36,44,65,
Pinus165,202,206 66, 67, 73, 75, 78, 100, 101, 106, 180, 181,
Pitar 118, 119 182,208
Platanichthys 60, 61 Predation 45, 46, 56, 61, 72, 78, 80, 81, 82,
Plectris 103 84,86,121
Pleopys 42, 111 Predator 39,45,46, 51, 61, 72, 77, 80, 81,
Pleoticus 134, 135, 152, 154, 177, 185, 191, 194 82, 84, 103, 104, 109, 114, 115, 116, 118,
Pleurodema 104 136,146,169,170,175,177
Pleuronectidae, pleuronectiformes 121, Preservation 3, 206, 207
123,131 Priacanthus 131
Pleurosira 35 Priapulus 120
Pleuroxus 40 Primnoella 120
Pleuston 40 Prionace 176, 191
Plover 63, 155, 158, 159, 169, 170 Prionotus 51, 52, 55, 57, 130, 131, 134, 135,
Pluchea 102, 163 188,191
Plume 39, 181 Proarna 103
Pluvialis 157, 158, 170 Proboscia 106
Podiceps 62 Proboscidactyla 110
Podon 42, 111 Procellaria 156, 177, 194
Poecilidae 57 Procellariiforme 154
Pogonias 57, 60, 131, 133, 169, 176, 183, 185, Production 1,28,29,3°,32,36,37,39,49,
190, 191, 195 65, 66, 67, 68, 69, 70, 77, 81, 82, 84, 97,
Subject Index 305

104, 106, 107, 108, 115, 116, 121, 125, 145, Rhyncalanus 111
167, 168, 171, 172, 179, 180, 181, 182, 200, Ritteriella 113
205,207 Rostratoverruca 120
Productivity 36,66,67,98,106,177,203 Runoff 1, 6, 12, 19, 21, 34, 51, 52, 56, 66, 73,
Promysis 42, 134 77, 88, 92, 95, 96, 97, 106, 107, 113, 117,
Prorocentrum 34, 35, 36,105, 106 119, 124, 132, 171, 172, 179, 182, 197, 198,
Protoperidinium 109 199
Protozooplankton 39, 109, 174 Ruppia 27, 33, 35, 45, 62, 67, 68, 69, 70, 71,
Psammobatis 142,143 78, 82, 83, 84, 85, 199, 200
Psammodius 103 Rynchops 62, 63, 157, 159
Pseudo-nitzschia 105, 106
Pseudoevadne 111 Sagitta 40, 42, 112, 113
Pseudorca 151, 152 Salicornia 25, 77
Pseudosida 40 Salinity 19, 20, 21, 23, 24, 25, 26, 27, 28,30,
Pseudosolenia 106 32,34, 35, 40, 42, 44, 47, 48, 49, 50, 52,
Pseudosphaeroma 80 53, 55, 61, 63, 67, 73, 76, 80, 86, 88, 107,
Pteria 119 110,122,123,124;"12-5,138,162,197,198
Pterocaulon 99 Salp 135
Pteroctopus 149 . Salpa 113
Pterodroma 155 Sandpiper 63, 158, 159, 170
Pteropod 110, 113, 175 Sapphirina 111
Pterosagitta 112, 113 Sarda 121, 123, 124, 126, 127, 128, 176
Pterygioteuthis 148 Sardinella 120
Puffinus 155, 156, 157, 177, 194 Sarsia 110
Satellite 1, 185, 205
Radiation 5, 202 Saurida 127, 130
Raja 142, 143, 144, 175, 176 Scaeurgus 149
Rajidae, rajiforme 141, 145 Scapteriscus 103
Ramnogaster 57,60,61 Scarites 103
Ray 141,142,143,169,172,174,185,190,195 School (=shoal) 59, 124, 126, 127, 159, 177,
Recruitment 44, 45, 49, 56, 72, 73, 80, 81, 184, 192, 194
85,125,143,145,188,193,195 Schroederichthys 143
Regression 13, 92, 161 Sciaenidae 51, 57, 58, 59, 61, 78, 85, 121,
Reproduction 1,32,42, 45, 48, 56, 88, 122, 122,131,133,134,152,154,159,172,174,
125, 126, 127, 132, 133, 134, 135, 136, 143, 183, 185, 187
145, 147, 151, 153, 183, 197 Scirpus 24,25,26,68,69,75,77,201
Rhinohatidae 145 Scomber 121, 123, 124, 125, 126, 128, 131, 176,
