1978 Book UpwellingEcosystems

Upwelling
Ecosystems
Edited by
R Boje and M. Tomczak
Contributors
R T. Barber J. R Bennett M. Blackburn D. Blasco
R Boje A. Bustamante D. H. Cushing L. Diester-
Haass M. Elbrachter M. Estrada P. M. Hargreaves
A. Herbland R W Houghton G. T. Houvenaghel
B. Koopmann R Margalef O. A. Mathisen
M. A Mensah T. T. Packard T. Rivera M. Sarnthein
H.-J. Schrader H. Thiel A. Thiriot J. D. Thompson
R E. Thome M. Tomczak R J. Trumble
S. W Watson T. E. Whitledge W S. Wooster
S. Zuta
With 132 Figures
Springer-Verlag
Berlin Hddelberg New York 1978
Dr. ROLF BOJE
lnstitut flir Meereskunde an der Universitat
Diistembrooker Weg 20, 2300 KiellFRG
Dr. MATTHIAS TOMCZAK

Department of Oceanography
The University of Liverpool
Bedford Street North
P. O. Box 147, Liverpool L69 3BX/GB
ISBN-13: 978-3-540-08822-6 e-ISBN-13: 978-3-642-66985-9

001: 10.1007/978-3-642-66985-9
Library of Congress Cataloging in Publication Data. Main entry under title: Upwelling
ecosystems. Bibliography: p. Includes index. 1. Marine ecology. 2. Upwelling (Oceano-
graphy). I. Boje, R, 1934-. II. Tomczak, M., 1941-. QH541.5S3U57. 574.5'2636. 78-15685
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2131/3130-543210
Preface
Upwelling areas are among the most fertile regions of the

ocean. In principle, upwelling is caused by the divergence
of the flow in the surface layer of the ocean which
arises as a consequence of a particular wind field, the
presence of a coastline, or other special conditions. Since
deeper oceanic layers are usually enriched wi th nutrients,
it is the permanent supply of nutrients which forms the
basis for the high producti vi ty of upwelling reg ions.
The study of upwelling and its consequences were, for
a long time, the task of individual scientists from all
disciplines of marine science. Today, it is perhaps the
branch of oceanography where interdisciplinary coopera-
tion has developed best. Becoming aware of the large
potential yield of upwelling regions, governments in-
creased the funds for upwelling research. With research
activities developed on a larger scale, interdisciplin-
ary cooperation became a necessity.
On the international level, several symposia documented
the rapid development. Three volumes reflect the results
of these scientific meetings (Rapp. Proc.-Verb. 159,
1970; Inv. Pesq. 35, 1, 1971; Tethys §.' 1-2, 1974).
The present book contains selected papers from the
Third Symposium on Upwelling Ecosystems, which was held
in Kiel in September 1975. Although the third of a
series of meetings, it was the first where the word
"ecosystem" stood in the title for a scientific program.
Moreover it clearly reflected the Tendency to unite the
efforts of physical, chemical, and biological oceano-
graphers, marine meteorologists, and marine geologists
into one common aim, the understanding of the process
of upwelling and its consequences.
The contributions to this volume were refereed by scien-

tists from the respective fields and revisen by the
authors. There are three papers of more general content,
two are on the application of ecosystem analysis in up-
welling studies (Boje and Tomczak; Margalef), one dis-
cusses the future of upwelling research (Wooster). As-
pects of upwelling ecosystems are treated, which con-
cern phytoplankton (Estrada; Elbrachter and Boje) , zoo-
plankton (Thiriot; Hargreaves; Packard et al.; Whit-
ledge), fish (Cushing; Mathisen et al.), benthos (Thiel),
and bacteria (Watson; Herbland). The conditions at the
coast of Ghana (Houghton and Mensah) and in the vicinity
of the Galapagos Islands (Houvenaghel) are described in
additional papers. Hydrographical aspects of upwelling
systems are covered in three contributions (Thompson;
Bennett, who discusses results from the Great Lakes;
Zuta et al.), while two papers deal with geological
results from upwelling areas (Diester-Haass; Koopmann
et al.) .
VI
The present debate on the future of the exploitation

of marine resourcesis,of course, having its impact on
marine science and especially on upwelling research,
which deals with one of the largest living resources
of the oceans. It is obvious that the proposals put
forward at the United Nations Conference on the Law
of the Sea concerning marine research have led to
different opinions, which are clearly reflected in the
way the research goals in upwelling studies are for-
mulated. We sincerely hope that this book may stimu-
late a large number of marine scientists to think cri-
tically about the objectives of upwelling research and
to dive into the study of upwelling ecosystems in or-
der to make optimum use of these immense marine re-
sources.
The editors wish to thank Professor Dr. Gotthilf Hempel,
who convened the symposium and supported the editing
work. The Third Symposium on Upwelling Ecosystems
was made possible by grants from the German Research
Society, from the Scientific Committee on Oceanic Re-
search (SCOR), the Intergovernmental Oceanographic
Commission (IOC) and the United Nations Educational,
Scientific and Cultural Organization (UNESCO). The
German Research Society funded the German research
projects, which form the basis of five of the contri-
butions.
Especially we wish to thank the Springer-Verlag for

their good cooperation and their interest in producing
this book.
Kiel, August 1978 R. BOJE

M. TOMCZAK
Contents
Ecosystem Research in Upwelling Regions
Ecosystem Analysis and the Definition of Boundaries

in Upwelling Regions
R. BOJE and M. TOI1C ZAK •••••••••.•••••••••••••••. 3
What is an Upwelling Ecosystem?

R. !'-lARGALEF (With 1 Figure) ••••••.••••.•••.••.•. 12
Mesoscale Heterogeneities of the Phytoplankton

Distribution in the Upwelling Region of NW Africa
~l. ESTRADA (With 9 Figures) ••••••••••.•••••••••• 15
On the Ecological Significance of ThaZassiosira par-

theneia in the Northwest African Upwelling Area
M. ELBRACHTER and R. BOJE (With 7 Figures) •••••• 24
Zooplankton Communities in the West African Up-

welling Area
A. THIRIOT •••••.•••••••••••••••••••••.••.•••••.• 32
Relative Abundance of Zooplankton Groups in the

Northwest African Upwelling Region During 1968 and
1972
P.M. HARGREAVES (With 8 Figures) •.••••..•••••••• 62
Mesodinium rubrum in the Baja California Upwelling

System
T.T. PACKARD, D. BLASCO, and R.T. BARBER (With 5
Figures) .••••••••••••• '. • • • • • • • • • • • • • • • • . . • • • • • • • 73
Regeneration of Nitrogen by Zooplankton and Fish

in the Northwest Africa and Peru Upwelling Eco-
systems
T.E. WHITLEDGE (With 1 Figure) ••••••••.•••••••.• 90
Upper Trophic Levels in Upwelling Areas

D.H. CUSHING (With 2 Figures) ••••••••••••••••••• 101
Food Consumption of Pelagic Fish in an Upwelling

Area
O.A. MATHISEN, R.E. THORNE, R.J. TRU~mLE, and
M. BLACKBURN (With 5 Figures) ••••••••••••••.•••• 111
Benthos in Upwelling Regions

H. THIEL (With 4 Figures) ••••..•••••..•••••••••. 124
Role of Bacteria in an Upwelling Ecosystem

S.W. WATSON (With 2 Figures) •.••.......•.••.•••• 139
VIII
Heterotrophic Activity in the Mauritanian Up-

welling in March 1973: Assimilation and Minerali-
zation of Amino Acids
A. HERBLAND (With 11 Figures) ...••....••........ 155
Physical Aspects and Biological Consequences of

Ghanaian Coastal Upwelling
R.W. HOUGHTON and M.A. MENSAH (With 13 Figures) 167
Oceanographic Conditions in the Galapagos Archi-

pelago and Their Relationships with Life on the
Islands
G.T. HOUVENAGHEL (With 15 Figures) ......•..•.•.• 181
Hydrographical Aspects of Upwelling Regions
Role of Mixing in the Dynamics of Upwelling Sys-

tems
J.D. THO~WSON (With 6 Figures) 203
The Circulation of Large Lakes

J .R. BENNETT (With 6 Figures) •...••..•......•.•. 223
Hydrologic Aspects of the Main Upwelling Areas

off Peru
S. ZUTA, T. RIVERA, and A. BUSTAMANTE (With 16
Figures) .........•......•.........••............ 235
Geological Aspects of Upwelling Regions
Sediments as Indicators of Upwelling

L. DIESTER-HAASS (With 14 Figures) .........•.... 261
Sedimentation Influenced by Upwelling in the Sub-

tropical Baie Du Levrier (West Africa)
B. KOOPMANN, M. SARNTHEIN, and H.-J. SCHRADER
(Wi th 7 Figures) ..•......•.•....•......•........ 282
General Aspects of Upwelling Research
Upwelling Research and Ocean Affairs

W.S. WOOSTER ..•.........•....•.....••••...•..... 291
Subject Index .•......•...............•.•..••.... 301

Contributors
Dr. R.T. BARBER, Marine Laboratory, Duke University,

Beaufort, NC 28516/USA
Dr. J.R. BENNETT, Department of Earth and Planetary

Sciences, Massachusetts Institute of Technology,
Cambridge, MA 02139/USA
Dr. M. BLACKBURN, c/o Roderick Mager, P.O.Box 774,

Friday Harbor, WA 98250/USA
Dr. D. BLASCO, Bigelow Laboratory for Ocean Sciences,

West Boothbay Harbor, ME 04575/USA
Dr. R. BOJE, Institut fUr Meereskunde an der UniverSitat,

DUsternbrooker Weg 20, 2300 Riel/FRG
Dr. A. BUSTAMANTE, Instituto del Mar del Peru, Casilla

3734, Lima/Peru
Dr. D.H. CUSHING, Fisheries Laboratory, Pakefield Rd.,

Lowestoft, Suffolk/GB
Dr. L. DIESTER-HAASS, Geologisches Institut der Univer-

sitat, Pleicherwall 1, 8700 WUrzburg/FRG
Dr. M. ELBRACHTER, Biologische Anstalt Helgoland, Lito-

ralstation, 2282 List, Sylt/FRG
Dr. M. ESTRADA, Instituto de Investigaciones Pesqueras,

Paseo Nacional, sin, Barcelona-3/Spain
Dr. P.M. HARGREAVES, Institute of Oceanographic Sciences,

Wormley-Godalming, Surrey GU 8 5 UB/GB
Dr. A. HERBLAND, Centre de Recherches Oceanographiques,

29, Rue des Pecheurs, B.P.V. 18-Abidjan/Republique
de Cote d'Ivoire
Dr. R.W. HOUGHTON, Lamont-Doherty Geological Obser-

vatory, Columbia Universit~ Palisades NY 10964/USA
Dr. G.T. HOUVENAGHEL, Laboratoire d'Oceanologie, Uni-

versite Libre de Bruxelles, 50 avo F.D. Roosevelt,
1050 Bruxelles/Belgium
Dipl.-Geol. B. KOOPMANN, Geologisch-Palaontologisches

Institut der Universitat, Olshausenstr. 40-60,
2300 Kiel/FRG
x
Dr. R. MARGALEF, Catedra de Ecologia, Facultad de

Biologia, Universidad de Barcelona/Spain
Prof. Dr. O.A. MATHISEN, Fisheries Research Institute,

University of Washington, Seattle, WA 98195/USA
Dr. M.A. MENSAH, Fishery Research Unit, P.O. Box B-62,

Tema/Ghana
Dr. T.T. PACKARD, Bigelow Laboratory for Ocean

Sciences, West Boothbay Harbor, ME 04575/USA
Dr. T.• RIVERA, Instituto del r-lar del Peru, Casilla

3734, Lima/Peru
Prof. Dr. M. SARNTHEIN, Geologisch-Palaontologisches

Institut der Universitat, Olshausenstr. 40-60,
2300 Kiel/FRG
Dr. H.-J. SCHRADER, Oregon State University, School

of Oceanography, Corvallis, OR 97331/USA
Dr. H. THIEL', Institut fUr Hydrobiologie und Fische-

reiwissenschaft, Palmaille 55, 2000 Hamburg 50/FRG
Dr. A. THIRIOT t, Station ZOologique, 06230 Ville-

franche sur Mer/France
Dr. J.D. THOMPSON, Environmental Models Branch, Numer-

ical Modeling Division, NSTL Station, MS 39529/USA
Dr. R.E. THORNE, Fisheries Research Institute, Uni-

versity of Washington, Seattle, WA 98195/USA
Dr. M. TOMCZAK, Department of Oceanography, The Uni-

versity of Liverpool, Bedford Street North, P.O.
Box 147, Liverpool L69 3BX/GB
Dr. R.J. TRUMBLE, Fisheries Research Institute, Uni-

versity of Washington, Seattle, WA 98195/USA
Dr. S.W. WATSON, Woods Hole Oceanographic Institution,

Woods Hole, MA 02543/USA
Dr. T.E. WHITLEDGE, Oceanographic Sciences Division,

Brookhaven National Laboratory, Upton, NY 11973/USA
Dr. W.S. WOOSTER, Institute for Marine Studies HA-35,

University of Washington, Seattle, WA 98195/USA
Dr. S. ZUTA, Instituto del Mar del Peru, Casilla 3734,

Lima/Peru
Ecosystem Research in Upwelling Regions
Ecosystem Analysis and the Definition of Boundaries in
Upwelling Regions
R BOJE and M. TOMCZAK
1. Introduction
An ecosystem, according to Smith (1970) is "a functional unit with re-

cognizable boundaries and an internal homogeneity". Ellenberg (197 3a)
describes an ecosystem as "an interacting system between organisms and
their inorganic environment which is open but has to a certain degree
the ability of selfregulation."
In the marine environment, shallow water ecosystems can be distinguished

from open ocean ecosystems, but i t is quite difficult to draw boundaries,
especially in' the open ocean. This is confirmed by Ellenberg (1973b)
in his article on the classification of the ecosystems of the world.
Upwelling regions extent from shallow to great water depths. With the
present knowledge it is not possible to decide whether they can be more
adequately described as an individual ecosystem or as part of the open
ocean ecosystem. The reason for this is that not enough is known on the
structure and function of upwelling regions as ecological unitsl, and
on the boundaries of the areas which have to be included in the analysis.
These considerations do not necessarily imply that the application of

ecosystem analYSis in upwelling research is impossible, because the
methods of ecosystem analysis can be used for the investigation of a
complete ecosystem as well as for the study of a part of it.
2. Choice of the Research Goal and Sensitivity Analysis
Before dealing with the application of ecosystem analysis in the in-

vestigation of upwelling areas, we have to decide what the goal of our
research is. This is necessary because any evaluation of the use of
ecosystem analysis in upwelling studies depends on the goal of the in-
vestigations.
We will assume in the following that our main interest is to achieve

optimum use of the living resources of upwelling regions. This research
goal is only one possible goal, but in the opinion of the authors the
only important one at present. There are of course other goals which
might stimulate the interest of researchers in the analysis of upwelling
areas.
The basic task thus would be to model the upwelling system in order
to be able to predict its behavior under varying boundary conditions.
Once an adequate model summarizing already known facts is developed,
1 R. Margalef gives his ideas on the structure and function of upwelling systems in
the next contribution.
4
we start observation campaigns in order to find the correct numbers

for the boundary conditions, as well as for the transfer rates between
each of the important variables of the system. Since the large number
of variables gives us a vast research program, we investigate first the
importance of the observations of each individual variable. The question
we ask is: to what degree of accuracy do we have to determine all phys-
ical, meteorological, biological and chemical quantities in order to
take into account their influence on the higher trophic levels of marine
life if we want to predict and monitor optimum yield.
The method we have to apply is sensitivity analysis. It must be stressed

that the application of this method is again based on the choice of our
research goal.
As an example, let us consider the individual variable vertical veloc-

ity. Suppose two upwelling theories have been developed which differ
markedly only in one pOint: vertical velocity should be 10- 3 cm s-l
according to the first theory, 10- 2 cm s-l according to the second.
If we deem it necessary to decide which theory is correct, we will have
to determine vertical velocity to at least 10- 3 cm s-l which will cost
us much work and money. If, however, we think our goal should be to
improve our knowledge on the possibilities of using the living resources
of upwelling systems, it is by no means clear whether the determination
of vertical velocity to that degree of accuracy is necessary at all.
A review of past upwelling studies gives the impression that not only
with regard to vertical velocity a lot of work has been invested in
studies which do not yield new information for our knowledge of trans-
fer rates and boundary conditions, especially as needed for the study
of living resources in upwelling areas.
3. The Boundaries of Upwelling Regions
Although at present it is not possible to decide whether an upwelling

region should be described as a complete ecosystem on its own or whether
it cannot be separated from the larger oceanic ecosystem, we would still
like to find some natural boundaries of upwelling regions. This would
enable us to apply the ecosystem approach if we could manage to describe
the conditions of input and output on the boundaries and, thus, would
give us quite a useful tool for our upwelling studies. Of course, this
approach is rather pragmatic; it opens, however, the possibility of
applying a powerful existing technique to our problem.
3.1 Hydrographical Aspects
Whatever our choice for the boundaries of an upwelling region might be,
one fundamental condition is that these boundaries take into account
the hydrodynamic structure of the ocean.
If we look at the wind field, which undoubtedly defines the oceanic

circulation to a large extent, there is no reason to consider any scale
smaller than the dimensions of the earth. However, since we are dealing
with processes in the ocean, restriction to the scale of the ocean's
dimensions is justified. In order to find smaller natural scales. we
define a coordinate system with axes orientated vertically, cross-
stream and down-stream, and consider the scales of the three axes sep-
arately. In coastal upwelling regions the main flow is meridional, and
the cross-stream component of flow is zonal; in equatorial upwelling
5
regions the main flow is zonal, and the cross-stream component is

meridional.
In the vertiaaL, the natural boundaries set by the ocean's dimensions

are the surface and the bottom. They represent discontinuities for all
properties; inflows and outflows may be minimized there. We may, never-
theless, consider smaller scales as alternatives. As can easily be
seen, the frictional or Ekman layer depth de - a very important natural
vertical scale - cannot help to reduce the vertical size of our system:
one of the most important processes of upwelling is the supply of nu-
trients from below the Ekman layer into it, i.e., an upwelling system
has to reach the surface, as well as to extend well below the Ekman
layer.
As far as the ocean bottom is concerned, there are two possibilities:

in shallow regions the "bottom Ekman layer" supplies an important per-
centage, if not the main bulk, of the onshore transport and thus has
to be included. In deep water i t is dynamically unimportant and very
faint. In homogeneous water the onshore transport is distributed over
all depths below the surface Ekman layer (Garvine, 1974) - again the
system should extend to the bottom. In stratified waters the onshore
flow below the surface Ekman layer is not constant, but decreasing
with depth, and vanishes at a vertical scale large compared to the
Ekman layer depth (Tomczak jr., 1970). This scale depends on the rate
of vertical diffusion (or, more precisely, on the vertical Prandtl
number) and the stratification, and may thus vary in time. Since under
natural conditions i t may be of the order of the ocean's depth d there
is no reason to prefer it to the topographic depth scale.
For practical reasons it might be a reasonable procedure to introduce

an artificial bottom at a depth d a < d as long as the condition d a »
de is not violated. Since from the physics of the ocean well-defined
conditions for inflows and outflows cannot be given in that case, it
should be avoided whenever possible. An important consequence is that
those parts of the upwelling models which describe the physics usually
extend much deeper than those which treat the biological and chemical
aspects (and usually can be restricted to a scale of several Ekman
layer depths.) An example for this situation is the model described by
Walsh (1975). In that model the physical quantities are described be-
tween the ocean surface and the ocean bottom, but all calculations of
chemical and biological quantities are performed for the upper 50 m
only. The influence of the velocity field below 50 m depth on the
distribution of physical quantities above 50 m is reflected in the
density distribution, the vertical mass transport at 50 m, and the rate
of diffusion at 50m. Since the model of Walsh does not consider a con-
tinuous stratification, the influence of the velocity field below 50
m can be parameterized in terms of vertical mass transport and diffusion
at 50 m, and the model is restricted to the layer 0-50 m in all quanti-
ties. In more sophisticated models which treat stratification as another
variable, the physical quantities below 50 m will have to be included
explicitly.
The situation with the aross-stream scales is similar but much more com-
plex. On one hand, all upwelling is an integral part of the oceanic
circulation, since i t is simply the result of horizontal divergence
of the surface current field (Tomczak jr., 1970). As far as the dynamics
of the deep interior oceanic regions is concerned, i t can even be said
that it depends to a large extent on the structure of the upwelling
regions, i.e., the interior geostrophic flow is driven by the upwelling
layers (Pedlosky, 1968). Any scale smaller than the width of the ocean
disregards this natural interdependence. On the other hand, for an up-
welling model we are not interested in the velocity field of the oceanic
6
interior, and provided the influence of the oceanic interior on the

upwelling dynamics is negligible or can be parameterized, we might be
able to define a smaller scale, which is restricted to the upwelling
region.
In classical analysis based on the assumption of constant mixing co-

efficients, the typical cross-stream scale of upwelling regions is
set by lateral friction. According to these studies, all changes of
physical quantities coupled with upwelling are confined to a strip of
the width of the frictional layer. If we can manage to formulate bound-
ary conditions at the oceanic edge of the frictional layer, we may
thus restrict our model to.a considerably smaller scale than the oceanic
one. Since the interdependence between upwelling regions and the ocean's
interior is important, these boundary conditions have to be the ex-
pression of the interdependence and cannot be formulated merely on the
basis of our general knowledge of upwelling. In coastal upwelling, for
example, the boundary conditions for meridional and zonal mass trans-
port and pressure gradients have to be consistent with the distribution
of mass transport and pressure known as the Sverdrup circulation (Gar-
vine, 1 974) •
Unfortunately, the simple idea of the classical studies does not fit
observational knowledge accumulated during recent years. From expedi-
tion work in several important coastal upwelling regions, i t is well
established today that a frontal zone exists at the off-shore edge of
the upwelling region which separates the upwelled water from water of
oceanic surface characteristics (Hagen, 1974). Both in-shore and off-
shore of the frontal zone all processes can be considered geostrophic
and highly diffusion-free, while the frontal zone itself is an area
which is highly governed by diffusion, and probably by nonlinear ad-
vection where generation of water masses takes place (Bang, 1973).
This leads us to a model where the cross-stream scale is set by the

distance of the frontal zone from the coastline, and the off-shore
boundary is given by the frontal zone itself. From the classical studies
we retain the condition that the boundary conditions have to be con-
sistent with the neighboring Sverdrup circulation. In addition, in
order to fix the inflow and outflow through the frontal zone, we thus
have to quantify the rate of diffusion through it. It can be hoped that
precise parameterization of this process will be possible in future
studies.
While these considerations guide us in our proper formulation of the

boundary conditions in most situations of coastal upwelling, there are
cases where a definite frontal zone does not exist. Moreover, in equa-
torial upwelling it is not yet clear whether a frontal zone forms part
of that upwelling system, or whether the classical idea of large-scale
diffusion with constant coefficients might be appropriate to equatorial
upwelling. In these situations, only oceanographic observations can
help us to fix the cross-stream width of the upwelling region. Looking
again at coastal upwelling as an example, we note that although the
direction of the surface flow - and qUite often its speed - does not
change very much over distances off-shore which well exceed the classi-
cal frictional scale, a change in hydrographic parameters such as tem-
perature and salinity is usually found. In fact, Gunther (1936) already
deduced from observations that adjacent to the Peru-Chile upwelling
surface current there exists a current of equal direction outside the
coastal upwe·lling region. By carefully evaluating the hydrographic
characteristics of this current, and comparing them with the charac-
teristics of the coastal upwelling region, we should be able to define
the correct cross-stream scale of the upwelling region.
7
The situation with the down-sti'eam scale is different for coastal and
equatorial upwelling regions. In equatorial upwelling, the scale set
by the dimensions of the ocean is smaller than the scale set by the
driving force (the wind), a situation similar to that found with the
cross-stream scale. In coastal upwelling, on the other hand, the width
of the Trades is definitely smaller than the width of the ocean, and
thus defines the largest down-stream scale. In practice these differ-
ences between equatorial and coastal upwelling are of no importance,
since we usually do not intend to include the areas far up-stream and
far down-stream into an upwelling model. In these areas the down-stream
flow is small - either developing or disappearing; vertical motion is
generated by larger-scale horizontal divergence which is part of the
large-scale circulation rather than the proper upwelling mechanism,
and the distinction between down-stream flow and cross-stream flow is
doubtful because of a diffuse current pattern in general.
Thus, from physical arguments a natural down-stream scale for applying

ecosystem modeling techniques cannot be defined.
3.2 Biological Aspects
The main biological characteristic of an upwelling area is the increase

in productivity caused by the fertilization of surface water through
the process of upwelling of deeper waters rich in nutrients. As in
other productive regions of the ocean, the high productivity seems to
be accompanied by phenomena like high dominance of species (= low di-
versity), and shortness of food chains.
An increase in productivity will of course not always be observed in

those upwelling areas for which hydrographical boundaries have been
defined. For example, the development of high phytoplankton concentra-
tions may often depend on an increase in stability of the water column.
In the following we will look at the boundaries of upwelling areas

using the criterion of increased primary and secondary production, be-
cause this consequence of upwelling biologically separates upwelling
regions from the open ocean region.
Considering the three axes suggested earlier we first have to deal with
the vertical extension of the upwelling system. This means deciding
whether our model should reach from the surface of the sea to its bot-
tom and include it, or not.
No problems of definition arise concerning shallow waters, because

the sea bottom and the overlying water are definitely part of the up-
welling system. Nutrients are set free from the sediment, larvae of
bottom animals are produced and feed in the water, increased sedimenta-
tion favors animal life at the sea bed, etc. Shelf areas and the shelf
break should therefore be included in a model which covers the complete
system.
It is not well known whether the higher productivity in the surface layers
causes an increase in the standing stock of the benthos in deeper waters
beyond the shelf break of upwelling areas. The deep sea bottom will
certainly have no influence on the water layers in the upwelling re-
gime. Therefore a model will be adequate which shows the dependence
of the deep sea bottom on the productivity in the surface layers of
upwelling regions. This model does not form a part of the upwelling
model.
8
To fix the aross-stream scale of the upwelling region and its seasonal
variation would require an intensive sampling program in space and time.
This has been accomplished so far only by the Calcofi program off Cali-
fornia and Baja California. Using these data, Cushing (1971) compares
surface temperature distribution with the amount of zooplankton biomass
off California and Baja California and estimates that the zone of higher
zooplankton production is 2.5 times wider than the zone of physical
upwelling as derived from the temperature distribution. This ratio is
employed by him to calculate fish production in upwelling areas from
estimates of primary and secondary production.
From a theoretical point of view, the zone of higher primary production

should end where either the nutrients are depleted or a region of hori-
zontal surface convergence is reached causing a sinking of phytoplank-
ton below the euphotic layer. Such an area of horizontal surface con-
vergence can be assumed to exist at the off-shore boundary for all wind
fields which occur in upwelling regions. The surface convergence will
be strong and concentrated in a narrow band if a front is present. In
case no frontal zone exists, a broad area of weak convergence can be
expected outside the frictional boundary layer.
This means that because nutrient limitation of primary productivity

has not been'observed in upwelling areas, we can assume that the bio-
logical cross-stream width of the upwelling area corresponds with that
derived from hydrographical considerations.
We have to add that in the case where a frontal zone can be found, the
level of production beyond the front might still be slightly higher
than in the open ocean. Apart from the exchange through the front,
upwelling which takes place beyond the front may be responsible for
it. With more data at hand i t may be necessary in future to extend a
model of the upwelling region beyond the front.
Concerning the doum-stream scale of the upwelling region, not much bio-
logical information is available. Cushing (1971) used the distribution
of the surface temperature for the determination of the down-stream
scale of the upwelling zone.
In general it seems that in the areas far up-stream and far down-stream,
no sharp gradients in productivity can be found. This is in accordance
with the hydrographical situation.
4. The Size of the Investigated Area
For the application of ecosystem analysis i t is necessary to define

the size of the investigated area also in the down-stream direction, in
order to be able to model the system and to study the boundary condi-
tions and the transfer rates between each of the variables.
According to Smith (1970), boundaries in ecosystem analysis should be

drawn where the discontinuities are most clear. If there are no dis-
continuities the author recommends "to locate the boundary where in-
flows and outflows are most similar".
The biological consequences of upwelling start with primary production

in the region of maximum upward velocity, when the conditions are favor-
able for phytoplankton development. Higher productivity begins inshore
in coastal upwelling and at, the equator in equatorial upwelling, and
ends when the region of convergence is reached, where sinking causes a
removal of phytoplankton from the euphotic layer.
9
From these considerations we deduce a suitable size of the area which

should be studied for the investigation of an upwelling system. While
the vertical and cross-stream scale of the suggested area corresponds
to the boundaries of the upwelling regions as defined before, the down-
stream scale is equal to the distance traveled by newly upwelled water
in down-stream direction while i t moves over one scale length in cross-
stream direction. The time equivalent to the distance traveled will be
of the order of one to several weeks which means that enough time is
available for the phytoplankton to develop dense populations.
The ratio of the cross-stream and down-stream scales corresponds to

the ratio of the cross-stream and down-stream current components. The
values of this ratio probably range from 0.1 to 1, and can be used to
characterize different upwelling regions.
5. Physical Measurements
If we accept the above definition of the size of an investigation area,

and if we recall our research goal as formulated earlier, we are able
to ask some questions with respect to future expedition work and theo-
retical analysis.
There is first the problem to what extent the investigated area is in-
fluenced by down-stream gradients. In the simplest case, which reduces
to a quasi-two-dimensional model, inflow and outflow balance each other
at both ends. This can occur when there is no down-stream variation in
bottom topography. Topographic variations seem to control the intensity
of the undercurrent to a large extent (Hurlburt, 1974), producing non-
zero integrated cross-stream transport locally. Of course this topogra-
phic influence is not restricted to the area investigated. In order to
be able to insert the correct numbers into the boundary conditions, i t
is necessary to know the topography of a considerably larger region.
Several oceanographic cruises have demonstrated (e.g., Shaffer, 1976)
that with a better knowledge of topography the cruise program could
have been much improved. Since topographic mapping is no difficult task
with present-day echo-sounding and navigation equipment, it should be
easy to comply with this result of sensitivity analysis.
Vertical velocity is often assumed to be an important variable for the

rate of primary production. Hence, we should determine vertical veloc-
ity, but to what decimal place? It has been the opinion for decades
that the rate of vertical transport (10- 4 , 10- 3 , 10- 2 or 10- 1 cm s-l?)
plays a key role in the determination of the productivity of the region.
However, it becomes evident today that upward transport of nutrients
is one prerequisite among others for the primary productivity of an up-
welling area; the development of phytoplankton blooms in time, however,
is determined nearly eXClusively by lateral advection processes and
frontal dynamics. It turns out that maybe all we have to know of the
vertical velocity is its sign and the times at which strong upward mo-
tion occurs. This is sufficient for the determination of the periods
when the surface water is nutrient-enriched. For the rest of the time,
knowledge of horizontal velocity and diffusion is much more important.
If this is true, extensive attempts to come to a determination of ver-

tical velocity (Tomczak jr., 1972) have been to a large extent super-
fluous for our resource-orientated goal.
As far as the present methods of measuring horizontal velocity are

concerned, we find that they are adequate for the purpose of an eco-
10
system model, but can be improved if the investigators are guided by

sensitivity analysis. In general we may say that knowledge of the cross-
stream component should be better than knowledge of the down-stream
component, since it is the former that defines the width of the eco-
system, and the typical life time for the phytoplankton populations.
On the other hand, determination bf the down-stream gradient of the
down-stream velocity component is quite important for some boundary
conditions. This leads us to the conclusion that, instrumental precision
being high enough for our purposes, the arrangement of current meter
arrays has to be improved in order to meet the needs of an upwelling
ecosystem model: the numerous investigations where several current
meter moorings are deployed in a small area quite close to each other
have to be followed now by oceanographic programs where current meter
moorings are set at distances in down-stream direction large enough
to enable us to evaluate down-stream gradients. Since small variations
in the gradient of the flow yield appreciable variations in the diver-
gence of the mass transport it ~an be expected that the determination
of the down-stream gradients will be one of the major problems in future.
6. Biological Measurements
The size of the investigated area as defined will make it possible to

study the primary production and standing stock of phytoplankton in up-
welling systems and the influence of the hydrographical conditions on
it. An important factor which one has to know is the grazing on the
phytoplankton by herbivores in the investigated area. Methods are ap-
parently becoming available now for these measurements (Boucher and
S amain, 1 9 74) •
While these investigations will be fundamental for an understanding of

the productivity of upwelling regions, there is no doubt that they
have to be supplemented by other studies. The size of the investigated
area and the corresponding time period do not allow the study of the
higher trophic levels because of the long generation times involved.
From this it results that two kinds of investigation will be needed in

future: small-scale studies orientated toward an understanding of the
primary productivity of upwelling areas and large-scale studies for
the analysis of the productivity of the higher trophic levels. It has
to be stressed that the large-scale studies should cover phytoplankton
surveys and hydrography as well, For example, i t was shown already that
only large-scale studies will allow the evaluation of down-stream gra-
dients in upwelling areas.
An assumption of all these considerations has been a quasi-steady state

of the upwelling. However, one of the major goals of ecosystem analysis
is to predict time changes. This contradiction is solved by noting that
time changes have different effects on the system, depending on which
parameters are changing in time. If a time change in an input parameter
causes some quantitative fluctuations of the variables, the ecosystem
model is formulated correctly. As an example, variations of the wind
will change the location of the frontal zone of the upwelling region -
which results in a translation or dilatation of the system but does
not effect its characteristics - and change the frontal gradient, i.e.,
alter one of the input parameters (diffusion), and cause quantitative
changes within the system, which may have biological consequences.
11
If, however, a time change in an input parameter causes a qualitative

change of the system, model breakdown occurs. As an example, consider
the situation of "El Nino". As long as the winds blow steadily, up-
welling is well described by our model. When the winds cease, the
eastern branch of the oceanic circulation shifts westward, and the
region covered by our model becomes stagnant (White and McCreary, 1974)
It is successively filled by warm water advected laterally. With our
ecosystem model we are unable to describe this process because we have
to include Rossby waves in our dynamics which results in a much larger
scale than our cross-stream scale. Small variations in the wind field,
on the other hand, will not alter the upwelling so drastically, and
can be described by our model within certain limits.
So far ecosystem models in upwelling research have concentrated on

small-scale studies and phytoplankton dynamics (Walsh, 1975). The next
step is the construction of an ecosystem model which deals with the
complete upwelling region and has a corresponding time scale of up to
several years. It is only this model that would fulfill the needs of
ecosystem analysis orientated towards the optimum use of the living
resources in upwelling areas.
References
Bang, N.D.: Characteristics of an intense ocean frontal system in the upwell regime
west of Cape Town. Tellus 25, 3, 256-265 (1973)
Boucher, J., Samain, J.-F.: L'activite amylasique indice de la nutrition du zoo-
plancton ; mise en evidance d'un rythme quotidien en zone d'upwelling. Tethys ~,
1-2, 179-187 (1974)
Cushing, D.H.: Upwelling and the production of fish. Adv. Mar. Bioi. ~, 255-334
(1971 )
Ellenberg, H.: Ziele und Stand der Okosystemforschung. In: Okosystemforschung. Ellen-
berg, H. (ed.). Berlin-Heidelberg-New York: Springer, 1973a, pp. 1-31
Ellenberg, H.: Versuch einer Klassifikation der Okosysteme nach funktionalen Gesichts-
punkten. In: Okosystemforschung. Ellenberg, H. (ed.). Berlin-Heidelberg-New York:
Springer, 1973b, pp. 235-265
Garvine, R.W.: Ocean interiors and coastal upwelling models. J. Phys. Oceanogr. i,
121-125 (1974)
Gunther, E.R.: A report on oceanographic investigations in the Peru coastal current.
Discovery Rep. 11, 107-276 (1936)
Hagen, E.: Ein einfaches Schema der Entwicklung von Kaltwasserauftriebszellen vor
der nordwestafrikanischen Kuste. Beitr. Meeresk. }1, 115-125 (1974)
Hurlburt, H.E.: The influence of coastline geometry and bottom topography on the
eastern ocean circulation. Florida State Univ. Techn. Rep., 103 pp (1974)
Pedlosky, J.: An overlooked aspect of the wind-driven oceanic circulation. J. Fluid
Mech. 32, 809-821 (1968)
Shaffer, G.: A mesoscale study of co as tal upwelling variabili ty off NW-Africa.
"Meteor" Forsch.-Ergebn. A 17, 21-72 (1976)
Smith, F.E.: Analysis of ecosystems. In: Analysis of Temperate Forest Ecosystems.
Ecological Studies 1. Reichle, D.E. (ed.). Berlin-Heidelberg-New York: Springer,
1970, pp. 7-18
Tomczak, M., jr.: Eine lineare Theorie des stationaren Auftriebs im stetig geschich-
teten Meer. Dt. Hydr. Z. 23, 214-234 (1970)
Tomczak, M., jr.: Problems of physical oceanography in coastal upwelling investiga-
tions. Geoforum ll, 23-34 (1972)
Walsh, J.J.: A spatial simulation model of the Peru upwelling ecosystem. Deep-Sea
Res. 22, 4, 201-236 (1975)
White, W.B., McCreary, J.P.: Eastern intensification of ocean spin-down: application
to El Nino, J. Phys. Oceanogr. i, 295-303 (1974)
What is an Upwelling Ecosystem?
RMARGALEF
As a tentative answer to the question posed by Boje and Tomczak, and

perhaps as a rejoinder to the views expressed by them, I would like
to present my opinions on the subject. Upwelling areas are the site of
processes of fertilization. They represent a local deformation of the
ecological fields. The deformation is the result of a stress applied
on a limited space. In the case of upwelling, the stress consists of
energy made available in such area by some mechanism. The energy is
degraded in relation with its transfer from the sea to the atmosphere
or vice versa. The exchange means a locally enhanced increase in en-
tropy. Careful models of the dependence of primary production on light,
nutrients, temperature, and so on, may be useful in many situations,
but in upwelling areas they may be replaced, probably with advantage,
by the simple dependence of primary production on the auxiliary energy
made available (Fig. 1). It is like in agriculture, where yield can be
simply related to the input of subsidiary energy (in machines, oil
power, fertilizers, irregation).
What is important is: (1) the amount of energy involved, and (2) the
extent of the area in which the exchange is concentrated, and how it
is distributed. The amount of energy is related to the size of struc-
tures that act as collectors (oceanic Circulation, tides, waves). The
localization and distribution in space of the availability of energy
depends on mechanisms of funneling. Energy is used in moving water
one way (advection) or both ways (turbulence) over a large spectrum
of dimensions. The partition between advection and turbulence is im-
portant. Also important is the creation of discontinuous structures
(cells, eddies) by amplification over shore and bottom topographies
of irregularities in the movement, as well as by local accelerations
and decelerations of the component of the movement of water directed
upwards.
At the level of the chemistry of water and nutrients, upwelling does

not consist simply of an inflow coming from depth. More realistically,
an upwelling area is characterized by a number of circulation cells,
in which a strong recycling takes place, kept active by a moderate
advection. In the core or at some strata of the circulation cells,
depletion of oxygen and of inorganic nitrogen compounds and metals
can happen.
The biological cycle in offshore or oceanic waters can be regarded as

almost closed, with a trend to minimize turnover. This is because bio-
genic elements when in particulate form travel down with increased
probability than when they are in solution. Such an ecosystem can be
modeled as a vertical prism or column of moderate and arbitrary section,
since exchange all around is symmetrical.
In an upwelling area no such model is acceptable, if some degree of

closure in the cycle of matter is desirable, since there is much ex-
change with neighboring areas, and the exchange is asymmetrical, with
input of nutrients in depth and export of organisms in surface. More-
over, an upwelling system, like any system under stress, works with
13
SINK Fig. 1. A rough sketch of
atmosphere/ hydroSPhere_ , - - - - - - ' - - - - - - - - - - .

t an upwelling ecosystem,
on an x, z plane. The up-
welling, at right, is the
InteraClion ,--,oc;,.;.;;u.;;.ss;:,.l,;...n9
"-0'-"_--r---=_";"",,:,,,,,:;:.:....j
result of local availabil-
.
ity of auxiliary energy
t
" memo,, " that, among other things,
enhances production. Some
SINK ---f'.~-------"-~--~ physical distributions
Instabilities In (above) are reflected on
vertICal speed the biological structure
(below). The interfaces
ampllflcalion of effects resulting (air-water, sediment) act
from shore and bottom topography as a "memoir" of past
events
loop In atmosphere
symmetric
p.!;: : ~~"l
exchange
~) ~ I I '
oxygen
UI;lf'-L ~ ~_-'~
depletl()(1
t
I "' memoir "'
here apphes_..".._ _ _ _ _ _ _ _ _ _ _.-i~~~$~~
jF(Chl. N. L)dzfj Bdz-mlll
loop In sediment
leaks; oxygen and nitrogen are passed to the atmosphere, and carbon,
phosphorus, and metals to the sediment. Such pathways or external loops
act as brakes on the ecological cycles. The area that has to be studied
or modeled to understand and represent the workings of the ecosystem
must be necessarily larger in an upwelling system than further offshore.
If we choose to draw surfaces of equal value for some of the descriptive
parameters of the ecosystem (density of cells, concentration of nutri-
ents, chlorophyll, etc . ), such surfaces become increasingly closer to-
wards the centre of upwelling. This is one representation of the de-
formation of ecological fields. The length of the unit gradient (the
distance between two isosurfaces in the map) is inversely related to
the extension of the system that has to be studied to achieve a uniform
degree of closure in ecological cycles.
Upwelling can be understood only in the frame of models covering a large

space, perhaps all the Ocean, where the upwelling centers appear as
nodal points of stress. The definition of boundaries is an academic
question. In practice the boundaries can be defined by selecting some
value or some gradient in a chosen parameter, including the intensity
of horizontal exchange as a possible parameter. In a provisory and
pragmatic way I the boundarie's can be placed where the usual methods
14
are not sensitive enough to reveal directional horizontal heteroge-

neities.
Upwelling ecosystems are essentially anisotropic systems, in every

aspect. If not limited to small areas, spectral analysis is essentially
poor, since any phenomenon propagating as waves has to increase wave-
length and reduce wave amplitude as it travels centrifugally from the
spots of maximal energy degradation. This can be applied also to any
sort of pattern resulting from interaction between organisms. Some
sort of "relativistic" criteria must be introduced in the analysis,
and the comparison between the different spatial compartments of the
models should make use of appropriate transformations in the scales of
time and of space. The structure (pattern) of communities could appear
as the result of a nonhomogeneous propagation of waves describing super-
posed phenomena, as water movement and population dynamics.
In terms that I have used often, but to which many of those present
would object, the coupling between an upwelling centre and the peri-
pheric region is a typical instance of an extended interphase between
a less mature subsystem (upwelling center) and a more mature subsystem
(oceanic area). It can be said that the more mature feeds on the less
mature, and that the less mature is also less efficient. Organization
is prevented 'from increasing in the upwelling areas by vertical move-
ment and by its variability, and foodchains are kept shorter. It can
be added that the distribution of the relative frequencies of the r-
and K-strategies among the populations of the area fits the same pat-
tern: in the upwelling center, less mature, opportunistic species, or
r-strategists, may have an advantage.
Mesoscale Heterogeneities of the Phytoplankton Distribution in
the Upwelling Region ofNW Africa
M.ESTRADA
1. Introduction
This communication deals with some of the results of a taxonomic anal-

ysis of phytoplankton samples taken during the cruise Atlor II of the
R/V Cornide de Saavedra, in March 1973. These samples were collected
every 30 min along transects between stations, from 3 m depth water
which was continuously pumped into the laboratories. At an average
speed of 8 knots, this represents a distance of about 7.2 km between
sampling points. Simultaneously, samples for photosynthetic pigment
determination were taken and particle counting with a Coulter Counter
was also done; information concerning these parameters has been given
by Estrada (1974) and Margalef (1974). A partial report of the analysis
of phytoplankton data from the oceanographic stations has already been
published (Margalef, 1975). Along many of the transects, temperature,
fluorescence and concentration of nitrate and nitrite were continuously
monitored and recorded by the minute by means of a data logger. Figure
1 shows the position of the stations and the traject of the ship.
2. Material and Methods
For phytoplankton determination, 150 ml of water were stored in glass

bottles and fixed with lugol solution. Counting was carried out with
an inverted microscope, using 100-ml combined chambers; the whole
chamber bottom was scanned under low magnification (x 100) to count
large-sized individuals of infrequent species, and three transects
were examined under high (x 400) magnification to record the smaller
organisms. Many forms, especially of naked nannoplankton, could not be
identified, and were given a code number or assigned to general groups
such as "flagellates" or "small dinoflagellates". Several samples were
discarded because of the presence of silicate precipitates that made
a reliable estimation of algal densities impossible.
As has often been discussed before (Margalef, 1973, 1975), methods of

phytoplankton counting suffer many limitations; besides difficulties
and mistakes in the iaentification and censusing of species (or of
taxonomical entities used as such), the high variability due to the
heterogeneity of phytoplankton distributions, and the statistical errors
inherent to the process of sampling small volumes have to be taken into
account.
A total of 213 samples were considered in the analysis. From the 350
species and other taxonomical entities recorded, 20 were chosen to
carry out a principal component analysis of the correlation matrix be-
tween species. Selection of a particular species was based on its pre-
sence in a high number of samples and on the reliability of its iden-
tification; evidently, many subjective considerations affect this pro-
cedure; however, several analyses made after changing some of the spe-
16
cies gave very similar results. A logarithm transform, X -+ log (X+1),

of the data was used.
Three different measures of correlation, based on the abundance of all

the species recorded, were calculated for every pair of successive
samples: the coefficient of linear correlation after making a logarithm
transform of the data; the rank correlation coefficient of Spearman
and the percentage of cornmon species in the total number of different
species present in the two samples. The three coefficients showed very
similar trends of variation, although their absolute values were dif-
ferent; therefore, only one of them, the rank correlation coefficient
of Spearman, was retained and utilized in the study. It should be noted
that low plankton abundance in a body of water will tend to lower the
observed correlation among subsamples, simply because of statistical
sampling errors; the calculated coefficients have no more than a re-
lative value.
Methods for the continuous analysis were essentially those described

in Ballester et al. (1972). The values of temperature, fluorescence,
nitrate and nitrite corresponding to the time of extraction of the
phytoplankton samples, were used to calculate their correlation with
chlorophyll concentration and cell numbers. All calculations were done
wi th the IBM 1130 of the Ins ti tuto de Investigaciones Pesqueras of
Barcelona.
3. Results and Discussion
As has already been shown by the study of the station samples (Margalef,
1975) and by the pigment and productivity data (Estrada, 1974), higher
concentrations of phytoplankton were found close to the coast, where
upwelling is more intense; offshore, another rich zone was located be-
tween 20 0 and 21 0 Nand 21 0 and 22 0 W (Fig. 1).
Fig. 1. Distribution of the number of phyto-

plankton cells per ml. Dots indicate ap-
proximately the position of the sampling
points; striped areas represent concentra-
tions higher than 100 cells/ml. Underlined
numbers indicate the stations; arrows show
the direction of the trajects
17
Table 1. Loadings of the 20 species on the first three principal components
Species Components
1 2 3
Brachydinium capitatum 0.845 -0.029 -0.280

Ceratium furca 0.185 0.370 -0.092
C. fusus 0.597 0.195 0.046
C. kOfoidi 0.651 -0.026 -0.114
C. tripos -0.163 0.317 -0.345
Exuviae Zla sp. -0.360 0.530 -0.142
Prorocentrum rostratum 0.815 0.054 -0.248
CoccoZithus huxZeyi 0.262 0.385 -0.303
HeZicosphaera sp. -0.416 0.490 -0.319
Syracosphaera sp. -0.244 0.447 -0.437
Arrrphora hyaZina -0.484 0.550 -0.210
Chaetoceros affinis 0.007 0.514 0.609
Chaetoceros didymus 0.041 0.363 0.724
PZanktonieZZa soZ 0.7'70 -0.064 -0.093
RhizosoZenia aZata 0.626 0.203 -0.030
Rh. fragiZissima -0.081 0.767 0.028
Rh. irrUJricata 0.497 0.414 0.061
Rh. stoZterfothi£ 0.419 0.567 0.078
ThaZassiosira partheneia 0.143 0.082 0.747
Coscinodiscus aZborani 0.123 0.362 0.187
The variance accounted for by the first three principal components was
21.7%, 15.5%, and 11.2%, respectively; these low values are typical
of the reticulate interrelations among densities of phytoplankton spe-
cies. Table 1 gives the correlation coefficients of the components with
the 20 species consi'dered.
As can be seen in Table 1, Braahydinium capitatum, Prorocentrum rostratum,
Ceratium kofoidi and PZantonieUa soZ show the highest loadings on the first
component; the more negative values correspond to Arrrphora hyaZina. The
second component is basically associated with several species of diatoms
and coccoli tophorids (RhizosoZenia fragiZissima, Arrrphora hyaZina, Rh. stoZter-
fothii, He Zycosphaera sp., Syracosphaera sp.) and with Exuviae Zla sp. ThaZas-
siosira partheneia, Chaetoceros affinis and Chaetoceros didymus show the highest
loadings on the third component. Figure 2 shows the geographical dis-
tribution of the values of the first and second components at the sampl-
ing points. Several general trends can be distinguished. The first com-
ponent shows a marked N-S gradient, corresponding to the main division
of the zone in northern and southern regions (Fraga, 1974; Fraga and
Manriquez, 1975; Margalef, 1975). The boundary between these zones is
given as the meandering front between North Atlantic Central Water and
South Atlantic Central Water; it must be pOinted out that these water
masses are subsuperficial, whereas the samples carne from superficial
water. The waters form the Canarias current in the North, and the tro-
pical water in the South from a thin layer, covering an extension of
the central waters which varies with time of the year. The gradient
in the first component reflects this subdivision, although with the
data available it is not possible to establish the existence of definite
associations between species and water masses; factors like the past
evolution of the hydrographic structures, probably have a decisive im-
portance in the actual distributions found. The same considerations
apply to the second component; as can be seen in Figure 2, this com-
ponent shows an E-W variation and can be globally associated with the
higher intensity of fertilization in the proxirnities of the coast, al-
though more detailed relationships cannot be established. The third com-
ponent reflects the importance of ThaZassiosira partheneia and other diatoms that
18
Fig. 2. Geographical distribution of values of the first (left) and second (right)
principal components
appear closely associated to it in the intense zone of upwelling located

off Cabo Blanco.
Superimposed on the general trends indicated by the components was a

great heterogeneity that resulted in highly jagged profiles of phyto-
plankton density along the transects. Figure 3 shows one example. Ob-
viously, these recordings cannot be interpreted as a static picture;
they reflect the combination of several important sources of variation
like the displacement of the vessel, the water movements, and the chan-
ges of the population with time. In general, peaks in phytoplankton
concentration presented a high proportion of diatoms in relation to
other groups; coccolithophorids were also very important in particular
regions. As can be expected from the exponential character of growth
processes, a pattern of patches in a background of lower abundances
was very apparent in the figures; Margalef (1976) has discussed this
problem in connection with other sets of data. This feature was re-
flected in the histogram of frequencies of the phytoplankton abundance
values (Fig. 4a). The distribution was very asymmetrical, with positive
skewness; a logarithm transform of the data made it closer· to normal
(Fig. 4b), although it was .still significantly different when it was
checked by a X2 test at the 95% confidence level.
When all the samples were pooled, phytoplankton density (measured by

the total number of cells per ml, chlorophyll concentration or fluo-
rescence) was positively correlated with nitrate plus nitrite concen-
tration and negatively with temperature (Table 2), as can be expected
from the general features of the region. However, at a local scale
(Figs. 5, 6, 7), different relationships can be found, depending on
the unknown past evolution of the water body involved. As can be seen
in these figures, variations in community composition were related to
changes in the relationships among the physicochemical variables re-
corded. A statistical analysis of these changes is difficult to make,
due to the need to average data over a determined space that must more-
over be changed according to each case. Of course, temperature and
19
'6r-----------------
chIOr.(1J9,/i)
'4
,2
'0
'6r 8
0.30/0665
'2 6
to
.
8 4
'1
6
4 2 l
2
..........
' 600 1
cells/ml
I:::::::::! D,atoms
'400
~ Dinoflagellates
CoccolithophOrids
,200 c:::J
Other groups
~ Detached coccolIths
1000
800
Fig. 3. Above, Chlorophyll concentration (Chlor.) and the relation between optical
densitie~pigment extracts at 430 and 665 nm (D430/D665)' Below, Population den-
si ties of several phytoplankton groups (cells/ml); "detached coccoliths" indicate
the value found by dividing the number of detached coccoliths counted by an accepted
average number of coccoliths per c e ll for each species; they were not further con-
sidered in the calculations. The segme nts between sampling points are proportional
to the actual distances traveled
nitrate plus nitrite were considered, only, as the available indicators

of the processes which had occurred, without direct causal impiications.
Figures 8 and 9 show another way of looking at the connection between
taxonomic composition and hydrographic features. Cruzado and Kelley
(1973) had published similar phase diagrams for fluorescence and nu-
20
30%
r--r--
20%
r--
0;
S-
.z-I'
+5.4s +4.8s +4.2s +3.6s +3.0s +2.4s +1.8s +1.2s +0.6s m -0.6s
1200 cellsjml 1000 800 600 400 200 o
30%
20%
+2.4s +1.8s +1.2s +0.6s m -2.4s -3.0s

12 In(cells/mll 10 9 8 7 6 5 4
Fig. 4. (a) Histogram of frequencies of the total number of cells per ml. (b) Histo-
gram of frequencies of the natural logarithm of the total number of cells per ml.
~, mean; ~, standard deviation
Fir Toe N03

40
.21 .. ~g-at/I
o
30
2
20
4
Figs. 5, 6, and 7. Above, Recordings
10 of temperature (tOe), fluorescence
6
(Flr., relative units) and nitrate'+
0
nitrite (N03, llg-at/l; the scale is
reversed) • Below, Spearman's rank
correlation coefficient (Rs) between
every pair of successive phytoplankton
samples. The scale of the abscissa
refers only to time of sampling.
Dotted areas indicate zones of changes
22 24 2 4 in the relationships between the
10. Mar. 1973 variables
21
Fig. 6. Legend see Fig. 5

Fir T' e N03
40 119- a l/l
.21
o
30
.20 2
20
4
.19
10
6
o As
6
4
2
0
-2
22 24 2 4 6 8 hours
11. Mar. 1973
Fir T'e N03

~g-al /I
50 .20 0
N03
40 2
~~
.19
30 4
20 Fir
6
Fir Fir
10 8
0 I ;:: I
11
10
4
2
~ .,
0
-2
I I
12 14 16 18 4 6 8 14 16 18 hours
18. Ma r. 1973 19. Mar. 1973
Fig. 7. Lege.nd see Fig. 5
Table 2. Correlation coefficients among temperature (tOe), nitrate + nitrite concen-

tration (N03) , logarithm of fluorescence [log (frl.)], logarithm of chlorophyll con-
centration [log (chlor.)] and logarithm of the total number of cells per ml [log
(cell/ml)]. The calculations were based on the 114 samples for which data on all the
variables considered in the table were available
N03 log (frl. ) log (chlor. ) log (cell/ml)
tOe -0.84 -0.52 -0.54 -0.47
N03 0.23 0.35 0.35
log (fIr.) 0.80 0.39
log (chlor.) 0.52
Fig. 8. Above, Number of cells
per ml (cell/ml) and Spearman's
rank correlation coefficient
(Rs) for the transect from
2300 h March 11th to 2200 h
March 12th; the segments between
sampling points are proportional
to the actual distances travel-
led. E31 indicates the position
f-"""l'E,..-..,1 1-1--=F--I of station 31. Below, Position
of the points belonging to the
~ ______L-____ ~I
100 km different patches in the space
defined by temperature and
ni trate + nitrite. The points
considered as belonging to each
unit patch are bracketed (up)
or enclosed (below) by a c~
tinuous line ~designat~
wi th a capital letter
O~~~~~~~=Z~~ __L-________~
18.5 19.5 20.5
Temperature 'c
Rs
cells/ml .4
:~~l·~
100 -.2
o -.4L....l'-l-l-,-+-I-I--I-~~-I--l--,r-i+--I-~I!!!!!O..-1j.J Fig. 9. Above, Number of cells
per ml (cell/ml) and Spearman's
rank correlation coefficient
B o IT (Rs) for the transect from
o 100 km
..J....____--',
L . . '_ _ _ _ _ _
1100 h, March 9th to 1600 h,
3.2..-------------------, March 11 th; the segments be-
Nitrate tween sampling points are pro-
1l9-at/1 portional to the actual distances
2.4 traveled. E45 and E19 indicate
the position of stations 45
and 19. Below, Position of the
1.6 points belonging to the dif-
ferent patches in the space
defined by temperature and
0.8 nitrate + ni tri te. The points
considered as belonging to each
unit patch are bracketed (up)
O~~=c~~~_L~L_____~ or enclosed (below) by a con-
18.5 19.5 20.5 tinuous line and designated
Temperature 'c with a capital letter
trient data. Although the variable space considered (temperature,

nitrate + nitrite) was very limited, it appeared that points that could
23
be considered as belonging to the same phytoplankton patch (designated

by a capital letter in the fi.gures) could be clustered together. Ob-
viously, a phytoplankton "patch" cannot be defined in a clear-cut way,
although it is possible to fragment a sequence of samples establishing
operational limits, based, as was done in this case, on the occurrence
of a given drop in the similarity coefficients.
A major problem in the interpretation of this type of data is the ab-

sence of information on evolution in time and the asynchrony of observa-
tions at different points in space. Although care has to be exercised
to avoid excessive generalizations, the association of qualitative and
quantitative changes in phytoplankton populations with changes in tem-
perature and temperature-nutrient interrelations, indicate that, at
the scale considered, biological patchiness can be related to hydro-
graphic phenomena.
AcknowZedgements. I thank Dr. R. Margalef for his suggestions and comments. M. Alca-
raz, F. Fernandez, Dr. G. Mateu, P. Rubies and A. Sanz helped in obtaining the phyto-
plankton samples. The chemistry group, led by Dr. F. Fraga, took care of nitrate
and nitrite analysis. A. Julia cooperated in the obtaining and handling of the data
logger recordings.
References
Ballester, A., Cruzado, A., Julia, A., Manriquez, M., Salat, J.: Analisis automatico
y continuo de lascaracteristicas fsicas, quimicas y biologicas del mar. Publs.
Tecnicas Patr. "J. Cierva" 1, 1-72 (1972)
Cruzado, A., Kelley, J.C.: CZntinuous measurements of nutrient concentrations and
phytoplankton density in the surface water of the western Mediterranean, winter
1970. Thalassia Jugoslavica 2., (1/2), 19-24 (1973)
Estrada, M.: Photosynthetic pigments and productivity in the upwelling region of NW
Africa. Tethys.§. (1-2), 247-260 (1974)
Fraga, F.: Distribution des masses d'eau dans l'upwelling de Mauritanie. Tethys 6
(1-2), 5-10 (1974)
Fraga, F., Manriquez, M.: Oceanografia quimica de la region de afloramiento del
noroeste de Africa. II. Campana "Atlor II", marzo 1973. Res. Exp. Cient. B/O Cor-
nide i, 185-218 (1975)
Margalef, R.: Fitoplancton marino de la region de afloramiento del NW de Africa. II.
Composicion y distribucion del fitoplancton (campana "Sahara II" del "Cornide de
Saavedra"). Res. Exp. Cient. B/O Cornide ~, 65, 94 (1973)
Margalef, R.: Distribution du seston dans la region d'affleurement du nord-ouest
de l'Afrique en mars 1973. Tethys.§. (1-2), 77-88 (1974)
Margalef, R.: Composic·ion y distribucion del fitoplancton marino en la region de
afloramiento del NW de Africa, en marzo de 1973. (Campana "Atlor II" del "Cornide
de Saavedra"). Res. Exp. Cient. B/O Cornide i, 145-170 (1975)
Margalef, R.: Distribucion horizontal del fitoplancton marino (1 a 10 km) ilustrada
con un ejemplo del area de afloramiento del NW de Africa. Mem. Real Acad. Sci.
Artes Barcelona 43 (5), 131-148 (1976)
On the Ecological Significance of Thalassiosira partheneia in the
Northwest African Upwelling Area
'"
M. ELBRACH1ERand R BOJE
"
1. Introduction
One of the most striking phytoplankton species in the Northwest African

upwelling area is the diatom ThaZassiosira partheneia Schrader. This diatom
forms colonies of up to 5 cm length, which are visible from the ship.
In the euphotic zone 10 to 50 or more colonies can be found in 1 I of
sea water. Each colony consists' of from several hundred to 25,000 cells,
each cell being between 7 and 15 ~ in diameter. The majority of the
cells have a diameter of 8 to 10 ~. Some 8 to 128 individual cells are
connected by a thicker central thread to chains similar to other ThaZas-
siosira species, but in addition, the many marginal threads cOming out
of the marginal strutted processes of each cell are twisted, and thus
connect one chain to the other, building a tube-like colony with a
diameter of up to 1 cm. Mucus, which was described for this species
in the original description (Schrader, 1972) was absent, as revealed
by light microscopy of living colonies on board the ship.
The contribution of this species to primary productivity in the North-

west African upwelling area is not known. During Mete0r cruise 36 (ex-
pedition "Upwelling 75") i t was possible to investigate this problem,
and to make some additional observations of ecological significance.
2. Material and Methods
The investigations took place during February, 1975 (Meteor cruise 36).
Samples were taken at the stations, which are shown in Figure 1. Water
samples were collected at depths which corresponded to 100%, 50%, 30%
and 10% of the incident light, and occasionally at greater depths. A
quantameter, constructed at the Institute of Physical Oceanography in
Copenhagen, Denmark, was used for the determination of the light levels
at local noon. The water bottles sampled 5 I and were made of dark
opaque plastic (Hydrobios, Kiel, FRG).
For productivity measurements four subsamples of 1 I each were filled

in glass cylinders. All colonies of Thalassiosira partheneia visible to
the naked eye were pipetted under dim light into a 120 ml glass bottle,
which was filled with 0.45 ~ filtered sea water from the same location.
As reference sample one bottle of each depth was filled with sea water
from which the colonies had been removed.
After inoculating the samples with 14C, the primary productivity was
measured from noon to sunset, using the simulated in situ technique.
Light levels were simulated by neutral density filters, and samples
were kept under in situ temperature. The incubator was covered by a
glass plate for protection against UV-light during the measurements
(Steemann Nielsen, 1974). Radioactive bicarbonate was purchased from
The International Agency for 14C-Determination in H¢rsholm, Denmark,
"'Member of the "Taxonomische Arbeitsgruppe"

25
Fig. 1. Location o f the stations

19' W
24'
UPWELLING '75
N N
Jan./Feb. 1975
23' :.:.: 23'

~6
::.:.::.:.:.:.:.:.:.:.:.:
22' .:::::::::::::::::::::::::::·:· 22'
~fj i;i';;
65 .:
i
. ..:::::::::::::::::::::::::.:.: 21 '
21' ':,'::::,':::.'::
.............
.:: ....:
where the Geiger Muller processing of the filters was done after the
cruise.
The filters used for the productivity measurements were mounted in

immersion oil on slides for microscopic observation . Cell numbers of
Tha"lassiosira part heneia - and if needed of Chae toceros radicans - were counted
with phase contrast at a magnification of 250 times. For counting,
every third strip of 200 ~ width was used. In samples with high numbers
of Chae toceros r adicans only two cross sections of 200 ~ width were eval-
uated. Living phytoplankton were studied at sea with a light microscope
(Wild M 20). Observation of living colonies of Th . partheneia was done
with bright field, dark field, or phase contrast, some samples were
stained with neutral red or erythrosine .
Isolated colonies of Th. partheneia were cultivated in 200 ml glass bot-

tles and in petri dishes. The glass bottles were exposed to daylight
in running sea water of surface temperature with a layer of at least
25 cm water and a glass plate above the samples. The petri dishes were
kept in an incubator (Memmert) at 16 0 to 1a o C and about 2000 lux light
intensity with a light period of 14 h and a dark period of 10 h. The
culture medium was that of Stosch and Drebes (1964), but the additives
were given in 1/10 of its original concentration.
For microbiological studies samples of Th . parth eneia were fixed with

freshly filtered 1% formaldehyde. The preparation for epifluorescent
microscopy followed a method described by Zimmermann and Meyer-Reil
(1974). For scanning electron microscopy (SEM) a suspension of colonies
was placed on Nuclepore membrane filters (Nuclepore, Pleasanton, CA,
USA) and treated with 10%, 20%, 30%... 96% ethanol followed by 25%,
50%, 75% and 100% frigen 113 for drying. The drying solution was re-
moved by suction with a small pressure differential . Dried samples
were coated with a thin gold film by sputtering (Hummer, Technics Inc.,
26
Alexandria, VA, USA) and examined with a Cambridge S 600 Stereoscan

(Cambridge Instr. Co., Cambridge, England).
3. Results
The values for the primary production are shown in Figures 2-7. The
results for the Thalassiosira fraction and for the reference sample are
7 6 12 2,7 3,8 2,4 12

mgC/m3 152 5 92 65 Il,85 78 1,1 mgC/m 2
2000
160 Stat.56 Oate:5.2. 1600
120 1200
80 800
40 400
0 0
30% 10%
light depth euph. zone
Fig. 2. Left, primary production in mg m- 3 per exposure time at different light
depths for the ThaZassiosira rartheneia fraction (black column) " the remainder
(hatched column) and the sum of both (white column), which is taken as 100%. Numbers
over the black columns, percent of ThaZassiosira production in relation to 100%.
Numbers over the hatched and the black columns: cell number 1- 1 x 104 ; upper line,
Thalassiosira rartheneia, lower line, Chaetoceros radicans. Right, primary production
in mg m- 2 per exposure time integrated over the euphotic zone, symbols as above
added and given as total production. The ThaZassiosira production as

percentage of total production is shown over the black columns. In ad-
dition, the cell number per liter of Th. rartheneia and, if present, of
Chaetoceros radicans are given as numbers over the reference sample as
well as over the ThaZassiosira sample. The percentage of Thalassiosira
production varies from 6% to 48% at different stations and light depths.
At station 56 (Feb. 5 and Feb. 6) and also at station 86, Feb. 14, the
diatom Chaetoceros radicans formed a dense bloom together with Th. rartheneia.
Therefore, the percentage of Thalassiosira production is relatively low.
In addition, microscopic observation revealed that the Thalassiosira
population was "aged": the colonies had a pale yellow-green color in-
stead of the normal green-brown. Furthermore, the colonies were not
regularly rounded, but disintegrated at the ends of the tubes. As a
consequence a high percentage of single cells or short chains occurred.
The photosynthetic activity of such single cells seems to be very low:

on station 65, at the 10% light depth 90,000 cells of ThaZassiosira per-
27
3 12,5 1 1,2 2J> 1,2 '1

140 100 3,8 150 1,5 90 1,2
mgC 1m 3 ",1 mg C/m2
100 1000
80 Stat.56 Date:6.2. 800
60 600
40 400
20 200
a
100% 30%
a
light depth euph.zone
9 0,8 5,6 11 19 3 10
0,1
mgC/m 2
500
St a 1.60 Date: 8.2. 400
20
10
formed a production of 1.36 mg C/m 3 • In contrast, in the reference

sample the production was only 2.98 mg C/m 3 although as many as 250,000
Thalassiosira cells, most of them being single cells or in short chains,
were present besides the other phytoplankton. If the single cells of
Thalassiosira showed the same production as cells in active colonies,
we would expect a value of 3.78 mg C/m 3, for Thalassiosira only, neglect-
ing the contribution to production by other phytoplankton species.
Thalassiosira partheneia is a diatom which is very sensitive to high light

intensities. This is demonstrated at stations 60 and 65, where primary
production of Thalassiosira was lower at the 100% light depth than at the
28
mgC/m 3 0.9 17 9,5 10 2,5 14 25 9 mgC/m 2

20 200
16 160
Sta t. 6 5 Da te:9. 2.
12 120
8 80
40
o
I ight depth euph.zone
mgC/m 3 6 30 17 16,5 50 10 16 57 mgC/m 2

500
400
Stat. 68 Date:l0.2.
300
200
100
50% light depth, although cell density dropped. In addition, culture

experiments with isolated colonies clearly showed light inhibition. In
cultures in 200 ml glass bottles covered by 25 cm sea water and a glass
plate, exposed to direct sunlight, the colonies changed their color
into yellow-green after half a day. After two days the colonies were
completely white and all cells were dead. In contrast, colonies culti-
vated with about 2000 lux in artificial light grew well; in addition,
their color remained green-brown and they formed new colonies.
A single colony of Th. paY'theneia represents a small biocoenosis. Light

microscopy at sea of living colonies of Th. paY'theneia revealed that in
29
2,7 13 10 13,5 9 9 7,2 10,5

mgC/m 3 200 1,9 180 3,3 180 4,2 190 4,5 mg C/m2
100 1000
Stat. 86 Oate:14.2.
80 800
60 600
40 400
20 200
48%
0 0
100"10
one single colony the flagellates Pronoctiluca phaeocysticoZa (Scherffel)

Pavillard (18 individuals), Amphidiniwn phaeocysticoZa Lebour (6 indi vid-
uals), Gyrrrnodiniwn heterostriatwn Kofoid et Swezy (9 individuals) and
GZenodiniwn sp. ( 1 2 individuals), up to 42 amoeba, the ci lia te EupZotes
sp. (5 individuals) and at least two other hypotrich ciliates each with
2 individuals could be seen to feed on Th. partheneia cells. In addition,
naviculoid, Nitzschia-like diatoms were commonly observed on the Th.
partheneia threads.
Epifluorescent and scanning microscopy showed that bacteria are present

on the living diatom cells as well as on the many threads coming out
of the strutted processes. On one colony of about 1000 Th. partheneia
cells about 20,000 bacteria were found. Whether bacteria are responsible
for the disintegration of the colonies to single chains or even to
single cells is unknown. Disintegration is a common phenomenon when
colonies age and when they are light-damaged. These single cells or
short chains are sometimes abundant at great water depths. We observed
up to 130,000 cells per I in 60 to 75 m water depth, most of them being
single cells.
4. Discussion
Normally there is no possibility in the measurement of primary pro-

ductivity to determine the contribution of one particular species of
the phytoplankton present. Selection of the large colonies of Th. par-
theneia, visible to the naked eye, gave the possibility of doing so.
Contamination of the ThaZassiosira samples with other algae was very low
at stations 60, 65 and 68 as revealed by direct microscopic cell counts.
At station 56 and at station 86 contamination with cells of Chaetoceros
radicans was considerable. As many as 50,000 single Chaetoceros cells were
present in the ThaZassiosira sample, which corresponds to about 3% of
the number of Chaetoceros cells present in the reference sample.
30
The number of short chains or single cells of ThaZassiosira, which was

overlooked during the selection procedure and, therefore, present in
the reference sample, was sometimes considerable although quite variable.
The calculated ThaZassiosira production is overestimated in the presence
of Chaetoaeros cells and underestimated if many single cells of ThaZas-
siosira are present.
If we assume that the overlooked ThaZassiosira fraction has the same
production capacity per cell as i t has in the selected samples we can
make the following calculations: for station 86, at the 30% light-level,
Chg.etoaeros cell contamination was about 2.3% when compared to the re-
ference sample, which represents a production of 1.48 mg C m- 3 • ThaZas-
siosira production after this correction would be 4.84 mg C m- 3 instead
of 6.32 mg C m- 3. In the reference sample the same cell number of Tha-
Zassiosira was present as in the ThaZassiosira sample. After this second
correction the ThaZassiosira production would be 9.68 mg C m- 3, which
is 14% of the total production at this light-depth, compared to the
value without correction of 9%. As mentioned earlier, there is strong
evidence that single cells do not have the same production capacity
as cells in active colonie~ (see results, station 65, 10% light level).
As a consequence of this and other calculations i t can be concluded
that the values given without correction are a good approximation to
production of Th. partheneia.
It is not known whether organisms other than colorless flagellates,

amoeba/and ciliates feed on ThaZassiosira. These organisms live wi thin
the healthy colonies and seem to feed on single cells. Some of these
organisms were first described as living in the colonies of the hapto-
phycean Phaeoaystis pouahetii (Hariot) Lagerheim, e. g., PronoatiZuaa phaeo-
aystiaoZa and Amphidiniwn phaeoaystiaoZa, which previously have been found
exclusively in this haptophycean. Gymnodiniwn heterostriatwn also feeds
effectively on Phaeoaystis pouahetii if available (Lebour, 1917; Elbrach-
ter, 1975) as do amoeba and ciliates.
Small naviculoids, in most cases Nitzsahia-like diatoms, which we found

in colonies of Th. partheneia have been observed to live regularly in
the colonies of Phaeoaystis pouahetii as well. Whether these diatoms be-
long to the same species must still be ascertained.
Bacteria are present on the threads of Th. partheneia, but may use these
threads only for attachment. It is know that bacteria attached to
particles use dissolved organic carbon (DOC) which is adsorbed to these
particles by non-biological processes. The close relation of the
threads to the living diatom cells which excret DOC, seems to be
favorable for bacterial nutrition. Another possibility is that the
treads of ThaZassiosira cells are used as substrate for growth. The
treads consist of chitan (McLachlan et al., 1965), which is similar
to chitin. Since some bacteria use chitin as substrate for growth,
they possibly might also be able to grow on chitan. B51ter and Meyer-
Reil (1974) demonstrated that in the Northwest African upwelling area
there are many bacteria present which use chitin for growth. Perhaps
bacterial activity is responsible for the disintegration of the colonies
into short chains or even single cells, especially during aging or light
damage. If that occurred then the bacteria would be very important
ecologically, making ThaZassiosira colonies available to the food web.
As far as we know, no organisms have been found to feed on the large
colonies of Th. partheneia with the exception of those living within
the healthy colonies.
After disintegration the short chains and single cells are a good diet
for filter feeders and cili,ates like tintinnids. It is assumed by many
31
authors that a considerable part of nanoplankton is transfered to large

herbivores or omnivores via tintinnids. Gold (1973, additional litera-
ture therein) demonstrated that tintinnids are very effective feeders.
Further investigations have to show whether there is a correlation be-
tween Thalassiosira colonies and the number of tintinnids.
Acknowledgments. The authors thank W. Weise for the pictures, obtained by epifluore-
scence staining and scanning electron microscopy. This research was supported by
the Deutsche Forschungsgemeinschaft.
References
Bolter, M., Meyer-Reil, L.-A.: Untersuchungen an Bakterienstammen aus dem Auftriebs-

gebiet vor der westafrikanischen Kuste: Taxonomie und Nahrstoffanspruche. Bot.
Mar. 12, 227-248 (1974)
Elbrachter, M.: Taxonomical notes on North Sea dinoflagellates I. Kieler Meeres-
forsch. 11, 1, 58-64 (1975)
Gold, K.: Methods for growing tintinnida in continuous culture. Am. Zool. ll, 203-208
(1973)
Lebour, M.V.: The peridiniales of Plymouth Sound from the region beyond the Break-
water. J. Mar. Biol. Assoc. Plymouth 11, 183-200 (1917)
McLachlan, J., McInnes, A.G., Falk, M.: Studies on the chitan (chitin: poly-N-acetyl-
glucosamine) fibres of the diatom Thalassiosira fluviatilis Hustedt. Can. J. Bot.
43, 707-713 (1965)
Schrader, H.-J.: Thalassiosira partheneia, eine neue Gallertlager bildende zentrale
Diatomee. 'Meteor'-Forsch.-Ergeb. D 10, 58-64 (1972)
Steemann Nielsen, E.: Light and primary production. In: Optical Aspects of Oceano-
graphy. Jerlov, N.G., Steemann Nielsen, E. (ed.). London-New York: Academic Press,
1974, pp. 361-388
Stosch, H.A. v., Drebes, G.: Entwicklungsgeschichtliche untersuchungen an zentrischen
Diatomeen IV. Helgolander Wiss. Meeresunters. l!, 209-257 (1964)
Zimmermann, R., Meyer-Reil, L.-A.: A new method for fluorescence staining of bacterial
populations on membrane filters. Kieler Meeresforsch. 30, 1, 24-27 (1974)
Zooplankton Communities in the West African Upwelling Area
A TIIIRIOT
The interest in zooplankton populations in upwelling areas is not very

new, but it has been increasing during the last ten years. The develop-
ment of pelagic and demersal fisheries in these areas is one of the
prime reasons for this interest, because it is essential to know the
possibilities and limits for exploitation of different populations of
commercial interest. Another reason which has motivated the scientific
research is a need to understand what mechanisms are involved in the
planktonic production.
Originally it was thought that the acceleration of phenomena by a high

and more or less continuous input of nutrients at the start of the
food webs would facilitate understanding them. If the simplification
of the trophic exchanges is not as obvious as supposed, these upwelling
areas are nevertheless extremely interesting fields of observation and
experimentation. The fact that we are facing reduced scales in both
time and space of trophic relationships is enough to justify such an
effort.
It is at present out of the question that we might realize a true syn-

thesis of all results, and that we might define secondary production
as well as the energetic budget of these systems. We are still in the
analytic phase. It seems more interesting to limit this review to the
geographical point of view, and to try to point out the state of our
research.
I have therefore chosen the African Atlantic which has been the object
of numerous oceanographic expeditions, and which is now the object of
a large number of national and international research programs all
concerned with the biological consequences of the upwelling phenomena.
However, some detailed studies from other areas have been included
where they help in understanding the African upwelling ecosystem. In
particular reference is made to numerous studies on Euphausia pacifica
off California and to the euphausiids of equatorial upwelling areas.
The upwelling of deeper and colder waters into the upper layers con-
stitutes a hydrological eveni~ that allows the sporadic occurrence and
in some cases the development, of cold and bathypelagic forms where
the tropical plankton would be expected if no upwelling took place.
Upwelling is also an ecological event in which the enriched waters
support blooms of primary consumers as well as phytoplankton blooms.
This stress results in a juvenilization of the ecosystem with abundant
popUlations with low diversities.
The seasonal ending of the phenomenon, or the exhaustion of the nu-

trients during the drift of water masses, induces a return to the ini-
tial conditions. This agrees with the theories of Margalef (1968).
I will not discuss here the faunistic characteristics of zooplankton

in the different areas of the African Atlantic, I will only refer to
tables of the dominant species of the prinCipal groups.
33
The description by quantitative and qualitative observation is the

first indispensable step in the analysis of the relationships between
the populations and the variations in environmental conditions. The
second step is the knowledge of the biology, from the chemical composi-
tion to the behavior, of the dominant or the most characteristic spe-
cies. This second step should be studied simultaneously with the first,
because it is one thing to study the characteristics of a species in
the laboratory or in an area at a given time, and quite another to
transfer these results to other areas, or to other conditions, or even
other species.
The first task should be the categorization of the different species

into trophic types which will then allow their classification into the
ecosystem. Although it might seem easy to classify the zooplankton
species into categories of herbivores, carnivores, and omnivores, it
has been shown that this is not the case, and very often this classifi-
cation has appeared extremely simplistic. Most of the species pass
from one trophic level to another, according to the various phases of
their biological cycle or the conditions of the environment. Other
elements necessary to understand the ecosystem come from the fact that
a species behaves at its level as if i t were a transferer of matter or
energy, and it, is therefore important to know for each "cell" the quan-
tity and quality of organic matter that is entering, being transformed,
and leaving the "cell". This is a very broad task, and each process
is influenced by the external parameters and by internal factors of
the animals. We are now at the very beginning of this phase and are
particularly concerned with the transfer of laboratory data to more
direct measurements of the metabolic characteristics of the animal in
its environment; but many problems are still arising at the level of
the methodology and the adaptation of the data, as well as of their
application.
I have chosen to discuss some of these studies, and have grouped the
animals from the systematic point of view, and also in terms of feeding
patterns whenever possible. Examples will be chosen first from the
African Atlantic zooplankton; studies of other upwelling areas which
possess identical species, or more or less similar species are then
considered.
Four categories will be analyzed:
1. the euphausiids, whose behavior in the adult stage causes them to

be included in the micronekton as omnivores; however, they also present
characteristics which demand that they be regarded as herbivores
2. some groups of animals with filtering habits of a more or less micro-

phageous nature; the position of these diverse animals in the pelagic
ecosystem and their trophic relationships in upwelling areas are not
well defined, e.g., radiolarians, foraminifera, thecosomatic pteropods,
appendicularians, thaliacea and cladocera; this paper will deal with
tunica tes only
3. groups or species strictly carnivorous, such as siphonophora, trachy-

medusa, ctenophora and chaetognatha
4. last but not least, the copepoda, the category which is at the same
time the most important and the most diversified.
34
1. Euphausiids
Principal references in African Atlantic: Boden (1954, 1955, 1961),

Nepgen (1957), Meira (1970), Baker (1970), Andreu-Puyal (1976). General
work: Mauchline and Fisher (1969).
1 .,1 General Remarks
The euphausiids constitute one of the key elements of the pelagic eco-
systems: they are primarily oceanic, but the genus Nyctiphanes and a
few other species extend their importance onto the continental shelf.
By their size and their behavior (vertical migrations) they are inter-
mediate between plankton and nekton. By their trophic value and their
biomass they are important in the diet of many pelagic fishes, and also
of some partially benthic animals.
They possess the ability to feed either by predation or by filtration.

They are usually omnivorous with a tendency towards either herbivorous
or carnivorous behavior, depending on the species or on the trophic
conditions of the environment.
Because of vertical migrations of many species they are able to trans-

port certain elements, and to establish relations between epipelagic
levels and meso- or bathypelagic levels.
Lastly they represent experimental material which is very interesting,

and many authors have attempted to keep them alive and watch them on
board oceanographic vessels. The first works in these fields, which
were essentially made on feeding patterns, have been summarized by
Nemoto (1971/1972).
It is very difficult to evaluate the real density of euphausiid popula-

tions in the sea, and all the research on their vertical or geographical
distributions is rather imprecise because of the possibility that some
of the animals have avoided the net, an imprecision that is dependent
upon the type of the net used. A great deal of research has been carried
out to evaluate these possible errors, and to discuss the advantages
of various sampling methods, for example Jerde (1967), Allen (1972),
Brinton (1962) and Roger (1971a). It seems obvious that we do not have
at the moment the correct means to evaluate abundances of euphausiids
and other micronektonic crustaceans. Consequently when comparing re-
sults, it is necessary to be very careful because of the wide variety
of methods used by various authors.
From the various works carried out in the African Atlantic i t is pos-
sible to establish a list of 43 species. Table 1 shows author by author
the area studied, the species present and an indication of their re-
lative abundance using 4 categories. I have included in this discussion
only those species which contributed more than 10% to the total euphau-
siid population in at least one area near the coast. Thus we have six
species in thegenus Euphausia, two in Nyctiphanes, and one in NematosceNs
which present, according to their abundance, very distinct latitudinal
distributions.
Euphausia Nyctiphanes Nematoscelis
Southern Africa E. lucens N. capensis

E. recurva
35
Euphausia Nyytir..hanes NematosceUs

In tertropical zone E. eximia
E. hanseni
E. tenera
Northern Africa E. krohnii N. cap ens is N. megaZops
N. couchii
Table 1. La ti tudinal variations of abundance of the most important species of euphau-

siids in the African atlantic
Area investigated 2 3 4 5 6 7 8 9
340S 220S 100S 28 0 N 170 N 23 0 N 300 N 300 N 35 0 N
32 0 S 31 0 S lOON 16 0 N 15 0 N 16 0 N 280 N 280 N
lSoE 16 0 E 10~ 16 0 w 230W 170 W 130W 28 0 W 100W
16 0 E 110E 14 0 W 160W 25 0 W 21 0 W 170W 290W 15 0 w
Euphausia Zucens xxxx xxxx

E. simi Us x
E. simiUs var. armata x x
E. recurva x xxx xxx x
E. eximia xxxx x
E. hanseni x x xxx x
E. ten era x xxx xx x
E. brevis xxx x x xxx x
E. mutica x x x x x
E. americana xx xxx xxx
E. gibboidEs x x x x x
E. pseudogibba x
E. hemigibba x xx x x xxx xxx x
E. krohnii xxxx xx xxx
Nyctiphanes cap ens is xxx xxx xxxx x
N. couchii xx
Thysanoessa gregaria x x x x x
T. parva x xxx
T. Zongicaudata x
NematosceZis megaZops x xx x xxx x x xxx
N. microps et N. a ttantica x x x x xxx x x
N. teneUa x x x x
Thysanopoda microphthaZma x x x
T. tricuspidata x x x x x
T. aequaUs x
T. subaequaUs x x xx
T. pectinata x
T. obtusifrons x x x
T. cristata x x
T. cornuta x
StyZocheiron carina tum x x x
S. Zongicorne x x x x x xxx xx x
S. eZongatum x x x xxx x
S. abbreviatum x x x x
S. maximum x x x x
S. affine x x x x x x x
S. suhmii x x x xx x
S. microph tha Zma x x
36
Table 1. Continued
Area investigated 2 3 4 5 6 7 8 9
34 0S 22 0S 100 S 28°N 17~ 23 0 N 30~ 300 N 35~
32 0S 31 0 S lOoN 16~ 15~ 16~ 28~ 28 0 N
18DE 160 E l00E 16°w 230W 170 W 130 W 280 W l00W
16DE 11 0 E 140W 160W 250W 210W 170 W 290W 150W
Nematobraehion fZexipes x x x x x
N. boopis x x x x x
N. sexspinosus x x x
Meganyetiphanes norvegiea x
Bentheuphausia amblyops x
1. Nepgen (1957); 2. Boden (1955); 3. Boden (1961); 4. Boden (1961); 5. Meira (1970);
6. Andreu-Puyal (1976), Thiriot (in press); 7. Baker (1970); 8. Weigmann (1974);
9. Thiriot (in press).
Datq on the euphausiids of the Guinean Gulf are lacking except the
very brief accounts of Boden (1961) and of Kinzer (1969). This lack of
data is important because it prevents us from studying the influence
of the oceanic upwelling (Guinean Dome, Angola Dome and, essentially,
the equatorial divergence) on the populations of this group, although
the richness of those areas compared to the neighboring areas has been
noticed (Kinzer, 1969).
1.2 Genus Nyetiphanes
The species of the genus Nyetiphanes are the only ones that are charac-
teristic of the neritic zone. N. eapensis which was thought to be ex-
clusively distributed along S;W. Africa (Mauchline and Fisher, 1969),
has been found in abundance off the Cape Verde Islands (Meira, 1970)
and particularly off Mauritania (Casanova, 1974a). Its occurrence was
noticed on the continental shelf of the Guinean Gulf, but without large
densities. N. eouehii takes over at the latitude of Morocco, which is
the southern limit of its geographical distribution. On the coast of
Peru and California the genus is represented by N. simplex. These are
essentially temperate or cold temperate forms very well adapted to
regions of cold water upwelling.
Typically omnivorous, these species are able to feed equally well on

benthic detritus as well as plankton and suspended organic matter
(Mauchline, 1967). Therefore i t is because of the general enrichment
of an area that upwelled waters induce the abundance of these species.
The omnivorous feeding habits of Nyetiphanes start as soon as the calyp-
topis larval stages are reached, and Le Roux (1973) has shown that the
growth obtained with a strictly phytoplanktonic diet is not as good
as that obtained with Artemia nauplii. Fowler et al. (1971) were able
to keep N. eouehii for 11 months with a mixed diet; they observed varia-
tion of molting frequency as a function of temperature and individual
size. The molting interval was from 4 to 9 days in the adult stages
during the experiment, and molting usually occurred (80% or 90%) during
the night. This represented an input of organic matter in the super-
ficial layer which is rather important. Jerde and Lasker (1966) also
experimented on the molting of N. simplex on board an oceanographic
vessel. They found a molting interval of about 5 days, and measured
values of molting weight o~ about 10% of the total dry weight of the
animal for individuals weighing between 0.300 and 0.800 mg (dry weight).
37
Like many other euphausiids the species of the genus Nyctiphanes may
occur in swarms (Cram and Schulein, 1974) or may show strong variations
in abundance in a particular area OVer periods of a few days, as shown
by Brinton (1962) for N. simplex in the Californian coastal waters (the
maximum concentrations he observed for the latter species being about
8000 1000 m- 3) •
1. 3 Gen us Nematosce lis
Nematoscelis is represented by three species on the African Atlantic

coast, but only N. megalops is of importance in zones influenced by up-
welling. N. megalops is an oceanic species from cold temperate waters;
it has been found off S.W. Africa but seems more abundant in the Morocco
and Mauritania areas. Mesopelagic during the day, it comes up towards
the surface during the night without becoming superficial (Andreu Puyal,
1976; Thiriot, personal observation).
It is an omnivorous species with carnivorous tendencies which are more

pronounced than for Nyctiphanes or Euphausia. This characteristic is partly
deduced from its diel behavior, but also from the study of stomach
contents and observations of cephalothoracic appendages (Weigrnann,
1970). Ponomareva (1971) indicated the lack of vertical migration for
N. gracilis and thus the lack of diel feeding rhythms.
In the east Pacific we find N. megalops along the Chili coast down to
about 30 0 S, whereas in the northern hemisphere the genus is represented
by a very closely related species, N. difficilis, from 50 0 N to 20 0 N in
the California current. The intertropical areas are populated by N.
gracilis (in the Pacific and the Indian oceans; Mauchline and Fisher,
1969). The zoogeography of the genus has been reviewed and elaborated
by Gopalakrishnan (1974). In the Indian Ocean N. gracilis can reach max-
imum concentrations of more than 100 individuals 1000 m- 3 in the Arabian
sea and the Bay of Bengal during the S.W. monsoon (April to October),
a report which confirms the observations of Weigmann (1970) and Pono-
mareva (1972).
Similar concentrations were found by Andreu Puyal (1976) for N. megalops

(100 individuals 1000 m- 3 ) and by Brinton (1962) for N. difficilis (500
1000 m- 3 ). Nemoto et al. (1972) evaluated the fecundity of N. difficilis
in the N.W. Pacific (41 0 N, 165 0 W). These authors established a linear
relationship between the number of eggs and the size of animal, and
measured the carbon content and the ratio of carbon to nitrogen; these
values were also measured for N. microps and N. atlantica; they had been
previously determined for N. difficilis by Hopkins (1968): the number of
eggs ranged between 150 and 425, and the total weight of eggs was about
20% of the dry weight of the animals.
1. 4 Gen us Euphausia
The studies of the species of this genus are particularly numerous and
include experimental and field studies. They principally concern the
antarctic species E. superba and the Pacific species E. pacifica. The
biogeography of the most abundant species off the African coast is
given in the monograph by Machline and Fisher (1969).
E. lucens is a southern hemisphere species abundant to the north of the

antarctic convergence between 30 0 S and 45 0 S.
E. recurva is found in the Atlantic, Pacific, and Indian oceans and is

defined as "biantitropical" in the Pacific (that is to say i t is local-
ized between 20 0 and 40 0 in the two hemispheres.)
38
E. exirrria is abundant in the California current, the Peru current and

the Pacific south equatorial current, but Boden (1961) also collected
large numbers of them in the Gulf of Guinea.
E. ha:nseni is found along the coas t of S. W. Africa and south of 20 0 N off

N.W. Africa (Weigmann-Haass, 1976).
E. tenera is a tropical and subtropical species in the three oceans.
E. krohnii is found in the north-eastern Atlantic and the Hediterranean

sea.
In the Pacific, north of 25 0 N, Brinton (1962) gives indications of the

abundance and distribution of E. pacifica, E. brevis, E. recurva, E. exirrria
and E. gibboides which have the highest population densities, higher
than 5000 individuals 1000 m- 3 , mainly near the north coast of Cali-
fornia. The other species reach maximum values between 500 and 5000
individuals 1000 m- 3 at a little distance offshore.
Brinton and Gopalakrishnan (1973) present quantitative distribution

maps during the two seasons that are established by the monsoon regime
in the Indian Ocean. In the upwelling areas of the Arabian sea and Bay
of Bengal one can collect the predominantly coastal species E. dis tin-
guenda (more than 5000 individuals 1000 m- 3) and E. diomedeae. Ponomareva
(1972) observes certain annual differences of dominance of some species
such as E. pseudogibba in the Arabian sea, and confirms the abundance
of the two other species.
Because of the abundance of E. krohnii in the N.W. African upwelling

area the work of Casanova-Soulier (1974) on this species in the Hediter-
ranean sea is important. These results concern essentially the biometry
(larval and adult), biogeography, bathymetric characteristics as well
as nutrition, reproduction, growth, and life cycle. E. krohnii seems
to be a widely spread species with a distribution similar to N. couchii
and M. norvegica, but more oceanic. Eurybathic between 2000 m and the
surface this species presents diel migrations to the surface, although
some individuals remain in deep water. These results are different
from the observations along the N.W. African shelf (Andreu Puyal, 1976;
Thi~iot, in press) and imply different behavior in an oligotrophic
zone compared to a rich one. E. krohnii does not seem to be influenced
by the divergence in the Ligurian Sea, but by other hydrological char-
acteristics of the Hediterranean Sea such as the Atlantic superficial
current (Casanova-Soulier, 1974).
The diet of this species in the Ligurian sea presents seasonal varia-
tions which reflect the variations of trophic conditions in this area;
during the spring the gut contents consist of phytoplankton and de-
tritus, and between 20% and 50% of zooplankton during the other seasons
(Casanova-Soulier, 1974). These results, and also some data concerning
N. couchii and N. megalops, are very interesting but it is difficult to
extrapolate to upwelling areas which have a very different trophic
cycle. Of particular interest is the very short period of enrichment
in the Hediterranean sea compared with the life cycle of euphausiids,
which in some is about two years long.
E. krohnii appeared in very large numbers in the superficial layers

during the night in the upwelling area off Rio de Oro and Mauritania
near the continental slope (more than 10000 1000 m- 3 ) and constituted
about 80% or 90% of total euphausiids (Andreu Puyal, 1976; Thiriot,
in press). This fact is important from a biogeographical point of view
because this species is, in these areas, at the southern boundary of
its geographical distribution.
39
1 .5 Selected Works on Euphausia pacifica
While our knowledge on the ecology of Euphausia krohnii in upwelling

areas is still limited, extensive studies on Euphausia pacifica provide
a rather comprehensive picture on distribution and life history of
this species under the upwelling conditions of California. Both species
and habitats are sufficiently similar to warrant the following, somewhat
detailed review of the information available on E. paaifiaa.
Experimental studies have shown the importance of the relation between

the size of individuals and their biological and physiological char-
acteristics. E. pacifica is one of the larger species of the genus, not
as large, however, as most of the antarctic species. The geographical
and vertical distributions, and the relation of this species with tem-
perature and salinity at different latitudes have been observed by many
authors.
Brinton (1962) notes offshore of California a population that has a

relatively homogeneous distribution and undergoes slow changes in abun-
dance. Komaki (1967) - to the contrary - notes swarms near the coast
of Japan during spring in the area where cold waters and coastal waters
are mixed. At this period euphausiids are fished by men and are eaten
by fishes (principally ScorrUJer japonicus), whales, and birds. In these
swarms more than 50% of the females are carrying spermatophores. Day
(1971), also studying E. pacifica, notes a greater number of individuals
in spring and autumn between 46 0 N and 55 0 N, but he does not report their
occurrence in swarms. E. pacifica is a major component (76%) of the bio-
mass of the macroplankton and micronekton at the limit of the continen-
tal shelf especially in the southern part. Alton and Blackburn (1972)
study the variations of the vertical distribution of Thysanoessa spinifera
(a neritic species) and Euphausia pacifica offshore of the state of
Washington, and show the increase of the population at the surface
during the night. Smiles and Pearcy (1971) observe the same thing along
the coast of Oregon~ the density of E. pacifica is higher inside 25 miles
than offshore. These authors compare the growth rate of this species
in different regions of the Pacific and note that growth is about twice
as fast near shore (with a shortening of the life cycle) than in the
N.W. Pacific. They attribute this growth to the high primary produc-
tion associated with the upwelling. The growth rate is equivalent to
a ·mean of 0.065 rnrn per day for the whole length of life with a maximum
of 0.095 rnrn per day for the juvenile stages. This rate is higher than
that found by Lasker (1966) in the laboratory (0.048 rnrn per day).
Lasker was the first to experiment on the metabolism of euphausiids.