Rhinobatos 142, 143, 144, 145, 174, 176, 190, 186, 191, 193
191, 192, 195 Scombroidei 121, 159
Rhinoptera 143, 144 Scrippsiella 105, 106
Rhithropanopeus 72, 78 Scyliorhinus 143, 145
Rhizoclonium 32, 33, 67, 83 Sea lion 63, 175, 194
Rhizoprionodon 142 Seagrass 43, 70, 82, 181, 182, 200
Rhizopsammia 120 Sediment 9, 12, 13, 16, 18, 20, 21, 22, 23, 25,
Rhizosolenia 34, 35, 105, 106, 108 27,33,35, 43, 46, 66, 70, 71, 72, 73, 75,
Rhodophyta 30 77,78, 80, 81, 82, 84, 86, 92, 94, 96, 103,
Rhopalodia 35 108,117,162,163,167,179,181,198,199,
Rhopalonema 110 200
306 Subject Index

Sedimentation 92, 108, 177, 181, 200 Sphyrna 128, 141, 143, 144. 174. 1]6, 193
Seed 27, 84. 98, 101, 104, 163 Sphyrnidae 145
Seine 58, 59, 61, 126, 143, 152, 183, 184, 185, Spider 77, 103, 104
186, 187, 192, 193 Spio ll5, ll6, 168
Selene 61 Spiochaetopterus ll9, 120
Semirossia 146, 147, 148 Spirula 148
Senecio 25, 100, 103, 163 Squaliforme 141
Sergestid 134 Squaliolus 141, 143
Serolis 119 Squalus 142, 143, 144, 145
Serranid 57, 121, 131, 136, 152 Squatina 142, 143, 144, 145, 159, 175, 176,
Sewage 23, 63, 198 185, 191, 192, 195
Shark 127,141,142,143,152,169,171,172, Squatinidae, squatiniforme 141, 145
174,175,176,184,185,186,190,192,193, Squid 126, 136, 155, 172, 174, 175, 177, 194
194,195 Squilla ll9, 134
Sheathbill154 Starvation 139
Shoal 16, 18, 27, 30, 34. 35, 36, 40, 42, 44, Stegosoma ll3
45, 56, 57, 73, 78, 80, 182, 199, 200, 207 Stenella 150,.151, 152
Shrimp 10, 40, 44, 47, 64, 84, 85, 132, 134, Steno 150, 152
136, 152, 154, 155, 177, 182, 183, 185, 186, Stercorariu5 157
190, 191,'193, 194. 195 Sterna 62, 63, 155, 157 159, 177, 194
Sigalion ll5, ll6, 117 Sternoptychidae 120
Sigambra44 Sthenelais ll6
Silicate 22, 23, 96, 97, 167, 180 Stheneloplis ll6
Silicoflagellate 106 Stilt 63, 155
Siluriforme 51 Stock 1, 3, 30, 85, 114, 125, 126, 131, 132, 133,
Simosa40 134,147,150,151,182,183,185, 188,190,
Siphonaria 88 193, 194, 195, 196
Siphonophore 42, llO, ll3 Stomatopod 135
Skate 141, 142, 143, 169, 172, 175, 194, 195 Stomias 120
Skeletonema 34, 35, 36, 106, 167, 169, 180 Stomiidae, stomiiformes 120, 121, 122
Skimmer 62, 154. 155 Stratification 20, 21
Skua 155, 157 Stromateu5 124. 127, 128
Skunk 104, 163 Strombidium 38, 39, 109
Snail 40 Stylocheiron 111
Soleidae 121 Stylochoplana ll5
Solen ll7, 177 Stylochus 86
Solenopsis 102 Stylocidaris 120
Sotalia 150, 152 Subantarctic Water (SAW) 1, 94, 95, 97,
South Atlantic Central Water (SACW) 105, 106, 107, 109, lll, 112, 113, ll4, 171,
94. 171, 173 172,173,174,184
Spartina 24, 25, 26, 44, 68, 69, 75, 77, 100, Substrate 16,30,32,35,40,44,45,67,70,
101,102,103,162,163,165,201 71, 72, 77, 80, 81, 82, 86, 88, 89, 100, ll6,
Spawning 52, 55, 81, 121, 122, 125, 131, 132, ll7, ll8, ll9, 120, 129, 131, 158, 161, 162,
liI3, 134, 135, 136, 137, 138, 147, 150, 154, 163
160, 171, 180, 181, 182, 188 Subtropical Convergence (SC) 1,3, 5,94.