In his important study (Lasker, 1966) he observed the length of the
intermolt period (between 3 to 8 days) according to the temperature
and the quantity of food~ the relation with temperature has been studied
further by Fowler et al. (1971). Lasker analyzes also the variations
of the rate of oxygeQ consumption as an index of energetic utilization
of carbon, similar to the first works on copepoda.
The molt represents between 4% and 14% of the dry weight of the animals~
these data confirm the preliminary observations (Lasker, 1964~ Jerde
and Lasker, 1966). Lasker measured the percentage of ingested carbon
which was assimilated by the animal during feeding. This assimilation
of carbon is about 30% during the fast growth of the larval and juvenile
phases and 6% during the slow growth~ the molt represents between 6%
and 11% and the respiration between 62% and 87%. These values are very
close to those found by Ponomareva, 67% for respiration, 9% for growth,
15% for molting, and 9% for egg production (in Lasker, 1966).
40
The size, the forms, and the quantity of particulate matter are im-
portant for the feeding of Euphausia, as well as the phase of the molting
cycle. Experiments have shown the possibility of selective predation,
and its high variability. Lasker estimates also the quantity of carbon
per day in relation to total weight; the requirements of the whole
population in the area studied represented about 3% of the daily primary
production. Nemoto (1968) studied the feeding pattern of some species
with different bathymetric distributions and different kinds of verti-
cal migrations; he measured the chlorophyll a and the pheophytin in
the stomach contents, the gut, and the fecal pellets. This work shows
the high herbivorous tendency of E. pacifica, the importance of vertical
migration in feeding of this species, and the importance of fecal pel-
lets for the transfer of pigments to the deeper waters. Fowler and
Small (1972) have also noted the importance of fecal pellets of Medi-
terranean euphausiids in the transport of organic matter and some min-
eral elements down to the sediments. With a sinking rate of 100 to 800
m per day, the fecal pellets sink faster than molted exoskeletons or
dead animals, and disintegrate more slowly.
The excretion rates of nitrogenous compounds (ammonia, arninoacids, urea,

or total nitrogen) have been measured by Jawed (1969) for E. pacifica.
Ammonia constitutes 72% to 87% of the total nitrogen excretory product.
The excretion rate changes in relation to the temperature; between 21
~g N per mg N body weight per day at 10 0 C and 12 ~g at 4 o C. When the
animal is starving, the utilization of its own proteins reaches about
2% per day. The QIO values of the different forms of nitrogen excretion
have been calculated and are between 2.3 and 3.0.
The measurement of the oxygen consumption rate and the evaluation of

the influence of external parameters upon respiration of the animals
is an important indication of the adaptative possibilities of different
species to variations in environmental conditions. Teal and Carey (1967)
have studied the influence of temperature and pressure on the respira-
tion of some euphausiid species. For the epipelagic species only tem-
perature has influence; an increase in pressure does not result in an
increase in respiration at low temperatures. The QIO ranges between
2.0 and 3.5. These experiments have been made on board oceanographic
vessels in tropical areas of the Atlantic and the Indian Oceans and
in the north Atlantic. The authors confirm MCLaren's hypothesis (Mc-
Laren, 1963) of the advantage from an energetic point of view of stay-
ing at low temperatures during the day. For E. hemigibba the experimental
curves show that this economy corresponds to the equivalent of the
energetic value of a 2-mg copepod. With another method of estimating
the respiratory rate (measurement of the activity of the electron trans-
port system) King and Packard (1975) obtain similar results, that is
to say, the nonsignificance of pressure upon the respiration of Euphau-
sia pacifica. Small et al. (1966) measured the respiration of E. pacifica
and Thysanoessa spinifera in the area of Oregon and show the importance
of the temperature effect (QIO = 2.11). The study of the QIO of animals
which have been kept at different temperatures and the comparison with
the natural populations (Small and Hebard, 1967) show that E. pacifica
is well adapted to temperatures between 50 and 10 o C, but not for 15 0 C.
Wi th these data Small (1967) tried to evaluate the "energetic flow"

through E. pacifica - a term more important than biomass for the study
of food webs. The author presents firstly the seasonal variations in
the composition of the population (percentages of three categories of
weight) and estimates the quantity of oxygen consumed, taking into
account the mean levels during the day and night, and transforms the
respiration into equivalent g cal per animal per day. The curve shows
an increase from January to June (animal growth) followed by a rapid
decrease (egg-laying and arrival of juveniles). The author arrives at
41
an energetic flow of 100 g cal per animal per year which corresponds
to the ingestion of 9 mg carbon per mg dry weight per year.
Paranjape (1967) studied the molting and the respiration of five species
of euphausiids (including E. paC'1~fica). The animals were caught near the
coast of British Columbia (surface temperature between 11 0 and 15 0 C).
The results obtained are close to the data above. However it is neces-
sary to notice that E. pacifica seems more eurythermal here than near
Oregon, with an upper limit of 20 0 C. The QIO of the respiration varied
between 2.21 (50 to 10 0 C) and 2.55 (10 0 to 15 0 C) and did not vary with
the size of animals. The ingestion of food for molting and respiration
corresponded to 0.022 mg C per mg dry weight per day. During the molting
he observed a decrease in the food uptake and an increase in respiration
(about 34%).
The different possibilities of adaptations presented by different geo-

graphical populations of E. pacifica have been studied by Gilfillan
(1972a), who observed the combined effects of temperature and salinity
on oxygen consumption. The coastal popUlations were able to tolerate
variations of temperature and salinity to a greater extent than the
oceanic populations. The lethal temperature was 24 0 C, the variations
of salinity alone had no effect on the respiration of the coastal pop-
ulations. The interaction of temperature and salinity produced seasonal
variations; summer populations tolerated wider variations than the
winter popUlations. The author studied an area corresponding to the
whole geographical distribution of E. pacifica along the American coast
between 30 0 and 55 0 N (Gilfillan, 1972b). Significant differences were
shown for the interactions: temperature-season, salinity-season, salin-
ity-season-area, but there is no indication of an evolution of the
physiological possibilities in relation to the latitude. This means
that the stress due to the temperature variations is suppressed by
the influence of an internal factor: genetic adaptation or acclimatiza-
tion.
1.6 Euphausiid Populations in the Equatorial Pacific
Before closing this chapter I wish to summarize the different works of

Roger (1971a,b; 1973a-f; 1974a,b; 1975) on the zoogeography, ecology,
biology, and trophic relations of Pacific and Indian euphausiids. This
author studied the vertical distributions in the equatorial currents
and especially the qualitative composition in the superficial layers.
E. diomedeae constituted more than 90% of the whole euphausiid population.
The study of the variations in the diversity index showed the difference
between the high values of the tropical areas and the low values of the
equatorial areas, especially in the eastern part located between the
Galapagos Islands and the Marquesas Islands. Along the equator this
index increased towards the west, and corresponded to the biomass dis-
tribution. Increased diversity was associated with the evolution of
the pelagic ecosystem resulting from the enrichment by the equatorial
divergence which is more important in the eastern part. This evolution
of diversity index has also been observed in other zooplanktonic groups
(Gueredrat et al., 1972).
In his ecological study Roger grouped the different species according

to their trophic affinities. The equatorial divergence area was char-
acterized principally by Nematoscelis teneUa, N. gracilis, Thysanopoda orien-
talis, T. monacantha, Euphav.sia diomedeae and StyZocheiron affine.
In his biological study the author noted the characteristics of repro-

duction, growth, and length of life of several species. The feeding
42
patterns were indicated from the stomach contents. We can notice that
N. gracilis is essentially carnivorous, but i t is able to supplement
its diet with phytoplankton; N. microps, N. teneUa and E. diomedeae are
typically omnivorous; E. paragibba and E. gibboides are strictly herbi-
vorous. These data confirm several studies cited above; similar feeding
tendencies are indicated at the level of the genus but in general they
are different from one species to another and also from one season or
one trophic condition to another. The diel variations of feeding, shown
by the percentage of empty and full stomachs, are different according
to the species, but thE~ total ingestion is more important during the
night and between 100 m and the surface. The percentage of carnivorous
species increases in oligotrophic conditions and, accordingly, we have
a large dominance of herbivores in the rich equatorial areas.
The author discusses also the predation of euphausiids by micronektonic

fish. The direct predation by tunas seems to be negligible because this
fish presents a daily and superficial feeding pattern in these areas.
These observations are different from those of Dragovitch and Potthoff
(1972), for instance, from the Gulf of Guinea.
The place of the euphausiids in the ecosystems of areas enriched by

upwelling is consequently particularly important because of the trans-
fer of energy to the higher trophic layers, especially fishes. It is
in general near the continental shelf that we observe the maximum con-
centrations; however, high concentrations have been noted in areas more
offshore, which are characterized by a dome or a divergence, but abun-
dances in these areas have been relatively less quantified.
2. Tunicates
Principal references in African Atlantic: Furnestin (1957), Van Zyl

(1960), Godeaux (1962), Godeaux and Goffinet (1968), Seguin (1973),
Vi ve s e t al. (1975).
It is difficult to have a good idea of the quantitative importance

of these groups in relation to the upwelling phenomenon and to know
their real place and influence in the food webs.
Heron (1972a, b) has shown that Thalia democratica has a very fast bio-
logical cycle: two days from oozoid to blastozoid, which corresponds
to an increase in the populations of between 1.6 to 2.5 per day. Fraser
(1961, 1962) notes some cases of very high densities of Salpa fusiformis
with important reductions of the other groups. The author attributes
these decreases either to the feeding competition or to the influence
of some products of the metabolism of salps. The suspension feeding
pattern and especially the herbivorous behavior of salps is well known.
Silver (1975) has shown that this filtration is not selective in the
case of Thalia democratica, Salpa fusifriirmis, and Cyclosalpa bcikeri in the
California current.
Many authors have tried to find relationships between blooms of salps

and phySical or biological characteristics of the environment. Braconnot
(1971) in the Mediterranean sea observed a seasonal increase in numbers
of Thalia democratica and Salpa fusiformis which took place just after the
spring bloom of phytoplankton. Offshore of Oregon Hubbard and Pearcy
(1971) have studied salps where Salpa fusiformis and Iasis zonaria could
be sampled in high concentrations throughout the year; maximum abun-
dances occurred during the upwelling season. However, one is struck
by important variations between the different years. Bhavanarayana and
43
Ganapati (1971) have tried to form groups of zooplankton species as-

sociated with the different species of tunicates in the western part
of the Bay of Bengal, and have tried to find relationships with the
different water masses. Silver (1975) tried in the California current
to characterize the waters where salps occurred in swarms (more than 1
animal m- 3 ) from the qualitative and quantitative composition of diatoms.
Significant differences were found between stations with swarms of S.
fusiformis, stations with swarms of T. democrotica, and stations with no
swarms. Silver thinks that there is a possibility of identifying the
habitats of these speCies whose ecological niches seem so difficult
to define.
These animals are called "opportunists" because they have the possibil-
ity of counter-balancing their competitive struggle with other herbi-
vores such as the copepods, by taking advantage of one of the fastest
reproductive cycles in the metazoa (Heron, 1972b). When good conditions
occur, the salps reproduce quickly; consequently the correlations with
copepods and euphausiids become negative. Silver (1975) summarizes also
the studies on predators of salps; these predators are few in number,
and this is another explanation for their very fast formation of swarms.
Tokioka (1960) studied the distributions of appendicularians and thali-

acea in the whole of the North Pacific Ocean, and tried to distinguish
water masses with dominant species in a manner similar to that done
for chaetognaths (Tokioka, 1959). The author found species which were
located in the enriched waters near the coasts (especially California
current) such as DoUoZum nationaUs and DoUoZetta gegenbauri, and forms
with a broad oceanic distribution; we can observe some species with an
increased abundance in the equatorial divergence such as OikopZeUPa
Zongicauda. In the same way the groups proposed by Bhavanarayana and
Ganapati (1971), in the Bay of Bengal, indicate that FritiHa:r>ia peUucida
is located within upwelled waters.
Table 2 summarizes the observations of appendicularians, salpids and

doliolids of the African Atlantic zooplankton.
Together with the Radiolaria, the Pteropoda Thecosomata and the

Cladocera the Tunicata present several similar characteristics:
1. cosmopolitan geographical distributions which correspond to very

different thermohaline characteristics
2. a life pattern that is neritic or epipelagic
3. a very high reproduction power
4. a suspension feeding pattern upon small particulate organic matter.
These ecological characters seem well adapted for high production in

coastal upwellings, but nevertheless these specles are rarely found
in high numbers in these areas. No clear relationship seems to exist
between abundance of these groups and phytoplankton blooms. We find
these species in greater abundance in other enriched areas. For in-
stance, DoUoZum nationaUs has been found in the part of the Mediterranean
sea influenced by the Rhone (Braconnot and Casanova, 1967).
It is essential that we increase our knowledge of the feeding patterns

of these animals, and state more precisely the physical and the bio-
logical properties of the waters where these swarms occur, and of the
organic matter in suspension.
44
Table 2. Summary of principal observations on tunicates in the African Atlantic
Morocco
Furnestin (1957) Thalia democratica Summer swarms, indicator of
slope waters
Salpa fusiformis Summer maximum in oceanic zone
Oikopleura dioica Neritic water and spring
maximum
O. longicauda Oceanic
Vives et al. (1975) Salps Rare
June-July Oikopleura dioica Maximum 10 animals m- 3
oceanic zone o. longicauda
o. fusiformis
Fritillaria borealis
F. pellucida 100 "
Rio de Oro, Mauritania

N.LO. Anon. (1968) Salpa maxima Swarms near Canary Islands
"Discovery" Jan.-April Pegea confoederata
Andreu-Puyal etal. (1975) Thalia democratica Swarms
oceanic zone Salpa aspera
March-September Doliolum nationalis Maximum 100 animals m- 3
Doliolum gegenbauri
Oikopleura fusiforwn:s
o. dioica
Jean Charcot (1975) o. longicauda Swarms
April
Dakar
Seguin (1973) Thalia democratica Rare
Oikopleura longicauda All the year, maximum 35% of
zooplankton
Doliolum nationalis Maximum 5% May-June and
October-November
Gulf of Guinea
Seguin (1973) Salpids 3.6% of annual zooplankton,
during warm season 55 m- 3
Thalia democratica
Appendicularians 3.4% of zooplankton, during
cold season 59 m- 3
Doliolids 0.01% of zooplankton
Mensah (1969) Salpids All the year, min. Oct.-Nov.
Appendicularians Minimum July maximum Aug.-Sept.
Bainbridge (1972) Oikopleura longicauda Maximum during cold season
Dolioletta gegenbauri
Thalia democratica Not clear
Binet (1970) Doliolids Max. during cold season 1000 m- 3
Salpids Swarms rather during warm season
Appendicularians Numerous, variations not clear
Neto and De Paiva (1966) Dolioletta tritonis Several max. all the year
Doliolum nationalis Absent during cold season
N.LO. Anon. (1968) Salpa cylindrica Swarm near 100 S
Godeaux and Goffinet Thalia democratica Maximum for temp. > 25 0
(1968) Salpa fusiformis
Salpa cylindrica Little variation
Godeaux (1962) Thalia democratica Not numerous
45
Table 2. Continued
Cap LOpez SaZpa cyZindPica Not numerous

DoZioZum nationaZis More abundant
D. denticuZatum
DoZioZetta gegenbauri
D. tri toois
South West Africa
Van Zyl (1960) ThaZia democratica
DoZioZum denticuZatum Maximum during warm season
DoZioZum nationaZis Maximum during cold season
Kollmer (1963) SaZpa fusiformis Swarm in oceanic zone chiefly
during spring and winter
Hart and Currie (1960) Salpids Swarms in Sept.-oct.
Stander and De Decker DoUoZum, SaZpa Increasing during a special
(1969) warm year
3. Carnivorous Species of Macrozooplankton
Although numerous authors have tried to categorize zooplankton into

suspension feeders, herbivores, or carnivores (Greze et al., 1969;
Greze, 1970; Bainbridge, 1972) i t is a very simplistic approach, and
not close to reality. In fact there are few species which have a strict-
ly carnivorous feeding pattern; included in this category are some
macroplanktonic forms such as Ctenaria, Siphonophora, the Medusa Liriope
tetraphyUa and chaetognaths.
There are few studies of the distribution of these groups in relation
to zooplankton biomass; also little is known of their biological,
ethological, and ecological characters in relation to the trophic
conditions of the environment.
3.1 Siphonophora
PrinCipal references in African Atlantic: Cervignon (1961), Margulis

(1971), Neto and Lourenco (1973), Pugh (1974).
Off the coast of S.W. Africa the Siphonophora are almost exclusively
located beyond the continental shelf (Unteruberbacher, 1964); the
principal species are Muggiaea aUantica, AbyZopsis tetragona, CheZophyes ap-
pendicuZata, and AgaZma sp.
In the Gulf of Guinea Binet (1970) found Siphonophora abundant on the
continental shelf of Gabon (up to 40 animals m- 3 ) from January to Sep-
tember, a period not correlated with a typical hydrological event. Neto
and De Paiva (1966) also found Muggiaea aUantica principally between May
and October. Bainbridge (1972) did not see quantitative differences
between the upwelling season and the subsequent period of stratified
waters near the Nigerian coast. Off the Ivory coast Seguin (1973)
observed that this group was not very important (0.7% of the total zoo-
plankton); siphonophores reached their maximum abundance in February,
wi th Diphyes bojani, CheZophyes appendicuZata, and AbyZopsis tetragona.
At Dakar they had two peaks of abundance (Seguin, 1973), in June and
October, with Lensia subtiZis" Muggiaea aUantica, CheZophyes appendicuZata and
Diphyes bojani. Off the Spanish Sahara and Mauritania, Muggiaea aUantica
46
and Chelophyes appendiculata were relatively abundant in February to March,

near the slope (Cervignon, 1961). In the Moroccan Atlantic Furnestin
(1957) observed Lensia conoidea, L. subtilis, Chelophyes appendiculata, and
Muggiaea atlantica without important seasonal variations.
In the oceanic zone we can note the studies of Neto and Lourenco (1973)
near the Cape Verde Islands and Pugh (1974) near the Canary Islands.
Margulis (1971) studied the different species of the genus Lensia in
the whole Atlantic Ocean, and observed maximum populations in the en-
riched areas at the boundary of the northern and southern Atlantic
gyres, as well as along the African coasts and the equatorial diver-
gence.
I am aware of only a few quantitative studies on Siphonophora and their

trophic importance, although Barham (1963) observed their importance
as constituents of the deep scattering layer.
3.2 The Trachymedusa Liriope tetraphylla
This species is cosmopolitan in warm waters, holoplanktonic and epipe-

lagic. Its maximum concentrations are always observed in the neritic
zone. Off S.W. Africa Kollmer (1963) found it, however, in high numbers
in oceanic waters, especially in the drift of the Benguela current.
In the Gulf of Guinea this species was very abundant along the Nigerian
coast (Kramp, 1955), rare in oceanic waters (Repelin, 1965; Seguin,
1973), and without clear seasonal variations along the Ivory Coast
(Binet, 1970).
Near Dakar this species was principally present between March and June,
that is to say during the second half of the upwelling season, with a
maximum in April (8% of the total zooplankton; Seguin, 1966). Off Moroc-
co Furnestin (1964) observed Liriope with other forms indicative of
slope waters. Goy and Thiriot (1974) studied its distribution in this
region during two seasons. More or less scattered in the whole area
(offshore included) between January and March, L. tetraphylla was con-
centrated in the neritic area during the summer, a season rich in
neritic zooplankton. The correlation between the occurrence of this
species and high planktonic biomass in upwelling areas is better than
for the chaetognaths or the Siphonophora.
Goy (1974) summarized the different observations on this species and

studied in detail its seasonal variations near Dakar. The maximum abun-
dance (about 4 animals m- 3 ) was found near the coldest waters. It is
well known that L. tetraphylla is a very active predator and feeds on
copepods as well as young fishes. Fraser (1969) studied another species
of the same size and found that one animal is able to eat between 100
to 200 young fishes per month. However, we do not have any quantitative
studies dealing with the feeding of Liriope.
3.3 Chaetognaths
Principal references in African Atlantic: Furnestin (1957, 1966), Al-

varino (1959, Heydorn (1959), Neto (1961), Ducret (1962, 1968), De
Saint Bon (1963a, b), Venter (1969).
Numerous studies have been made on the chaetognaths of the African

Atlantic, essentially from a biogeographical point of view. The studies
result mostly from occasional cruises, and we have few observations
of seasonal variations of chaetognaths, and their relationships with
47
upwelling. Moreover, there is some confusion over the systematics of

some species from upwelling areas.
In the Cape area of S.W. Africa several authors have noted the influence
of the Indopacific populations, represented by S. pacifica, S. bedoti, S.
robusta, S. neglecta and S. regularis (Heydorn, 1959; Furnestin, 1966;
Ducret, 1968; De Decker, 1973). These species can be transported north-
ward by the Benguela current to 15 0 S (Ducret, 1968; for S. bedoti).
The very important upwelling off the Cape of Good Hope does not seem
to increase the number of any particular species (De Decker, 1973).
Masson (in De Decker, 1973) observed however, in the superficial layers,
some bathypelagic species like EUkrohnia hamata and S. decipiens. S. friderici,
normally a neritic species drifts off in the N.W. direction during the
maximum of upwelling season (August-Oct.). S. decipiens is also considered
as a good indicator of upwelled waters by Sund (1961) in the Costa
Rica dome and by Haq et al.(1973) south of Pakistan. In the Benguela
current the dominant species are S. tasmanica (Neto, 1961; Venter, 1969)
and particularly S. bierii Neto, 1961; Ducret, 1968). Near the coast
important populations of S. friderici and S. setosa have been observed
(Neto, 1961).
S. tasmanica is a cold temperate species occurring throughout the world

ocean; its distribution appears continuous along the African Atlantic
coast (Alvarino, 1969) in the upwelling areas. This species is partic-
ularly abundant off Mauritania, and off the southern part of Morocco.
It is a form noted for its high individual biomass and its lipid stores
under optimal trophic conditions (i.e., high number of copepods).
S. bierii is well represented along the west coast of the two Americas,
especially in the Peru current and the California current (Bieri,
1959). In the African Atlantic this species has been collected in the
Benguela current, the Guinean Dome and the coastal upwelling of Senegal
and Mauritania (Furnestin, 1966; Casanova, 1974b).
From the Gulf of Guinea to the Cape Verde Islands one may observe in
great numbers the neritic and subneritic species Krohnitta pacifica, S.
tenuis, S. friderici, S. hispida, and S. enflata. S. hispida seems dominant
off the Ivory Coast during the little and the great cold seasons (De
Saint Bon, 1963a). The abundance of this species increased more quickly
in relation to the increase of the prey than did S. enflata and S. fride-
rici.
In conclusion, and with some assumptions, we can classify the species

of Atlantic chaetognaths into three categories according to their re-
lationships with upwelling:
1. The species normally meso-or bathypelagic that come to the surface

wi thout increasing their numbers; EUkrohnia hamata, E. fowleri, Sagitta de-
cipiens, S. neodecipiens, S. lyra, and S. planctonis.
2. The subsuperficial or mesopelagic species rare in stable conditions

but abundant in upwelling areas; S. bierii, and perhaps Krohnitta sub-
tilis and Pterosagitta draco.
3. The epipelagic or subsuperficial species which increase as do other

carnivorous species (Binet, 1968; Goy and Thiriot, 1974), and which
are often located far off the center of upwelling; S. tasmanica, S. his-
pida, S. enflata and S. friderici.
These observations are not sufficient, and it is necessary to study

with better sampling methods the relation of the cold upwelled waters
48
and the enriched drift currents. Studies of the differences of feeding

patterns (size of prey, etc.), metabolism (lipid storage or not), thermal
characteristics, and the possibilities of quick responses to the varia-
tions of the environmental conditions (growth and reproduction) are
also very important to increase our understanding.
4. Copepods
Principal references in African Atlantic: Marques (1953, 1957, 1958),

Vervoort (1963, 1965), Unteruberbacher (1964), Binet and Dessier (1971).
Bainbridge (1972), Binet et al. (1972a-c), Roe (1972a-d), Vives (1974,
1975) •
This group of holoplanktonic Crustacea constitutes the most important

element of the zooplankton in all seasons and all areas, in number as
well as in biomass. We may consider that it represents about 70% or
80% of the number of mesoplanktonic animals. During the rich upwelling
season along the Nigerian coast, the copepods represented 92% of the
zooplankton (Bainbridge, 1960). However,some exceptions occur especial-
ly in neritic areas. For instance in the Angola area off Baia Farta
the copepods constituted only 44% of the total annual zooplankton (Neto
and De Paiva, 1966).
In general, copepods are the first group to increase after enrichment

by upwelling. The number of species is very high [more than 200 in the
most complete studies, such as Vervoort (1963 and 1965), Vives (1975)
in Mauritania and Rio de Oro area, or Roe (1972a-d) near the Canary
Islands.] This group is also very diversified in biological, ethological,
and ecological characteristics. The trophic enrichment of one area in-
creases the dominance of only a few species. At the beginning of the
enrichment the total biomass increases along with the development of
cold and herbivorous fauna and some bathypelagic species. At the end
of the cold period the specific diversity increases, the strictly car-
nivorous species appear with the omnivorous ones, and we pass progres-
sively to a tropical or subtropical fauna in which only few species
reach 10% of the total zooplankton.
The copepods of the African Atlantic have been studied by many authors.
Table 3 summarizes their principal conclusions concerning the dominant
species during the upwelling season and during the other seasons.
Off S. and S.W. Africa the qualitative composition and seasonal varia-
tions of copepods have been studied by Kollmer (1963), Unteruberbacher
(1964) and De Decker (1973). Along the coast of Angola we have the in-
ventory of Marques (1953, 1957, 1958), and the description of the sea-
sonal cycle by Neto and De Paiva (1966). In the Pointe Noire area Binet
(1970), Binet et .al. (1971, 1972a, b), and Roux et al. (1973) have
tried to form groups of species using multivariate analysis. In this
area 11 groups can be distinguished, but the cold fauna and the warm
fauna are not well separated. This very complicated ecosystem is the
result of mixing and seasonal influences of different kinds of waters:
warm and stratified waters, waters with low salinity, or a branch of
the Benguela current.
On the contrary along the coasts of Nigeria, Ghana, and the Ivory Coast,
the inshore upwelling season is well marked. The composition of the
groups near Abidjan can be established with the same mathematical method
(Binet et al., 1972b). The two first axes defined by the analysis are:
first, a thermal axis which isolates the fauna of the cold season, and
49
Table 3. Principal species of copepods in African Atlantic zooplankton
During upwelling season During other seasons
Morocco Acartia cZausi, CZausocaZanus, CZausoaaZanus, Oithona, Oncaea,

Vives (1975) Oithona nana, Oncaea, Temora Aaartia, CaZoaaZanus styUremis,
st;yUfera, ParacaZanus par>Vus, PZeuromaIm1a, ParaaaZanus parvus
Centropages t;ypicus
Rio de Oro CaZanoides carinatus, Temora NannoaaZanus minor, NeoaaZanus
Mauritania st;yZifera, ParacaZanus par>Vus, graaiUs, Oithona frigida. Rhin-
Vives (1974,1975)
CtenocaZanus vanus, Metridia aaZanus aornutus, EuaaZanus sub-
Binet (1973) Zucens, Acartia danae, Oncaea tenuis, PZeuromamma robusta,
aurta, CaZanus heZgoZandiaus, EuahireUa rostrata, UndinuZa
Centropages ahierahiaes vuZgaris
Oithona
Dakar CaZanoides aarinatus, Euca- NannoaaZanus minor, UndinuZa
Gaudy and Seguin
Zanus attenuatus, E. arass us , vuZgaris, Cen tropages furaatus,
(1964)
EuchireUa rostrata, Euchaeta EucaZanus monaahus, E. piZeatus,
hebes, Candaaia bipinnata E. subtenuis. ParaaaZanus parvus,
P. aauZeatus, Temora st;yUfera,
T. turbinata, Euahaeta marina
Port. Guinea ParaaaZanus arassirostris,
Marques (1955, Euterpina aautifrons, Oithona
1957, 1958, 1961) breviaornis, O. nana, O. simpZex
Ivory Coast CaZanoides aarinatus, EucaZanus Temora st;yUfera, Onaaea media,
Binet et al. (1972
monaahus, CtenoaaZanus vanus, Coryaaeus, ParaaaZanus parvus,
a-c)
Agetus Zimbatus, Centropages Oithona pZumifera, NannoaaZanus
Seguin (1973) ahierahiae, Temora turoinata, minor, Coryaaeus speaiosus,
T. st;yZifera, Onaaea aurta, NeoaaZanus. UndinuZa vuZgaris,
L.uaiautia MaaroseteZZa graaiZis
Nigeria CaZanoides aarinatus, Para- Temora turbinata, Coryaaeus
Bainbridge (1960)
aaZanus parvus, Temora st;yU- giesbreahti, Onaaea venusta,
Longhurst and
fera, EucaZanus piZeatus, E. Oithona pZumifera. CZausoaaZanus
Bainbridge (1964)
monaahus, Onaaea venusta, furaatus
Euahae ta marina
Gabon CaZanoides aarinatus, Onaaea UndinuZa vuZgaris, EucaZanus pi-
Binet et al.
mediterranea major, EucaZa- Zeatus, E. subtenuis, ParaaaZa-
(1972a-c)
nus monaahus, E. arassus, nus aauZeatus. CZausoaaZanus
Coryaaeus afriaanus, Saphirina furaatus, Euchaeta marina,
nigromaauZata Centropages furaatus
Angola CaZanoides aarinatus Euterpina aautifrons, Oithona
Neto and De Paiva nana, Temora tubinata, Para-
(1966)
aaZanus parvus
S.W. Africa CaZanoides aarinatus, Metridia ParaaaZanus parvus. P. arassi-
Kollmer (1963)
Zucens, Centropages braahiatus, rostris, Oithona simi Us, Nan-
RhinaaZanus nasutus, EucaZanus noaaZanus minor, Aetidius
eZongatus armatus
second a gradient from inshore stations at the end of the cold season
to the offshore stations. At the offshore station of Abidjan (Ibanez
and Seguin, 1972~ Seguin, 1973) the results were not as clear, and
only CaZanoides aarinatus was well isolated. The comparison between the
mathematical analysis of the zooplankton found off the Pointe Noire
50
area and Abidjan area clearly demonstrates that the Pointe Noire region
is not a real upwelling area.
The copepods of Mauritania and the Spanish Sahara area are known from
the works of Binet (1973) and Vives (1974, 1975), and by recent experi-
mental studies of the CINECA program (Cooperative Investigations of
the Northern part of the Eastern Central Atlantic) which will be con-
sidered in the last paragraph.
Calanoides carinatus was the most characteristic copepod of the cold rich
upwelled waters from area South Africa to the Rio de Oro. Bainbridge
(1972) thinks this species plays the same role in the African ecosys-
tem as Calanus finmarchicus in the North Atlantic or Calanoides acutus in
the South Atlantic. This species is large, herbivorous, and is able
to store lipids; it appears in the inshore waters during maximum prim-
ary production. In the warm and stratified waters it is a meso- or
bathypelagic species (Vives, 1975), and the major part of the popula-
tion is at copepodite stage V (Bainbridge, 1960; Binet and Suisse De
Sainte Claire, 1975). Longhurst (1967) has made the same observations
for Calanus helgolandicus off Baja California. Bainbridge (1972) thinks
that the blooms of big diatoms in upwelling areas are better for the
development of C. carinatus than for the other herbivorous copepods which
have filtering apparatus more suitable for smaller particles - especial-
ly the tropical and subtropical species which are generally small
forms. Binet and Suisse De Sainte Claire (1975) have elaborated the
relationship between C. carinatus and the temperature in the inshore
waters of the Ivory Coast. They have found a correlation between the
number of individuals and the water temperature of the preceding fort-
night, also a correlation between the temperature and the mean cephalo-
thoracic length. Vives (1975) has shown well the correlation between
the depth of the maximum of C. carinatus and the depth of the cold up-
welled waters.
Among the most common genera are ParacaZanus (P. parvus, P. crassirostris),
Temora (T. styZifera, T. turbinata), EucaZanus (E. subtenuis, E. piZeatus, E. mona-
chus), Oithona (0. nana), Centropages (C. chierchiae, C. typicus), Acartia (A. danae,
A. cZausi), Oncaea (0. venusta) and NannocaZanus (N. minor).
5. Experimental Studies in the African Atlantic
5.1 Biological Observations Following a Drogue
During a cruise of the R.V. Capricorne in March to April 1972, a drogue

was followed for nine days from the center of cold upwelled water
(Herbland et al., 1973). Primary and secondary productions and regen-
eration of nutrients were measured. The secondary production was eval-
uated by the variations of biomass (dry weight). The time between the
phytoplankton bloom and the maximum of zooplankton abundance was only
one day; after that a synchronism was observed between variations of
chlorophyll a and zooplankton biomass. The simultaneous decreases of
phytoplankton and zooplankton was explained by predation: grazing by
copepods on phytoplankton, predation by anchovy on copepods. The daily
rate of increase of the zooplankton biomass was estimated to be 28%.
In fact this experiment poses many questions, especially methodological,
because such a short time between phytoplankton and zooplankton maximum
cannot be attributed only to secondary production. Oxygen consumption
and the excretion of phosphorus and different forms of nitrogen have
been measured by experiments, and the production: biomass ratio has
been calculated from the carbon equivalent of the respiration and the
S1
dry weight. This ratio, which is a productivity ratio, has a maximum

value of 48%, which corresponds to the optimal phase of the growth
of crustaceans in the laboratory. At the beginning of the experiments
grazing was estimated to be 6% of the primary production and to be 260%
at the end. The excretion of the mesozooplankton is important and cor-
responds to 40% of the production of ammonium (Le Borgne and Binet,
1974) .
The progression of copepod populations during the same experiment

(Binet, 1973) showed typical variations of the specific diversity index
and of the percentages of carnivores and herbivores. The author called
herbi vores: Calanoides carinatus, Paracalanus parvus, and Ctenocalanus vanus;
carn i vore s : Oi thana nana and Oncaea curta; omnivore s: Cen tropages chi erchi ae .
The feeding patterns of the other copepods were not specified. The
number of species (62) is high in the upwelled water at the beginning,
but it decreases as the total biomass increases (especially the number
of herbivores), and finally we observe a new increase in the number
of species by new carnivores and omnivores entering the system.
S.2 Respiration and Excretion of the Zooplankton in the Mauritania

and Rio de Oro Area
Le Borgne (1973) studied the relationships between respiration and ex-

cretion of a zooplankton population, excluding the carnivores. The
ratios between the different products of excretion gave indications
of the nature of the oxidized substance, and of the rate of nitrogen
and phosphorus assimilation. The zooplankton was sampled in three dif-
ferent areas: in cold upwelled water, in the maximum of the phytoplank-
ton bloom and in the maximum of zooplankton biomass. The ratios in
these three zones were not significantly different. The proportion of
nitrogen and phosphorus excreted as organic compounds was about 50%.
~~ese products correspond to a lipid-carbohydrate catabolism, which is
the mark of well-fed animals. The author discusses the differences
between these results and previous ones, and thinks the most important
reason for the differences is methodological.
Smith and Whitledge (1977) also measured the biomass, nitrogen excre-
tion and oxygen consumption north of Cap Blanc during the cruise Joint
I (March-April 1974). The authors distinguished four size categories
by sieves (in: Blackburn, 1975). Animals larger than 1 mm were dominant
(SO to 80% of dry weight) off the slope and animals between 102 ~ and
SOS ~ were the most abundant (62 to 88%) in the inshore area. Near the
coast the cyclopoid and harpacticoid copepods were dominant (Euterpina,
Oithana, Oncaea and Corycaeus), further offshore they found chiefly cala-
noids (Acartia, Centropages, Paracalanus, Lucicutia and numerous copepodi tes) •
The excretion decreased as the size of animals increased. In the coastal
area it was the excretion by the smallest animals which was the most
important (0.107 ~g-at NH4-N mg dry weight- 1 h- 1 or a total regenera-
tion of 7.S mg-at NH 4 -N m- 2 day-l in the coastal area and 4.1mg-at in the
oceanic area). This quantity corresponds to about SO% of the primary
production needs, which is similar to the results of other studies of
upwelling in the Pacific Ocean (Whitledge and Packard, 1971; Whitledge,
1 9 72; Jawed, 1 973) .
The importance of zooplankton excretion to primary production has been

known for several years and the first results were summarized by Ketchum
(1961). The mineral parts are immediately assimilated by phytoplankton,
and the appropriate mechanisms may be very fast. High nutrient concen-
trations due to zooplankton excretion have been found in several African
Atlantic areas: S.W. Africa (Calvert and Price, 1971), Mauritania
(Herbland et al., 1973), and Morocco (Grall et al., 1974).
52
5.3 Respiration and Excretion of Zooplankton in the Morocco Area
There are two studies in this area, Champalbert and Gaudy (1972) and
Nival et al. (1974), the first before the upwelling season (in January
to February), and the second in. July during the period of maximum pro-
duction.
Champalbert and Gaudy (1972) chose copepods of different bathymetric

levels: hyponeuston (Anomal-ocera patersoni, Pontel-l-opsis viUosa, P. regaUs,
Lahidocera woUastoni and PonteUa l-o biancoi); epiplankton (Cenhoopages typicus,
Temora styUfera, Cal-anus hel-gol-andicus and Acartia cZausi); meso- and bathy-
plankton (Pl-euromamna xiphias, P. ahdominaUs, Euohaeta acuta and Undeuohaeta
pl-umosa). Examination of the metabolism curves in relation to temper-
ature gives some indication of different patterns of adaptation ac-
cording to geographical distribution, vertical migration, and feeding
pattern. In particular the study of the QIO shows the poor adaptation
to cold waters of some species from the hyponeuston. Euohaeta acuta
yielded highly variable results in concordance with its carnivorous
feeding pattern, and Pl-euromamma xiphias had a relatively constant QIO'
which suggests its adaptation to vertical migration. With this method
it is possible to determine the optimal thermal conditions for each
species in this area, and to estimate the possibilities for each spe-
cies to increase or not in cold upwelled waters. These authors observed
a high correlation between the respiration rate and the weight of an-
imals. The temperature 18 0 c appears to be optimal for this area.
Nival et al. (1974) studied the morphological characteristics (mean

size of Temora styUfera according to trophic conditions and geographical
location) and the biochemistry and metabolism of different species.
Experiments on respiration showed an increase of the respiratory rate
between 13 0 and 20 0 C with a QIO moving from 1.8 to 5.3. Above 20 0 C a
regulation of the respiratory rate was observed for Acartia cl-ausi and
Centropages typicus, but many other species showed a decrease. The effect
of starvation has been studied for Temora styUfera. Respiration and ni-
trogen excretion decreased in the first days, the dry weight decreased
a few days after; the ratio O:N, constant at the beginning of the ex-
periment, decreased after the 4th day, and by the 7th day the copepods
used only their proteins for survival. The study of the metabolism of
Cal-anus hel-gol-andicus sheds some light on the energetic aspects of vertical
migration. It is possible to show that there is an optimal length of
time needed in the superficial layer for the animal to fulfill its
energetic requirements. This time depends on the food concentration
at the surface, the filtration rate, the decrease of its ingestion
rate over a period of time, aqd also the temperature difference between
the surface and the deepest level reached by the animal during the
day.
5.4 Measurement of the Respiratory Activity by Study of the E.T.S.
Knowledge of oxygen consumption rate is essential if one is to under-

stand secondary production and the transfer of energy by zooplankton.
Packard (1971) uses the measurement of activity of those intracellular
enzymes which control respiration, that is to say the Electron Trans-
port System (E.T.S.). This technique presents many practical advantages;
principally, it avoids the experimental difficulties of measuring and
interpreting the respiration rate of animals in a plankton sample.
This method has been used in different upwelling areas: Peru, Baja
California, Costa Rica Dome, and N.W. Africa. The pressure effect (King
and Packard, 1975) and the temperature effect (Packard et al., 1975a)
have been studied and comparisons between enzyme activity and oxygen
consumption have been made on several species (Owens and King, 1975).
53
Pressure has no influence on this enzyme activity; on the contrary,

temperature has an important influence, but the animals possess some
means of compensation for the decrease of metabolic activities at low
temperatures. In poikilotherms these mechanisms are principally bio-
chemical changes, especially involving the concentrations of active
enzymes. Packard et al. (1975a) have studied the E.T.S. activity of
zooplankton groups in the N.E. Pacific at various temperatures between
20 and 27 0 C.
The different studies on the correlation between the measure of E.T.S.
activity and oxygen consumption (King and Packard, 1975; OWens and
King, 1975) indicate that the E.T.S. is a good estimation of the max-
imum potentialities of biological oxygen consumption. This potentiality
may be more than 100% of the oxygen consumption measured. The ratio of
respiration rate: E.T.S. activity has been calculated for CaZanoides
aarinatus near Cap Blanc (P ackard e t al., 1974), and has been used to
estimate the total consumption by the zooplankton in this area. In the
first 50 m the authors estimated rates between 146 and 412 ~l 02 h- 1
m- 3 in the Mauritania area, and on the same cruise (Cineca-Charcot II,
February-March 1971) in the Morocco area during the nonupwelling season
they found rates between 30 and 250 ~l 02 h- 1 m- 3 • The curves clearly
show decreasing values with depth. During the cruise Joint I (Packard
et al., 1975b) near Cap Blanc in 1974, the maximum of the E.T.S. ac-
tivity corresponds to the class 200-500 ~, as does the maximum of ex-
cretion (Smith and Whitledge, 1977).
The authors compared their E.T.S. results with measurements from dif-
ferent geographical areas. The highest values were found in the Baja
California area and the lowest in the Costa Rica Dome area.
5.5 Zooplankton Feeding Estimated by Measurement of the Digestive

Enzyme Activities
Boucher and Samain (1974) have proposed the use of amylase activity
of zooplankton (total sample or isolated species) as an index of feed-
ing. This assumption is verified by the study of the relationships be-
tween enzyme activity and trophic behavior. The feeding pattern, eco-
logical conditions, and starvation influence the specific activity of
amylase. The correlation between amylase activity and biomass of phyto-
plankton is most significant. This relationship is best observed during
the night, and it is clearer in the superficial layer (0-100 m) than
in the deeper layers. This fact suggests a daily rhythm of feeding be-
havior. Boucher and Samain (1974) studied the rhythm in the Moroccan
upwelling area. The samples were taken at three depths, once every 4 h
during three days at different stations. The increase of the amylase
specific activity was synchronous at the three depths. This rhythm,
however, did not correspond to vertical migration. The variations of
the qualitative composition of the population, and the regulation of
the digestive enzyme activities in relation to the quantity of avail-
able food explain part of this rhythm (Boucher and Samain, 1975). Using
the method described (Samain and Boucher, 1974) it is possible to mea-
sure the activity of an individual Arromia. Boucher and Samain (1975)
extended this study during a cruise (Cineca-Charcot V, April 1974) to
the upwelling area near Cap Blanc,. where they found a significant
correlation between the amylase specific activity and the grazing index
of Lorenzen.
The specific activity of proteases has also been measured. The most im-
portant species appear to have omnivorous feeding patterns, but the
Cladocera have carnivorous tendencies (Podon intermedius); the copepod
Temora styUfera seems to be herbivorous. Comparison of the digestive
54
enzyme activity between Temora stylifera and CaZa:noidEs carinatus shows

some differences which can be explained by examining their feeding
mechanisms .
6. Conclusions
This review of the principal groups of zooplankton in upwelling condi-

tions, their biological characteristics and their behavior, illustrates
the results that come from the use of different research methods.
The qualitative analysis of the populations remains a very important

step in the understanding of an ecosystem. In my opinion the biological,
ecological, ethological, and physiological characteristics of the dif-
ferent species are too little known to allow one to consider the whole
zooplankton populations without a corresponding qualitative study.
Moreover for a specific population it is important to know the sizes
of individuals, the sex ratio, the stage of life cycle, and the bio-
chemical composition.
In the case 'of quantitative studies, i t is necessary to make an effort

toward the adoption of sampling equipment and procedures that take
into account the behavior (and especially the avoidance of nets) of
different groups.
Another essential aspect of the animals' characteristics concerns the

feeding behavior of the principal species: size of the prey, rhythms,
influence of environmental conditions, and interspecific competitions.
It is obvious that the herbivorous tendencies of some zooplankton fa-

cilitate, as a first step, the general increase of the biomass, but
the place of the microphageous groups in this scheme is not well known.
We are at the very beginning of our efforts to establish new ways of

estimating secondary production and productivity. The transfer of energy
inside the marine food chains, the differences in the values from dif-
ferent areas, and the real quantitative consequences of the enrichment
from upwelling are still very poorly understood.
The Crustacea euphausiids and particularly the copepods are the most
important groups which react to the upwelling conditions, but they are
not the only ones. Within these two groups, the remarkable increase
in the numbers of zooplankton derives from increased abundance of only
a few species. For instance in the African Atlantic we have especially
Euphausia Zucens, E. k:t>ohnii, and Nyctiphanes capensis from the euphausiids;
the genus EucaZa:nus, Temora, Oncaea, Centropages, and Acartia, and principally
the species CaZanoidEs carinatus from the copepods. In other groups some
species such as DolioZum nationalis, OikopZeum dioica, and o. Zongicauda, SaZpa
fusiformis, and Thalia dEmocratica are capable of high production rates in
upwelling conditions.
It is difficult to define the specific composition of an upwelling eco-

system. Upwelling is a hydrological and ecological event which modifies
a previous regime. The qualitative consequences will not be the same
if we speak about an oceanic area (dome, equatorial divergence) and
about a coastal upwelling. In all zones the resulting populations are
dependent on the biogeographical associations of the area. In coastal
conditions i t is also difficult to make real boundaries between benthos,
plankton and nekton, and in these conditions "upwelling ecosystem" is
not a very correct concept.
55
However, we can state some very simple rules about the variations of
the diversity or of the percentage of herbivorous forms during the
evolution of upwelled waters. All new elements on the biology, ethology
or physiology of the principal species will increase our understanding
of the zooplankton dynamics of these productive areas.
Research on metabolism seems at present the best way to distinguish
species which have very close biological or ecological characteristics,
but which do not respond with the same intensity and the same speed
to the improvement of trophic conditions and to the variations of
hydrological factors.
Acknow~edgments. The author wishes to thank Dr. Boyd for his help in this transla-
tion. Thanks are also due to Drs. Blackburn, Packard, Vives and Whitledge for sending
their papers in press and those of their associates, and to Drs. Boucher, Dallot
and Vives for helping in the preparation of the respective parts: digestive enzymes,
chaetognaths, and copepods.
References
Allen, G.H.: Notes on sampling Amphipods and Euphausiids in the north Pacific Ocean.
Trans. Am. Fish. Soc. 101 (4), 577-582 (1972)
Alton, M.S., Blackburn, C.J.: Diel changes in the vertical distribution of Euphau-
siids Thysanoessa spinifera and Euphausia pacifica in coastal waters of Washington.
Calif. Fish Game 58 (3), 179-190 (1972)
Alvarino, A.: Los Quetognatos del Atlantico, distribucion y notas esenciales de
sistematica. Trabhs Inst. esp. Oceanogr. 37, 290 pp (1959)
Andreu-Puyal, 'P.: Contribucion al estudio de las Eufausiaceos de los alrededores de
cabo Blanco (N.W. de Africa). Res. Exp. Cient. B/O Cornide 5, 77-111 (1976)
Andreu-Puyal, P., Santamaria, G., Trepat, I.: Distribution de-quelques groupes de
zooplancton (Euphausiaces, Pteropodes-Heteropodes, Salpides, Doliolides et Apen-
diculaires) dans Ie N.O. africain. Communication, Cons. perm. Int. Explor. Mer
CM/L 16 (unpublished manuscript, 1975)
Bainbridge, V.: Occurrence of Ca~anoides carinatus in the plankton of the Gulf of
Guinea. Nature (Lond.) 188 (4754), 932-933 (1960)
Bainbridge, V.: The zooplankton of the Gulf of Guinea. Bull. Mar. Ecol. 8, 61-97
(1972) -
Baker, A. de C.: The vertical distribution of euphausiids near Fuerteventura, Canary
Islands ("Discovery" SOND Cruise, 1965). J. Mar. BioI. Ass. U.K. 50 (2), 301-342
(1970)
Barham, F.G.: Siphonophores and the deep scattering layer. Science (N.Y.) 140,
826-828 (1963)
Bhavanarayana, P. V., Ganapati, P.N.: Species groups among pelagic Tunicates in the
western part of the Bay of Bengal. Mar. BioI. 11 (2), 173-177 (1971)
Bieri, R.: The distribution of the planktonic Chaetognatha in the Pacific and their
relationship to the water masses. Limnol. Oceanogr. 4 (1), 1-28 (1959)
Binet, D.: Aperyu sur les variations saisonnieres du zooplancton et plus particuli-
erement des Copepodes du plateau continental de Pointe Noire (Congo). Document
scientifique Pointe Noire, ORSTOM 8, 107 pp (1970)
Binet, D.: Note sur l'evolution des populations de Copepodes pelagiques de l'upwel-
ling mauritanien (mars-avril 1972). Document Sci. Abidjan, ORSTOM! (1), 77-90
( 1973)
Binet, D., Dessier, A.: Primieres donnees sur les Copepodes pelagiques de la region
congolaise. I. Liste des especes et notes ecologiques. Cah. ORSTOM Serie Oceanogr.
2. (4), 411-457 (1971)
Binet, D., Dessier, A.: Premieres donnees sur les Copepodes pelagiques de la region
congolaise. III. Diversite specifique. Cah. ORSTOM Serie Oceanogr. 10 (3), 243-250
(1972)
56
Binet, D., Dessier, A., Gaborit-Rezzouk, M., Roux, M.: Premieres donnees sur les
Copepodes pelagiques de la region congolaise. II. Analyse des correspondances.
Cah. ORSTOM Serie Oceanogr. 10 (2), 125-137 (1972a)
Binet, D., Gaborit-Rezzouk, M., Roux, M.: Copepodes pelagiques du plateau ivoirien.
Utilisation de l'analyse des correspondances dans l'etude des variations saison-
nieres. Document scientifique Abidjan, ORSTOM 3 (1), 47-79 (1972b)
Binet, D., Suisse De Sainte Claire, E.: Le copepO"de planctonique Calanoides carinatus.
Repartition et cycle biologique au large de la Cote d'Ivoire. Cah. ORSTOM Serie
Oceanogr. 11 (1), 15-30 (1975)
Blackburn, M.: Distribution of zooplankton biomass by sizes and taxa during Joint I
expedition. Preview of preliminary scientific results. Document CUEA 26,1-26,9
(1975)
Boden, B.P.: The euphausiid crustaceans of the Southern African waters. Trans. R. Soc.
S. Afr. 34 (1), 181-243 (1954)
Boden, B.P.: Euphausiacea of the Benguela current first survey RRS William Scoresby
March 1950. Discovery Rep.~, 337-376 (1955)
Boden, B.P.: Euphausiacea (Crustacea) from tropical west Africa. Atlantide Rep. ~,
251-262 (1961)
Boucher, J., Samain, J.F.: L'activite amylasique indice de la nutrition du zooplanc-
ton: mise en evidence d'un rythme quotidien en zone d'upwelling. Tethys ~ (1-2),
179-188 (1974)
Boucher, J., Samain, J.F.: Etude de la nutrition du zooplancton en zone d' upwelling
par la mesure des activites enzymatiques digestives. In: 9th Eur. Mar. Bioi. Symp.
Barnes, H. (ed.). Aberdeen: Aberdeen Univ. Press, 1975, pp. 329-341
Braconnot, J.C.: Contribution a l'etude biologique et ecologique des Tuniciers pela-
giques Salpides et Doliolides. I - Hydrologie et ecologie des Salpides. Vie Milieu
22 (2 B), 257-286 (1971)
Braconnot, J.C., Casanova, J.P.: Sur le tunicier pelagique Doliolum nationalis en
Mediterranee occidentale (campagne du P.T.T. sept. oct. 1958). Revue Trav. Inst.
(scient. tech.) pech. Marit. 11 (4), 393-402 (1967)
Brinton, E.: Variable factors affecting the apparent range and estimated concentra-
tion of Euphausiids in the North Pacific. Pacif. Sci. 16 (4), 374-408 (1962)
Brinton, E., Gopalakrishnan, K.: The distribution of Indian Ocean Euphausiids. In:
Biology of the Indian Ocean. Zeitzschel, B. (ed.). Berlin-Heidelberg-New York:
Springer, 1973, pp. 357-382
Calvert, S.E., Price, N.B.: Upwelling and nutrient regeneration in the Benguela cur-
rent, October 1968. Deep Sea Res. ~, 505-523 (1971)
Casanova, J.P.: Biomasse et distribution du zooplancton dans l'upwelling mauritanien
(Resume). Tethys 6 (1-2), 320 (1974a)
Casanova, J.P.: Rep-artition quantitative et composition du zooplancton dans les "eaux
guineennes et tropicales" du N .0. africain (compagne Cineca aout 1973). Communica-
tion, Cons. perm. into Explor. Mer CMIL 3 (unpublished manuscript, 1974b)
Casanova-Soulier, B.: Les Euphausiaces de Mediterranee (systematique et developpe-
ment larvaire. Biogeographie et biologie). These Doc. Sci. Univ. Aix-Marseille,
380 pp. mimeo (1974)
Cervignon, F.: Descripcion y consideraciones sobre los Sifonoforos de las costas
occidentales de Africa recogidos en las campanas del "Costa Canarias". Investig.
Pesq. 18, 8-31 (1961)
Champalbert, G., Gaudy, R.: Etude de la respiration chez les Copepodes de niveaux
bathymetriques varies dans la region sud marocaine et canarienne. Mar. Biol. ~
(2), 159-169 (1972)
Cram, D.L., Schulein, F.H.: Observations on surface shoaling cape hake off South
West Africa. J. Cons. Perm. Int. Explor. Mer 35 (3), 272-275 (1974)
Day, D.: Macrozooplankton and small nekton in the coastal waters off Vancouver Is-
land (Canada) and Washington, Spring and Fall of 1963. Spec. scient. Rep. Nat.
Mar. Fish. Serv., Fisheries 619, 1-94 (1971)
Decker, A. De: Agulhas Bank plankton. In: Biology of the Indian Ocean. Zeitzschel,
B. (ed.). Berlin-Heidelberg-New York: Springer, 1973, pp. 189-219
Dragovich, A., Potthoff, T.: Comparative study of food of skipjack and yellowfin
tunas off the coast of west Africa. Fishery Bull. Fish. Wildl. Servo U.S. 70 (4),
1087-1110 (1972)
57
Ducret, F.: Chaetognathes des campagnes de l'Ombango dans la zone equatoriale afri-
caine (1959-1960). Bull. Inst. fro Afr. noire 24 Ser. A 2, 331-353 (1962)
Ducret, F.: Chaetognathes des campagnes de l'Ombango dans les eaux equatoriales et
tropicales africaines. Cah. ORSTOM Serie Oceanogr. ~ (1), 95-141 (1968)
Fowler, S.W., Small, L.F.: Sinking rates of Euphausiid fecal pellets. Limnol. Oce-
anogr. 17 (2), 293-296 (1972)
Fowler, S.W., Small, L.F., Keckes, S.: Effects of temperature and size on molting
of Euphausiid Crustaceans. Mar. Biol. l! (1), 45-51 (1971)
Fraser, J.H.: The oceanic and bathypelagic plankton of the north east Atlantic and
its possible significance to fisheries. J. Mar. Res. ~, 1-48 (1961)
Fraser, J.H.: The role of Ctenophora and Salps in zooplankton production and standing
crop. Rapp. P.-v. Reun. Cons. Perm. Int. Explor. Mer 153, 121-123 (1962)
Fraser, J.H.: Experimental feeding of some Medusae and Chaetognaths. J. Fish. Res.
Bd Can. ~ (7), 1743-1762 (1969)
Furnestin, M.L.: Chaetognathes et zooplancton du secteur atlantique marocain. Revue
Trav. Inst. (scient. tech.) pech. Marit. 21 (1-2), 1-356 (1957)
Furnestin, M.L.: Indicateurs planctoniques dans la baie Iberomarocaine. Revue Trav.
Inst. (scient. tech.) pech. Marit. 28 (3), 257-264 (1964)
Furnestin, M.L.: Chaetognathes des eaux africaines. Atlantide Rep. 9, 105-135 (1966)
Gaudy, R., Seguin, G.: Note sur ,la repartition annuelle des Copepodes pelagiques
des eaux de Dakar. Recl Trav. Stn Mar. Endoume 34 (50), 211-217 (1964)
Gilfillan, E.: Reactions of Euphausia pacifica from oceanic, mixed oceanic-coastal
waters of British Columbia to experimental changes in temperature and salinity.
J. Exp. Mar. Biol. Ecol. 10, 29-40 (1972a)
Gilfillan, E.: Seasonal and:latitudinal effects on the responses of Euphausia paci-
fica to experimental changes of temperature. In: Biology and Oceanography of the
Northern North Pacific Ocean. Takenouti, A.Y. (ed.). Tokyo: Idemitsushoten, 1972b,
pp. 443-463
Godeaux, J.: Tuniciers pelagiques. Result. Sci. Exped. oceanogr. belg. Atlant. S 1
(7),3-32 (1962)
Godeaux, J., Goffinet, G.: Donnees sur la faune pelagique vivant au large des cotes
du Gabon, du Congo et de l'Angola. Annls Soc. R. Zool. Belg. 98 (1), 49-86 (1968)
Gopalakrishnan, K.: Zoogeography of the genus Nematoscelis. Fish. Bull. Fish. Wildl.
Servo U.S. 72 (4), 1039-1074 (1974)
Gcy, J.: Note sur les Hydromeduses dans les eaux tropicales et subtropicales. Bull.
Mus. Natn. Hist. Nat., Paris, Ecologie generale 11, 333-343 (1974)
Goy, J., Thiriot, A.: Distribution d'esp€ces planctoniques carnivores dans la region
de l'Atlantique marocain. Resultats des campagnes Cineca-Charcot I, II et III.
Tethys ~ (1-2), 291-302 (1974)
Grall, J.R., Laborde, P., Le Corre, P., Neveux, J., Treguer, P., Thiriot, A.: Char-
acteristiques trophiques et production planctonique dans la region sud de l' Atlan-
tique marocain. Resultats des campagnes Cineca-Charcot I et II. Tethys ~ (1-2),
11-28 (1974)
Greze, V.N.: The biomass and production of different trophic levels in the pelagic
communities of south seas. In: Marine Food Chains. Steele, J.H. (ed.). 1970, pp.
458-467
Greze, V.N., Gordejana, K.T., Shmeleva, A.H.: Distribution of zooplankton and bio-
logical structure in the tropical Atlantic. In: Proc. Symp. Oceanography Fisheries
Resources Tropical Atlantic. Paris: UNESCO, 1969, pp. 85-90
Gueredra t, J.A., Grandperrin, 'R., Roger, C.: Di versi te specifique dans le Pacifique
equatorial. Evolution de l'ecosysteme. Cah. ORSTOM Serie Oceanogr. ~ (1), 57-69
(1972)
Haq, S.M., Ali Khan, J., Chugta, S.: The distribution and abundance of zooplankton
along the coast of Pakistan during postmonsoon and premonsoon periods. In: Biology
of the Indian Ocean. Zeitzschel, B. (ed.). Berlin-Heidelberg-New York: Springer,
1973, pp. 257-272
Hart, T.J., Currie, R.I.: The Benguela current. Discovery Rep. 31, 123-298 (1960)
Herbland, A., Le Borgne, R., Voituriez, B.: Production primaire-;-secondaire et re-
generation des sels nutritifs dans l'upwelling de Mauritanie. Document scientifique
Abidjan, ORSTOM! (1), 1-75 (1973)
Heron, A.C.: Population ecology of a colonizing species the pelagic Tunicate Thalia
democratica. I - Individual growth rate and generation time. Oecologia 10 (4),
269-293 (1972a)
58
Heron, A.C.: Population ecology of a colonizing species the pelagic Tunicate ThaZia
democratiaa. II - Population growth rate. Oecologia 10 (4), 294-312 (1972b)
Heydorn, A.E.F.: The Chaetognatha off the west coast of the Union of South Africa.
Invest. Rep. Div. Fish. Un. S. Afr. 36, 1-56 (1959)
Hopkins, T.L.: Carbon and nitrogen content of fresh and preserved NematosceZis dif-
ficiZis,a Euphausiid Crustacean. J. Cons. Perm. Int. Explor. Mer ~ 300-304
(1968)
Hubbard, L.T., Pearcy, W.G.: Geographic distribution and relative abundance of Sal-
pidae off the Oregon coast. J. Fish. Res. Bd Can. 28 (12), 1831-1836 (1971)
Ibanez, F., Seguin, G.: Etude du cycle annuel du zooplancton d'Abidjan. Comparaison
de plusieurs methodes d'analyse multivariable: composantes principales, correspon-
dances, coordonnees principales. Investig. Pesq. 36 (1), 81-108 (1972)
Jawed, M.: Body nitrogen and nitrogenous excretion in Neomysis rayii and Euphausia
pacifica. Limnol. Oceanogr. 1i (5), 748-754 (1969)
Jawed, M.: Ammonia excretion by zooplankton and its significance to primary produc-
tion during summer. Mar. Bioi. 23, 115-120 (1973)
Jerde, C.W.: A comparison of Euphausiid shrimp collections made with a micronekton
net and a one me.ter plankton net. Pacif. Sci. 21 (2), 178-181 (1967)
Jerde, C.W., Lasker, R.: Molting of Euphausiid shrimps. Shipboard observations.
Limnol. Oceanogr. ~ (1), 120-124 (1966)
Ketchum, B.H.: Regeneration of nutrients by zooplankton. Rapp. P .-v. Cons. Perm.
Int. Explor. Mer. 153, 142-147 (1961)
King, F.D., Packard, T.T.: The effect of hydrostatic pressure on respiratory electron
transport system activity in marine zooplankton. Deep Sea Res. E, 99-105 (1975)
King, F.D., Packard, T.T.: Respiration and the activity of the respiratory electron
transport system in marine zooplankton. Limnol. Oceanogr. 20, 5, 849-854 (1975)
Kinzer, J.: Quantitative distribution of zooplankton in surface waters of the Gulf
of Guinea during August and September 1963. Proc. Symp. Oceanogr. Fish. Res. Trop.
Atlantic-Results ICITA and GTS (Abidjan 1966). Paris: UNESCO, 1969, pp. 231-240
Kollmer, W.E.: Notes on zooplankton and phytoplankton collections made off Walvis
Bay. Investl Rep. Mar. Res. Lab. S.W. Afr. 8, 1-78 (1963)
Komaki, Y.: On the surface swarming of Eupha~iid Crustaceans. Pacif. Sci. 21 (4),
433-448 (1967)
Kramp, P.L.: The Medusae of the tropical west coast of Africa. Atlantide Rep. l'
242-324 (1955)
Lasker, R.: Molting frequency of a deep-sea Crustacean Euphausia pacifica. Nature
(Lond.) 203, 96 (1964)
Lasker, R.~eeding, growth, respiration and carbon utilization of a Euphausiid
Crustacean. J. Fish. Res. Bd Can. ~ (9), 1291-1317 (1966)
Le Borgne, R.: Etude de la respiration et de l'excretion d'azote et de phosphore
des populations zooplanctoniques de l'upwelling mauritanien (mars-avril 1972).
Mar. Bioi. 19, 249-257 (1973)
Le Borgne, R., Binet, D.: Biomasse, production et excretion du zooplancton de l'up-
welling de Mauritanie, evolution des populations de Copepodes. Tethys 6 (1-2),
321-322 (1974) -
Le Roux, A.: Observations sur le developpement larvaire de Nyctiphanes ooWJhii au
laboratoire. Mar. Bioi. 22, 159-166 (1973)
Longhurst, A.R.: Diversity and trophic structure of zooplankton communities in the
California current. Deep Sea Res. 14, 51-63 (1967)
Longhurst, A.R., Bainbridge, V.: The coastal oceanography of western Nigeria with
an appendix on zooplankton. Bull. Inst. fro Afr. noire 26 Ser. A 2, 337-402 (1964)
Margalef, R.: Perspectives in Ecological Theory. Chicago-London: Univ. of Chicago
Press, 1968
Margulis, R.Ya.: Distribution of Siphonophores of the genus Lensia in the Atlantic.
Oceanology 11 (1), 80-84 (1971)
Marques, E.: Copepodes marinhos de Angola. Trabhs Miss. Bioi. marit. ~, 87-126
(1953)
Marques, E.: Copepodes da Guine portuguesa (coligidos pela missao geohidrographica
da Guine). Ans Junta Investig. Ultramar 10 (4), 23-47 (1955)
Marques, E.: Copepodes dos mares de Angola. II - Ciclopoida e Harpacticoida. Trabhs
Miss. Biol. marit. 20, 131-150. (1957)
Marques, E.: Sobre Copepodes marinhos de Angola. Trabhs Miss. Bioi. marit. 24, 197-
222 (1958)
59
Marques, E.: Copepodes da Guine Portuguesa. III - Contribuicao para 0 seu conheci-
mento. Mem. Junta Investig. Ultramar ~, 41-57 (1961)
Machline, J.: Feeding appendages of the Euphausiacea. J. Zool. 153, 1-43 (1967)
Machline, J., Fisher, L.R.: The biology of Euphausiids. Adv. Mar. Biol. 2,1-454
(1969)
McLaren, I.A.: Effects of temperature on growth of zooplankton and the adaptative
value of vertical migration. J. Fish. Res. Bd Can. 20, 685-727 (1963)
Meira, C.: Contribuiyao para 0 estudo dos Eufausiaceos do arquipelago de Cabo Verde.
Notas Cent. BioI. Aquat. Trop. 12, 1-25 (1970)
Mensah, M.A.: Zooplankton occurrence over the shelf of Ghana. In: Proc. Symp. Oce-
anography Fisheries Resources Tropical Atlantic. Paris: UNESCO, 1969, pp. 241-254
Nemoto, T.: Chlorophyll pigments in the stomach of Euphausiids. J. oceanogr. Soc.
Japan 24 (5), 253-260 (1968)
Nemoto, T.: History of research into the food and feeding of Euphausiids. Proc. R.
Soc. Edinb. 73, 259-265 (1971/72)
Nemoto, T., Kamada, K., Hara, K.:-Fecundity of a Euphausiid Crustacean NematosceZis
difficiZis in the North Pacific Ocean. Mar. BioI. 14 (1),41-47 (1972)
Nepgen, C.S.: The Euphausiids of the wesb coast of South Africa. Investl Rep. Div.
Fish. Un. S. Afr. 11, 1-36 (1957)
Neto, T.S.: Quetognatas dos mares de Angola. Trabhs Cent. Biol. Pisco 31, 1-60 (1961)
Neto, T.S., Lourenco, L.: Sifonoforos Calicoforos do arquipelago de Cabo Verde. Notas
Cent. BioI. Aquat. Trop. 33, 1-55 (1973)
Neto, T.S., Paiva, I. De: CicIo annual do zooplancton colhido na Baia Farta em 1960.
Notas mimeogr. Cent. BioI. aquat. trop. 2, 1-61 (1966)
N.I.O. Anonymous: RRS Discovery cruise 21 report January-April 1968. Upwelling off
N.W. Africa and plankton distribution 11 0 N 20o W. N.I.O. Cruise Rep. Ser. Cr ll,
30 pp. mimeo. (unpublished manuscript, 1968)
Nival, P., Malara, G., Charra, R., Palazzoli, I., 'Nival, S.: Etude de la respiration
et de l'excretion de qu~lques Copepodes planctoniques dans la zone de remontee d'eau
profonde des cotes marocaines. J. Exp. Mar. BioI. Ecol. 15, 231-260 (1974)
Owens, T.G., King, F.D.: The measurement of respiratory electron transport system
activity in marine zooplankton. Mar. Bio!. lQ.. (1), 27-36 (1975)
Packard, T.T.: The measurement of respiratory electron transport system in marine
phytoplankton. J. Mar. Res. 29, 325-344 (1971)
Pachard, T.T., Devol, A.H., King, F.D.: The effect of temperature on the respiratory
electron transport system in marine plankton. Deep-Sea Res. ~, 237-249 (1975a)
Packard, T. T., Harmon, D., Boucher, J.: Respiratory electron transport acti vi ty in
plankton from upwelled waters. Tethys 6 (1-2), 213-222 (1974)
Packard, T.T., Wold, E., Abrahamson, J.,-Blasco, D.: Biological recipes for a coastal
upwelling ecosystem. Preview of preliminary scientific results, Document CUEA
20,1-20,10 (1975b)
Paranjape, M.~.: Molting and respiration of Euphausiids. J. Fish. Res. Bd Can. 24,
1229-1240 (1967)
Ponomareva, L.A.: Circadian migrations and feeding rhythm of some Indian Euphausiids
species. Oceanology 11 (2), 226-231 (1971)
Ponomareva, L.A.: Quantitative distribution of Euphausiids in the Indian ocean.
Oceanology ~ (4), 576-581 (1972)
Pugh, P.R.: The vertical distribution of the Siphonophores collected during the SaND
cruise 1965. J. Mar. Bio!. Ass. U.K. 54,25-90 (1974)
Repelin, R.: Quelques Meduses de l'ile d'Anno Bon (Golfe de Guinee). Cah. ORSTOM
Serie Oceanogr. l (1), 73-79 (1965)
Roe, H.S.: The vertical distributions and diurnal migrations of Calanoid Copepods
collected on the SaND cruise 1965. I - The total population and general discussion.
J. Mar. BioI. Ass. U.K. ~ (2), 277-314 (1972a)
Roe, H.S.: II - Systematic account: families Calanidae up to and including the Aetidae.
J. Mar. BioI. Ass. U.K. ~ (2), 315-343 (1972b)
Roe, H.S.: III - Systematic account: families Euchaetidae up to and including the
Metridiidae. J. Mar. BioI. Ass. U.K. ~ (3), 525-552 (1972c)
Roe, H.S.: IV - Systematic account of families Lucicutiidae to Candaciidae,the re-
lative abundance of the numerically most important genera. J. Mar. BioI. Ass. U.K.
g (4), 1021-1044 (1972d) .
60
Roger, C.: Les Euphausiaces du Pacifique equatorial et sud-tropical: zoogeographie,

ecologie, biologie et situation trophique. These Doc. Sci. Univ. Aix-Marseille,
331 pp. mimeo. (1971a)
Roger, C.: Distribution verticale des Euphausiaces dans les courants equatoriaux de
l'Ocean Pacifique. Mar. BioI. 10, 134-144 (1971b).
Roger, C.: Recherches sur la situation trophique d'un groupe d'organismes pelagiques
(Euphausiaces). I - Niveaux trophiques des especes. Mar. BioI. ~, 312-316 (1973a)
Roger, C.: II - Comportements nutritionnels. Mar. BioI. 18, 317-320 (1973b)
Roger, C.: III - Potentiel alimentaire du groupe. Mar. BioI. 18, 321-326 (1973c)
Roger, C.: IV - Relations avec les autres elements du micronecton. Mar. BioI. ~,
54-60 (1973d)
Roger, C.: V - Relations avec les Thons. Mar. BioI. 12J 61-65 (1973e)
Roger, C.: VI - Conclusions sur les roles des Euphausiaces dans les circuits tro-
phiques de l'Ocean Pacifique intertropical. Mar. BioI. 19, 66-68 (1973f)
Rober, C.: Influence de la phase et de l'eclairement lunaire sur les repartitions
verticales nocturnes superficielles de Crustaces macroplanctoniques (Euphausiaces).
Cah. ORSTOM Serie Oceanogr. ~ (3), 159-171 (1974a)
Roger, C.: Repartitions bathymetriques et migrations verticales des Euphausiaces
dans les zones de p~che au thon du Pacifique sud tropical. Cah. ORSTOM Serie Oce-
anogr. 12 (4), 221-239 (1974b)
Roger, C.: Rhythmes nutritionnels et organisation trophique d'une population de
Crustaces pelagiques (Euphausiaces). Mar. BioI. ~ (4), 365-378 (1975)
Roux, M., Gaborit, M., Binet, D.: Copepodes pelagiques du plateau ivoirien. II -
. Utilisation de l'arbre de longueur minimum dans Ie classement des recoltes. Docu-
ment scientifique Abidjan, ORSTOM 4 (2), 1-16 (1973)
Saint Bon, M.C. De: Les Chaetognathe~ de la Cote d'Ivoire (especes de surface).
Revue Trav. Inst. (scient. tech.) P~ch. Mar.n (3), 301-346 (1963a)
Saint Bon, M.C. De: Complement a l'etude des Chaetognathes de la Cote d'Ivoire
(especes profondes). Revue Trav. Inst. (scient. tech.) P~ch. Mar. n (4), 403-415
(1963b)
Samain, J.F., Boucher, J.: Dosage automatique et simultane de l'amylase et des pro-
teines du zooplancton. AnnIs Inst. Oceanogr., Monaco 50 (2), 199-205 (1974)
Seguin, G.: Contribution a l'etude de la biologie du plancton de surface de la baie
de Dakar. Etude quantitative, qualitative et observations ecologiques au cours d'un
cycle annuel. Bull. Inst. fro Afr. noire 28, Ser. Al, 1-30 (1966)
SegUin, G.: Cycles compares du zooplancton dans l'ouest africain et la Mediterranee
sud-occidentale. Etude systematique, quantitative et ecologique. These Doc. Sci.
Univ. Lille, 168 pp. mimeo (1973)
Silver, M.W.: The habitat of SaZpa fusiformis in the California current as defined
by indicator assemblage. Limnol. Oceanogr. 20 (2), 230-237 (1975)
Small, L.F.: Energy flow in Euphausia pacifica. Nature (Lond.) 215, 515-516 (1967)
Small, L.F., Hebard, J.F.: Respiration of a vertically migrating marine Crustacean
Euphausia pacifica. Limnol. Oceanogr • .!b 272-280 (1967)
Small, L.F., Hebard, J.F., McIntire, C.D.: Respiration ion Euphausiids. Nature (Lond.)
1.D, 1210-1211 (1966)
Smiles, M.C., Pearcy, W.G.: Size structure and growth rate of Euphausia pacifica
off the Oregon coast. Fish. Bull. Fish. Wildl. Servo U.S. ~ (1), 79-86 (1971)
Smith, S.L., Whitledge, T.E.: The role of zooplankton in the regeneration of nitrogen
in a coastal upwelling system off northwest Africa. Deep Sea Res. 24, 1, 49-56
(1977)
Stander, G.H., De Decker, A.H.B.: Some physical and biological aspects of an oceano-
graphic anomaly off S.W. Africa in 1963. Investgl Rep. Div. Fish. Un. S. Afr. ~,
1-46 (1969)
Sund, P.N.: Some features of the autecology and distributions of Chaetognatha in
the eastern tropical Pacific. Bull. Inter-Am. Trop. Tuna Comm. 2- (4), 315-340
(1961 )
Teal, J.M., Carey, F.G.: Effects of pressure and temperature on the respiration of
euphausiids. Deep Sea Res. 14, 725-733 (1967)
Thiriot, A.: Les Euphausiaces, resultats preliminaires. In: Resultats des campagnes
Cineca-Charcot V et Capricorne 74 03, Publ. CNEXO (in press)
Tokioka, T.: Observations on the· taxonomy and distribution of Chaetognaths of the
north Pacific. PubIs Seto Mar. BioI. Lab. 7 (3), 349-356 (1959)
61
Tokioka, T.: Studies on the distribution of Appendicularians and some Thaliaceans

of the north Pacific with some morphological notes. Publs Seto Mar. Biol. Lab. 8
(2), 351-444 (1960)
Unteruberbacher, H.K.: The Pilchard of S.W. Africa. Zooplankton studies in the waters
off Walvis Bay with special reference to the Copepoda. Investl Rep. Mar. Res. Lab.
S.W. Afr. 11, 1-42 (1964)
Venter, G.E.: The Pilchard of S.W. Africa. The distribution of some Chaetognaths
and their relation to hydrographical conditions with special reference to the S.W.
African region of the Benguela. Investl Rep. Mar. Res. Lab. S.W. Afr. 16, 1-71
(1969)
Vervoort, V.: Pelagic Copepoda I - Copepoda Calanoida of the families Calanidae up
to and including Euchaetidae. Atlantide Rep. 2, 77-194 (1963)
Vervoort, V.: II - Copepoda Calanoida of the families Phaennidae up to and including
Acartiidae, containing the description of a new species of Aetideidae. Altantide
Rep. ~, 9-216 (1965)
Vives, F.: Le zooplancton et les masses d'eau des environs du Cap Blanc. Tethys ~
(1-2), 313-318 (1974)
Vives, F.: Zooplancton et hydrographie des environs du Cap Blanc (N.O. de 1 'Afrique) .
Communication, Cons. perm. into Explor. Mer CMIL 10 (unpublished manuscript, 1975)
Vives, F., Santamaria, G., Trepat, I.: El zooplancton de los abrededores del estrecho
de Gibraltar en junio-julio de 1972. Result Exped. cient. Buque oceanogr. "Cornide
de Saavedra" 4, 7-100 (1975)
Weigmann, R.: Zur Okologie und Ernahrungsbiologie der Euphausiaceen im Arabischen
Meer. Meteor Forsch. Ergebn. D 2, 11-52 (1970)
Weigmann, R.: Untersuchungen zum Vorkommen der Euphausiaceen im Bereich der GroBen
Meteorbank. Meteor Forsch. Ergebn. D 12, 17-32 (1974)
Weigmann-Haass, R.: Verbreitung und Verteilung der Euphausiacea (Crustacea) im Auf-
triebsgebiet vor NW-Afrika. "Meteor"-Reise 19 (1970) und 26 (1972). Meteor Forsch.
Ergebn. D 11, 62~72 (1976)
Whitledge, T.E.: Excretion measurement of nekton and the regeneration of nutrients
near Punta San Juan in the Peru upwelling system derived from nekton and zooplank-
ton. Ph. D. Thesis Univ. Washington Seattle, 114pp. (1972)
Whitledge, T.E., Packard, T.T.: Nutrient excretion by anchovies and zooplankton
in Pacific upwelling regions. Investig. Pesq. 35, 243-250 (1971)
Zyl, R.R. van: A preliminary study of the Salps and Doliolids off the west and south
coasts of S. Africa. Investl Rep. Div. Fish. Un. S. Afr. 40, 1-31 (1960)
Relative Abundance of Zooplankton Groups in the Northwest
African Upwelling Region During 1968 and 1972
P.M. HARGREAVES
The main object of this paper is to present data obtained during bio-
logic sampling in an upwelling area. The distribution and relative
abundance of taxa are discussed and where possible indications of some
dominant species are given.
1. Methods
In January-February 1968 and July 1972 sampling programmes were carried

out in the upwelling region oft the northwest coast of Africa. RRS Dis-
covery station positions are shown in Figures 1 and 2, respectively.
All stations worked in 1968 and under discussion were positioned in-
shore of the 170-m depth contour and were completed within a period of
10 days. For the pupose of this discussion these stations have been
grouped according to their geographic location: group 1 stations 6547-
6549, group 2 6560-6564, group 3 6565-6566, 6582-6583, group 4 6585-
6588, group 5 6621-6622, 6631-6632.
Sampling during 1968 was carried out using a depth-monitored (Bowers

and Tucker, 1962), obliquely towed ring net (N113) with a 1-m 2 mouth
opening and a mesh size of 0.33 mm (Foxton, 1969), which is designed
to catch zooplankton. The sampling depth extended from the surface to
just above the sea floor. Observations made using a temperature/salin-
ity/depth recorder indicated that upwelling was active throughout the
area with strongly mixed water inshore. The sea surface temperatures
tended to fluctuate but ranged from a minimum of 15.7 o C farthest in-
shore to approximately 17~OoC at stations over the 100-m depth contour.
Phytoplankton concentrations, particularly-at some stations in groups
1 and 5, suggested that the area had been enriched for a con~iderable
time. One station, over the 100 m depth contour, was monitored for
14 h to obtain temp./sal. data. Here intermittent but strong hydrologic
fluctuations correlated with internal waves that were thought to be
of importance in vertical mixing, were apparent (National Institute of
Oceanography, Cruise Report 21, 1968).
The 1972 sampling programme in this area was conducted jOintly by the
National Institute of Oceanography and by Liverpool University, Depart-
ment of Oceanography (N.I.O. Cruise Report 53, 1973). Five separate
lines of stations were worked seawards of the shelf from the 50 m to
the 2000-m depth contour. Nets were fished at 4 stations on each line:
line 1 stations 7973-7979, line 2 7982-7988, line 3 7991-7999, line
4 8001-8008, line 5 8012-8020. The sampling was completed within a
period of 9 days.
A depth-monitored, obliquely towed rectangular midwater trawl (RMT 1)

(Baker et al., 1973), the mesh and mouth area of which are the same as
those of the N113, was fished from the surface to within about 5 m of
the sea floor or to a maximum of 1000 m, whichever was the shallower.
The temperature and salinity- measurements indicated that upwelling was
63
Fi g . 1. RRS Discovery
2(fw Stat ion positio ns,
Jan u ary-February 19 6 8 .
Gp .1-5 = groups 1- 5
iI ~47
~ i' .~~~9
GP. 2 ~
\ '£61
'" 562
/ 6563
$./ .6564
25 GP.3 " ~S65
I 658 t.
---L- 658 6566
GP.4 , I65860 ~'65 5
() I .~
\ ) f
I :1 i.
6
J,
A
BARBAS
) GP. 06621
I '~22
2 2(f
MAURITANIA
C",VERDE IS. • Station positions
.""
JAN - fEB 1968
I'
r-/o ~
15
occurring within the region, but was strongest off Cape Bojador (line
1). The survey showed a decrease in temperature towards the coast with
the isotherms generally following the edge of the shelf. The sea sur-
face temperature ranged from a minimum of 17.0oC (inshore on line 1)
to between 1S.SoC and 20.0oC at various stations over the 1000-m depth
contour (Hughes and Barton, 1974).
2. Results
Wet displacement volumes of hauls, most of which consisted almost en-

tirely of zooplankton, were standardised. In some hauls, particularly
at group 1 and group 5 stations, large amounts of phytoplankton were
present . Most of this was removable and is not included in the volumes
given. However mucilaginous remnants sometimes remained in the samples
64
Fig. 2. RRS Discovery

20'W Station positions,
3et 300 July 1972
LINE 2
8001
LINE 4 8003
C. VEROE 15. Station positions

'I JULY 1972
./
""""'0
and so figures giving zooplankton volume should be regarded as approx-

imations.
Vol. of catch (ml) x depth of haul (m)

Vol. (ml) beneath
Mouth area of net (m 2 ) x distance of tow (m) 1 m2 sea surface
This estimate of volume (Fig. 3) therefore relates to total biomass

caught at each station and is not indicative of relative density. In
1968 the greatest volumes of zooplankton were obtained at some stations
in groups 1 and 4 (> 20 ml beneath 1 m2 ) and one station in group 5
(> 30 ml beneath 1 m2 ). Phytoplankton was particularly abundant at
group 5 stations near Cape Blanc. In 1972 zooplankton volumes were
greatest at four offshore stations on lines 4 and 5 (> 38 ml beneath
1 m2 ). Low volumes tended to occur at inshore stations in groups 1 and
3 (1968) and at the inshore stations on lines 1 and 5 (1972).
All samples were fractionedto a varying extent, depending on the vol-

ume of the catch, by means of a Folsom Plankton Sample Splitter (McEwen
65
VOLUMES OF ZOOPLANKTON Fig. 3. Wet displacement

volumes of zooplankton
Jan-Feb 1968 July 1972 (ml) beneath 1 m2 sea
surface calculated from
Gpl Line 1 hauls during 1968 and
1972 .using nets Nl13
and RMT 1. Note change
of scale for sea floor
soundings at 100 m
Line 2
Line 3
Line 4
line 5 (79)
50 0
Sea floor soundings (metres)
et al., 1954), and then analysed. Individuals from each major zooplank-
ton group were counted, their total in the sample estimated, and this
figure expressed as a percentage of the estimated sum total of all in-
dividuals in the various major zooplankton groups, thus giving a rough
indication of the numerical importance of the various groups. This
method has some limitations for it cannot take into account the relative
biomass of the various planktonic groups; however, due to net selection,
individuals sampled were relatively small in size, many being larval
forms, and so there was relatively little disparity in size of specimens.
At all except two inshore stations cope pods accounted for more than 40%
of the total number of individuals in the haul and often exceeded 60%.
66
COPEPODA Fig. 4. Numbers of

m
Copepoda beneath 1 2
Jan-Feb 1968 July 1972 sea surface calculated
> Gpl Linel
from hauls during 1968
and 1972. Note change
soundings at 100 m
Line 2
line 3
Line 4
(>40.000) (40.000)
line 5
( >60.000) (~.ooo)
2100 1700 1300900 500 lqO 50 0

:
Euphausiids accounted for more than 40% only at the inshore station in
group 5. Cladocerans exceeded 40% of the haul at each of the two in-
shore stations on line 4. The large volumes of zooplankton obtained
seawards of the shelf on lines 4 and 5 in 1972 were accounted for mainly
by copepods and a few adult and sub-adult fish and crustaceans.
It is too early to comment in detail on all the dominant species within

the animal communities, because many groups have yet to be analysed
to species level. Those zooplankton groups which appear to be widely
distributed, the further investigation of which should lead to a clearer
understanding of the factors influencing animal communities in this
region, include - copepods, euphausiids, decapods, fish, siphonophores,
67
EUPHAUSIACEA Fig. 5. Numbers of Eu-

phausiacea beneath 1 m2
Jan -Feb 1968 July 1972 sea surface calculated
> Gpl linel
and 1972; ( .... ) ~ nil.
Note change of scale for
sea floor soundings at
100 m
line 2
line 3
line 4
line 5
2100 1700 1300 900 500 lyO 50 o

ostracods, appendicularians, and chaetognaths. Those identified to

species level were euphausiids, siphonophores, and ostracods and some
decapods, mysids, and amphipods.
Maximum numbers of copepods (> 40,000 beneath 1 m2 ) were found to occur

at the offshore station in group 1 and at each of the two offshore sta-
tions on lines 4 and 5 (Fig. 4).
Euphausiids showed wide variability in distribution (Fig. 5). Maximum

numbers occurred inshore and offshore and could not be correlated with
particular geographic locations. Numbers tended to be low at line 1
stations. A neritic species, Nyctiphanes capensis (Brinton, 1962; Meira,
68
DECAPODA
Fig. 6. Numbers of Deca-
poda beneath 1 m2 sea
Jan -Feb 1968 July 1972 surface calculated from
hauls during 1968 and
Gpl line 1 1972. Note change of
3000 scale for sea floor
soundings at 100 m
2000
1000
Gp2 Line2
3000
2000
1000
Gp3 Line 3
...
Q)2000
c:
::>
~1000
Z
Gp4 Line 4
3000
2000
1000
Gp5 line 5
3000
2000
1000
500 100 50 0 2100 1700 1300 900 500 100 50 0

1970), was found to occur in all hauls in 1968 sometimes accounting

for more than 97% of the total euphausiid catch. In 1972 this species
was absent or rare at stations on lines 3 and 4; elsewhere its abundance
was variable. Two other species of euphausiids which are numerically
importan t in the area are Euphausia krohnii and Nematosce lis megaZops.
The majority of decapods were sub-adult or larval. Numbers in excess

of 2000 beneath 1 m2 occurred only at the two inshore stations on line
4 (Fig. 6). Abundance was particularly low on line 1. Many of the deca-
pod larvae have yet to be identified to species level but results so
far indicate that many are benthic in origin. Brachyuran zoea larvae
were found to be widely distributed, accounting for a relatively large
69
SIPHONOPHORA Fig. 7. Numbers of

Siphonophora beneath
Jan -Feb 1968 July 1972 1 m2 sea surface cal-
Gp1 linel culated from hauls
during 1968 and 1972;
( .... ) = nil. Note change
soundings at 100 m
Line 2
Line 3
Line 4
Line 5
o 2100 1700 1300 900 500 100 50 0
proportion of the total decapods, except at line 1 offshore stations

where they were rare. Abundance levels tended to be greatest at inshore
stations on line 4. Carideans accounted for 3-70% of the total decapods.
Adult specimens of Acanthephyra purpU:t'ea were present at some offshore
stations on lines 1 and 3. Adult SysteUaspis debiUs were present at some
offshore stations on lines 1, 2, 4, and 5. In general, hauls from in-
shore stations contained a higher percentage of small early stage cari-
dean larvae than did those from offshore stations. Pagurids and gala-
theids were found mainly inshore, numbers were variable. Penaeid and
serges tid larvae tended to be more abundant at the northerly stations.
Sub-adult Sergestes (Sergestes) atlanticus were present at one offshore
station in 1972.
70
Fish abundance showed very little correlation with distance from the
shore. In 1968 numbers exceeded 370 beneath 1 m2 only at group 1 off-
shore stations. In 1972 abundance beneath 1 m2 was variable especially
at stations on lines 2-5 where they ranged from 1 inshore on line 5
to approximately 780 offshore on line 4. At stations on line 1 numbers
tended to be low. Fish eggs were present at many stations.
Siphonophores were also found to be patchy in distribution (Fig. 7).