Spermatophyte 27, 67 95, 98, 105, ll4, 127, 129, 130, 135, 150,
Spheniscus 155 171,172,174,193,194
Sphizogenius 103
Subject Index 307

Subtropical Water (STW) 42, 94, 95, 98, Thunnus 127, 128, 150, 175, 176, 187, 191,
106,107,109,112,121,123,127,129, 135, 193
171,172 Thyrsitops 121, 123, 127, 128, 130, 131
Sulculeolaria 110, 113 Thysanoessa 111
Supratidal 114, 115, 169 Thysanopoda 111
Surfzone 94, 105, 108, 113, 114, 115, 116, 117, Thysanoteuthis 148, 149
124,125,159,166,167,168,169,17°,179 Tide 19, 63, 66, 69, 78, 100, 114, 116, 138,
Surirella 35 158, 159, 162, 165, 170, 207
Suspended matter 21, 23, 97, 98, 181, 199, Tintinnid 39, 109, 174
200 Tintinnopsis 39, 109
SymbolophoTUs 128 Tivela 168, 169
Symphurus 51, 52, 55, 57, 175 Todarodes 148
Sympterigia 142, 143, 144, 175, 176, 177 Torpediniforme 141
Synagrops 52, 55, 130, 136 Torpedo 143
Synedra 35, 82, 83 Toxic 25, 50, 108, 167, 179, 198
Syngnathidae 85 Trachinotus 57, 64, 123, 124, 169
Syngnathus 51, 52, 55, 56 TrachuTUS 123,124,-125,126,127,128,130,
Synidotea 44, 115, 116, 117 131,135,136,175,176,186, 191, 192, 193,
195
Tachytes 103 Transgression 13, 14, 92, 161, 165, 202
Tagelus 71, 79, 80 Transport 1, 12, 16, 22, 42, 50, 52, 82, 92,
Tanaid 43, 45, 72, 78, 81, 86 94, 95, 98, 106, 121, 124, 161, 162, 164,
Tanais 43, 72, 78, 79, 86 165, 171, 179, 181, 183, 198, 202
Teleost 50, 123, 129, 130, 131, 139, 152, 153, Trawl 58, 59, 61, 125, 127, 130, 134, 135, 141,
154,174,184,185,186,187,189,190,192, 142,143,172,177,183,184,185,186,187,
194,195 188,189,19°,192,193,194,195
Tellina 117 Tremoctopus 149
Temora 111, 174 Trichiuridae 61, 121, 131
Temperature 5, 20, 21, 27, 28, 32, 34, 35, Trichiurus 51, 52, 53, 54, 55, 56, 57, 58, 61,
36,42,44,47,49,50,55,66,67,68,86, 64,121,123,127,128,130, 131, 134, 153,
96,99, 100, 102, 103, 107, 110, 122, 124, 154,172,174,175,176,185,191,19 2,193
125, 126, 129, 135, 138, 146, 181, 192, 193, Trichodesmium 108
207 Triglidae 121, 123
Terebellides 119 Trochocyathus120
Terebra 117 Trophic 37, 39, 46, 70, 71, 72, 73, 80, 82,
Tern 62, 154, 155, 156, 157, 159, 170 85, 88, 104, 113, 145, 168, 170, 171, 174,
Terpsinoe 35 177,203
Tetragonoderus 103 Tropical Water (TW) 94, 95, 96, 105, 106,
TetraptuTUs 128, 193 '107,111,112, 123,143,150,171,172,173,
Thais 88 184
Thalassionema 106 Tuna 127, 128, 141, 150, 159, 171, 177, 184,
Thalassiosira 106 187,193,194
Thalia 113 Tunicate 110, 112
Thaliaceae 113, 114, 175 Turbelarian 71, 115
Thermocline 95, 96, 113 TUTTitopsis 110
Thermohaline 95, 96, 124 Tursiops 63, 64, 151, 152, 195
Threskiornitidae 62 Turtle 150, 151, 154
Thronistes 103 Typha 24, 102, 163
308 Subject Index

Ulothrix 32, 33, 67 Wave 14, 15, 16, 18, 19, 20, 23, 26, 29, 33,
Ulva 88 88,94, 116, 158, 164, 165, 166, 167, 170,
Ulvaria 86 199,202
Umbrina 51, 52, 55, 57, 58, 59, 60, 129, 130, Wind 5, 16, 19, 20, 21, 23, 26, 28, 34, 36, 40,
131,134,136,174,176,177,183,185,187, 44, 66, 78, 85, 88, 94, 95, 96, 98, 100,
188, 189, 191, 195 103, 155, 158, 161, 162, 163, 164, 165, 166,
Upwelling 95, 96, 98, 106, 107, 124, 129, 167,179,199,202, 207,208
135, 171, 172
Urophycis 51,57, 121, 130, 131, 136, 176, 177, Xantophyta 30, 32
178, 188, 191 Xiphias 128, 150, 152,175, 176, 187, 191, 193
Yoninae 115
Valenciennellus 120
Vaucheria 32, 33,67 Zanclea 110
Verruca 120 Zannichellia 27, 67
Vigna 25 Zapteryx 142, 144
Vinciguerria 120, 122 Zeidae 121
Vosseledone 146, 147, 149 Zenopsis 122, 130
Ziphius 151
Wader 63, 154, 155, 156, 157, 158 Zoanthidea 120
Water leVel, seawater level 2, 13, 16, 19, Zooplankton 40, 41, 42, 43, 61, 66, 71, 73,
20,24,26, 28, 75, 77, 78, 91, 92, 115, 158, 110,111,112, 113, 114, 146, 155, 168, 169, 170,
161, 197, 202 175,177, 180, 181, 199