Numbers exceeded 800 under 1 m2 only at 4 stations. In 1968 Muggiaea
atZantiaa, a neritic species, proved dominant; i t accounted for more
than 78% of the total siphonophores from each station. Bassia bassensis,
a widespread shallow-living species normally found further offshore,
was rare as was Diphyes dispar, a species which is widely distributed
and which is found in offshore oceanic water (Pugh, 1974). In compari-
son, in 1972 MU{Jgiaea atZantiaa was most abundant only at the inshore
station on lines 1, 3, and 5 where i t accounted for more than 80% of
the total siphonophores. This species was also abundant on the whole
of line 4, extending to the offshore stations. Bassia bassensis was com-
mon at each of the three offshore stations on lines 1 and 2. Small
numbers of Diphyes dispar were present at some offshore stations. These
three species have been previously recorded off the Moroccan coast
(Furnestin, 1957).
Ostracod distribution was somewhat variable with abundance levels tend-

ing to be highest at all offshore stations in 1972 but especially on
lines 4 and 5. This contrasts with the lower figures recorded for most
inshore stations in 1968 and 1972 (Fig. 8). A wide variety of species
was present in many hauls, the most common being Conahoeaia eZegans, C.
spinirostris, and C. aurta, all shallow-living species (Angel and Fasham,
1975). Not unexpectedly shallow-living species were dominant nearshore,
and on the average a higher proportion of deeper water species occurred
offshore especially at the deeper stations beyond the shelf break worked
in 1972.
In 1968 mysids were absept or rare except at group 1 offshore stations

(> 300 under 1 m2 ), which were sampled at night. The dominant species
here was Leptomysis graaiZis, a neritic species. In 1972 only one haul,
nearshore on line 2 at night, contained a large number of mysids (> 1000
under 1 m 2 ). Here two neritic species Leptomysis Zingvum and Gastrosaaaus
ar. normani were dominant. These species have previously been recorded
off the coast of Morocco (Furnestin, 1959). Many benthic samples ob-
tained in 1972 contained large numbers of mysids, and i t is probable
that the presence of this group in the epipelagic and mesopelagic region
at night may be correlated with diurnal vertical migration.
The largest catch of amphipods (> 900 under 1 m2 ) was taken at a sta-
tion on line 3 at night where the sounding was 100 m. Here approximately
98% of the amphipod catch was comprised of sub-adults of a species
similar to A-tyZus vedZomensis, a European coastal species (Schellenberg,
1942) •
Chaetognath distribution was variable. Figures exceeded 1000 beneath

1 m2 at twelve stations. Abundance levels could not be correlated with
geographic location.
Appendicularians tended to be abundant at one group 3 station and some

onshore stations on lines 2 and 3, all within the 100 m depth contour,
with values in excess of 2000 under 1 m2 • Elsewhere numbers were re-
latively low.
Salps and doliolids were nQt abundant, with numbers in excess of 1000
under 1 m2 only at one station in group 2 and at two stations on line
4. Numbers were particularly low in the Cape Blanc area.
71
OSTRACODA Fig. 8. Numbers of

Ostracoda beneath 1 m2
Jan-Feb 1968 July 1972 sea surface calculated
Gpl Line 1 and 1972; ( .... ) = nil.
3000 Note change of scale
for sea floor soundings
2000 at 100 m
1000
3000
-- Gp 2 Line 2
2000
1000
1--.
N
Gp3 Line 3
E
~3000
....
~2000
c
:::>
.,;1000
0
Z
Gp4 Line4
3000
2000
1000
..
Gp5 Line 5
3000
2000
1000
500 100 50 o 2100 1700 1300 900 500 190 50 o

Sea floor sound i n9s (metres)
Cladocerans were sparse in 1968 except inshore at some group 2 to 4

stations where numbers exceeded 2000 beneath 1 m2 • This group was also
sparse at stations on lines 1 and 2 in 1972, but on most inshore sta-
tions on lines 3 to 4 numbers ranged from> 1000-21,000 beneath 1 m2 •
Few cirripedian larvae were found in 1968 or in 1972 except at one sta-
tion in group 2 and one in group 3 where numbers exceeded 900 beneath
1 m2 •
Medusae were not abundant in 1968 or in 1972. Maximum numbers occurred

at a group 1 station.
72
3. Discussion
It seems apparent that to clarify the situation with regard to the

animal communities it will be necessary to continue to identify most
of this material to the species level. Results obtained so far are en-
couraging in that they are clearly indicative of the ecologic varia-
tions existing between inshore and offshore stations. For instance
ostracods, although variable in number, ,,;hen analysed to species level
showed changes in species composition from onshore to offshore with
the total number of species decreasing shorewards. Similarly data on
the decapod distribution in general indicate variation in species com-
position between inshore and offshore stations. In addition, preliminary
comparisons of maturity stages in a few of the decapod groups pOint to a
size difference existing between inshore and offshore specimens. Even-
tually it should be possible to speculate on the relationships between
hydrologic conditions and breeding cycles. Similarly further identifica-
tion of species should eventually enable us to comment on possible
trophic levels.
Acknowledgments. I wish to thank colleagues at the lOS Wormley who have assisted
in the collection and identification of specimens.
References
Angel, M.V., Fasham, M.J.R.: Analyses of the vertical and geographic distribution
of the abundant species of planktonic ostracods in the North East Atlantic. J.
Marine BioI. Assoc. U.K. 22, 709-737 (1975)
Baker, A. De C., Clarke, M.R., Harris, M.J.: The N.I.O. combination net RMT 1 + 8
and further development of the rectangular midwater trawl. J. Marine BioI. Assoc.
U.K. 53, 167-184 (1973)
Bowers, R., Tucker, M.J.: The N.I.O. Depth Telemeter. N.I.O. Internal Report A. 16 p.
6 (1962) (Mimeo)
Brinton, E.: The distribution of Pacific euphausiids. Bull. Scripps Inst. Oceanog.
Univ. Calif. ~ (2), 51-270 (1962)
Foxton, P.: SOND cruise 1968. Biological sampling methods and procedures. J. Marine
BioI. Assoc. U.K. 49, 603-620 (1969)
Furnestin, M.-L.: Chaetognaths et zooplankton du secteur atlantique marocain. Rev.
Trav. Inst. Pech. Marit. 11 (1 et 2), 1-356 (1957)
Furnestin, M.-L.: Mysidaces du plankton marocain. Rev. Trav. Inst. Pech. Marit. 23
(3), 297-316 (1959)
Hughes, P., Barton, E.D.: Physical investigations in the upwelling region of North-
West Africa on R.R.S. Discovery Cruise 48. Tethys 6 (1-2), 43-52 (1974)
McEwen, G.F., Johnson, M.W., Folsom, T.R.: A statistical analysis of the performance
of the Folsom plankton sample splitter based upon test observations. Arch. Met.
Geophys. Bioklim. (Ser. A) 2, 502-527 (1954)
Meira, C.: Contribuicao para 0 estudo dos eufausiaceas do arquipelago de Cabo Verde.
Notas do Centro de Biologia aquatica tropical ~, 1-27 (1970)
National Institute of Oceanography: R.R.S. Discovery Cruise 21 report Jan.-April
1968. UpVlelling off North West Africa and plankton distribution at 11 oN 20OW.
N.I.O. Cruise Report 21 (1968) (Mimeo)
National Institute of Oceanography: R.R.S. Discovery Cruise 48 report. UpVlelling
off the coast of N.W. Africa. N.I.O. Cruise Report 53 (1973) (Mimeo)
Pugh, P.R.: The vertical distribution of the siphonophores collected during the
SOND cruise 1965. J. Marine BioI. Assoc. U.K. 54, 25-95 (1974)
Schellenberg, A.: Krebstiere oder Crustacea IV: Flohkrebse oder Amphipoda. Die Tier-
Vlelt Deutschlands Jena 1QJ 1-252 (1942)
Mesodinium rubrum in the Baja California Upwelling System
T. T. PACKARD, D. BLASCO, and R T. BARBER
1. Introduction
Mesodiniwn ruhrwn (Lohmann) Hamburger and Buddenbrock is a holotrichous

ciliate, occasionally responsible for nontoxic red waters in every
major ocean except the Indian. Unlike most other ciliates, M. rubrwn
and some members of the genera Prorocwn and Strorribidiwn (Blackbourn et
al., 1973) contain chloroplas ts. Prorodon and strorribidiwn contain chloro-
plasts, only, and occasionally have been found without them. M. ruhrwn,
however, also contains mitochondria in addition to the chloroplasts;
furthermore, the organism has never been observed without the chloro-
plasts (Taylor et al., 1971). These chloroplasts were thought to be
associated with quasi-independent cryptomonads (Parsons and Blackbourn,
1968; Barber et al., 1969), but the absence of nonciliate nuclei and
the thylakoid characteristics of the chloroplasts indicate that the
chloroplasts are not associated with endosymbiotic algae, but are
functioning as ciliate organelles. How they became associated with
the ciliate is unclear. The vestigial condition of M. ruhrwn's cytostome
suggests that new algae cannot possibly be ingested in each ciliate
generation, although they might be ingested at less frequent intervals
during some unknown stage of M. ruhrwn's life cycle (Taylor et al.,
1969, 1971). Regardless of their origin, the chloroplasts contain
cryptomonad-type pigments, they photosynthesize, and give M. ruhrwn
the characteristics of a phytoplankter. Ryther (1967), Parsons and
Blackbourn (1968), and Barber et al. (1969) measured chlorophyll a and
c in M. ruhrwn, Ryther (1967) and Barber et al. (1969) measured light-
stimulated 14C-uptake, and Barber et al. (1969) demonstrated 02 pro-
duction and 3 2 p assimilation. Margalef (1956) found higher 02 concentra-
tions in Mesodiniwn blooms than outside them. Parsons and Blackbourn
(1968) and Barber et al. (1969) observed several accessory pigments
in M. ruhrwn, namely, a-carotene, a phycoerythrin with a unimodal peak
at 542 nm, and an unidentified orange pigment. The identification of
a-carotene and the orange pigment corroborates the cryptomonad origin
of the chloroplasts.
The distribution of M. rubrwn, although widespread, appears to be con-

fined to bays and fjords or to upwelling in coastal regions. No sight-
ings have been made farther than 180 km offshore. When sighted, the
organism has been identified as Halteria ruhra (Lohmann, 1908), M. ruhrwn,
or Cyclotrichiwn mezmieri (Powers). The use of Halteria rubra was discon-
tinued soon after 1911 (Hamburger and Buddenbrock, 1911; Fenchel,
1968a) but the confusion between M. rubrwn and C. mezmieri was dispelled
only recently by Fenchel (1968a) and Taylor et al. (1971) who concluded
after a review of earlier descriptions and detailed cytologic analysis
that the organisms were conspecific. They recommended the use of the
taxonomically correct name, Mesodiniwn rubrwn. Despi te the errors in
nomenclature, many of the early records contain useful information.
Powers (1932), identifying the ciliate as C. meunieri, describes patches
and windrows of red water in Frenchman's Bay, Maine following a period
of unusually warm summer weather. In the same season and along the
same coast of the Gulf of Maine, McAlice (1968) observed C. mezmieri
74
concentrated in numbers as high as 2.2 x 10 6 cells/liter and associated

wi th the diatom, Cosainodisaus aentraUs, the copepod naupli us of Eu:ryte-
mora herdJnani and two cladocerans, Evadne nordJnanni and Podon Zeuka:rti. Both
authors noted the extreme fragility of the organism, finding i t both
difficult to preserve and difficult to observe alive. This problem
was also encountered by Hart (1934) who observed red-water blooms of
M. ruhrwn in a rich assemblage of diatoms and copepods during the aus-
tral winter along the Atlantic coast of South Africa. He noted that
rotifers were feeding on the ciliates and that a green scum apparently
caused by disintegrating M. ruhrwn accumulated at the tide line of the
adjacent shore. Further evidence of the role of M. ruhrwn .in marine
food chains was cited by Clemens (1935) who observed red crystalline
styles in clams and oysters following blooms of M. ruhrwn along the
eastern shore of Vancouver Island. As with the blooms along the Maine
coast, these occurred after several weeks of calm, sunny weather. Bary
and Stuckey (1950) recorded that blooms of C. meunieri occurred after
similar weather conditions in Wellington Harbour, New Zealand, and
Fenchel (1968a) suggests the same cause for an M. ruhrwn bloom in Isef-
jord, Denmark. This ciliate bloom and others that have occurred in
Isefjord do not adversely affect the marine fauna, in fact, they pro-
vide food for at least one group of zooplankton. The mysids, Praunus
fie:x:uosW3 and P. inermis, have been seen feeding at the borders of the
red clouds of the M. ruhrwn blooms (Rasmussen, 1973). Kuenzler and Bar-
ber (personal communication) observed a rich zooplankton assemblage
congregating in a similar fashion around the edges of a C. meunieri
bloom in the equatorial front between the Tropical Surface Water and
the Subtropical Surface Water (Wyrtki, 1967). In the ria de Vigo,
Spain (Galicia) a strong shallow thermocline developes after warm
weather, encouraging not only Mesodiniwn, but also GonyauZax to bloom
concurrently, in different parts of the ria. Margalef (1956) concluded
that these ciliate blooms represent a typical final stage of species
succession in the ria.
Upwelling systems evidently stimulate the development of M. ruhrwn blooms.

These environments offer abundant nutrient supplies and both diverging
and converging surface waters. Through a combination of its autotrophic
capaci ty and its mobility, M. ruhrwn can exploit these waters when other
organisms are washed out. Ryther (1967) and Barber et al. (1969) found
C. meunieri blooms at the northern boundary of the Peru coastal, current.
In both cases the blooms were nontoxic and confined to the upper part
of the euphotic zone (1-3 m). Ryther (1967) found a bloom in a small
upwelling region where the density of the seawater and the nutrient
concentration were higher than they were in the adjacent oligotrophic
seawater. The two blooms described by Barber et al. (1969) were less
closely associated with freshly upwelled seawater. Fonds and Eisma
(1967) found M. ruhrwn blooming under upwelling conditions within 11 km
of the Dutch coast. The bloom was associated with a rich diatom flora
of SkeZetonema aostatwn and Chaetoaeros sp.·
Individuals of M. ruhrwn range in size from 20-50 ]lm. The cells are
ringed with cilia, ovoid in shape, and have an average cell volume of
1.8 x 10 3 ]lm 3 (Fig. 1 and Fenchel, 1968a). Although they never have
been observed in mitosis, M. ruhrwn cells could have a generation time
of 5 h if the volume-dependent growth relationship in benthic ciliates
can be applied to them (Fenchel, 1968b). They swim rapidly, rotating
as they dart about on zig-zag tacks, attaining speeds of 0.6 to 2 rnm/s
(Powers, 1932; Bary and Stuckey, 1950). They are positively phototactic,
but shun strong light (Hart, 1934), seeking a somewhat lower, albeit
optimum light level (Bary and Stuckey, 1950). This characteristic sug-
gests that the vertical distribution of M. ruhrwn might evince a sub-
surface maximum, although as Taylor et al. (1971) discovered there is a
dearth of data to support this prediction.
75
Fig. 1. Sketches of Mesodinium rubrum.

A D (A) Power's (1932) diagram of CycZO-
trichium meunieri from the Maine coast;
(B) Hart's (1934) drawing of Mesodinium
rub rum from the red water in Simons
Bay, South Africa; (C) Fonds and Eis-
rna's (1967) Mesodinium puZex "rubrum"
from the Dutch coast; (D) Bary and
Stuckey's (1950) CycZotrichium meu-
nieri from Wellington Harbour, New
Zealand; (E) Mesodinium rub rum from
English Bay, Vancouver, B.C. (Taylor
et al., 1971). The large arrow in the
lower left corner of this panel in-
dicates the direction of travel and
the rotary component of the organism's
motion. The organelles are identified
as follows: a, cirri; b, equatorial
ciliary band; £, chlor~plast complex;
d, macronucleus; e, micronucleus; f,
mltr ochondrion; g~ vacuole; h, tri~ho
cysts; i, rudime~tary cytostZme. The
last pa~el (F) is from a photograph
of CycZotrichium meunieri from Val-
paraiso Bay, Chile (Avaria, 1970).
Each panel has been redrawn from the
originals
The tolerance of the organism to variations in temperature and salinity

is exceptional: Blooms have been found in waters ranging in salinity
from 4%0 to 35%0 and ranging in temperature from 2 0 to 20 0 C (Taylor
e t al., 1971).
This paper describes observations of phytoplankton - numerical abundance

and vertical distribution as well as carbon and nitrogen assimilation
rates in seawater inhabited by M. rubrum along the open coast of western
Baja, California.
2. Methods
This study was made between Punta Asuncion and Punta Abreojos (Fig. 2)
during the spring of 1973. It was a project of the Coastal Upwelling
Ecosystems Analysis (CUEA) program, an organization sponsored by the
U.S. National Science Foundation and dedicated to further the under-
standing of the biologic, chemical, and physical phenomena associated
with coastal upwelling. A partial description of these phenomena during
the spring of 1972 off Punta San Hipolito has been given by Walsh et
al. (1974). The stations occupied in 1973 were in the vicinity of this
point (Fig. 2).
Seawater samples were collected between 0630 and 0830 (local time) in
30-liter Niskin bottles with a Rosette sampler from the depths to which
100%, 50%, 25%, 10%, and 1% of the incident light penetrated. These
depths are referred to throughout this paper as light depths. From each
Niskin bottle, subsam~les were drawn for analyses of inorganiC nutrient
salts [P0 4 - 3 , N0 3 , NH 4 , and Si(OH)4]' chlorophyll, particulate nitrogen
(PN) , particulate carbon (PC), nitrate reductase activity (NR) , respi-
ratory electron transport activity (ETS) , and phytoplankton taxonomic
76
Fig. 2. Station locatio n s f o r

R/ V T.G. Thompson cruise 78
(MESCAL II )
IS'
BAJA
CALIFORNIA
•
46
27·
79-.
27
00' ~.':;'.:. 00'
• 7381 '
6,26,38
53,67, 78,82
4S'
Pocific Ocean
30' 30 '
4S ' 30' IS' 114·00' 4S ' 30'
composition. Separate casts, with 6-liter Niskin bottles arrayed at

10-m intervals between the sea surface and 50 m, were made for salinity
and temperature measurements. Data for intermediate depths (i.e., 3,
7, 16 m, etc.) were obtained from STD profiles.
The concentration of nutrient salts was determined by the method of

Armstrong et al. (1'967), within an hour of the sampling time. The
samples were stored at 0-4 o C in the dark for part of that time inter-
val.
Chlorophyll was measured by the absorbance method (UNESCO, 1966) and

the fluorescence method of Holm-Hansen et al. (1965). Throughout this
paper these chlorophyll methods will be identified by UNESCO and FLUOR,
respectively. The particulate nitrogen and carbon were determined by
the gas biuret method of Pavlou et al. (1974) and Menzel and Vaccaro
(1964) . NR and ETS were determined by the methods of Eppley et al .
(1969) and Owens and King (1975); the calibration factor for the NR
method was 390 nm N0 2/OD unit. The phytoplankton samples were preserved
with a few drops of a solution prepared by adding 20 g Na acetate to
100 ml of Lugol solution (9 M K1 saturated with 1 2 ), The samples were
stored in the dark for 2 to 12 months and were enumerated after sedi-
mentation with the aid of an Utermohl inverted microscope (Utermohl,
1931; Blasco, 1971; Margalef, 1975) .
3. Resul t s
Me s odinium ruh r um was observed off the western coast of Baja, California
during the months of April and May of 1973. Blooms appeared to be con-
fined to Hipolito Bay (stations 73, 79, 81) on the south side of Hipo-
lito Point (Fig. 2), although on two occasions red-water patches were
sampled 5 km off the point (stations 38 and 67X). When the sea was calm
and the wind velocity was nearly zero, the ferruginous blooms occurred
77
in irregularly shaped patches (stations 38 and 73). When the wind blew
at 10-20 knots they aligned themselves in windrows at 100-m intervals
(stations 67 and 67X) and when the wind blew stronger than 30 knots
the blooms were despersed in low numbers throughout the water column
(stations 6, 26, 46, 53, 78, and 82). When conditions favored patch
formation, the seawater between the patches was devoid of M. rubrurn.
Stations 67 and 67X were made within several hundred meters of each
other, yet because the former was taken in between the patches, not a
cell of M. rubrurn was observed in a 100-ml sample. Within the patches,
the ciliates could be observed in densities as high as 538 x 10 3 cells/
I (station 73, bucket sample). In the routinely taken phytoplankton
samples the ciliates occurred in lower numbers, possibly as a result
of mixing in the 30-liter Niskin sampling bottles. At station 73,
when the count from the bucket sample gave 538 x 10 3 cells/I, a sample
from the large Niskin bottles gave 71 x 10 3 cells/I. Regardless of the
source of the sample, cell densities of M. rubrurn were low. For compari-
son, McAlice (1968) found M. rubrurn cell densities varying from 0.21
to 2.2 x 10 6 cells/l in the Gulf of Maine and Fenchel (1968a) found
values varying from 177 to 254 x 10 6 • A summary of the relative cell
counts at the "Mesodinium" stations (Nos. 38, 73, 79, and 81) are shown
in Table 1, and the vertical distributions of these counts against a
Table 1. Cell counts (cells/ml of seawater) of M. rubrurn from the upwelled waters
off western Baja, California
Light
depth Station
(%) 38 73 79 81
100 23 71 176.2 3
50 76 236.3 3
25 9.4 28.3 2
10 0 24.4 64.2 0
1 9.4 0
background of the other four components of the phytoplankton population

are shown in Fi~ure 3. The cell numbers at these stations ranged from
a low of 2 x 10 cells/l at station 81 to a high of 236 x 10 3 cells/l
at station 79. With the low cell densities observed at station 81, i t
qualifies only marginally as a "lilesodinium" station, which was sur-
prising because station 81 was located at the maximum in the chlorophyll
field (Kelley et al., 1975) in Hipolito Bay and on the previous day
many blooms were observed at this location. At the two stations at
which M. rubrurn was abundant and at which the data suite was complete
(stations 38 and 79) a subsurface maximum at the 50% light level char-
·acterized the vertical profiles. At the 1% light level the organisms
were represented in low numbers. This distribution is understandable
in view of the findings of Bary and Stuckey (1950) that M. rubrurn is
positively phototactic, seeking an optimum light level somewhat diffuse
and somewhat lower in intensity than the maximum incident light. The
other components of the phytoplankton community in the region of Punta
San Hipolito included small flagellates, dinoflagellates, coccolitho-
phorids, diatoms, and silicoflagellates, in that order of relative
numerical abundance (Table 2). When M. rubrurn was present, the relative
importance of the coccolithophorids and the diatoms was reversed. On
these occasions M. rubrurn ranked after the dinoflagellates in the sur-
face waters and before them at the 50% light depth. At the 1% light
level the diatoms were only slightly more important than the silico-
flagellates. The species composition remained relatively constant
throughout the entire period of study. The dinoflagellate population
78
,\ \ /
I j /\
I I.0X.
III
a::
I&J
t- 10
I&J
~
:x: 15
t-
o.
I&J
0 m
. ••
e 010
o A
F
20
•• 0
oe F
38 79 81
25 L-____~______~--J
o 100 200 o 100 200 o
C ELL COUNTS ( mI-I )
Fig. 3. Cell-count depth profiles of Mesodinium rubrvffi (m), dinoflagellates (Df) ,

diatoms (D), coccolithophorids (C), and small flagellate; (F) at 3 MESCAL II stations.
Red water-was detectable by eye -;nly at Station 79 -
Table 2. Phytoplankton numerical abundance from seawater inhabited by M. rubrum and

from seawater not inhabited by M. rubrum
Seawater inhabited by M. rub rum Seawater not inhabited by M.rubrum

rubrvffi
(sts. 38, 73, 79 and 81) (Sts. 67, 78, and 82)
Surface 50% light 1% light Surface 50% light 1% light
level level level level
Small flagellates 42 ± 10 38 ± 13 39 ± 9 56 ± 8 48 ± 4 69 ± 7
Dinoflagellates 23 ± 6 23 ± 13 37 ± 17 29 ± 11 16 ± 8 18 ± 9
M. rubrum 21 ± 20 24 ± 26 4 ± 8
Coccolithophorids 5 ± 5 7 ± 8 7 ± 6 12 ± 13 32 ±13 12 ±
Diatoms 9 ± 7 7 ± 7 13 ± 11 3 ± 4 3 ± 0.1 5 ± 4
Silicoflagellates 0.4±0.2 0.3±0.1 0.4±0.4 1.4 ± 2 ± 1 ±
Average values and standard deviations in percentages are given.
was dominated by Ceratium fW'ca and C. dens, Diplopsalis minor, Peridinium

trochoideum, P. depressum, Glenodinium sp., and Gymnodinium sp. The diatom
population was dominated by Coscinodiscus excentricus, Lauderia annulata,
Rhizosolenia delicatula, R. Stolterfothii, Schroederella delicatula, Thalassi~aema
nitzschioides, Thalassiosira rotula, and PlanktonieUa sol. Cyclococcolithus
fragilis dominated the coccolithophorids although at stations 81 and 82
Helycosphaera sp. became abundant. The absence of major differences in
the floral composition between waters inhabited by M. rubrum and the
surrounding waters is in agreement with the observations of Holm-Hansen
et al. (1970) on a pigmented ciliate bloom in the Bering Sea.
Table 3. summary of hydrographic, chemical, biomass, and enzyme data from the seawater in which Mesodinium rubrum did not occur
Light Hydrography Chemistry Biomass Enzyme activity

depth Depth Salinity Temp. Density P04 Si(OH)4 N03- NH4+ Chlorophyll
(%) (m) (%0) (oC) (at) UNESCO FLUOR. PN PC NR ETS
(]lg/l) (]lg- (]lg/ (ng-at N03- (]ll 02 h- l e l )
at/I) 1) N h- l e l )
Station 67
100 0 33.82 15.24 25.03 0.61 4.65 1. 67 0.48 1.88 1.48 2.48 68 7.74
50 5 0.62 4.65 1. 73 0.40 1. 39 1.85 2.16 147 6.47
25 10 0.61 4.58 1.72 0.40 1.42 2.37 2.51 139 6.55
10 17 33.84 15.18 25.06 0.61 4.58 1.90 0.39 1.40 2.00 2.26 135 6.83
34 0.88 5.85 4.86 0.94 0.96 1.04 2.55 129 5.60
Station 71
100 0 33.81 15.60 24.94 0.50 3.91 0 0.49 1.42 1.55 2.25 81
50 3 0.50 3.96 0.06 0.43 1. 34 1. 33 2.67 70
25 8 0.57 4.04 0.12 0.40 1.63 2.18 60
10 13 33.81 15.60 24.94 0.56 3.80 0.29 0.39 1.40 1.48 2.08 53
1 28 33.78 15.40 24.96 0.54 3.66 0.35 0.38 1.51 1.48 1. 92 59
Station 78
100 0 33.83 16.47 24.76 0.45 1.95 0 0.35 0.12 0.30 21 3.58
50 78 33.83 16.45 24.77 0.45 2.03 0 0.36 0.23 1.20 26 4.80
25 15 33.82 16.21 24.81 0.47 2.37 0 0.38 0.24 0.30 1.12 41 4.28
10 25 33.80 15.72 24.91 0.50 2.85 0 0.37 0.85 1.06 1.98 60 3.42
1 49 33.82 12.66 25.57 1.62 14.21 9.26 0.45 0.62 0.36 1.29 34 8.25
Station 82
100 0 33.85 14.76 25.16 0.82 10.80 7.16 0.61 1. 91 1.55 2.29 167 20.4 24.42
50 4 33.83 14.70 25.16 1. 27 11.05 7.66 0.77 2.03 1.55 1. 65 193 13.7 16.42
25 9 33.84 14.64 25.18 1.12 10.77 7.39 0.68 1.36 1. 70 2.27 171 24.24
10 16 33.86 14.60 25.20 1.05 10.71 7.70 0.65 3.57 1. 70 2.37 141 24.84
28 33.45 13.81 25.25 1.11 12.27 9.12 0.73 0.73 0.81 1.23 51 2.7 13 .13
All the stations were taken off the western coast of Baja California during the R/V T.G. Thompson cruise, MESCAL II.
--.J
1.0
()O
0
Table 4. Summary of the hydrographic, chemical, biomass, and enzyme data from the seawater in which Mesodinium rubrum occurred
Light Hydrography Chemistry Biomass Enzyme Acti vi ty

depth Depth Salinity Temp. Density PO Si(OH)4 N03 - NH4+ Chlorophyll PN PC NR ETS
(%) (m) (%0) (OC) (at) (]Jg-at/l) UNESCO FLUOR. (]Jg- (]Jg/ (ng-at N03- (]Jl 02 h- 1 1- 1 )
(]Jg/l) at/l) 1) N h- 1 1- 1 )
Station 38
100 0 34.18 14.31 25.51 1.72 15.44 9.85 0.47 5.57 3.35 3.64 251
50 3 9.87 - 6.84 5.98 3.49 364
25 6 10.22 6.22 6.93 3.48 335
10 8 34.09 13.90 25.52 2.29 16.44 10.82 0.32 6.00 6.21 3.48 261
1 18 34.10 13.70 25.57 11.82 5.38 5.50 3.32 230
Station· 73
100 0 33.99 14.38 25.34 1.43 16.59 13.28 0.46 2.18 1.18 3.38 295
50 3 1.50 16.68 13.67 0.43 3.59 2.18 3.89 254
25 6 5.30 2.89 8.56 379
10 9 33.98 14.01 25.42 1.48 16.47 13.87 0.51 12.34 3.82 196
1 18 34.00 13.90 25.45 1.48 16.61 14.28 0.51 2.60 - 3.44 153
Station 79
100 0 33.94 14.18 25.35 1.22 15.67 6.53 0.53 8.31 4.30 121
50 3 1. 25 15.50 6 .• 12 0.57 15.76 4.78 5.98 141
25 6 1.25 15.64 6.53 0.55 8.08 2.87 4.42 100
10 9 33.94 14.11 25.36 1.26 16.24 6.62 0.54 11.75 4.78 8.08 338
1 18 33.94 14.07 25.37 1.30 15.48 7.60 0.58 1.75 1.67 2.59 129
Station 81
100 0 33.92 14.28 25.31 2.07 51.9 26.38
50 3 1.04 29.2 20.61
25 6 1. 70 30.9 24.20
10 10 33.92 14.26 25.31 2.00 32.6 21.22
1 21 33.86 14.17 25.31 1.92 28.9 25.79
All the stations were taken off the western coast of Baja California during the R/V T.G. Thompson cruise, MESCAL II.
81
A summary of all the hydrographic, chemical, biomass, and metabolic

data from seawater not inhabited by M. ruhrwn and from seawater in-
habited by M. ruhrwn, is presented in Tables 3 and 4. When the hydro-
graphic and chemical data are averaged (Table 5), i t becomes apparent
that M. ruhrwn is associated with relatively high-density, freshly up-
w~lled seawater (crt = 25.38 ± 0.09) with a temperature of 14.28 ± 0.08 0 C
Table 5. Mean hydrographic and chemical characteristics of the surface waters (0 m)

at stations from which M. ruhrwn was found and not found
Stations at which Stations from which M. rubrwn

M. ruhrwn was found could not be found
(Nos. 38, 73, and 79) -(Nos. 67, 71, 78, and 82)
Hydrography:
Salinity (%0) 34.01 ± 0.12 33.83 ± 0.02
Temp. (oC) 14.28 ± 0.08 15.52 ± 0.72
Density (Ot) 25.38 ± 0.09 24.97 ± 0.17
Chemistry:
P04-3 (]lg-at/l) 1.46 ± 0.25 0.60 ± 0.16
Si(OH)4 (]lg-at/l) 15.90 ± 0.61 5.33 ± 3.82
NO"3 (]lg-at/l) 9.89 ± 3.38 2.21 ± 3.39
NH;t (]lg-at/l) 0.49 ± 0.04 0.48 ± 0.11
and a salinity of 34.01 ± 0.12%0. The seawater at other times was char-
acterized by relatively lower density (crt = 24.97 ± 0.17), lower salin-
ity (33.83 ± 0.02%0), and higher temperature (15.52 ± 0.72 0 C). A com-
parison of the nutrient salt concentration wi thin and outside M. ruhrwn
water reveals that the M. ruhrwn water was rich in phosphate (1.5 ± 0.3
]lg-at P04-3 -P/1) , silicic acid [15.9 ± 0.6 ]lg-at Si(OH)4-Si/ll and
nitrate (9.9 ± 3.4 ]lg-at N0 3 -NIL). By contrast the seawater not in-
habi ted by M. ruhrwn was poorer in nutrient salts by a factor of 3 or
4. The ammonium concentration in both types of water was 0.5 ]lg-at
NHt - Nil. The biologic properties of the M. ruhrwn water are averaged
in Table 6. This seawater was distinguished from the surrounding waters
by elevated values of chlorophyll, PN, PC, NR, and ETS. For example,
the PC and PN were twice as high in M. ruhrwn water as outside it -
238 ]lg Cil and 4.1 ]lg-at Nil (within) as compared to 97 ]lg Cil and
2.1 ]lg-at Nil (outside). The chlorophyll (UNESCO) was 5 x higher in
M. ruhrwn water. None of these differences was as great as the carbon,
nitrogen, and chlorophyll differences that Holm-Hansen et al. (1970)
observed in the Bering Sea ciliate blooms. The high value of specific
NR (26.6 ± 2.1 ng-at N0 3 - N h- 1 ]lg chlorophyll-l) is of particular
interest, not only because of its magnitude, but because it provides
additional evidence of M. ruhrwn's autotrophic capacity.
Table 7 presents the results of a search for potential indices of M.

rubrwn. Because the organism is highly motile and because its peripheral
membrane system does not incorporate silicon, i t is possible that the
ratios of PC to PN, nitrate to silicic acid, and silicic acid to PC
are altered in the red-water patches. The N03:Si(OH)4 ratio should
be depressed from differential uptake and should show a continual
decrease with time. The Si(OH)4:PC ratio should be elevated in the
blooms. The ratios of these variables as well as the ratios of chloro-
phyll (UNESCO) to chlorophyll (fluorescence), PC-Chlorophyll (UNESCO),
and PC-chlorophyll (fluorescence) exhibit differences between the M.
rubrwn water and the surrounding seawater, but only the difference in
the PN-chlorophyll (UNESCO) ratio is significant at the 95% level of
82
Table 6. Mean biologic characteristics of the surface seawater inhabited by M. rUbrum
Seawater inhabited by Seawater not inhabited by

M. rUb rum M. rUbrum
(Sts. No. 38, 73, and 79) (Sts. No. 67, 71, 78, and 82)
Chlorophyll:
UNESCO (Jlg/l) 7.0 ± 4.8 1.4 ± 0.7
FLUOR. (Jlg/l) 3.1 ± 1.8 1.2 ± 0.6
PN (Jlg-at/l) 4.1 ± 1.1 2.1 ± 0.5

PC (Jlg/l) 328 ± 92 97 ± 65
NR (ng-at NOS h- 1 1- 1 ) 40.1 ± 16.0 17.1 ± 4.7
ETS (Jll 02 h- 1 1- 1 ) 23.5 ± 4.1 10.6 ± 8.2
NR/Chl-F (ng-at NOS h- 1 26.6 ± 2.1 6.7 ± 3.1
g-l)
ETS/Chl-F (Jll O2 h- 1 1- 1 ) 16.3 ± 5.0 11.4 ± 6.5
Table 7. Potential indices of M. rUbrum
Surface waters Surface waters Station 73, Station 79

containing from other 100% light 50% light level
M. rUbrum stations (Nos. level (M. (M. rUbrum
(Sts. 38, 73, 67, 71, 78, rUbrum com- comprised 50%
79, and 81) and 82) prised 40% of of the popula-
the population) tion)
Chl-u/Chl-F 1.92 ± 0.73 1.24 ± 0.59 1.85 3.3

(6) (7)
PC/Chl-u 54.6 ± 45.9 86.9 ± 46 135 9

(Jl9C/Jlg Chl) (6) (7)
PC/Chl-F 93.3 ± 83.0 80.7 ± 31.0 250 30

(JlgC/Jlg Chl) (6) (8)
PC/PN 5.8 ± 2.5 4.4 ± 2.9 7.27 1.96

(atom/atom) (5) (7)
PN/Chl-U 0.83 ± 0.48 1.41 ± 0.40

(Jlg-at N/Jlg) (5) (6)
N0"3/Si (OH)lf 0.61 ± 0.2 0.42 ± 0.27 0.80 0.39

(5) (5)
Si (OH) If/PC 1.01 ± 0.4 0.74 ± 0.24 0.67 1.32

(Jlg-at/JlgC) (5) (8)
When appropriate, mean values, standard deviations, and the number of measurements
are given. Values are calculated from data in Tables 3 and 4 at the 100% and 50%
light depths.
confidence. The Si(OH)lf:PC ratio is elevated as predicted, but the

N03:Si(OH)lf ratio is also elevated, contrary to prediction. The de-
parture from 1.0 of the ratio between the two chlorophyll measurements
suggests that the chlorophyll methods detect different entities. The
effect of this departure on the chlorophyll:carbon ratio is relatively
83
unimportant outside the M. l'ubl'wn water, but wi thin it there is a 71 %

difference. This could cause a major error in any future prediction
of carbon from chlorophyll measurements made in M. l'ubl'wn blooms. The
nitrogen:chlorophyll ratio in chemostat-grown phytoplankton (Caperon
and Meyer, 1972) decreases with increasing growth rate. The average
ratio in the bloom waters was 0.83 ± 0.48 ~g-at N/~g chlorophyll while
outside the M. l'ubl'wn water it was 1.41 ± 0.40 ~g-at N/~g chlorophyll,
suggesting a relatively high growth rate in the bloom. It should be
remembered that this ratio will be enhanced by nonphytoplankton PN.
This fraction can be determined by calculating the cell nitrogen from
the chlorophyll (Table 6) using the PN:chlorophyll ratio (0.38 ~g-at
N/~g chlorophyll) of Eppley et al. (1971) and Estrada and Vallespin6s
(1975). The calculation gives 2.7 ~g-at Nil and 0.5 ~g-at Nil for the
phytoplankton cell nitrogen wi thin and outside patches of M. l'ubl'wn
water. The nonphytoplankton PN would then represent 34% of the total
PN wi thin the M. l'ubl'wn water and 75% outside it (Table 6).
The effect of incubation on the measurement of C-uptake was investigated
at Station 79. Mesodinium was abundant, comprising 28% of the phyto-
plankton cells at the surfaqe and 53% at the 50% light depth. The ex-
periment was made with samples from each light depth. 14C uptake was
measured after both 6-h and 24-h incubation. At the 100%, 50%, 25%,
10%, and 1% light depths, the results from the 6-h incubation experi-
ment gave rates of 17, 61, 36, 12, and 0.5 ~g C h- 1 1-1, respectively.
The 24-h ex~eriment yielded lower values of 3, 3.5, 2.0, 1.5, and 0.1
~g C h- 1 1- , respectively. These results suggest that cell disinte-
gration depressed the productivity measurements in the 24-h incubation
experiment.
Table 8. Measurements of the standing stock and physiology of M. l'ubl'wn

Station
67-X 73
ETS activity (~l 02 h- 1 1- 1) 652 1450
Nitrate reductase activity 11 5.14
(~g-at N03 -N h- 1 1- 1)
14C uptake (~g C h- 1 1- 1) 1232 468
Particutale organic carbon 16.1 3.67
(mg/l)
Chlorophyll (~g/l) 88.7
Each subsample was taken from the portion of a bucket sample in which the organism
was most highly concentrated.
On two occasions bucket samples from the surface waters of a bloom

were taken (Stations 67X and 73); the results are shown in Table 8. In
the sample from Station 67X the particulat~carbon (PC) concentration
was 16.09 mg/l. The carbon fixation rate was 1.232 mg C h- 1 1-1, mea-
sured over a 30-min incubation period. Based on these measurements
the turnover time for PC was 13.06 h and the carbon uptake velocity
was 0.077 h- 1 indicating that the bloom was healthy and growing rapidly.
The ETS activity and the NR activity were also measured. The ETS activ-
ity was 652 ~l 02 h- 1 1-1. Normally, a sea-surface sample exhibits
activity with a range of 1-10 ~l 02 h- 1 1-1; thus by comparison, this
Mesodinium bloom was catabolically, extraordinarily active. Respiration
can be calculated from this ETS activity. Using OWens and King's (1975)
value of 0.5 for the respiration:ETS activity ratio and Mullin and
84
Brook's (1970) conversion factor between 02 consumption and CO 2 evolu-

tion of 0.43, we arrive at a respiratory C02 production rate of 140
~g C h- 1 1- 1 • The respiratory turnover number, based on this value and
the PC concentration, is 114.9 h. The ratio of carbon fixation to carbon
lost in respiration is 8.8, indicating that the respiration is 11% of
the photosynthesis. If the carbon uptake rates are considered as gross
production rates and corrected for respiratory losses (1232-140
1092 ~g C h- 1 1- 1 ), the carbon turnover time becomes 14.7 h and the
uptake velocity becomes 0.068 h- 1 •
The NR activity was 11 ~g-at N0 3 - N 1- 1 h- 1 , two orders of magnitude

higher than a normal sea-surface NR value. Neither protein nor par-
ticulate nitrogen was measured in this sample, but by assuming a partic-
ulate C:N ratio of 7:1 or 8:1 by weight (Strickland et al., 1969) one
can calculate the PN to be 2.3-2.0 mg/l or 161-142 ~g-at/l. Taking 150
~g-at/l (mean value) as the PN value for the bucket sample, one can
calculate a specific uptake rate for nitrogen of 0.073 h- 1 and a turn-
over time of 13.6 h. Both values are remarkably close to the uptake
rates and turnover time for carbon.
In the second bucket sample (Station 73) M. rubrwn comprised 46% of the
phytoplankton at the surface and 20% at the 10% light level. Since these
are large organisms (40 ~m diameter) their contribution to the com-
munity biomass is even greater than their numbers suggest. The PC in
the sample was 3.67 mg/l, the mean of 4 measurements. The chlorophyll
was measured 3 times, giving a mean value of 88.7 ~g/l. The PC:chloro-
phyll ratio was 41.4, a low value indicating high photosynthetic po-
tential. The 1 4 C uptake was measured (four times) after a 1-h incuba-
tion period, giving a mean value of 468 ~g C h- 1 1-1. Division by the
PC value yielded a gross specific uptake rate of 0.127 h- 1 , which is
higher than the value found at Station 67X. The carbon turnover time
based on this rate is 7.8 h. The carbon assimilation ratio was 5.2 ~g
C h- 1 ~g ChI-I, a high value, but within normal range. The NR and the
ETS activities were 5.14 ~g-at N03 - N h- 1 1-1 and 1450 ~l 02 h- 1 1- 1 ,
respectively. As calculated from the ETS activity the respiratory C02
production was 312 ~g C h- 1 1- 1 • The respiratory turnover number was
11.8 h, much faster than it was at Station 67X. The ratio of carbon
fixed to carbon respired was 1.49 indicating that respiration was 67%
of the photosynthesis. Correcting the carbon productivity for respira-
tion, as was done before, the carbon turnover time and the specific
uptake rate become 24 hand 0.042 h- 1 , respectively. These high values
suggest that the organism were stressed or that their photosynthetic
capacity was impaired. The specific respiration rate was 0.085 h- 1 ;
at Station 67X it was 0.0087 h- 1 , an order of magnitude lower. The PN,
as calculated from the PC, ranged from 32.4-36.9 ~g-at N 1- 1 • Using
this value and the NR level of 5.14 ~g-at N0 3 - N one can calculate
a range for the specific assimilation rate of 0.139 to 0.159 h- 1 • The
mean of these values is within 17% of the gross specific carbon-uptake
rate (0.127 h- 1 ) •
An aqueous extract of a macerated M. rubrwn sample from Station 73

(bucket sample) yielded a dark orange solution after centrifugation.
The absorption spectrum of this pigment at pH 8.0 showed two distinct
peaks at 680 nm and 440 nm with shoulders at 350, 380, and 500 nm,
suggesting that the pigment is the same as the orange pigment that
Parsons and Blackbourn (1968) described. The 440 and the 680 nm peaks
are also suggestive of a-carotene and chlorophyll a (Govindjee and
Braun, 1974).
85
4. Discussion
Our results and those of other authors (Fenchel, 1968b; Taylor et al.,
1969) demonstrate the extraordinary autotrophic and motile capacities
of M. rubrum. They suggest that this ciliate is a nitrate reducer and
thus. a relatively unique member of the animal kingdom. The nitrate-
reducing capacity further suggests that M. rub rum does not need to live
by heterotrophy and/or by phagotrophy because its nitrogen requirements
could be met through auto trophy . Furthermore, if M. rub rum has the ca-
paci ty to reduce NO"3 and if M. rubrum inhabits NO"3-rich seawater, as we
have found (Table 4), then a symbiotic relationship between an autotroph
and a heterotroph becomes less advantageous for the autotroph. This
reasoning thus supports the findings of Taylor et al. (1971) that the
erstwhile symbiotic cryptomonads of M. rubrum are, in fact, only chloro-
plasts and mitochondria, which, although functional, are not associated
with nonciliate nuclei.
The role of M. rubrum in upwelling ecosystems is uncertain, but is po-

tentially important. From its traits of mobility and autotrophy one
can hypothesize about its spatial distribution. In a section across
the continental shelf and slope, the circulation of the seawater in
these ecosystems frequently assumes a series of longitudinal cells
parallel to the coast in which nutrient-rich seawater ascends along
the bottom in one cell and along an overlying pycnocline in a second
cell (Hopkins, 1974; Hagen, 1975; SCOR WG 36, 1975). Upon reaching the
sea surface these upwelled waters are transported offshore, with a
longshore component, until they are forced to submerge beneath ascend-
ing waters of lower density associated with the offshore circulation
cell. During their descent, the waters of the inshore cell maintain
their longshore velocity component so that the stream lines of this
cell resemble a helix. In an upwelling system of this structure, M.
nbrum, because it is mobile and positively phototactic, would be con-
centrated in the inshore converging surface waters of the front between
the cells (Fig. 4). If the upwelling system is comprised of cold-water
Wind
Fig. 4. A hypothetical east-

west cross section through
a coastal upwelling system
showing three circulation
cells. The wind is blowing
to the south (G) inducing
a surface Ekman flO\, offshore
that sinks under the ascend-
ing lighter waters of seaward
cells in frontal regions.
M. rub rum maintains itself
against the descending cur-
rent and is concentrated on
the inshore side of these
fronts (the northern hemi-
sphere situation)
86
•
40
...
0 5 10 15
...
0 5 10 15 III
a:
w
I-
W
::E
:r
I-
, ...
0-
W
0 5 10 15 0 5 10 15 c
•
I o 5 10 15 0 5 10 15
DISTANCE OFFSHORE (Km)
A B
Fig. 51 and II. I, 1-6. Sequential development of various states of an upwelling

system in which blooms of Mesodinium rub rum may develop. Dashed lines represent iso-
pycnals, strippled zones represent nutrient-rich water, solid black patches demark
the blooms of M. rubrum or fast-swimming dinoflagellates, and the black and white
striped patch demarcates a bloom of diatoms and/or dinoflagellates. The continental
margin is located on the left of each panel. (1) A cross section of stratified coastal
waters before the onset of upwelling. To simplify the situation, coastal currents
are neglected. (2) strong upwelling with the incursion of nutrient-rich water into
the euphotic zone, a prebloom condition. (3) Weak upwelling in which the nutrient-
rich water does not break into the euphotic zone. Fast-swimming autotrophs can ex-
ploit this condition by photosynthesizing at the surface during the day and taking
up nutrients in the subsurface waters after their carbohydrate supply has been re-
plenished. (4) An upwelling situation in which a diatom and/or dinoflagellate bloom
is in progress. (5) Weak upwelling situation with low insolation. The surface waters
have been stripped of their nutrients by the previous bloom making the situation
suitable for fast-swimming autotrophs (as in panel No.3). (6) A partially relaxed
upwelling system with an incursion of warm oligotrophic surface waters overriding
previously upwelled water. If the nutrient-rich subsurface waters are within the
87
plumes spreading seaward from small sources (Walsh, 1972), M. ~rum

should occur at the convergence zone on the inside edge of the front
between the cold plume waters and the relatively warm surrounding
waters (Fig. 5). The C. meunieri bloom that Ryther (1967) observed off
the coast of Peru occurred on this side of a front. On other occasions
when the upwelling is weak or relaxed, the water column is strongly
stratified, and the current velocities at the surface are low, M. rUbrum
may bloom by exploiting both the nutrient-rich waters below the pyc-
nocline and the euphotic zone above it (Fig. 5). In this situation
its competitors would be reduced to highly mobile dinoflagellates,
such as GonyauZax polyedra. Because of its potential in exploiting these
condi tions, M. rUbrum may lengthen the time and space domain of photo-
synthesis in an upwelling ecosystem and thus would increase the ef-
ficiency with which the ecosystem cycles carbon, nitrogen, and solar
energy through the biosphere. In effect, it would increase the pro-
ductivity life span with respect to the circulation life span of the
upwelling ecosystem. Furthermore, by exploiting the otherwise unpro-
ductive zones of an upwelling system M. rUbrum would augment the dis-
semination of the products of primary productivity to the surrounding
oceanic ecosystem. To substantiate its potential importance to upwel-
ling ecosystems more effort should be devoted toward understanding
the physiology, biochemistry, and ecology of Mesodinium rUbrum.
Acknowledgments. We thank T. Hopkins for his helpful discussions on upwelling circu-
lation and B. Ives and L. Pukulis for their aid in preparing the manuscript. This
work was supported by NSF grants Nos. DES 75-19025 and GX-33502 (CUEA-12 and 15).
Contribution No. 75018 from the Bigelow Laboratory for Ocean Sciences, west Boothbay
Harbor, Maine 04575.
References
Armstrong, F.A.J., Stearns, C.R., Strickland, J.D.H.: The measurement of upwelling

and subsequent biological processes by means of the Technicon Auto Analyzer and
associated equipment. Deep Sea Res. 14, 381-389 (1967)
Avaria, S.: Observaci6n de un fen6meno de marea roja en la bahia de Valparaiso. Rev.
Biol. Marine Valparaiso 14, 1-5 (1970)
Barber, R.T., White, A.W.,Siegelman, H.W.: Evidence for a cryptomonad symbiont in
the Ciliate, Cyclotrichium meunieri. J. Phycol. 5, 86-88 (1969)
Bary, B.M., Stuckey, R.G.: An occurrence in Wellington Harbour of Cyclotrichium
meunieri Powers, a ciliate causing red water, with some additions to its morphology.
Trans. Roy. Soc. N.Z. 78, 86-92 (1950)
Blackbourn, D.J., Taylor, F.J.R., Blackbourn, J.: Foreign organelle retention by
ciliates. J. Protozool. 20, 286-288 (1973)
Blasco, D.: Composition and distribution of phytoplankton in the region of upwelling
off the coast of Peru. Invest. Pesq. 35, 61-112 (1971)
Caperon, J., Meyer, J.: Ni trogen-limi ted" growth of marine phytoplankton. I. Changes
in popUlation characteristics with steady state growth rate. Deep-Sea Res. 19,
601-618 (1972)
Clemens, W.A.: Red "water bloom" in British Columbia waters. Nature (London) 135,
473 (1935)
daily swimming distance of the autotrophs they can bloom. II, 1, 2. (1) A map showing
the plume-like shape of the isotherms as they intersect the sea surface in a coastal
upwelling situation. (2) A cross section along the A-B transect (from panel No.1)
showing the zones at the edge of the upwelled water that are favorable for blooms of
fast-swimming autotrophs
88
Eppley, R.W., Coatsworth, J.L., Solorzano, L.: Studies of nitrate reductase in marine
phytoplankton. Limnol. Oceanog. 14, 194-205 (1969)
Eppley, R.W., Rogers, J.N., McCarthy, J.J., Sournia, A.: Light/dark periodicity in
nitrogen assimilation of the marine phytoplankters SkeZetonema costatum and Coe-
coZithus huxlei; in N-limited chemostat culture. J. Phycol. !.J 150-154 (1971)
Estrada, M. y Vallespinos, F.: Consideraciones estadisticas sobre algunos parametros
oceanograficos en la region de afloramiento del NW de Africa. Resultados Exp.
Cient. B/O Cornide i, 131-143 (1975)
Fenchel, T.: The ecology of marine microbenthos. III. The reproductive potential of
ciliates. Ophelia 2, 123-136 (1968a)
Fenchel, T.: On "red water" in the Isefjord (inner Danish waters) caused by the
ciliate Mesodinium rubrum. Ophelia 2, 245-253 (1968b)
Fonds, M., Eisma, D.: Upwelling water as a possible cause of red plankton bloom along
the Dutch coast. Neth. J. Sea Res. 1., 458-463 (1967)
Govindjee and Braun, B.Z.: Light absorption, emission and photosynthesis. In: Phys-
iology and Biochemistry, Botanical Monograph, Stewart, W.D.P. (ed.). Berkeley:
Univ. California Press, 1974, 10, pp. 346-390
Hagen, E.: On the question of the cross circulation in the coastal upwelling areas.
CUEA Newsletter i (3), 27-32 (1975)
Hamburger, C., Buddenbrock, W.V.: Nordische Ciliata mit AusschluB der Tintinnoidea.
Nord. Plankt. 12, 1-152 (1911)
Hart, T.J.: Red "water-bloom" in South African seas. Nature (London) .!li, 459-460
(1934)
Holm-Hansen, 0., Lorenzen, C.J., Holmes, R.W., Strickland, J .D.H.: Flurometric de-
termination of chlorophyll. J. Cons. Perm. Int. Explor. Mer. 30, 3-15 (1965)
Holm-Hansen, 0., Taylor, F.J.R., Barsdate, R.J.: A ciliate red tide at Barrow,
Alaska. Marine Biol. 2, 37-46 (1970)
Hopkins, T.S.: The circulation in an upwelling region, the Washington Coast. Tethys.
~ (1-2), 375-390 (1974)
Kelley, J.C., Whitledge, T.E., Dugdale, R.C.: Results of sea surface mapping in the
Peru upwelling system. Limnol. Oceanog. 20, 784-794 (1975)
Lohmann, H.: Untersuchungen zur Feststellung des vollstandigen Gehaltes des Meeres
an Plankton. Wiss. Meeresunters. N.F. Abt. Kiel 10, 129-370 (1908)
Margalef, R.: Estructura y dinamica de la "purga de mar" en la Ria de Vigo. Inv.
Pesq. 2, 113-134 (1956)
Margalef, R.: Composicion y distribucion del fitoplancton marino en la region de
afloramiento del N.W. de Africa, en marzo de 1973 (Campana, Atlor II, del Cornide
de Saavedra). Res. EXp. Cient. B/O Cornide i, 145-170 (1975)
McAlice, B.J.: An occurrence of ciliate red water in the Gulf of Maine. J. Fish.
Res. Bd. Canada 25, 1749-1751 (1968)
Menzel, D.W., Vaccaro, R.F.: The measurement of dissolved organic and particulate
carbon in the ocean. Limnol. Oceanog. ~, 138-142 (1964)
Mullin, M.M., Brooks, E.R.: Growth and metabolism of two planktonic, marine copepods
as influenced by temperature and type of food. In: Marine Food Chains. Steele, J.
H. (ed.). Berkeley: Univ. California Press, 1970, pp. 74-95
Owens, T.G., King, F.D.: The measurement of respiratory electron-transport-system
activity in marine zooplankton. Marine Biol. 19, 27-36 (1975)
Parsons, T.R., Blackbourn, D.J.: Pigments of the ciliate Mesodinium rubrum (Lohmann).
Neth. J. Sea. Res. i, 27-31 (1968)
Pavlou, S.P., Friederich, G.E., Macisaac, J.J.: Quantitative determination of total
organic nitrogen and isotope enrichment in marine phytoplankton. Anal. Biochem.
61, 16-24 (1974)
Powers, P.B.A.: CycZotrichium meunieri sp. nov. (protozoa, ciliata); cause of red
water in the Gulf of Maine. BioI. Bull. ~, 74-80 (1932)
Rasmussen, E.: Systematics and ecology of the Isefjord marine fauna (Denmark).
Ophelia ll, 1-507 (1973)
Ryther, J.H.: Occurrence of red water off Peru. Nature (London) 214, (5095), 1318-
1319 (1967)
SCOR Working Group 36. Report of the second meeting on coastal upwelling processes,
Kiel, F.G.R., 24-28 June 1974. Proc. Sci. Comm. Oceanic Res. 10, 120-128 (1975)
Strickland, J.D.H., Holm-Hansen" 0., Eppley, R.W., Linn, R.J.: The use of the deep
tank in plankton ecology. Limnol. Oceanog. li, 23-34 (1969)
89
Taylor, F.J.R., Blackbourn, D.J., Blackbourn, J.: Ultrastructure of the chloroplasts

and associated structures within the marine ciliate MesodiniW7l rubrW7l (Lohmann).
Nature (London) 224, 819-821 (1969)
Taylor, R.J.R., Blackbourn, D.J., Blackbourn, J.: The red water ciliate MesodiniW7l
rubrW7l and its "incomplete symbionts". A review including new ultrastructural ob-
servations. J. Fish. Res. Bd. Canada 28, 391-407 (1971)
UNESCO: Determination of Photosynthetic pigments in seawater. Monogr. Oceanog.
Methodol. 1-, 1-69 (1966)
Uterm6hl, H.: Neue Wege in der quantitativen Erfassung des Planktons. Verh. Int.
Verein. theor. angew. Limnol. Oceanog. ~, 567-597 (1931)
Walsh, J.J.: Implications of a systems approach to oceanography. Science 176, 969-975
(1972)
Walsh, J.J., Kelley, J.C., Whitledge, T.E., Macisaac, J.J., Huntsman, S.A.: Spin-up
of the. Baja California upwelling ecosystem. Limnol. Oceanog. 19, 553-572 (1974)
Wyrtki, K.: Circulation and water masses in the eastern equatorial Pacific Ocean.
Int. J. Oceanol. Limnol. !, 117-147 (1967)
Regeneration of Nitrogen by Zooplankton and Fish in the
Northwest Africa and Peru Upwelling Ecosystems
T.E. WIDTLEDGE
1. Introduction
The availability of nutrients and light constitutes the dominant con-

trolling factors of the levels of primary production in the ocean. In
the lower latitudes where most coastal upwelling areas are located,
the amount of light is seldom below the critical level to inhibit pro-
ductivity, so nutrients are often the limiting factor in phytoplankton
growth (Koblentz-Mishke et al., 1970). Nutrients utilized in primary
productivity are derived from two sources in upwelling areas. Nutrients
are introduced to the euphotic zone from the depths by the physical pro-
cesses that create upwelling, and nutrients are recycled by biologic or-
ganisms that inhabit the area. Nitrate introduced into the euphotic zone
by upwelling supports "new productivity", while ammonium and other excre-
tory products regenerated by zooplankton and nekton support "regenerated
productivity" (Dugdale and Goering, 1967).
Intuitively one would think that the process of upwelling would provide
the major amount of nutrients used in production processes and that
regeneration would contribute only a small amount. However, estimates
of regenerated and new production using 15N isotope techniques have
shown that recycled ammonium may fulfil nearly half of the daily nitro-
gen requirement of phytoplankton, and upwelled nitrate may provide the
other half (Dugdale and Goering, 1970; Whitledge, 1972). Ammonium is
a preferential source of nitrogen to fulfil phytoplankton nitrogen re-
quirements (Conway, 1973) so in oceanic areas where relatively high
concentrations of ammonium occur the regenerated productivity is nearly
equal to new productivity. The process of upwelling with the high ver-
tical advection provides an efficient mechanism for maintaining a re-
generated nitrogen pool in the euphotic zone even if the organisms
excrete the ammonium at depths deeper than those at which phytoplank-
ton growth occurs. The upwelling process itself returns this ammonium
to the productive euphotic region.
Regeneration studies on zooplankton and nekton have been carried out

in the upwelling areas of Northwest Africa on the Joint I cruise and
of Peru on the PiscO cruise. The two areas contain nekton populations
that can potentially graze on phytoplankton. The Peru upwelling eco-
system is inhabited by the anchoveta, EngrauUs ringens Jenyns, the domi
nant nekton component. Cushing (1971) estimated that the anchoveta
population is 10 times that of all other species in all seasons. The
estimated stocks of anchoveta during the middle 1960s were in the range
of 20 million metric tons. This large biomass of fish therefore is
likely to exert a large influence through grazing and excretion. The
Northwest Africa upwelling area fish are composed of several demersal
and pelagic species. The amount of fish stocks off Northwest Africa
is uncertain but a rough estimate based on catch data (Gulland, 1970)
suggest that they are about an order of magnitude smaller than Peru
stocks.
91
2. Methods
For excretion experiments off Northwest Africa, specimens of Diplodus

senegalensis Cadenat, PageUus couperi Dieuzeide, Cantharus cantharus Lin-
naeus, and Pomadasys incisus Bowdich were captured by a bottom trawl.
The animals were allowed to acclimate for several hours in a holding
tank supplied with surface seawater. Healthy specimens were then trans-
ferred to an experimental tank that had previously been cleaned, rinsed
with ethyl alcohol, flushed, and filled with seawater. Additional ex-
periments were carried out on animals that had been starved for one
and two days. Samples for nutrients, dissolved organic nitrogen, and
urea were collected every 10 min for as long as 3 h. Zooplankton for
the Joint-I experiments were collected with vertical net tows of a
102-~ mesh net and were later fractionated into four size classes.
The Peruvian anchoveta were captured in a 9 m2 dip net attached to the

ship's crane. The specimens were kept in a holding tank for 2 h before
they were used in experiments. Individual fish were placed in one-
liter plastic tubes and incubated in tanks: with circulating surface
seawater. The excretion data were calculated from the change in con-
centration of ammonium in the plastic tubes.
The zooplankton biomass was estimated during the Pisco cruise using
100-m vertical net tows with 240-~ mesh netting (Walsh et al., 1971).
The laboratory studies on Engraulis mordax Girard were conducted in a

32-liter circular chamber similar to that described by Lasker (1970).
The specimens in these experiments had been in captivity for at least
2 weeks. Nutrient samples were collected from the chamber when the
fish were introduced and every 10 min thereafter for 70 min. The ex-
periments were stopped before the oxygen concentration went below 50%
saturation.
3. Northwest Africa
The biomass of pelagic fish stocks in the Northwest Africa upwelling

region estimated using acoustic and trawl methods was 60 g wet wt m- 2 ,
about 20% of the anchoveta biomass off Peru. Thorne et al. (1977) sug-
gest that Sardina pilchardus Walb contributes most of the biomass (50 g
wet weight m- 2 ) over the midshelf area and large concentrations Trachurus
(80 g wet weight m- 2 ) were located at the shelf break and upper con-
tinental slope. The demersal fish stocks were dominated by the family
Sparidae inshore, and at midshelf a mixture of Sparidae, Sciaenidae,
Congridae, and Pomadasidae families were found, having a mean biomass
of 2.2 g wet weight m- 2 (Haedrich et al., 1974). Biomasses for cephalo-
pods and shrimp (Plesionika spp.) were also found to be 0.9 and 1.4 g
wet weight m- 2 , respectively.
The Northwest African fish nutrient excretion rates were approximately

linear with time. The ammonium excretion of freshly caught Diplodus
senegalensis Cadenat ranged from 1.03 to 1.44 ~g N mg dry wt- I day-I.
Specimens starved 1 day had a mean excretion rate of 0.90 and those
starved 2 days had a mean rate of 0.64 ~g N mg dry wt- I day-I. The
highest ammonium excretion rate, 4.6 ~g N mg dry wt- I day-1, was ob-
served for SardineUa (Whitledge, 1972). The urea excretion rates ap-
peared similar in the well-fed and starved specimens. Excretion rates
were measured for some organisms other than fish that were captured in
the trawl nets. The ammonia excretion of cephalopods, determined in a
92
manner similar to that of the fish excretion experiments off Northwest

Africa, had a mean rate of 0.7 ~gN mg dry wt- 1 day-l. The excretion
rates of shrimp were not measured but the rate was estimated to be
about 2 ~g N mg dry wt- 1 day-l based on published excretion rates of
2 ~g N mg dry wt- 1 day-l of Euphausia pacifica Hansen (Jawed, 1969) and
a 2.9 ~g N mg dry weight- 1 day-l excretion rate of the red crab (PZeu-
roncodes pZanipes Stimpson) (Walsh et al., 1977).
Using the geographic distribution of the fish biomass (Haedrich et al.,

1974; Thorne et al., 1977), and excretion measurements for pelagic and
demersal specimens, the regeneration of ammonium by Northwest Africa
fish was calculated for the shelf area « 200 m) to be 3.0 mg-at m- 2
day-l (Table 1) and 1.8 mg-at m- 2 day-l (Table 2) for the offshore
area (> 200 m).
Table 1. Ammonium and nitrate budget for Northwest Africa shelf « 200 m), mg-at
m- 2 day-l
Excretory input
Pelagic fish 2.0

Demersal fish and cephalopods 0.06
Shrimp 0.03
Total 3.0
Zooplank ton 3.9-7.6

Mean zooplankton 5.1
TOTAL AMMONIUM INPUT 8.1
Phytoplankton uptake
Ammonium 11.0
Nitrate 10.0
TOTAL UPTAKE 21.0
Table 2. Ammonium and nitrate budget for Northwest Africa slope (> 200 m), mg-at
m- 2 day-l
Excretory input
Pelagic fish 1.8

Demersal fish 0
Total 1.8
Mean zooplankton 5.4
Ammonium 11.0
Nitrate 19.0
TOTAL UPTAKE 30.0

93
The zooplankton biomass over the shelf in Northwest Africa had a mean
value of 2.7 g dry wt m- 2 , and 10.4 g dry wt m- 2 for the offshore area.
This distribution of zooplankton dry weight biomass is different from
wet weight biomass values (Blackburn, 1976) and may be due to wet weight
to dry weight differences resulting from changes in the taxonomic com-
position. Nutrient excretion studies on the zooplankton captured in
the 102-~ mesh nets were combined with biomass data to estimate zoo-
plankton regeneration over the shelf and the offshore areas. The in-
shore re~ion over the shelf was calculated to have a mean of 5.1 mg-at
m- 2 day- of ammonium regenerated by zooplankton while regeneration
in the offshore area was 5.4 mg-at m- 2 day-I (8mi th and Whitledge,
1977). Of the total estimated ammonium regeneration over the shelf,
zooplankton accounted for 63% of the recycled nitrogen and nekton con-
tributed the remaining 37%.
Phytoplankton uptake of nitrate and ammonium in the shelf region, as

measured by I5N-labeled substrates by Dugdale and MacIsaac (fersonal
communication), was estimated to be 10 and 11 mg-at m- z day- • The re-
lease of 8.1 mg-at NH~-N m- 2 day-I in the shelf area by pelagic animals
produces 73% of the measured ammonium nitrogen uptake and 38% of the
combined nitrate-ammonium nitrogen uptake. Other sources such as benthos
(Rowe et al. r 1977) and bacteria, which are not included in this esti-
mate, apparently supply the remaining 27% of the measured phytoplankton
ammonium requirement.
The size of an individual zooplankter increased offshore of the shelf

as did the total zooplankton biomass. Zooplankton contributed 75% of
the total ammonium regenerated over the slope and the fish contributed
the remaining 25%. Uptake of nitrate and ammonium by phytoplankton in
the offshore region was 19 and 11 mg-at m- 2 day-I (Dugdale and MacIsaac,
personal communication). Offshore the ammonium input can account for
65% of the measured ammonium uptake and 24% of the combined nitrate-
ammonium nitrogen uptake. The difference between phytoplankton uptake
of ammonium and the apparent ammonium production by zooplankton and
fish probably cannot be contributed by the benthos or bacteria because
the water is much deeper over the slope and the euphotic zone is rela-
tively isolated from the bottom compared to the shelf region. Other
zooplankton species not adequately sampled with the 102-~ nets (euphau-
sids, etc.) could produce the difference in the offshore area since
they have been reported in abundance in the area (Casanova, 1974).
4. Peru
The excretion and regeneration studies performed on the Pisco cruise con-
centrated on the Peruvian anchoveta, EngrauUs ringens Jenyns. The mean
ammonium excretion rate for E. ringens was 1. 7 ~g NH~-N mg dry wt- I
day-I. A similar value was obtained for the ammonium excretion of E.
mordax Girard in laboratory studies. The mean Peruvian anchoveta specimen
dry weight was 1.7 g and contained 11.0% of dry weight as nitrogen
(McCarthy and Whitledge, 1972). The mean percent body nitrogen excreted
per day was 9.1.
To assess the significance of regeneration and upwelling as sources of

nutrients for primary production in the Peru upwelling area the quantity
of regenerated nutrients was calculated for comparison with nutrient
demands. The regeneration estimate of anchoveta in the Peru upwelling
region was made by combining fishery catch statistics with nutrient
excretion measurements collected on the Pisco cruise. Using an area of
82,317 km 2 (Wooster and Reid, 1963) and a depth of 40 m for the normal
94
depth ef an ancheveta scheel (Jerdan, 1971), the velume ef the upwelliqg

regien can be cemputed as 3.3 x 10 12 m3 • Ancheveta biemass estimates
fer the Peru regien are all near 20 x 10 6 metric tens (Gulland, 1968~
Ryther, 1969~ Cushing, 1971), but nermal lesses by fishing and bird
predatien weuld decrease the ancheveta biemass to' abeut 12 x 10 6 metric
tens in March 1969 when PiscO' data were cellected. Using these figures
and assuming that the fish are evenly distributed over the entire up-
welling area in a 40-m layer, there weuld be 3.6 g wet wt ef ancheveta
mi 3 at the end ef March. This biemass is abeut 5 times the estimated
pelagic biemass eff Nerthwest Africa. Using a facter ef 0.255 fer wet
weight to' dry weight cenversien (Whitledge, 1972) there are 0.9 g dry
weight fish m- 3 , which is equivalent to' 0.5 fish m- 3 • The ammenium ex-
cretien rate ef 1.7 ~g N mg dry wt- 1 day-l cerrespends to' a regenera-
tien rate ef 4.5 mg-at NH4-N m- 2 day-l in the 40-m layer er 1.9 mg-at
m- 2 day-l in the euphetic zene ef appreximately 17 m depth.
The mean zoeplankten biemasses measured were 39.1 mg dry weight m- 3
fer day samples and 46.1 mg m- 3 fer night samples (Walsh et al., 1971).
These values are similar to' the mean zeeplankten biemass fer Nerthwest
Africa. Zeeplankten ammenium excretien measurements ef 10.4 ~g N mg
dry wt- 1 day-l collected fer Cal-anus ahiZensis Redsky en Piquere cruise
to' Peru (McCarthy, 1971) were used in regeneratien estimates fer zee-
plankten. At' least 50% ef the biemass ef zeeplankten cellected in all
net tews using 100-~ mesh was larger that 505 ~ (University ef Cali-
fornia, 1971). The calculated ammonium regeneratien rates ef zeeplank-
ten using the mean and maximum biemasses were 3.1 and 12.1 mg-at N m- 2
day-l respectively in the upper 100 m and 0.5 and 2.0 mg-at m- 2 day-l
in the upper 17 m. The mean biemass ever the entire 100-m water celumn
was used in the budget table because all the regenerated ammenium weuld
eventually (less than 10 days) enter the euphetic zene as a result ef
vertical advectien.
Zeeplankten biemass samples were net cellected in discrete depth inter-
vals en the PiscO' cruise, sO' previeus estimates ef zeeplankten regenera-
tien in Peru (Whitledge, 1972~ Walsh, 1975), were calculated en a per
unit velume basis. This precedure assumes a hemogeneeus distributien
ef zeeplankton and certainly underestimates the ammenium preductien
by zeeplankten in the euphetic zene.
Nitregen uptake by phyteplankten was estimated using rates ef uptake
ef 15N-labeled ammenium and nitrate (University ef Washingten, 1970).
The mean ammenium uptake measurement frem 24 samples cellected threugh-
eut the study area frem the surface to' the 1% incident light level was
6.5 mg-at NH4-N m- 2 day-I. The mean measured nitrate uptake en statiens
where ammenium was measured was 13.9 mg-at N0 3 -N m- 2 day-I.
The requirements ef primary preductivity and inputs frem regeneratien
ef inerganic nitregen cempeunds are summarized in Table 3. Regeneratien
by zeeplankton and nekten supply 7.6 mg-at m- 2 day-I, which is 117%
ef the 6.5 mg-at m- 2 day-l required by phyteplankten.
The ancheveta and zeeplankten in the Peru upwelling ecesystem centribute
59% and 41% respectively fer a teta1 ef 117% ef the phyteplankten am-
menium requirements, while the nekten and zeeplankten eff Nerthwest
Africa preduced abeut 27% and 46% fer a tetal ef 73% ef the ammenium
required. The much higher nekten biemass and similar zeeplankten bie-
mass in the Peru ecesystem cempared to' the Nerthwest Africa upwelling
ecesystem makes the ancheveta an impertant cempenent in the Peru up-
welling ecesystem (Whitledge and Packard, 1971). Cemparing ecesystems,
the relative ameunt ef primary preductien, supperted by ammenium excre-
tien ef the ancheveta is more than twO' times the ameunt supperted by
Nerthwest African fish. The ammenium input by zeeplankten and nekten
95
Table 3. Nitrogen budget in the euphotic zone of the Peru upwelling region, mg-at
m- 2 day-l
Excretory input
Anchove ta 4 .5
Zooplankton
Upwelled nitrate 23.8
TOTAL INPUT 31. 4
Ammonium 6.5
Nitrate 13.9
TOTAL UPTAKE 20.4
off Peru produce 37% of the combined nitrate-ammonium uptake by phyto-

plankton.
5. Discussion
The large contribution of nutrients to primary productivity in the

Peru upwelling system by the anchoveta implies that the anchoveta must
consume a large portion of the primary production to maintain rapid
growth and excretion rates. If growth and excretion are assumed to be
the two main pathways of assimilated nitrogen, and the other miscel-
laneous losses are small, it is possible to estimate ingestion (Fig. 1)
Assimilation
Ingestion
Fig. 1. Simplified pathway of ingested nitrogen through zooplankton or nekton
as growth and excretion/assimilation efficiency. The anchoveta spawn

during the austral winter in the months June through September and
the young fish enter the fishery at about 6 months of age (Paulik,
1970). The average increase in fish biomass is calculated to be about
1.0 g wet wt m- 3 month-I. Converting to a dai11 basis the anchoveta
nitrogen biomass increases 0.06 ~g-at 1- 1 day- . This 6-month rate was
96
assumed to approximate a linear increase based on von Bertalanffy growth

curves for the anchoveta (Schaefer, 1967) and monthly growth curves for
the Pacific sardine (Sardinops eaerul.ea Girard) (Lasker, 1970). Table 4
presents the calculated daily nitrogen ingestion and per cent primary
production grazed using an 83% assimilation efficiency found for the
Pacific sardine (Lasker, 1970).
Table 4. Daily anchoveta growth, ingestion, and phytoplankton grazing, losses per
unit volume of water in the Peru upwelling region
Peru Pacific Japanese Panama

anchoveta sardine anchovy anchoveta
growth growth ingestion growth
estimate a estimateb estimate C estimated
Anchoveta biomass
j.lg-at N 1- 1 10.8 10.8 10.8 10.8
Daily anchoveta
N growth j.lg-at 1-1 day - 0.06 a 0.15b 0.10 0.08d
day-l
Daily anchoveta total

N excretion j.lg-at 0.67 0.67 0.67 0.67
1-1 day-l
Daily anchoveta
N assimilation 0.73 0.82 0.77 0.76
j.lg-at r 1 day-l
Daily anchoveta
N ingestion 0.88 1.01 0.93 0.91
j.lg-at 1- 1 day-l
Daily phytoplankton
N production 1.63 1.63 1.63 1.63
j.lg-at 1- 1 day-l
Per cent daily

phytoplankton 54.0 60.8 57.1 55.9
N grazed
Per cent anchoveta

body N ingested day-l 8.2 9.2 8.6 c 8.4
Per cent anchoveta

growth day-l 0.56 1.40 0.92 0.77
Per cent growth

efficiency 6.7 15.2 10.6 9.1
(growth/ingestion)
Estimates are based on data from Lasker (1970) , Takahashi and Hatanaka (1960) , and
Smayda (1966) .
Other ingestion estimates were calculated using growth rates of the

Pacific sardine (Sardinops eaerul.ea Girard) (Lasker, 1970) and Panama an-
choveta (Cetengraul.is mystieetus GUnther) (Smayda, 1966). A fourth inges-
tion estimate for the anchoveta was derived from an ingestion estimate
of the Japanese anchovy (Engraul.is japonieus Ternrninck et Schlegel) (Taka-
97
hashi and Hatanaka, 1960; Leong and O'Connell, 1969). The four ingestion
estimates ranged from 0.88 to 1.01 pg-at N 1-1 day-I. These grazing
rates represent 54% to 61% of nitrogen production. The phytoplankton
nitrogen eaten per unit of anchoveta body nitrogen each day was 8.2%
to 9.2%.
The ingestion calculations are very sensitive to the volume of the up-
welling area. The biomass, growth, assimilation efficiency, and excre-
tion rates are comparable to previously reported or commonly accepted
values. However, estimates of the areal extent of the Peru upwelling
region range from 6 x 10 4 to 5 x 10 5 km2 • The three estimates of Wooster
and Reid (1963), Ryther (1969), and Jordan (1971) are all in the range
of 6 to 9 x 10 4 km2 but the area given by Cushing (1971) is more than
5 times that of the others. Jordan (1971) places the anchoveta mainly
within 90 km of the coast with a total upwelling area of 8.6 x 10 4 km2 •
The larger area estimated by Cushing (1971) would increase the upwelling
volume and decrease the anchoveta biomass m- 2 so that the ingestion
rate of the anchoveta per unit volume would be lowered to about 10%
of the daily phytoplankton nitrogen production. This would allow zoo-
plankton to be included as an intermediate step in the food web pro-
vided that phytoplankton productivity remained as high throughout
Cushing's large area as was measured in the nearshore region on the
Pisco cruise. Zooplankton production estimates were not made on the
Pisco cruise, but zooplankton biomass from settling volumes indicates
that the anchoveta could live only one day on zooplankton as its only
source of nourishment. It is unlikely that the regeneration time of
zooplankton would be short enough to supply all the food necessary for
the anchoveta.
From this analysis of ingestion it seems possible that anchoveta could

switch from a phytoplankton to a zooplankton diet as seasonal changes
in the upwelling occur. This corresponds to the more recently available
literature on anchoveta feeding (Rojas de Mendiola, 1971; Rojas de
Mendiola anQ Ochoa, 1973), which can be interpreted that the anchoveta
is really an omnivore in contrast to previous reports that showed that
the anchoveta was probably a herbivore (Rojas de Mendiola, 1958). The
data collected during the first part of the Pisco cruise show the high
nutrient and chlorophyll concentrations confined to within 60 kID of
the coastline. Offshore of that point low open ocean concentrations of
nutrients and chlorophyll were present. This small region of productive
waters would at that time compare favorably with the volume of the up-
welling area used by Ryther (1969) and would concentrate the anchoveta
biomass in the productive nearshore region. With the anchoveta biomass
in a small area, more than 50% of the phytoplankton productivity would
be needed to satisfy their food requirements, so the Peru upwelling
ecosystem is probably a two-step food chain during this time.
During the last part of April on the Pisco cruise high nutrient and
chlorophyll concentrations were measured offshore to at least 100 kID
on the R/V T.G. Thompson and 110 km on the R/V Gosnold; later in the
year high chlorophyll concentrations as far as 260 km offshore were
observed (Guillen and Izaguirre de Rondan, 1973). During the austral
fall or winter the upwelling ecosystem apparently can be as large as
Cushing (1971) suggested, and there might be enough food to support
the phytoplankton-zooplankton-anchoveta food chain. The offshore en-
largement of the high nutrient and high productivity areas that are
associated with seasonal changes in the upwelling system conform with
the model of Walsh (1975), so it appears that both grazing calculations
using excretion data and an ecosystem model predict that the Peruvian
anchoveta can consume both zooplankton and phytoplankton.
98
The similarities of regeneration processes in the Peru and Northwest

African upwelling regions are appa~ent in the data showing that a sub-
stantial percentage of primary production is derived from regenerated
nitrogen even though relatively large amounts of nitrate are introduced
in the euphotic zone by upwelling. The differences, however, revolve
around the organisms responsible for recycling the largest percentage
of nitrogen in each system. In Peru the anchoveta produces the largest
amounts of ammonium while zooplankton and micronekton in Northwest
Africa appear to fulfil that role. This shift in dominance in regenera-
tion of nutrients from fish to zooplankton populations when going from
a low latitude to a higher latitude upwelling area may be another in-
dicator that nekton are replaced by zooplankton as phytophagous orga-
nisms in the more variable environments, as suggested from other anal-
yses (Walsh, 1976).
Acknowledgments. This research was supported by Grant GX 33502 of the National Science
Foundation as part of the International Decade of Ocean Exploration Coastal Upwelling
Ecosystem Analysis (CUEA) program. Preparation of the manuscript was also partially
supported by the United States Energy Research and Development Administration. I
would like to acknowledge the helpful comments by Creighton Wirick and Dr. Maurice
Blackburn and the valuable discussions with Drs. David Cushing and John Walsh con-
cerning the thoughts presented in this paper.
This work was performed, in part, under the auspices of the United States Energy
Research and Development Administration (ERDA).
References
Blackburn, M.: Distribution of zooplankton biomass in an upwelling area off Northwest

Africa: Preliminary results from Expedition Joint I. CUEA Newsletter ~ (1), 13-19
( 1976)
Casanova, J.P.: Biomasse et distribution du zooplankton dans l'upwelling Mauritanien.
Tethys i, 320 (1974)
Conway, H.L.: The uptake and assimilation of inorganic nitrogen by Skeletonema costa-
tum (Grev.). ph.D. Thesis. Seattle: Univ. of washington, 1973, p. 125
Cushing, D.H.: Upwelling and the production of fish. Advan. Marine BioI. ~, 255-334
(1971)
Dugdale, R.C., Goering, J.J.: Uptake of new and regenerated forms of nitrogen in
primary productivity. Limnol. Oceanog. 11, 196-206 (1967)
Dugdale, R.C., Goering, J.J.: Nutrient limitation and the path of nitrogen in Peru
Current production. In: Anton Bruun RepOrt No.4. Texas: A & M Press, 1970, pp.
5.3-5.8
Guillen, 0., Izaguirre De Rondan, R.: Distribution of Chlorophyll a in the Peru
coastal current. In: Oceanography of the South Pacific 1972. Fraser, R. (ed.).
Wellington: New Zealand National Commission for UNESCO, 1973, pp. 387-395
Gulland, J.A.: Population dynamics of the Peruvian anchoveta. FAO Fish. Tech. Pap.
72, 29 pp. FAO Rome (1968)
Gulland, J.A.: The fish resources of the ocean. FAO Fish, Tech. Pap. 97, 425 pp.
FAO Rome (1970)
Haedrich, R.L., Blackburn, M., Brulhet, J.: Distribution and biomass of trawl-caught
animals off Spanish Sahara, West Africa. Friday Harbor Joint I Working Conference.
21 pp. (1974) (Unpubl. manuscript)
Jawed, M.: Body nitrogen and nitrogenous excretion in Neomysis rayii Murdoch and
Euphausia pacifica Hansen. Limnol. Oceanog. l!, 748-754 (1969)
Jordan, R.: Distribution of anchoveta (Engraulis ringens J.) in relation to the en-
vironment. Invest. Pesq. 35, 113-126 (1971)
Koblentz-Mishke, O.J., Volkovinsky, V.V., Kabanova, J.G.: Plankton primary production
of the world ocean. In: Scientific Exploration of the South Pacific. Wooster, W.S.
(ed.). Nat. Academy Science, 1970, pp. 183-193
99
Lasker, R.: Utilization of zooplankton energy by a Pacific sardine population in the

California Current. In: Marine Food Chains. Steele, J.H. (ed.). Berkeley: Univ.
California, 1970, pp. 265-284
Leong, R.J.H., o'Connell, C.P.: A laboratory study of particulate and filter feeding
of the northern anchovy (EngrauZis mordax). J. Fish. Res. Bd. Canada 26, 557-582
(1969)
McCarthy, J.J.: The role of urea in marine phytoplankton ecology. Ph.D. Thesis. San
Diego: Univ. California, 1971, p. 165
McCarthy, J.J., Whitledge, T.E.: Nitrogen excretion by anchovy (EngrauZis mordax
and E. ringens) and jack mackerel (Trachurus symmetricus). Fish. Bull. 1QJ 395-401
(1972)
Paulik, G.J.: The anchoveta fishery of Peru. Center for Quantitative Science in
Forestry, Fisheries and·Wi~dlife. Seattle, Univ. Washington, 1970, Pap. 13, p. 79
Rojas de Mendiola, B.: Breve estudio sobre la variacibn cualitativa anual del planc-
ton superficial de la Bahia de Chimbote. Bol. Cia. Adm. Guano ~ (12) 7-17 (1958)
Rojas de Mendiola, B.: Some observations on the feeding of the Peruvian anchoveta
EngrauZis ringens J. in two regions of the Peruvian coast. In: F~rtility of the
Sea. Costlow, J.D. (ed.). New York: Gordon and Breach Science Publ., 1971, pp.
417-440
Rojas de Mendiola, B., Ochoa, N.: Observations on the food and feeding habits of the
anchovy EngrauZis ringens Jenyns made during cruise 6908-09. In: Oceanography of
the South Pacific. Fraser, R. (ed.). Wellington: New Zealand National Commission
for UNESCO, 1913, pp. 373-397
Rowe, G. T., Clifford, C.H., Smith, K.L.: Nutrient regeneration in sediments off Cap
Blanc, Spanish Sahara. Deep-Sea Res. 24, 1, 57-63 (1977)
Ryther, J.H.: Photosynthesis and fish production in the sea. Science 166, 72-76
(1969)
Schaefer, M.B.: Dynamics of the fishery for the anchoveta EngrauZis ringens, off
Peru. Boletin Instituto Del Mar Del Peru 1, 192-303 (1967)
Smayda, T.J.: A quantitative analysis of the phytoplankton of the Gulf of Panama III.
General ecological conditions, and the phytoplankton dynamics at 80 45'N, 79 0 23'W
from November 1954 to May 1957. Inter-Amer. Trop. Tuna Comm. Bull. 11, 355-612
(1966)
Smith, S.L., Whitledge, T.E.: The role of zooplankton in the regeneration of nitrogen
in a coastal upwelling system off Northwest Africa. Deep-Sea Res. 24, 1, 49-56
(1977)
Takahashi, M., Hatanaka, M.A.: Experimental study on utilization of food by young
anchovy EngrauZis japonicus Temminck et Schlegel. Tohoku J. Agr. Res. 11, 161-170
(1960) -
Thorne, R.E., Mathisen, O.A., Trumble, R.J., Blackburn, M.: Distribution and abun-
dance of pelagic fish off Spanish Sahara during CUEA expedition. Deep-Sea Res.
24, 1, 75-82 (1977)
University of California, Institute of Marine Resources: Research on the marine food
chain; progress report July 1970-June 1971 Part III A Data record, cruise Piquero,
leg 7, section 2, the plankton p. 126. Scripps Inst. Oceanog. (1971) (Unpubl.
manuscript IMR 71-10)
University of Washington: Biological production in upwelling ecosystems. Pisco data
report. Part I: hydrography and productivity 97pp. Dep. Oceanog. (1970) (Unpubl.
manuscript special report 42)
Res. 22, 201-236 (1975)
Walsh, J.J.: Models of the sea. In: The Ecology of the Seas. Cushing D.H., Walsh,
J.J. (eds.). Oxford: Blackwell Scientific Publ., 1976, pp. 388-407
Walsh, J.J., Kelley, J.C., Dugdale, R.C., Frost, B.W.: Gross features of the Peruvian
upwelling system with special reference to possible diel variation. Invest. Pesq.
35, 25-42 (1971)
Walsh, J.J., Whitledge, T.E., Kelley, J.C., Huntsman, S.A., Pillsbury, R.D.: Further
transition states of the Baja California upwelling ecosystem. Limnol. Oceanog. 22,
264-280 (1977)
Whitledge, T.E., Packard, T.T.: Nutrient excretion by anchovies and zooplankton in
Pacific upwelling regions. Invest. Pesq. 35, 243-250 (1971)
100
Whitledge, T.E.: Excretion measurements of nekton and the regeneration of nutrients

near Punta San Juan in the Peru upwelling system derived from nekton and zooplank-
ton excretion. PH.D. Thesis. Seattle: Univ. of Washington, 1972, p. 115
Wooster, W.S., Reid, J.L.: Eastern boundary currents. In: The Sea. Hill, M.N. (ed.).
New York: Interscience, 1963, Vol. 2, pp. 253-288
Upper Trophic Levels in Upwelling Areas
D.H. CUSHING
1. Introduction
In an earlier paper (Cushing, 1971), an analogy was drawn between the

product~on cycle in upwelling areas and that in temperate waters. Pro-
duction stops in winter in middle and high latitudes and, when the
spring increase gets under way, the algae grow more rapidly than do
the herbivores because the herbivores do not start to reproduce until
there ia enough algal food available. The delay between the start of
algal production and the appearance of sufficient grazing capacity to
reduce it affects the amplitude and spread of the production cycle al-
though both are modified by other factors, for example, the rate of
change of the algal reproductive rate. In contrast to the spring out-
burst in temperate waters, that of low latitude seas is continuous and
of low amplitude because the delay period may be so short as not to
exist at all.
In upwelling areas, the rising water originates from a depth of about

200 m. Because the smaller herbivores do not migrate below about 100 m
(Cushing, 1951) the upwelling copepods are adults. As the water rises
in the euphotic layer, the rate of production increases exponentially
much as i t does in temperate waters during the spring outburst. Hence,
there is a delay between the production of algae and that of herbi-
vores, which is why the production cycle in upwelling areas resembles
that in higher latitudes.
2. Structure of the Upper Trophic Levels
In a valuable paper, Ursin (1973) showed that the predator/prey ratio

in weight is 100:1 (or 4.6:1 in length). A completely predatory food
chain might be arranged in length ratios as follows:
100.0 cm Cod Tuna

21.4 cm Herring Sardine
4.6 cm Young sandeel Myctophid
1.0 em Euphausid Sagitta
0.2 cm Cal-anus Cal-anus
Within this structure, some real predatory food chains may be discerned
but herring or sardines may eliminate a step by filtering their food
from the water.
Each predator takes its prey at its burst speed (of about 10 lengths/s)
and the prey tries to escape, also at burst speed. Attack speed is
greater than cruising speed by a factor of at least 3.3 and if the
length difference between predator and prey is greater than 15 (4.6 x
3.3), attack is not needed; -herring of 25-30 cm in length can take
Cal-anus of 0.2-0.3 cm in length without needing to accelerate. However,
102
to obtain an adequate daily ration many more encounters are needed. A

cod of 100 cm in length may need up to 3% body weight/day, which is
3 whiting of about 20 cm in length. A herring of 25 cm in length weigh-
ing 100-150 g may need up to 5% body weight/day, or more than 1,000
CaZanus/day (Savage, 1937; Cushing, 1964). A large proportion of the
day available for feeding must be spent in filtering because Battle
et al. (1936) established that the herring visually select their food.
A filterer uses its white muscle for escape only, whereas a predator
(at an intermediate trophic level) uses i t for attack as well. If her-
ring eat euphausids, as they do, they must attack, and if cod eat
euphausids, as they do, they do not need to.
To take a thousand CaZanus/day, the herring must eat one every 1 1/2
min and their cruising speed must be restrained. The reduced speed
generates the aggregations by which the herring find their food. If
they were distributed evenly such speed restriction would make the
filterers more vulnerable to predators. So they may shoal to obtain
the greatest quantity of food at the least risk of predation. Table
is a summary of the differences between a predator and a filterer.
Table 1. Characteristics of filterer and predator
Filterers Predators
Encounters/day 10
(Predator/prey) in weight 1~
Speed difference
(predator/prey) 15 or more About 5:1
Attack speed needed No Yes
White muscle used For escape only For attack and escape
Pelagic feeding Aggregate in feeding Follow aggregations
Shoaling? Yes No
There is a group of animals that may be described as top filterers

(analogous to top predators) such as baleen whales, basking sharks, and
manta rays. They are larger than any predator, they all have few off-
spring (which implies that they do not depend on the predatory death
of numerous offspring to stabilize their numbers), and they do not
have to maintain an attack or escape speed for any reason. Their daily
ration depends on food density and the product of baleen area (or its
equivalent) and speed. Filterers like herring have to devote a pre-
dominant part of their body weight to escape (or sometimes for attack).
The top filterers can grow enough to carry disproportionately large
filters.
It is no accident that filterers are found in upwelling areas and in

those moderate and high latitudes where the density of food organisms
allow high encounter rates. In the deep ocean, where the food particles
are dispersed, fish do not shoal, there are no top filterers, and the
food chain has q more predatory character. Ryther (1969) suggested that
there were probably five links in the food chain in the oceanic province.
By using a filtering habit, herring and sardines eliminate one link
from a purely predatory chain and so there should be fewer links in
upwelling areas and in temperate latitudes than in the deep ocean.
103
From a literature survey, Longhurst (1971) suggested that the sardine-

like fishes were predominantly phytophagous, which allowed Ryther to
explain the reduced number of links in the food chains of upwelling
areas at least as compared with the fully predatory chain in the deep
ocean. If the reduced number of links can be explained by the filtering
habit, the question arises whether the sardine-like fishes are really
phytophagous.
3. Do Some Clupeoids Feed Predominantly on Diatoms and Dinoflagellates?
There are three methods of estimating the quantity of food in the guts
of fishes from the gut contents: (a) the method of occurrences, in
which the number of identified organisms is recorded; (b) the points
method, in which the number is weighted by arbitrary sizes (effectively
surfaces) under microscopic examination; (c) the method of gut volumes,
in which the number of identified organisms is weighted by thei.r vol-
umes.
Most sardines and sardinellas have a finer mesh of gill rakers than do
the herring-like fishes, which means that if they pursue the smaller
copepods and the larval plankton animals, as they do, they are bound
to collect the larger diatoms on the gill rakers; the longer gut (Yama-
shita, 1957) of SClX'dinops meZanostiata Ternrninck and Schlegel suggests
that the phytoplankton collected in this way is in fact used, as in
the truly phytophagous animals such as Brevoortia and CetengrauZis mysti-
aetus GUnther. Yoshida (1955) says that Sardinops meZanostiata strains
the plankton by swimming with an open mouth.
An argument on the feeding of Sardinops aaeruZea Girard was conducted many

years ago. Lewis (1929) examined 208 fish in each month in the region
of San Diego; he compared numbers per stomach with numbers in the water
and concluded that the sardine fed on copepods as well as on algae.
Parr (1930) used Lewis's material to show that although 7.35 1 were
filtered to obtain the diatoms, 64.66 1 were filtered to obtain the
dinoflagellates; hence the sardines selected the dinoflagellates in
preference to the diatoms. The maximum range of variation for the dia-
toms in the guts was 1:8.10 5 , that for the dinoflagellates was 1:750,
and for copepods it was 1:87. Parr concluded that the sardines sustained
an interest in copepods and therefore collected the algae inadvertently.
Hart and Wailes (1932) examined the food of Sardinops aaeruZea of British
Columbia between 1927 and 1930 by estimating the occurrence of copepods
and algae in the guts; from 275 stomachs 13.5-36.6% of occurrences
were copepods. Because the method of occurrences was used, their con-
clusion was that the animals fed predominantly on algae. However, the
copepods are 3 to 4 orders of magnitude larger than the biggest diatoms,
and the proper conclusion is almost certainly that the fish fed pre-
dominantly on copepods. Radovich (1952) carne to the latter conclusion,
but it was Hand and Berner (1959) who put the point properly, as shown
in Table 2.
Yamashita (1957a) examined the gut contents of 1315 fishes of the spe-
cies Sardinops meZanostiata and tabulated the quantities (Tables 3 and 4).
In numbers, there are more animals than algae in the sardines' guts,
and when the weights of the animals eaten are taken into account the
food must be considered predominantly zooplanktonic. The same conclu-
sion emerges from Yarnashi ta" s analysis of the gut contents of the Ja-
panese anchovy EngrauZis japoniaa Houttuyn.
104
Table 2. Food of Sardinops caeruZea (571 fishes), (Hand and Berner, 1959)
Average number Organic matter

(mg dry weight)
Diatoms 1.14, 10 1.77

Dinoflagellates 3.30, 10 0.70
Small copepods 6.70, 10 25.64
Large copepods 20 3.40
Euphausiids 2 1.80
Chaetognaths 9 0.90
Fish eggs 7 0.70
Table 3. Gut contents of Sardinops meZanosticta (Yamashita, 1957b)
February March April May June July
Diatoms 2 3 3
Dinoflagellates 2 2
Tintinnids 3 1
Copepods 2 3 2 2
Cladocera 2 1
Schizopoda
Amphipoda
Larval plank ton 2 2
r,lacroplank ton
1 = few; 2 = moderate number; 3 abundant.
Table 4. The weight of gut contents of SardineZla longiceps (from Kagwade, 1964)
Allocated Weight in mg by age group

sizes
I II III
Biddulphia 1.10 5 \13 0.53 0.58

Ditylum 1.10 5 0.03 0.25
Bac terias trum 1.10 5 0.08 0.32
Planktoniella 1.10 5 0.13 0.38
Aulacodiscus 1.10 5 0.76 0.60
Coscinodiscus 1.10 5 18.10 46.95
Thalassiosira 5.10 3 0.17 0.91
Skeletonema costatum 2.10 2
Fragilaria oceanica 0.5.10 3 13.42 6.34
Thalassionema nitzschoides 1.5.10 3
Tha ls siothrix 1.10 3 7.70 9.10
Pleurosigma 5.10 2 0.01 0.02
Nitzschia 5.10 2 0.10 0.10
Surirella 5.10 2
Trichodesmium 5.10 2 0.02
Proromicans 1.10 3
Dinophysis 1.10 5 1.89 5.68
Glenodinium 1.10" 0.04 0.14
Peridinium 1.10" 0.05 0.12
Geratium 1.10 5 0.39 0.90
Tintinnids 1.10 5 0.34 0.72
43.74 73.13
105
Table 4. Continued
Allocated Weight in mg by age group

sizes
I II III
Copepods 5 llg 69.50 137.00

Evadne 20 llg 14.00 10.00
Crustacean eggs 1 llg 1.56 1.00
Nauplii llg 1. 70 1.30
Zoae 50 llg 5.00 10.00
Bivalve larvae 20 llg 2.00 15.00
l: 2.00 106.76 159.30
A number of authors have examined the gut contents of the Indian oil
sardine, Sardinella longieeps Valenciennes (Venkataraman, 1960; Dhulkhed,
1962; Bensam, 1964; Kagwade, 1964; Noble, 1965; and a number of earlier
writers). Kagwade's paper gives numbers of food items in the guts by
genera (usually), by months, and by age groups in a large number of
fishes. Certain months were chosen when the guts were full (February,
May 1959; August to October, 1959; May to July 1960; April and August
1961). The sizes of organisms cannot be allocated very well because
they are classed as, for example, Coseinodiseus spp, copepods (which
could include any species or developmental stage). The sizes allocated
are given in Table 4. Sizes of the algae were taken from Lebour (1930)
on the assumption that cells in upwelling waters were the same size
as those in temperate waters. Weights of animals were taken from Bogo-
rov (1959) and scaled down by an order of magnitude; e.g., "copepods"
were one tenth of the weight of Psewioealanus.
The total estimated quantity during a period of full stomachs was about
0.15-0.23 g when the observed average quantity was 0.29 g; the sizes
of animals have probably been underestimated by about one-third (algal
sizes are probably about right). The quantity of animal food in the
guts was greater than the quantity of algal food by a factor of 3 to
4.
De Mendiola (1971) examined the food of Engraulis ringens Jenyn, the Peru-
vian anchoveta, and showed that numbers of algae predominated in samples
from north of Chimbote. When the numbers of algae were raised by their
sizes, the average quantity in each gut was 0.065 mi. Weights for length
were taken from Jordan (1974) and used to construct Table 5.
Table 5. The daily ration in ml/day of the anchoveta Engraulis ringens
Length Weight Ration as % body weight per day

1% 3% 5%
10.0 cm 7.0 g 0.07 0.21 0.35

12.5 cm 13.2 g 0.13 0.40 0.66
15.0 cm 24.2 g 0.24 0.72 1.21
The daily ration as percentage of body weight for a fish as small as

the anchoveta is likely to be nearer 5% than 1%. If the mean length
were 12.5 cm, a reasonable daily ration would be 0.66 ml, which is 10
times the weight of algae in the guts. In numbers, the quantities of
algae in the guts of the anchoveta do not differ much from the algal
quantities in other sardine-like fishes that feed on copepods. In the
106
samples taken from south of Chimbote, there were many copepods in the
guts. However, in the samples from north of Chimbote there were hardly
any copepods in the guts. The simplest conclusion is that the anchoveta
north of Chimbote were not feeding avidly where in fact most of the
spawning occurs.
Davies (1957) conducted a thorough investigation of the food of the
South African pilchard (Scwdinops oaeHata Pappe) using the points method
(i.e., weighting the numbers by the surfaces of food items); 1664 fish
were examined for a period of four years. The main result is shown in
Table 6.
Table 6. The proportion of zooplankton in the guts of the South African pilchard
Sardinops oaeHata
% Zooplankton Fullness index (out of 10)
January 62 6
February 39 4
March 22 5
April 29 3
May 31 2
June 35 3
July 64 2
August 24 3
September 10 4
October 8 3
November 45 3
The period of most intense feeding was in January to March. Davies'

conclusion was that the proportion of phytoplankton to zooplankton in
the guts was 2:1. If the proportions were weighted by volume rather
than by surface, as suggested by the points method, then the proportion
of zooplankton in Table 6 would be much higher.
If the sardine-like fishes did feed predominantly on algae, they would
have to swim very slowly (0.2 lengths/s, Yoshida, 1955) to strain the
algae from the water efficiently~ Such a speed would be an order of
magnitude slower than the normal cruising speed of 1-3 lengths/so The
filtering area of the Japanese sardine is 7.2 cm 2 (Yoshida, 1955), so
the fish would filter about 1 limine Very dense quantities of algae
amount to about 1 mg/l, which means that the fish would have to filter
for 8-16 hid to obtain 0.5-1.0 g/day. Such fishes would have to shoal
to avoid predation, and the shoals would have to be large and dense at
such low speeds. In fact, the Peruvian anchoveta is dispersed in the
surface layer day and night in the major spawning area north of Chim-
bote (Jordan, 1974), perhaps only feeding lightly, as suggested from
Table 5.
There is one phytophagous clupeoid CetengruuUs mystiaetus, which lives
in the gulfs of the eastern tropical Pacific. Its food consists pre-
dominantly of diatoms, which are crushed in a gizzard-like organ. In
this respect, it resembles the menhaden (Brevoortia tyrannus Latrobe),
which also feeds on algae in the silt on the bottom. However, the
clupeoids without gizzards have no ahatomic adaptation for crushing
diatoms. Another argument for phytophagy concerns the length of gut;
an abnormally long gut is needed to digest algae. The guts of animal-
feeding clupeoids are about half the length of the body; those of the
supposedly phytophagous ones such as sardines and anchovies are about
the same as the body length (Yamashita, 1957b): that of the menhaden
107
is 4 to 5 times the body length (Browne Goode, 1884), so the clupeoids

of upwelling areas are not as well adapted to eating algae as the men-
haden because they lack gizzards and have shorter guts. The question
asked is: Do some clupeoids feed predominantly on diatoms and dino-
flagellates? The answer is that the sardine-like fishes in the upwel-
ling areas probably feed predominantly on small copepods and planktonic
larvae although during the period of first feeding they probably depend
completely on algae (reference from Tim Wyatt). Larger diatoms and
dinoflagellates are taken inadvertently in the act of feeding (and are
made use of, witness the somewhat lengthened gut). If the answer is
right, Ryther's (1969) conclusion that a link in the food chains in
upwelling areas was eliminated by phytophagous clupeoids was wrong.
4. Upper Trophic Levels in Upwelling Areas
If the sardine-like fishes in upwelling areas feed predominantly on

·copepods the resemblance between the upwelling ecosystem and that of
temperate waters is strengthened. In temperate waters the predominant
fish are cod', herring, mackerei, and sprat. The "cod" represents a num-
ber of predatory gadoids of different sizes, ling, saithe, and blue
whiting in deep water over the edge of the shelf, cod and haddock on
the top of the shelf, and whiting in shallower water. The three pelagic
species also live in different depths of water, mackerel at the shelf
edge, herring on the top of the shelf, and the sprat inshore. In the
major upwelling areas, there is only one major gadoid, the hake, which
lives inshore when young and beyond the shelf edge when adult. The
continental shelves in the major upwelling areas are usually rather
narrow and one gadoid species occupies it; in the Atlantic north of
40 0 N the shelves are much broader and there is more than one species.
There are three pelagic species in the upwelling areas: jack mackerel,
sardines, and anchovies; like the mackerel, the jack mackerel tends
to live at the shelf edge or beyond it, but the other two species live
further inshore and may even spawn in the upwelled water. The major
fishes in temperate waters and in upwelling areas may be grouped as
follows:
Temperate Upwelling
"Cod" Hake
Mackerel Jack mackerel
Herring Sardine; anchovy
Sprat
Whiting
Haddock Cod
,p<!
r ."....
Hmi.J\aCkerel
"'.... ~"i I , . / /'"
,
I ~
1-I
I
smal];. ~ ~ Largel,,/ Euphausids
Copepods Copepods
---------------Depth-------------)~ Fig. 1. The temperate food chain
The pelagic fish feed on zooplankton. Sprats feed on small copepods

such as Aacwtia or PseudoaaZanus, herring feed on CaZanus, as do mackerel,
108
Hake Fig. 2. Food chain in upwel-
~
ling areas
Anchovy \ Jack
;rdin\ __ __ _ __ _ ;-;,. Mackerel
La~
Copepods
\
Copepods
Large ---
Copepods
I
Euphausids
---------------Depth----------------------------?>
and both can feed on euphausids, as indeed do cod. Anchovies and sar-
dines feed on larval copepods (as well as phytoplankton inadvertently) ,
and the jack mackerel feed on copepods as well as on euphausids from
time to time.
5. Discussion
On the basis .of some of the papers quoted here, Longh urs t (1971) con-
cluded that the sardine-like fishes in upwelling areas were primarily
phytophagous and Ryther (1969) suggested that the high fish production
there was due to the shortened food chain. Implicit in the argument was
the fact that the fishery for the Peruvian anchoveta was much larger
than any other because the fish appeared to feed only on algae.
In this paper an analogy has been drawn between production in temperate

waters and that in upwelling areas. Before they were fished out, the
stocks of herring in the North Sea and in the Norwegian Sea each yielded
anrrually up to 1.0 or 1.5 million tons, respectively; the total catches
of Hokkaido herring and the Japanese sardine would have amounted to
2 or 3 million tons had they been exploited at the same time. Let us
assume that the area of production in which the herring feed is about
100,000 km 2 , both in the western North Sea and off the polar front in
the Norwegian Sea. It might be a reasonable area to apply to the Peru
upwelling area. The production cycle supporting the 2 groups of fishery
in the eastern Atlantic and in the western Pacific is the typical tem-
perate outburst that lasts for 2 to 3 months at the most.
In quantity produced, the most important pOint about the production

cycle is not the intensity of production in gCjm 2 jday, but its duration
(Cushing, 1971). The intensity in upwelling waters and in temperate
seas ranges from 1 to 3 gCjm2jday, but the durations in different re-
gions vary considerably. The period of upwelling off Peru lasts for a
long time, perhaps for 8 or 9 months. Perhaps this is why catches off
Peru are 3 to 4 times greater than those of pelagic fishes in temperate
waters where the production cycles last for only 2 or 3 months. Catches
off California and in the Canary Current are less, perhaps because the
periods of upwelling are shorter and in themselves tend to be inter-
mittent.
There remains an important point: north of Chimbote, the Peruvian an-

choveta feeds lightly on algae, and Whittledge (this meeting) has model-
led the food chain processes on this basis. Also larvae of such clupe-
oids use algae as their first food. The anchoveta spawns in areas of
phytoplankton on which they feed; when the resulting larvae start to
feed they feed on the algae and then probably progress to zooplankton,
which emerge from the phytoplankton patches. Thus the observed feeding
of the spawning clupeoids on algae allows the larvae and growing post
larvae to be in the right place to exploit the smaller zooplankton.
109
References
Battle, H.I., Huntsman, A.G., Jeffers, A.M., Jeffers, G.W., Johnson, W.H., McNairn,
N.A.: Fatness, digestion and food of Passamaquoddy young herring. J. Biol. Bd Can.
2, 401-429 (1936)
Bayliff, W.H.: The food and feeding habits of the anchoveta Cetengraulis mystiaetus
in the Gulf of Panama. Bull. Inter-Amer. Trop. Tune Commn. 7, 399-459 (1963)
Bensam, P.: Differences in the food and feeding adaptations b~tween juveniles and
adults of the Indian Oil sardine, Sardinella longiaeps Velenciennes. Indian J.
Fish. A. 11, 1, 377-390 (1964)
Bogorov, B.G.: On the standardisation of marine plankton samples. Int. Rev. Hydrio-
biol. 44, 621-642 (1959)
Brown Goode, G.: The Fisheries and Fishery Industries of the United States. I. Natural
History of Useful Aquatic Animals, pp. 895 (1884)
Cushing, D.H.: The vertical migration of planktonic crustacea. Biol. Rev. 26, 158-192
(1951)
Cushing, D.H.: The work of grazing in the sea. In: Grazing in Terrestrial and Marine
Environments. Crisp, D.J. (ed.). Oxford: Blackwell, 1964, pp. 207-225
Cushing, D.H.: upwelling and the production of fish. Advan. Marine Biol. ~, 255-334
(1971)
Davies, D.H.: The Sout African pulchard (Sardinops oaellata) . Preliminary report on
feeding off the West Coast, 1953-1956. Div. Fish. Invest. Report, S. Africa 30,
40 (1957)
Dhulked, M.H.: Observations on the food and feeding habits of the Indian Oil sardine,
Sardinella longiaeps Val. Indian J. Fish. A. 9, 1, 37-47 (1962)
Fish, G.R.: The food of Tilapia in East Africa.-uganda. J. 19, 85-89 (1955)
Hand, C.H., Berner, L.: Food of the Pacific sardine (Sardinops aaerulea). F. U.S.
Fish Wildlife Servo Fish Bull. 60 (No. 164) 175-184 (1959)
Hart, J.L., Wailed, G.H.: The food of the pilchard, Sardinops aaerulea (Girard) off
the coast of British Columbia. Contrib. Can. Biol. Fish. 7, 247-254 (1932)
Hardy, A.C.: The herring in relation to its animate enviroTI;ent. I. The food and
feeding habits of the herring with special reference to the East Coast of England.
Fishery Invest. London Ser. 2, 7, 3, 53 (1924
Jager, B. de: Synopsis on the biology of the South African pilchard, Sardinops oael-
lata (Pappe). Proc. World Sci. Meeting. Biol. Sardines, F.A.O. Fish. Biol. Symposes.
7, 97-114 (1960)
Jo;dan, R.: Biology of the Anchoveta. I. Summary of our present knowledge. lDOE
Workshop El Nino Guayaquil, p. 21 (1974) (Draft publ.)
Kagwade, P.V.: The food and feeding habits of the Inidan Oil sardine, Sardinella
longiaeps Valenciennes. Indian J. Fish. A. 11, 1, 345-370 (1964).
Lebour, M.V.: The planktoniC diatoms of northern seas. Ray Soc. p. 244 (1930)
Lewis, R.C.: The food habits of the California sardine in relation to the seasonal
distribution of microplankton. Bull. Scripps Instn Oceanog. Tech. Ser. l, 155-180
(1929)
Longhurst, A.R.: The clupepod resources of tropical seas. Oceanog. Marine Biol.
Ann. Rev. ~, 349-385 (1971)
Mendiola, B.R. de: Some observations on the feeding of the Peruvian anchoveta Engrau-
lis ringens J. in two regions of the Peruvian coast. In: Fertility of the Sea.
Costlow, J.D. (ed.). New York, London, Paris: Gordon and Breach Sci. Publ., 1971,
Vol. 2, pp. 622, 417-440
Noble, A.: The food and feeding habits of the Indian Oil sardine, Sardinella longi-
aeps Valenciennes at Karwar. Indian J. Fish. A + B. 12, 1, 77-86 (1965)
Parr, A.E.: Is the presence of phytoplankton in the stomach contents of the California
sardine caused by special pursuit or merely due to incidental ingestion? Ecology
XI, 2, 465-468 (1930)
Radovi~h, J.: Food of the Pacific sardine, Sardinops aaerulea from Central Baja Cali-
fornia and Southern California. Calif. Fish Game 38, 575-585 (1952)
(1969)
Savage, R.E.: The food of North Sea herring 1930-1934. Fishery Invest. London Ser.
II, 15, ~, 60 (1937)
110
Ursin, E.: On the prey size preferences of cod and dab. Medd. Danm. Fisk.-og Havun-
ders. N.S. ~, 85-98 (1973)
Venkataraman, G.: Studies on the food and feeding relationship of the inshore fishes
off Calicut on the Malabar coast. Indian J. Fish. ~, ~, 275-306 (1960)
Weihs, D.: Hydromechanics of fish schooling. Nature (London) 241, (5387), 290-291
(1973)
Yamashita, H.: Relations of the foods of sardine, jack mackerel, and so on, in the
waters adjacent to west Kyushu. Bull. Seikai Reg. Fish. Res. Lab. 11, 45-53 (1957a)
Yamashita, H.: On the relation between the food and the shape of the intestines of
sardine, jack mackerel, mackerel and their kindred species found in the west coast
of Kyushu. Bull. Seikai Reg. Fish. Res. Lab. l!, 56-68 (1957b)
Yoshida, Y.: Relation between the sardine and the food plankton.II. On the feeding
mechanism of Sardinops me Zanosticta. Bull. Japan. Soc. Scient. Fish ~, ~, 467-470
(1955)
Food Consumption of Pelagic Fish in an Upwelling Area
O.A MATHISEN, RE.1HORNE, RJ.1RUMBLE, and M. BLACKBURN
1. Introduction
In recent years, studies of upwelling areas have evolved from a purely

descriptive mode to a process-oriented study with the underlying ra-
tionale that the relationships between several simultaneous processes
can explain the productivity levels of the system. Traditionally, the
high fish production characteristic of upwelling areas has been seen
as the immediate and direct consequence of high primary production
followed by high secondary production, all generated initially by the
nutrient supp~y brought up during the physical upwelling process. An
implicit assumption is that food supply per se is the mechanism which
regulates numbers of the populations utilizing this supply.
Food limitation as a control for adult fish has been questioned by

Gulland (1971), Murphy (1973), and Steele (1974) since, in those cases
where fish have been reduced to a very low level through fishing, the
structure and size of the next lower trophic level has not changed
drastically. In the Antarctic the population of krill has increased
somewhat today, evidenced in part by more numerous older age groups
and higher average life span than during the thirties before the stocks
of baleen whales, notably blue and fin whales, were reduced to a sub-
sistence level (Laws, 1974). In other cases the disappearance of the
Atlanto-Scandian stock of herring in the North Atlantic and Norwegian
sea, left no demonstrable changes in the CaLanus population.
Neither has the growth rate, as measured by the parameters Loo or k in

the Von Bertalanffy growth equation undergone more than a slight in-
crease when values determined for the exceptionally large 1904 year
class are compared with those associated with the present day's almost
extinct year classes of herring in northern waters (Mathisen, unpub-
lished). The size of recruited fish may change, but adult (post-recruit)
growth continues at a nearly constant rate, which Cushing (1973) inter-
prets to mean that density-dependency occurs during larval and juvenile
stages.
If adult fish are independent of their food supply then the require-
ments for food of the fish population should be considerably lower than
the production of the trophic levels on which it feeds. In order to
examine this hypothesis there must be estimates of the ichthyobiomass,
its bioenergetic requirements, and of the food production. Recent tech-
nical advances, especially in rapid three-dimensional measurements of
pelagic fish distributions, permit a closer exploration of food re-
quirements utilizing data acquired during the Joint I expedition. In-
tensive physical and biological measurements were made in a small, 40
by 100 km area established within the general area of observation
bounded by 21 0 00'-22 0 20' Nand 17 0 05'-17 0 40' W off the coast of the
Spanish Sahara.
112
2. Estimation of Nekton
2.1 The Hydroacoustic System
The system consists of: an instrumented towed body, towing cable and
winch; two echo sounders, one operating at 120 kHz and the other at
38 kHz; the control electronics; and a computer system for data pro-
cessing. The towed body is a Braincon, type 438, 1.3 m V-fin. At speeds
between 2 and 6 m s-l the body maintains attitude ± 30 in both pitch
and roll. The body is towed with Amergraph Type 2J35RL armored cable
which is hair-faired. The winch provides storage for the cable and
power to raise and lower the V-fin. An elastic shock cord assembly at-
tached to a stopper in the cable is used to damp out variations in
towing force encountered in rough seas.
The instrumentation in the towed body includes a calibration oscillator

to provide measurement of system gain, a signal steering filter, and
two signal transducers. The 120 kHz transducer is a 10-cm PZT-4 ceramic
disc, while the 38 kHz transducer is a 25-cm parabolic reflector with
a PZT-4 spherical transducer at the focal point of the parabola. Both
transducers have 9 0 -10 0 nominal beam widths. Shipboard electronics
control the echo sounders, generate control signals for the computer,
and monitor and display calibration and dynamics data sent from the
towed body.
The computer system processed data from the two sounders in real time
and calculated densities of pelagic fish. The system consists of three
major sections: signal conditioning elec.tronics, analog to digital con-
verter, and the central processor. The signal is first amplified to a
level proper for optimal processing, then detected and low-pass fil-
tered at a cut-off frequency corresponding to the transmitted pulse
length. The analog to digital converter consists of an eight-channel
mUltiplier, a 12-bit A-D converter, a programmable clock, two external
event inputs, and the computer interface. The central processor is a
Digital Equipment Corporation PDP 11/45 with a floating point proces-
sor, 24 K memory, two 1.2 M word disc storage units, line printer,
card reader and TTY for operator's console. The hydroacoustic data are
integrated in user-selected depth and time intervals, and corresponding
fish densities were calculated from input calibration and published
target strength information. The computer can access data files for
further reduction and manipulation of data using FORTRAN language pro-
grams.
2.2 Survey Procedures
Underway mapping took place continuously in the restricted survey areas

during the experiment, but acoustic data were collected only from March
31 to May 10, 1974. The transect lines were athwart the coastline and
extended from the 50-m isobath to a few miles beyond the edge of the
coastal shelf. Some replication of the transects took place but no real
systematic statistical design was followed. A large share of the map-
ping took place in the vicinity of 21 0 40' N where a number of observa-
tional buoys and current meters were anchored (Fig. 1).
The V-fin was towed over the stern at a depth of about 10 m and a speed
of about 4 m s-l. Soundings were made once per s, alternating between
sounders. Echo intensities were measured and integrated generally over
a 10-min period and within 20 to 140 m divided into 10-m thick depth
strata. For each period an average fish density was computed and ex-
pressed in g m- 3 using the acoustic calibration data for the equipment
113
2~·r-'---r-~--'---r--r--~~r-~--~~ Fig. 1. Map of the study area vIi th acous-

tic course lines in the sector 21°30' N
to 21 0 40' N, . Joint I, April-May 1974
I
N
:::;!Jf!!!f!!!?:
...... .
~
-:
".:.::::;:::::;:::::
::::f~:~.~
0-
2' "Oo'I...-....L...-:'-=7.:30
-- . -'---....L...---'w'---'----'---,...J 00.:.:..:..:.u.__-'--.....J
7.:.1:
and assuming that a kilogram of fish presents a target strength of -31

dB at 120 kHz and -32 at 38 kHz. Only data from night surveys have been
used because schooling density during daytime presents a nonlinear
acoustic reflection (Mathisen et al., 1974).
2.3 Analysis of Acoustic Data
An immediate objective was to derive a measure of total biomass for

pelagic fish and their food consumption. The available statistic was
in the form of individual density estimates calculated as total bio-
mass under 1 m2 surface area averaged over 7.5-min transects or about
a linear distance of one mile. Because such observations were made
sequentially, they are serially correlated, which adds another compo-
nent to the variance of the mean, proportional to the magnitude of the
serial correlation (Nickerson and Dowd, 1973). This correlation can
be measured by the auto-covariance and a few examples are given in
Figure 2, which shows that the auto-covariance ranges from 25% to 50%
for a lag factor of one, and is negligible for lags of second or higher
order. Therefore, it was omitted in this exploratory study. Physically
this can be interpreted to mean that the schools of fish present in
the pelagic zone had a linear dimension generally less than the tran-
sect distance of about one mile. A frequency distribution of observed
density estimates confirms this (Fig. 3). The majority of the density
estimates are low ones with no particular peak for the few and scattered
high values. This is further illustrated in Figure 4 which is an attempt
to determine the locations of high densities of fish. Aggregations of
fish were identified by subtracting 50 g m- 2 from the raw data which
exaggerated the patchiness. Further, the presence of a peak value is
a temporal feature, and peaks change in value and position during single
or adjacent nights. There is therefore, no real basis for a stratifica-
tion which will reduce the variance. However, division has been made
on a time-area basis for reasons related to the production pattern
generated by the upwelling. Four time periods were established on the
basis of increases and decreases in peak values of biomass: 6-9 April,
22-24 April, 25 April-2 May, and 3-6 May. The spatial division was
114
1600 Fig_ 2. Autocovariance series for hydroacoustic

800
0 • . ••••••••• • ••• • data, Joint I, 1-2 May 1974
-800 •
-1600
24()()
~ 1600 •
~ 800 ••
~ or-~-----..~~~--~~·----~·~.~
g -800 •••• •••••
~-I600LL~~~~~~~~~~~~~
:::l
< ~r------------------------,
3200
2400
1600
•• • •
800
•
0 • •
-800
., •• ••••••
-1600LL-L~-L~~-L~-L~~~~-L~
o 5 10 I~ 20
LAG (7lh minule In larva IS )
2rolr-----r----,r----,-----,-----.----~----_r----.
200
Fig. 3. Frequency distribu-

tion for acoustic measure-
ments of nekton standing
stock (all values at 25 g
m- 2 intervals), 21 0 30'-21 0 40'N,
0-25 125-1ro 2ro-275 375-4()() 500-525 625-650 750-775 875-900 ",000
INTERVALS (G/M2) Joint I, April-May 1974
g4B(!
731 , ,' 520
/ / /J
1 / L n
Jl 1JL
L
()()
. 1<0'- 17',.
If .l'
,..-
~ ~-
()()
-"" T ~ /' /' 17':I'tI,
,
()() I L L.£ ~
JII' /' L/ L L L L L oG' I/ Jo'" Il'
oc /'" L/ L L .L L I/ LL I
17 ·~1
,.
L L
0
I' .,.q,q. .... ... ~
.... L ,r...·
I' f'~~q. f' ",tf' qq.
'"
....
~
.,,'"
~.,. ~""()()
:1.~'"
~.,.
,I!,.~"'"
~
,0;; .~ 'I.') ",. 'I.'" ,'I. ,'I. 200
(M) (M) ,
Fig. 4. Time series of nekton standing stock (raw data reduced by 50 g m- 2 ) for
transects between 21 0 30' Nand 21 0 40' N, Joint I, April-May 1974
115
5. Mean (X) vs variance (S2) estimated from raw

-="'-'-"::f=-'"or time-area strata of Joint I. Line has a slope
2 3 4
LOG (X)
based on total depth with the inner shelf area extending to 17 0 15' W,
the outer shelf from here to 17 0 25' W, and the last area including the
shelf break out to 17 0 35' W.
A mean was found for each one of these cells and combined without
weighting, since both areal extent and time periods were almost equal.
The strongly skewed distribution of the independent variable neces-
sitated a normalizing transformation before a linear model could be
written. Certain properties of the observed means and variances sug-
gested a simple transformation of the variables.
If the values of the stratum means are plotted against the correspond-
ing variance, these points fall almost in a straight line with zero
intercept and slope 2 (Fig. 5),
log s~ = 2 log
~
x.
~
or that the standard deviation is proportional to the mean for which

a log transformation is appropriate.
Table 1 lists the cell means and associated variances of the log-trans-
formed data. Several levels of resolution of nekton are possible. At
the first level is a single biomass estimate averaged over time and
area. Since this was derived on log-transformed data, retransforming
gives the geometric mean of the sample. This underestimates the arith-
metic mean, which provides a consistent and unbiased estimate of the
population mean, although it is not the most efficient statistic. The
same statement holds for the variance of the original sample which will
not efficiently estimate the population variance.
Finney (1941) has treated the problem of converting means in logarithms
to mean in the original scale together with the variances which have
been simplified by Embody (1952), who developed the following approx-
imate formulas.
10glOXA = xT + 1.15129 s~
s
A
= xAJ 102 • 302585 s~ - 1
116
Table 1. Mean values of nekton biomass in the area 21°30' N-21040' N and calculated
after transformation Yijk = log (xijk + 1)
Area/time 2 3 4
I x 1.413 1.282 1.024 0.849

s 0.42 0.31 0.55 0.77
n 17 30 32 16
II x 1.127 0.935 1.111 1.551

s 0.67 0.50 0.52 0.79
n 47 28 33 26
III x 1.660 1.307 1. 125 1.236

s 0.76 0.82 0.84 0.66
n 34 29 35 21
mean after transformation from logarithmic to arithmetic scale
mean of data transformed to logarithms with base 10
standard deviation for data transformed from logarithmic to

arithmetic scale
s = standard deviation data transformed to logarithms with base

T 10
The values in Table 2 are derived from those in Table 1 after trans-
formation from the logarithmic to the arithmetic mode. The grand mean
is useful for a calculation of total food requirements over a long
period, on the assumption that fish are able to survive and grow with-
out continuous feeding by integrating the available food over time. At
the next level of resolution, one can make the same consideration on
a smaller area and time basis, once an expression of the bioenergetic
demand of ichthyonekton in this area has been developed.
Table 2. Mean values of nekton biomass in the area 21°30' N-21040' N after trans-
formation from logarithmic to arithmetic scale
Area/time 2 3 4
I xA 40.56 23.56 22.66 35.69

SA 51.05 18.90 45.47 185.66
n 17 30 32 16
II xA 42.28 15.80 24.21 184.83

SA 130.05 26.44 40.61 948.40
n 47 28 33 26
III xA 213.00 117.28 85.67 54.30

SA 973.80 674.11 550.29 165.74
n 34 29 35 21
The overall mean is 57.70 g m- 2 with an estimated standard deviation of 10.31 g m- 2 .
2.4 Bioenergetic Demand
Both Winberg (1954, 1961, 1962) and Ivlev (1960, 1961a, b) have proposed
schemes for partitioning ration which have been used to establish bio-
117
energetic budgets for wild populations (Mann, 1965, 1969). The use of
Winberg's balanced equation has been criticized by Warren and Davis
(1967) for its lack of flexibility and its inability to account for
nutrition and activity levels of the fish. They suggested a more complete
partitioning of the total ration into its components which has been
followed here with some small extensions.
R =G + S + T + A + L + P + M
Miscellaneous losses, M, include sloughed scales, mucus secretion,
osmotic and ion balance costs which all have been considered negligible
and omitted in this. case. Reproductive loss, P,represents the energy
required for gonad development which accounts for only a small propor-
tion of the total ration. Bomb calorimetry indicates P = 1%-3% of the
total ration (Mann, 1965; Lasker, 1970). Most energy associated with
reproduction would be for sexually related activities such as migration,
nest building, or parental care. Therefore P itself is small and has
been omitted.
Fecal and excretory losses as a function of ration size are difficult
to include accurately in the bioenergetic equation because both change
wi th quality 'as well as quantity of ,diet. For this study both sources
will be combined in an expression of loss; and for lack of specific
information, the most common value in the literature L = 0.2 R will be
used.
Growth G can be measured directly as body weight added per time. Lasker
(1970) estimated that growth of California sardine accounted for 0%-
20% of annual ingested energy, being highest for young of the year and
dropping with age; this range is probably typical of expected growth
for most fish. With data from Larraneta (1960), an exponential model
was applied for estimating the monthly growth of Sardina pitchardus. Daily
growth ranged from 0.12% to 0.4% of body weignt during the growing
season. Walsh (1975), using a Von Bertalanffy growth-in-weight model,
calculated that Peruvian anchovy add about 0.74% body weight per day
during the mos.t favorable upwelling. Thus, daily growth may be in the
range of 0.5 to 1.0% per day or in terms of calories 0.005 W (C:l) to
0.01 W (C:l).
The real cost of living for fish is metabolism which requires 60%-90%
of assimilated energy. Three components can be identified: standard or
basal metabolism, T; specific dynamic action, S; and activity, A. A
true basal rate for fish cannot be measured because of spontaneous
movement. Therefore, standard metabolism is considered to be the con-
sumption rate of 02 by an unfed fish which is moving as little as pos-
sible. Standard metabolism is a function of both body weight and tem-
perature, commonly expressed as T = aw Y where Y usually is set equal
to 0.8 according to Winberg (1954) who also gives a value of 0.3 for
the coefficient a, at 20 0 C which corresponds to a = 0.0242 at 16 0 C and
when converted to calories per hour. For temperature characteristics
like those of the Northwest African Upwelling, T = 0.0242 Wo.s kcal/
day.
Specific dynamic action, S, is primarily the cost of deaminating pro-
tein, but includes some cost of dearninating fats and carbohydrates and
digesting and storing food. For a constant composition diet, S is di-
rectly proportional to diet but varies with type of food. The specific
dynamic action of completely catabolized protein is about 30% of caloric
value (Brody, 1945; Kleiber" 1961). Since the diet is not entirely pro-
tein, a more reasonable value is 20% of the ration.
118
computations of energy required by activity A have been made by Ivlev

(1960) and Brett (1965). The speed at which fish actually swim in nature
is at best difficult to determine. Thus, expected ratio is a function
of swimming speed, but energy losses increase exponentially with swim-
ming speed. At lower speeds, the difference between maximum oxygen in-
take and oxygen used for swimming limits the amount of ration which
can be digested. Since an activity model is not complete, maximum en-
ergy expenditures will be considered to range from equal to twice stan-
dard metabolism and the lower limit most applicable in this case.
The bioenergetic equation is set up for individual fish and can be ap-
plied to a natural population if numbers of fish at various sizes, and
their mortality rates, are known. In the case of Northwest Africa, we
do not have numbers, population parameters, size or age. Even species
composition is circumstantial (Thorne et al., 1977; Blackburn and Nel-
len, 1976). Since only estimates of average standing stock (W) are
available, the equations must be expressed in a per weight basis. In
the_case of standar~ metabolism, for example, T = awy becomes T/W =
aWY 1. Thus (T/W) (W) (time) = standard metabolic requirements for a
long time period. The general equation on a weight basis then becomes
R:W = G:W + S:W + T:W + A:W + L:W + P:W + M:W.
Substituting maximum values derived previously leads to:
R:W = 0.01 + 0.3 R:W + T:W + 2 T:W + 0.2 R:W + 0 + 0

0.5 R:W = 0.01 + 3 R:W = 0.01 + 3(0.0242 W-· 2 )
R:W = 0.02 + 0.1432 W-· 2
Substituting the lower values leads to:
R:W 0.005 + 0.2 R:W + T:W + T:W + 0.2 R:W + 0 + 0

R:W = 0.008+ 0.0806 W-· 2
For a fish weighing 100 g the limit ranges from R:W 0.0770 to 0.0404.
2.5 Food Production
Production as such was never measured during Joint I. Zooplankton mea-

surements included only standing stock determined from 160 net tows
along the buoy line at 21 0 40' N (Blackburn and Nellen, 1976). Nets were
hauled vertically from the bottom or 200 m depth, depending on water
depth, to the surface. Two 60-cm diameter, conical nonclosing Bongo
nets, mounted side by side, were used. Both nets had uniform 102 ~
mesh size. The combined catch of both nets was divided into four equal
aliquots, two of which were filtered through 1050 ~, 505 ~, 223 ~ and
102 ~ filters. Wet weights were determined from one filtered aliquot
and reported as weight per meter squared.
Because of the scarcity of observations, time periods 2 and 3 were

combined. The raw data were first log-transformed and a mean found
which was transformed back into the arithmetic mode in a manner anal-
ogous to the treatment of acoustic density estimates. The results are
given in Table 3.
However, standing stock measurements are not particularly useful in

this case where a measure of daily production is needed. In the absence
of rates, an alternative is to calculate the daily food demand as a
fraction of the standing stock of zooplankters. Such ratios of produc-
tion to biomass have been summarized (Mullin, 1969) for a number of
localities in a review of indirect production measurements. The estab-
119
Table 3. Average zooplankton standing stock (g m- 2 surface) observed April 6 through

May 6, 1974 a
Area/time 2 + 3 4
I 39.78 36.86 59.41

II 57.40 48.12 62.44
III 60.10 101.58 120.64
The overall mean is 71.45 g m- 2 with an estimated standard error of 7.07 g m- 2 .
a Data have not been corrected for a small phytoplankton contamination.
lished ranges can then be compared with the actual values obtained
from the Joint I area.
2.6 Food Consumption Relative to Food Production
Upwelling is strongest during spring and early summer when northerly

winds blow along the coast. During summer an oceanographic front moves
into the area from the south bringing warm, tropical water. The con-
tinental shelf is wide and flat: The shelf break is very pronounced at
about 100 m depth, and is located 40 to 50 km offshore. Upwelling may
occur at the shelf break as well as inshore along the coast. The
strength of the wind varies markedly over a period of several days,
and fluctuations in the intensity of upwelling result. Primary produc-
tion is lower than for other upwelling areas, on the order of 2 g C
m- 2 day-l compared to > 5 g C m- 2 day-l for Peru or Baja California
(Walsh, personal communication). Sand from the African desert and a
deep, wind-stirred, mixed layer combine to keep primary production low.
Small zooplankton « 500 ~) predominate on the shelf and consist of
benthic as well as pelagic forms; seaward of the shelf break, zoo-
plankton larger than 500 ~ become most common (Blackburn and Nellen,
1976). Fish in the area during upwelling are mostly sardines and an-
chovy over the continental shelf, and horse mackerel at the shelf break
and upper continental slope (Blackburn and Nellen, 1976; Blackburn,
1975). Sardines eat mostly small zooplankton, while horse mackerel
prefer the larger ones. However, since there exists a lag in the pro-
duction of food and reaction of nekton, an overall mean value is a
useful statistic for comparisons between different areas separated
in space.
Starting then with an overall mean of 57.70 ± 20.62 g m- 2 and using the
equation derived for nekton energy requirements, an average biomass
of 58 g m- 2 composed of fish 100 g in weight, would need about 2.3 g
of food per day. The average standing stock of zooplankton observed
during Joint I was 71.45 g m- 2 • In order to satisfy this demand, a
production to biomass ratio (P:B ratio) of 0.033, which is well within
the range of P:B ratio summarized by Mullin (1969), was found. This
calculation assumes that all fish food is zooplankton, which is doubt-
ful for Northwest African sardine (Blackburn and Nellen, 1976).
At a higher level of resolution, each cell provides a preliminary look

at changes in standing stock and in food consumption. Calculated food
consumption for the time-area strata and P:B ratios necessary to meet
this demand are presented in Table 4, together with turn-over time of
the zooplankton population. Interpretation of the stratum data is dif-
ficult because the time period involved, usually 2-3 days, may not be
long enough for the fish to react and subsequently re-distribute them-
120
Table 4. Total daily ration, R, requirements in calories for standing stock of fish,
production biomass ratio of zooplankton matching daily food requirements, P:B and
turn-over rate
Area/time 2 + 3 4
I R 1.639 0.934 1.442

P:B 0.041 0.025 0.024
Days 24.4 39.5 41.2
II R 1. 708 0.808 7.467

P:B 0.030 0.017 0.120
Days 33.3 59.5 8.4
III R 8.605 4.100 2.194

P:B 0.143 0.040 0.018
Days 7.0 24.8 55.0
selves. The range of P:B ratios, however, indicates maximum conditions

encountered, which can then be compared to P:B ratios in the literature.
Mullin's (1969) summary lists production estimates mainly for temperate
and subarctic' areas, many of which are averaged over a year. P:B rates
of 0.01-0.08 predominate. Two high P:B ratios for Joint I were 0.12
and 0.143 (Table 4) but the first one can be explained due to a tem-
porary but strong influx of fish. Even so, this ratio falls within
ranges given in the literature. Mullin lists an estimate by Smayda
(1966) for the upwelling period in the Gulf of Panama for P:B = 0.29
or 0.98, depending on assumptions.
3. Discussion
Evidence has been presented to support the hypothesis that food is not
directly controlling the size of adult or recruited fish populations
in the Northwest African Upwelling System, although it may determine
their distribution. Estimates of the potential food production were
always greater than the daily ration required by the pelagic fish popu-
lations, and in most instances greatly in excess. This excess is a
necessity for any validation of the hypothesis, even in a tentative
manner, because of the great sampling error involved. The basic bio-
mass estimate of fish was given with a preCision of ± 30%, while the
range of possible bioenergetic estimates varied by a factor of two.
Estimates of the standing stock of zooplankton were the most consistent,
but the productivity could only be gauged by inference from observa-
tions elsewhere. .
Since the studies were conducted in April-May, during or shortly after
maximum reproduction, it is to be expected that peak biomass of fish
would occur in late summer or early fall at a time when food produc-
tion is less than in spring. It is quite conceivable that food for the
adults and recruited part of the various populations then assumes the
role of control mechanism.
However, the suggestion that density-dependent mortality, caused by
food availability and predation, largely determines survival during the
larval drift stage is stea4ily gaining support. Experimental studies on
feeding, digestion, and starvation of the fish larvae point out the
effect of food. Blaxter and Eempel (1963) introduced the concept of
"point of no return" for herring larvae. This is the time available for
121
the larvae for the first feeding and initial survival. Beyond this
time limit, reckoned from hatching, and temperature-dependent, the
larvae is still alive but too weak to feed. Since food appears in patch-
es, food-related survival can be treated as a stochastic process for
an encounter of the fish larvae as predator, and zooplankton as prey.
This probability is conditioned by many other factors. Some species

have a very small gap and require very small food organisms for their
early survival. Development of swimming ability, of visual and olfactory
perception and general filtering ability are rapid after hatching and
improve success in early feeding.
If one accepts the hypothesis that food availability exerts control

of numbers during larval drift, then the high fish production in an
upwelling system should primarily be looked on as a probability density
function with the mean expectation proportional to the magnitude of the
production. Thus, when a series of upwelling systems are ranked ac-
cording to fish production, their expected means of the probability
of larval survival should follow the same ranking.
Nonupwelling systems, predominantly those of boreal or arctic character,

have probabi~ity density functions of larval survival of an opposite
type with low mean values. While probability of success generally is
the rule for upwelling systems, failure is the normal event in a system
such as that involving the Atlanto-Scandian stock of herring. The famous
exception with the 1904 year class which dominated the total catches
for twenty years after recruitment is well known. During the last two
decades there have been only two to three strong year classes. Although
there has been a slight increase in a parameter like L , compared with
those determined for the 1904 year class, these changes indicate a very
low density-dependent control on the recruited stage consi.dering at
least a 300-fold range in numerical abundance.
In every instance, it appears that a regulation of numbers occurs at

the larval stage. It is not clear whether the probability of an en-
counter between predator and prey is all denSity-dependent or only
partially denSity-dependent.
The problems raised here need a solution before a complete modeling

of the entire upwelling system is possible. While estimates of biomass,
bioenergetic demand and food production can be improved by increased
effort, larval feeding behavior must be studied in the laboratory. How-
ever, the larval food base, extent, and distribution of food patches,
and sizes of their components can only be studied in the field, and
suitable techniques are still lacking. The time lag existing between
a spin-up event in an upwelling system and the consumption of produced
food is so large that all the finer details of hydrographic conditions
causing the upwelling become obliterated by integration over the lag
time. The present upwelling studies originating in the first production
steps result in a pyramidal hierarchy. A diametrically opposite approach
commencing with fish production would result in a downward-extending
pyramid in which only the integrated values of hydrographic features
remain. Because of the difference in time domain, no immediate com-
promise appears between these two approaches to describing an upwelling
system.
122
References
Blackburn, M.: Summary of existing information on nekton of Spanish Sahara and ad-
jacent regions, northwest Africa. CUEA Technical Report 8, 1975, 49 pp
Blackburn, M., Nellen, W.: Distribution and ecology of pelagiC fish studied from eggs
and larvae in an upwelling area of Spanish Sahara. NOAA Fishery Bulletin 74, 4,
885-896 (1976)
Blaxter, J.H.S., Hempel, G.: The influence of egg size on herring larvae. J. Cons.
Int. Explor. Mer 28, 211-240 (1963)
Brett, J.R.: The relation of size to rate of oxygen consumption and sustained swim-
ming speed of sockeye salmon (Oncorhynchus nerka). J. Fish. Res. Bd. Can. 22 (6),
1491-1501 (1965)
Brody, S.: Bioenergetics and Growth. New York: Reinhold Publishing Co., 1945, 1023 pp
Cushing, D.H.: Recruitment and parent stock in fishes. Washington: Sea Grant Publica-
tion 73-1, 1973, 197 pp
Embody, D.R.: Analysis of variance calculations as applied to creel census data.
Idaho Department of Fish and Game. 1954, 29 pp
Finney, D.J.: On the distribution of a variate whose logarithm is normally distrib-
uted. Suppl.J. R. Statis. Soc. 2 (2), 155-161 (1941)
Gulland, J.A.: Ecological aspects of fishery research. Adv. Ecol. Res. 2, 115-176
(1971)
Ivlev, V.S.: Method of estimating the food utilized by growing fish. Z. Fisch. ~ (3/4),
281-289. Fish. Res. Bd. Can. Transl. Series No. 374, 1961 (1960)
Ivlev, V.S.: Experimental ecology of the feeding of fishes. New Haven: Yale University
Press, 1961a, 302 pp
Ivlev, V.S.: On the utilization of food by plankton-eating fishes. Trudy Sevasto-
poloskoi Biologicheskoi Stantsii 14, 188-201. Fish. Res. Bd. Can. Transl. Series
No. 447 (1961b)
Kleiber, M.: The Fire of Life, an Introduction to Animal Energetics. New York: John
Wiley and Sons, Inc., 1961, 454 pp
Larraneta, M.G.: Synopsis of biological data on Sardina pilchardus of the Mediter-
ranean and adjacent seas. Proc. World Scie.nt. Meeting on the Biology of Sardines
and Related Species, 1960, Vol. II, FAO, Rome, pp. 137-173
Lasker, R.: Utilization of zooplankton energy by a Pacific sardine population in the
California Current. In: Marine Food Chains. Steele, J.H. (ed.). Berkeley: Univer-
sity of California Press, 1970, pp. 265-284
Laws, R.: MS. Population Dynamics and Ecology of Marine Animals. Working Paper No.
7. UN/FAO/ACMRR Ad Hoc III (1974)
Mann, K.H.: Energy transformation by a population of fish in the River Thames. J.
Anim. Ecol. 34 (2), 253-275 (1965)
Mann, K.H.: The dynamics of aquatic ecosystems. Adv. Ecol. Res. ~, 1-81 (1969)
Mathisen, O.A., O.J. ¢stvedt, Vestnes, G.: Some variance components in acoustic
estimation of nekton. Tethys 6 (1-2), 303-312 (1974)
Mullin, M.M.: Production of zooplankton in the ocean: The present status and prob-
lems. Oceanogr. Mar. BioI., Annu. Rev. 2, 293-314 (1969)
Murphy, G.I.: Clupeoid fishes under exploitation with special reference to the Peru-
vian anchovy. Tech. Rept. No. 30, University Hawaii, Hawaii Inst. Mar. Biol.,
1973, 73 pp
Nickerson, T.B., Dowd, R.G.: Design and operation of survey patterns for demersal
fishes using the computerized echo counting system. Paper No. 19, Symposium on
Acoustic Methods in Fisheries Research. Bergen (1973)
Smayda, T.J.: A quantitative analysis of the phytoplankton of the Gulf of Panama.
III. General ecological conditions, and the phytoplankton dynamics at 8 0 45' N,
79 0 23' W from November 1954 to May 1957. Bull. Inter-Am. Trop. Tuna Comm. !l (5),
355-612 (1966)
Steele, J.H.: The Structure of Marine Ecosystems. Cambridge: Harvard University
Press, 1974, 128 pp
Thorne, R.E., Mathisen, O.A., Trumble, R.J., Blackburn, M.: Distribution and abun-
dance of pelagic fish off Spanish Sahara during CUEA Expedition JOINT I. Deep-Sea
Res. ~, 1, 75-82 (1977)
123
Res. ~ (4), 201-236 (1975)
Warren, C.E., Davis, G.E.: Laboratory studies on the feeding, bioenergetics, and
growth of fish. In: The Biological Basis for Freshwater Fish Production. Gerking,
S.D. (ed.). Oxford: Blackwell Scientific Publications, 1967, pp. 175-214
Winberg, G.G.: Rate of Metabolism and Food Requirements of Fishes. Nauch. Trudy
Belorussk. gos. Univ. V. I. Lenina Minsk, 253 pp. Fish. Res. Bd. Can. Transl. Series
No. 194 (1956)
Winberg, G.G.: New information on metabolic rate in fishes. Voprosy Ikhtiologii l
(18), 157-165. Fish. Res. Bd. Can. Transl. Series No. 362, 1961 (1961)
Winberg, G.G.: The energy principle in studying food associations and the productiv-
ity of ecological systems. Zool. Zhur. ~ (11), 1618-1630. Fish. Res. Bd. Can.
Transl. Series No. 433 (1962)
Benthos in Upwelling Regions
H.THIEL
Investigations on the benthos in upwelling areas are briefly reviewed,

with the main emphasis on standing stock studies. As expected, the
benthic standing stock is high beneath highly productive surface waters.
However, when the oxygen content of the near-bottom water is depleted
by intensive degradation of organic matter, hydrogen sulfide may de-
velop in the sediment and reduce or even obliterate the faunal standing
stock.
Preliminary results on meiofauna densities and on the concentration of

chloroplastic pigments in surface sediments from the upwelling region
off Northwest Africa and from areas with less upwelling off Portugal
and off Morocco are presented (Fig. 3). The meiofauna was sampled on
three transects (Fig. 2, A-C) from the continental shelf down to the
foot of the continental slope. In the upwelling area a decrease of
rne_idlfaunal abundance from the shelf (80 m) to the upper slope (200 m)
is most pronounced (Fig. 3). An increase to 400 m (transect C) and
to 1200 m (transect B) is followed by decreasing densities to bathyal
depths. The minima at 200 m depth may be artifacts due to the chosen
sampling depth, but low numbers in the samples of the upper continen-
tal slope are real. These correspond well to the minima encountered
for chloroplastic pigments (Fig. 4), although these measurements were
made on samples from two other transects (1 and 2) off Spanish Sahara
three years later. On transect 2 the highest concentration was found
at 1000 m-depth. While the meiofauna of the three transects shows the
same ranges in densities, the values for chloroplastic pigments differ
by two orders of magnitude between the transects off Cape Blanc and
off Morocco, with the values for transect 1 falling between these ex-
tremes. The concentrations of chloroplastic pigments reflect the pri-
mary production in surface waters due to availability of nutrients.
However, the meiofauna data do not show such a clear relationship,
possibly due to the geographic position of the transects or to the
feeding effects of macrofauna. The corresponding minima of pigments
and meiofauna at 200 m occur within the depth range of strongest cur-
rent activity on the upper continental slope, where sedimentation rates
are presumably low, so that the food income for the infauna is limited.
From these preliminary results it is concluded that:
1. benthos investigations will contribute to the understanding of ocean-

ographic processes in upwelling regions;
2. such studies should consider the total benthos and should also in-
clude measurements of community oxygen consumption, nutrient recycling,
and sediment properties;
3. the benthos of upwelling and nonupwelling regions must be compared.

However, the investigated regions should be subjected to at least sim-
ilar climatological conditions.
125
1. Introduction
Plankton production has been studied in upwelling areas for half a

century. These investigations were intensified with the increase of
commercial fisheries during the last 25 years and with the large-scale
pelagic ecosystem research initiated only ten years ago (Mittelstaedt,
1972; Weichart, 1974), whereas benthic research in upwelling areas is
still in its infancy. The few papers presenting quantitative data are
confined to macro-infauna standing stock, but this single component
cannot be regarded as representative of the total benthos. In general
food availability to bottom organisms is the determining factor for
benthos standing stock (e.g., Filatova, 1969; Sanders and Hessler,
1969; Thiel, 1975) and for the size structure of deep-sea benthic in-
faunal communities, which exhibit a decrease of average organism size
with increasing depth (Thiel, 1975). High sedimentation of organic
matter is characteristic of upwelling areas, and high standing stock
and production of the benthos can therefore be expected.
The benthos from upwelling regions is not well known. Earlier investiga-
tions concentrated on the understanding of species composition and zoo-
geography, while aspects of standing stock and production were intro-
duced much later. The present paper will point to some of the qualita-
tive results, and it will summarize our knowledge of quantitative data,
including new findings from off West Africa. Finally, it will discuss
how benthic research can contribute to an understanding of upwelling
processes.
2. Benthic Communities in Upwelling Areas
2.1 General Description
Distribution and diversity of organisms are influenced by upwelling

processes. The shift of isotherms within the eastern boundary currents
in both hemispheres toward the equator allows cold-water species to
penetrate to lower latitudes. Ekman (1970) describes the conditions
for the different upwelling areas, with the tropical zone between the
20 0 C isotherms being narrower on eastern than on western sides of the
oceans. A zoogeographic analysis of the fishes of northwestern Africa
is given by Maurin (1968), who recognizes an Atlanto-Mediterranean and
a Guinean region, with two subregions of transitional character, the
Saharian and the Senegalian, respectively. The area off Cape Blanc
and the Banc d'Arguin constitutes the boundary between these subtropical
and tropical regions. Knowledge of benthic organisms is still limited,
and no marked zoogeographic zones are clearly distinguishable. Much
information on the different macrofauna groups has been collected in
the report on the results from the Danish expedition to West Africa
with the R.V. Atlantide, during the years 1945-1946 (e.g. Kirkegaard,
1959, on sedentary polychaetes). Tlirkay (1975, and unpublished results,
pers. comm.) describes the region off Cape Blanc as inhabited by mixed
temperate and tropical faunal components of the Decapoda Reptantia.
Similarly, Gosselck (1974) recognizes a transitional zone in the same
area. Other upwelling regions exhibit comparable faunal changes.
Sanders (1968, 1969) compared diversities of the macrofauna from dif-

ferent ocean regions and depths. For the upwelling area off Southwest
Africa he found the lowest diversity at a depth of 102 m, where the
near-bottom water has a very low oxygen concentration due to high sedi-
mentation rates and to incomplete degradation of organiq matter. While
126
only a few species can live under low oxygen conditions, the number of
individuals is fairly high. A similar situation was also observed off
Peru by Frankenberg and Menzies (1968) and by Rowe (1971a, b).
2.2 Abundance of Fauna and Oxygen Conditions

General agreement exists among benthologists that macrofauna standing
stock reflects the level of production in the overlying water masses
(Filatova, 1969; Sanders and Hessler, 1969; Thiel, 1975). Plankton and
benthos are related through the transport of food particles from the
surface waters to the ocean bottom, and we therefore expect a higher
benthic standing stock in upwelling areas than that beneath less pro-
ductive waters. Rowe (1971a) compared macrofauna standing stock below
the water column in the highly productive region off Peru with that in
the area of low pelagic production in the Gulf of Mexico (Fig. 1). The
6
z
o
m
~ 5
u
o
Z
<t
1-4
:::c
C)
ii:i
3: 3
1-'
w
~
-
N
:::E2
C)
:::E
Fig. 1. Decrease with depth of macrofauna

wet weight and carbon from two transects
2 3 4 5 6 off Peru and a transect in the Gulf of
DEPTH IN KILOMETERS Mexico (redrawn from Rowe, 1971b)
regression lines between water depth and wet weight or organic carbon
of macrofauna standing stock clearly reflect the difference in surface
productivity between these two regions (Rowe, 1971a; Rowe and Menzel,
1971). Further examples of macrofauna numbers and standing stock are
presented by Rowe (1971b) and Rowe et al. (1974). All papers exhibit
a similar relation between plankton production and benthos standing
stock.
Smith et al. (1974) compared total oxygen demand of sediments by in
situ measurements under the California Current off Baja California
(upwelling) and from the sewage-enriched region in the New York Bight
(outwelling). In both these organic-rich areas, biological and chemical
oxygen demands were significantly higher than in other comparable areas,
and chemical oxidation gained in importance over the biological degrada-
tion processes. Pamatmat (1971) presented a few values from the Peru
upwelling areas, where the bottom water layer exhibits low oxygen
127
concentration, and chemical oxidation exceeds biological oxygen consump-

tion.
A large standing stock of benthic organisms is not found in all up-

wel~ing regions. This can be explained by depauperate oxygen conditions
of interstitial and near-bottom water, due to high production of organic
matter in the euphotic zone and to incomplete degradation processes on
the bottom. High contents (5-26%) of organic matter in sediments of
upwelling areas are summarized by Diester-Haass (this volume). In con-
trast, much lower levels of organic matter (0.5-2%) are found in the
sediments of nonupwelling regions.
The depth of the oxygen minimum layer in the water column off the west
coast of India is changed by the monsoon regime (Banse, 1968), the
oxygen-poor water being shifted up the continental shelf during the
southwest monsoon. This has pronounced effects on the fisheries, de-
creasing total catches of fish and prawns from June to September by
more than 75% (Sankaranarayan and Qasim, 1968). For many sedentary or-
ganisms, the changing oxygen saturation of the water will be the limit-
ing factor. Low macrofaunal standing stock was found by Frankenberg
and Menzies (1968) and Rowe (1971a, b) off Peru in depths of 100-500 m,
where the oxygen minimum layer of the water masses impinges on the
slope. Khusid (1974) refers to similar unpublished results on the macro-
fauna obtained by Savilov, Moskalev and Zevina, and he reports low num-
bers of foraminifers in the sediments of the upper slope off Peru.
Gallardo (1963, in Sanders, 1968) observed less than 5% oxygen satura-
tion and decreased macrofauna densities in upwelling regions off north-
ern Chile, while Sanders (1968) found low numbers of individuals and
species on the Southwest African shelf, where oxygen saturation was
less than 2%. A low standing stock in the same area was also described
by Hart and Currie (1960). Varved sediments, indicating lack of bio-
turbation due to anoxic conditions, have been reported from off South-
west Africa, the Gulf of California, from off Peru, and the west coast
of India (see Diester-Haass, this volume, for further literature).
Macrofaunal biomass is food limited in most parts of the ocean (Fila-

tova, 1969; Sanders and Hessler, 1969) as is total benthic standing
stock (Thiel, 1975). However, high sedimentation rates of organic mat-
ter may cause oxygen values to approach zero, resulting in decreasing
faunal abundance.
3. Recent Quantitative Studies of the Benthos in the East Atlantic
While in earlier investigations the benthic communities of widely sep-

arated oceanic regions, characterized by different fertility of the
sea, were compared, a new approach was made in my own upwelling studies.
To understand the influence of upwelling processes on the benthos,
standing stock or production should be compared in regions of different
upwelling intensities but under equivalent climatic and oceanographic
conditions. However, the possibilities for undertaking such comparisons
are limited by the fact that prominent upwelling areas are situated
within arid climatic regions, while districts without upwelling usually
occur under the influence of humid zones and of river outflow.
3.1 Meiofauna Abundance
In preparation for an intensive benthos investigation, a pilot study

was run in the Cape Blanc upwelling region (Meteor cruise 26, 1972).
128
20 0
II
~1 __________________ ~ ____ ~~~~ __ ~~~~ ________________ ~ ·30
2~~ --~~~~~--~--~------------------ -+------------------+W~
Fig. 2. East Atlantic Ocean showing the six transects where sediment samples for
meiofauna (A-C) and for chloroplastic pigments (~) were taken
129
depth[m]
<0
0
/0
Meteor .19-
Portugal \ \
00
I
500
\0
Meteor.2S"
Westafrica
1000
\0 0/
1500
o o
2000
o 0
I
I
I
2500 I
I
I
I
I
3000 I o
I
3200 I
3800J
B-?>
I 111111111111111
A~ 10 3C ~ 10 4
I I I I I I 1111 I I I 1111111 I I "11111
Fig. 3. Numbers of meiofauna/10 cm2 (4-cm layer) from three transects in the Eastern
Atlantic: transect A off Cape Sines, Portugal, Meteor cruise 19, 1970; transect B
off Villa Cisneros, Spanish Sahara, ~ cruise 26, 1972; transect C off Cape Blanc,
Spanish Sahara, Meteor cruise 26, 1972
Transect C was located south of Cape Blanc heading southwest, while a

second transect (B) was run ,off Villa Cisneros with a similar depth
range (Fig. 2; Table 1). The results on meiofauna numbers are presented
in Figure 3, together with data from Meteor cruise 19, 1970, when sampl-
130
ing was accomplished on a transect (A) off Cape Sines (Portugal) (Ta-
ble 1).
Table 1. Transects off Portugal and off West Africa: Meiofauna transects A-C, chloro-
plastic pigments, and photosled transects 1-3
Cruise Time Location Trans- Depth Position of shallow station,

ect range position of deep station
(m) oN OW
Meteor 19 March Cape Sines, A 149 37°38.3' 08°58.5'
1970 Portugal 3809 37°44.8' 10°31.5'
Meteor 26 March Villa Cisneros B 64 24°03.8' 16°16.5'

1972 Spanish Sahara 3008 24°52.1' 17°35.9'
cape Blanc C 40 20°43.5' 17°10.8'

Mauretania 3104 19°50.9' 19°11.1'
Meteor 36 February Cape Blanc 2 85 21°18.5' 17°22.5'

1975 Spanish Sahara 2998 21°36.9' 18°42.1'
Cape Pefia 85 25°11.9' 15°48.2'

Grande 946 25°34.6' 16°0.2'
Spanish Sahara
March Cape Blanc du 3 188 33°28.5' 08°51.0'

1975 Nord, Morocco 2991 33°44.8' 09°15.8'
Meiofauna densities from the three transects show a remarkable simi-

larity, with values of about 10 3 individuals beneath 10 cm 2 sediment
surface (10 6/m 2 ). Transects A and B exhibit an expected decline between
500 and 3000 m, while numbers in transect C apparently increase again
from 2000 to 3000 m. In the upper parts of these curves transects B
and C show minima at about 200 m, and transect A exhibits an increase
between 149 m and 461 m.
Results for shelf and upper .continental slope depth, especially the
marked minimum of meiofauna abundance at 200 m, may demonstrate the
local sedimentation conditions of organic matter. The undercurrents
in upwelling regions flow along the upper continental slope, and it
may be assumed that this water movement prohibits food particles from
settling in these regions, thereby preventing high meiofaunal densities.
This hypothesis is in agreement with measurements of the undercurrent
by Mittelstaedt (1976, earlier lit.), who found the strongest flow,
with up to 30 cm/s, in a depth of 200-300 m. Similarly, Horn and Meincke
(1976) describe highest tidal activity in the range of the upper slope.
In this regicn, the change in the grain size distribution of the sedi-
ments is at a maximum (Lange, 1975) and the abundance, dominance, and
diversity of foraminifers alter to a greater degree than on the shelf
or deeper down the slope (Pflaumann, 1975). A distribution of macro-
fauna similar to that of the meiofauna in transects Band C was en-
countered by Boucher and Glemarec (1974) for macrofauna off southern
Morocco. The biovolume increases from shallow depth to 50-60 m by a
factor of 10, an intermediate minimum is observed at a depth of 100
to 150 m, ~d high values are found again around 400 m. Differences
in sedimentation conditions of organic matter, coupled with currents,
seem to account for this situation. The small increase of meiofauna
density from shelf to upper slope in transect A may be due to tidal
currents on the outer shelf. Relict sediments, more or less mixed with
131
recent materials, cover this zone on a world-wide scale (Kudrass, 1973;

for further literature, Calvert and Price, 1971; Marchig, 1972; Miro
Orell, 1973), indicating limited sedimentation of hydrodynamically
"lighter" particles. Siedler and Seibold (1974) measured current speeds
of 35 crn/s (maximum hourly mean over a 6-day period, February, 1967)
5 m above the bottom in a depth of 165 m on the Cape Sines (A) trans-
e.ct.
Comparing the three transects over their total depth range, the differ-
ences in meiofauna numbers were not sufficiently clear to support the
use of benthos as an upwelling indicator. Transect B off Villa Cisneros
and transect C off Cape Blanc may be too close together (200 nautical
miles), to allow for well-defined differences in plankton production
averaged over the year, and in food transport to the bottom. In con-
trast, the distance between these two transects and the one from Cape
Sines may have been too wide, i.e., a distance of 1000 nautical miles.
The Cape Sines transect is located in an area heavily influenced by
the discharges of water and nutrients from the river Tejo. This was
demonstrated by Kudrass (1973) with samples from the same transect
worked in 1967. The terrigen.ous component of the sand fraction amounted
to 17-42% of the surface sediments down to 1000 m, while less than 3%
were found in comparable materials off Cape Mazagan (Morocco). In ad-
dition nutrients are transported into the euphotic zone by upwelling
caused by the southwardly directed surface current and, in connection
with this, by upwelling south of Cape Roca. Thiede (1971) cited tem-
perature differences for nearshore regions off Portugal that were 4-7 0 C
lower than those in offshore areas. Thus, primary production off south-
ern Portugal may be similar to that in the upwelling regions off Spanish
Sahara and Mauretania, resulting in similar levels of benthos standing
stock.
In this paragraph meiofauna abundance is related to food availability

as the only factor. However, other influences may exist, and there is
at least one further explanation for the similarity of meiofauna num-
bers in the three transects. Feeding pressure by sediment-feeding macro-
fauna may reduce the meiofauna populations to a greater extent in areas
where macrofauna densities are higher, i.e., in upwelling areas. There-
fore, comparison of faunal abundance should cover the total benthos.
3.2 Chloroplastic Pigments
To provide a first quantitative insight into the sedimentation of or-

ganic matter, the determination of chlorophyll a and its derivative
pheopigment was performed. Pigment contents were determined with a
Turner fluorometer, after treatment of 2-ml sediment samples with MgC03
and glass spheres in a cell homogenizer and subsequent solution of the
pigments in acetone (Lorenzen, 1967). Chlorophyll a is altered by its
exposure to acid in the gut of organisms feeding on phytoplankton.
Pheophorbide a develops from chlorophyll by removal of the phytol chain
and by the loss of Mg via chlorophyllide a or if the Mg is released
at first, via pheophytin (Shuman and Lorenzen, 1975).
During Meteor cruise 36, February-March 1975, samples were taken from
three transects (Fig. 2; Table 1). Transect 2 is the southernmost one,
situated in the central upwelling area off Cape Blanc (Spanish Sahara) ,
from which much information on upwelling exists. Transect 1 lies 250 sm
to the north, off Cape Perra Grande, where upwelling is weaker.
Precipitation in the Sahara amounts to less than 250 rom per year and
fresh-water run-off is practically zero in the regions where transects
1 and 2 were taken. Annual primary production is estimated to be 0.583 g
132
C m- 2 day-l for the areas of Cape Blanc-Nouakchott, 0.447 g C m- 2 day-l

for B. de Gorrei to Cape Blanc and 0.306 g C m- 2 day-l for B. de Garnet
to B. de Gorrei (Schulz and Kaiser, 1974). A further decrease in primary
production is to be expected further north off Cape Pena Grande (trans-
ect 1). Transect 3 was positioned off Cape Blanc du Nord, Morocco
(termed Cape Mazagan in some publications), from which geological data
were available from Meteor cruise 8, 1967 (Giesel and Seibold, 1968;
Closs et al., 1969; Kudrass, 1973; Thiede, 1971). Morocco belongs to
the subtropical region with a precipitation of 250-500 rom per year
(c.f. Portugal 500-1000 rom) and with the rainy season during the win-
ter. Several rivers discharge their waters into the ocean, increasing
the concentration of nutrients and thus of primary productivity. Fur-
ther south along the Morrocan coast, upwelling intensity increases
toward the Canary Islands. Therefore, transect 3 is a compromise be-
tween low upwelling intensity and climatic conditions, with an unknown
amount of nutrients washed into the ocean by river discharge.
Marked differences between the transects were found (Fig. 4) in the

2-cm 2 sediment samples taken down to 5 cm depth. Transect 2 differs
from transect 3 by approximately two orders of magnitude, while trans-
ect 1 falls in between. Values of chloroplastic pigments measured by
Boucher and Glemarec (1974) for the area off southern Morocco lie be-
tween the figures for transects 1 and 3. The curves of Meteor transects
1 and 2 show a parallel course in the depth range of 80 to 400 m, with
minimum values at 200 m. A similar increase in chloroplastic pigments
from shallow depths to 80-100 m was found by Boucher and Glemarec
(1974). The values decrease down to their 200-m depth station, and no
deeper stations were sampled. Despite the differences in space and
time, the curves for chloroplastic pigments (Fig. 4) exhibit a striking
resemblance to those found for the meiofauna (Fig. 3). Explanations
for these similarities can be made using the same arguments as above
for meiofauna distribution. The prevailing current system, with the
strongest intensity of the undercurrent at a depth of 200-300 m, is
believed to be responsible for a low sedimentation rate of organic
matter, indicated by the minimum content of chloroplastic pigments
at these depths. A detailed description of chlorophyll a and pheopig-
ment contents in the upper sediment layers will be presented elsewhere.
3.3 Benthic Megafauna
A photos led (Thiel, 1970) was used to obtain some information on the
fishes and on the invertebrate megafauna, including the larger filter
feeders and predators, which are not reliably sampled by grabs. How-
ever, the photographs also provide some indication of the nature of the
sediment type and of its structure, data which are also useful in the
interpretation of upwelling processes. In transects 1 and 2 the sedi-
ments are coarse, and larger particles, such as shells and gorgonarian
pieces, are seen on the surface. The two transects differ markedly in
surface structure. Transect 1 shows, at a depth of 80 m, some smaller
mounds and craters, while in transect 2 the sediment surface is char-
acterized mainly by ripple marks. These are well developed and regular
in the depth range of 60-80 m, but down to a depth of 100 m the crests
are less regular and rounded off.
Within the visible fauna the sea pen, Pennatu7,a rubra (identified by Dr.
M. Grasshoff, Frankfurt), is the most ~rOminent species on transect 1.
On the average, one individual per 3 m (the area covered by each photo-
graph) is seen together with a few small fishes. With an average of
8 sea pens and with a maximum of 20, the frames from transect 2 show
a higher density of sea pe~s with larger individuals. The mobile fauna
is mainly shrimps, with up to three specimens per 3 m2 • Transect 3 ex-
133
dept hIm)
£/0
J \0
<x>oo 0 0 o~
!
0 08
JI
500
'Meteor' 36
transect _
00 0
1000
I
00
I
co
00 0
0
I
/
o 0
0 0
00
2000 0
2500
00
3000 00
Fig. 4. Concentration of chloroplastic pigments from three transects in the eastern

Atlantic, Meteor cruise 36, 1975: transect 1 off Cape Pena Grande, Spanish Sahara;
transect 2 off Cape Blanc, Spanish Sahara; transect 3 off Cape Mazagan, Morocco.
Chlorophyll a and pheopigment were measured in sediment samples with a surface area
of 2 cm2 and a depth of 5 cm; the concentrations are expressed in ~g g-1
hibits a fine-grained sediment with a smooth surface. There are a few

mounds and craters and some Lebensspuren. Animals are rare; single
echinoids, asteroids, and some fishes were photographed, but there were
no filter feeders. A detailed quantitative analysis of the sled photo-
graphs has yet to be made.
134
Benthic associations characterized by filter feeders were already de-

scribed from the upwelling areas off Northwest Africa. Off Cape Barbas
Newton et al. (1973) found the bivalve Pinna ramuZosa at a depth between
50 and 60 m with densities of 50-100 individuals/m2. In the same region,
I obtained a few grab samples from 40-57 m depth containing 1600 speci-
mens/m 2 of Branchiostoma senegaZense (Acrania) with a wet weight of 450
g/m 2 at depths between 25 and 30 m, decreasing to only a few individuals
at 40 m and deeper. Off Spanish Sahara this population extends over
approximately 240 miles (23-26 0 W).
The coarser sediments, photographed on transects 1 and 2, indicate

stronger currents. These in turn are capable of particle transport as
a food source for filter-feeding epifauna. Observations on sediments
and fauna are in accordance with each other, and both are related to
the currents and the upwelling processes.
4. Relations Between Upwelling Processes and Benthic Standing Stock

and Production
World-wide studies on the distribution of benthos standing stock have

revealed a relation to primary and secondary production in the overly-
ing water masses and to the transport of organic matter to the sedi-
ments as well as its input into the sediments. Therefore, high benthic
standing stock, as a first approximation for benthic production, is to
be expected in upwelling areas. The data so far available confirm this
conclusion. The question to be posed in this context is: How can ben-
thos res~arch contribute to the understanding of upwelling processes?
An important link in upwelling ecosystems is the total primary produc-

tion per year, which in itself is difficult to assess. Nutrient flow
into surface waters and subsequent primary production are variable in
space and time. Long-term research programs are needed to investigate
upwelling pulses, nutrient enrichment, horizontal transport of the up-
welled water, primary production, and the time intervals between plank-
ton blooms. Single upwelling events, including biological processes in
the surface water layers, are of the duration of several days to a
few weeks. Year-round investigations are necessary for elucidation of
total primary production per year.
Compared to the short life cycles of plankton organisms, benthic ani-

mals exhibit a much longer life span. Life cycles of the benthos from
upwelling areas are not known, but i t can be deduced from species liv-
ing in other regions. For the macrofauna, one, two, or even several
years life span is a conservative estimate, while the meiofauna may
have life spans of a few months to one year or longer. Benthic pro-
cesses are, on the average, more continuous and thus integrate the
short time pulses of plankton production and subsequent food sedimen-
tation. Benthos production and standing stock as a first approximation,
therefore, will allow an estimate of long-term primary production and
upwelling intensity.
This interpretation is oversimplified, and thus, several restrictions

should be considered. Upwelling associated with strong winds may result
in primary production not equivalent to the nutrient transport. The
drift of the water is too fast and the water sinks by downwelling be-
fore the nutrients are optimally used. In some cases, river discharge
brings nutrients into the ocean and primary production then will be
higher than that equivalent to upwelling.
135
Similarly, the biological processes involved in the transfer of organic

matter between primary production and benthic utilization are not the
same in all regions. Sinking of organisms and detritus is a slow pro-
cess in itself, but it is faster when the organic matter is concentrated
into fecal material by fishes, crustaceans, and other planktonic or-
ganisms. Blackburn and Thorne (1974) described vertical migration of
nekton, particularly of the red crab PleUl'oncoilEs pZanipes, and of the
anchovy EngrauZis mordax from the upwelling off Baja California. Down-
ward migration includes the fast transport of fecal material to the
bottom. In many animals, feeding on high food concentrations results
in low energy uptake efficiency and, consequently, in high-energetic
fecal material. High densities of plankton-feeding fishes, indicative
of short food chains (Ryther, 1969), are reported from all upwelling
areas. This indicates a rapid transfer in the plankton-nekton food
chain, and a fast transport of organic matter with high energy content
down to the sediment surface. Degradation of organic matter, so far,
was considered to be independent of its production. However, Nakajima
and Nishizawa (1972) observed a higher decay deeper in the water column,
where they measured the higher concentration of organic matter in sur-
face waters. If these findings are applicable to other areas, the higher
production in upwelling regions should be combined with relatively
high degradation rates during sinking, in comparison to less productive
waters. Absolute input of organic matter to the sediment would be great-
er in regions of higher production levels. However, the ratio of pro-
duction to sedimentation would decrease. Hargrave (1972) tried to re-
late oxygen uptake at the sediment surface to primary production. Again,
benthic metabolism increased with increasing primary production. How-
ever, with an increase in the depth of the mixed layer, both grazing
and recycling are enhanced in the pelagic system and relatively less
energy is transfered to the bottom.
In addition to biological processes, physical conditions are also im-

portant in the connection between surface production and benthos. Off-
shore downwelling fronts are described by Fraga (1974) and by Walsh
(1976). The transport of organic matter into deeper layers by down-
wardly directed water movement has to be considered. All the sinking
processes have additional horizontal components due to surface currents
directed toward the equator and to the poleward-flowing undercurrents.
Organic particles may settle far away from their area of origin. In
turn, physical conditions influence the energetic status of the par-
ticles, because the time necessary for degradation changes according
to horizontal transport, water depth, and drifting time.
In conclusion, variability of food available to the benthos is con-

trolled by a number of physical and biological factors. However, the
general distribution of benthos standing stock does reflect the gross
pattern of production in ocean surface layers. This implies that phys-
ical conditions and differences in degradation are of minor importance.
Benthic standing stock is indicative of primary production in arid
climates, and thus also of biologically effective upwelling. Some ex-
amples, covering a few steps in the food chain, exist for similar re-
lationships. Barber (pers. corom.) found a good correlation between
photosynthesis and fish yield in upwelling regions. Peterson (1972)
compared yearly catches of the dungeness crab (Cancer magister) to upwel-
ling indices (m 3 x s-l x 100 m- 1 coastline) along the west coast of
the United States for a period of more than 20 years. He found a good
relationship between upwelling and crab catch per year, taking into
account a time lag of 1.5 years for northern California and Oregon,
and of 0.5 years for Washington. Further observations of the shrimp
PandaZus jordani and of the razor clam SiZiqua patuZa, similarly affected
by upwelling, are cited by Peterson and Miller (1975).
136
Hargrave and Peer (1973), in their studies on the benthos of Canadian

east coast inshore waters, tried to relate macrofauna standing stock
to average chlorophyll concentrations in the water masses. They could
not detect a strong correlation between macrofaunal biomass and average
yearly chlorophyll concentration, but they were able to demonstrate
this correlation for the time of the spring plankton bloom (March-May).
Later in the year sinking may have been inhibited by a discontinuity
layer. In upwelling regions, each upwelling event may initiate another
"spring" bloom. The differences in chloroplastic pigments, contained
in the sediments of the transects off Northwest Africa (Fig. 4), are
an indication of this phenomenon. Similar trends in population den-
sities of benthic organisms in the Northwest African upwelling region
have been encountered by Rliger (1975) and Weyland (pers. corum.) for
bacteria and by Gaertner (pers. corum.) for fungi.
This report on quantitative observations of benthic organisms and some

sediment characteristics from upwelling regions shows clearly the limited
amount of information available. However, results to date on the benthos
as an indicator of upwelling allow to assume that biological processes
in the water column are integrated by the benthos over longer time pe-
riods. Benthos standing stock and production are regarded as presen-
tative of the intensity of primary production in surface waters includ-
ing upwelling regions.
Benthos research in upwelling and comparable non upwelling areas must

be intensified if this part of the upwelling ecosystem is to be under-
stood. The total benthos, its activity measured by oxygen consumption,
ATP, ETS, and its productivity, as well as sediment properties and
nutrient recycling, should be investigated within one geographical re-
gion under at least similar climatic conditions, but differing in up-
welling intensity, to establish basic data for the comparison of widely
separated upwelling regions.
AcknowLedgments. Thanks are due to 'Dr. R. Boje, Kiel, for advice and technical sup-
port for the determination of chloroplastic pigments and to my technical staff for
the tiring task of sorting the meiofauna samples. I am grateful to Dr. F. Nichols,
San Francisco, and Dr. A. Rice, Wormley, whose criticism and linguistic help resulted
in a much improved paper.
This research was supported by the Deutsche Forschungsgemeinschaft (Schwerpunktpro-
gramm: Auftriebsphanomene im Meer) and by the Sonderforschungsbereich 94 - Meeres-
forschung Hamburg.
Note: Literature and results obtained after December 1975 are not included in this
contribution.
References
Banse, K.: Hydrography of the Arabian Sea Shelf of India and Pakistan and effects
on demersal fishes. Deep-Sea Res. 12, 45-79 (1968)
Blackburn, N., Thorne, R.E.: CompOSition, biomass and distribution of pelagic nekton
in a coastal upwelling area off Baja California, Mexico. Tethys~, 281-290 (1974)
Boucher, D., Glemarec, M.: Donnees preliminaires sur le benthos de la cote sud de
l'Atlantique Marocain. Tethys 6, 29-32 (1974)
Calvert, S.E., Price, N.B.: Recent sediments of the South West African shelf. In:
Delany, F.M. (ed.): The geology of the East Atlantic continental margin. 2. Africa.
Rep. No. 70/16, Inst. geol. ScL, p. 175-185 (1971)
Closs, H., Dietrich, G., Hempel, G., Schott, W., Seibold, E.: "Atlantische Kuppen-
fahrten 1967" mit dem Forschungsschiff "Meteor" - Reisebericht. "Meteor" Forsch.
Ergebn. A~, 1-71 (1969)
Diester-Haas, L.: Sediments as indicators of upwelling: In: Upwelling Ecosystems.
R. Boje, M. Tomczak (eds.). Berlin-Heidelberg-New York: Springer 1978
137
Ekman, S.: Zoogeography of the Sea. London: Sidgwick and Jackson, Ltd., 1970
Filatova, Z.A.: Quantitative distribution of deep-sea benthic fauna. In: Deep-Sea
Bottom Fauna, Pleuston. Zenkevich (ed.). Tikii Okeana: BioI. Tikh. Okean., 1969,
7 (2), pp. 234-252
F~ga, F.: Distribution des masses d'eau dans l'upwelling de Mauritanie. Tethys 6,
5~10 (1974) -
Frankenberg, D., Menzies, R.J.: Some quantitative analyses of deep-sea benthos off
Peru. Deep-Sea Res. ~, 623-626 (1968)
Giesel, W., Seibold, E.: Sedimentechogramme vom iberomarokkanischen Kontinentalrand.
"Meteor" Forsch. Ergebn. C .!., 53-75 (1968)
Gosselck, F.: A quantitative survey of the macrobenthos communities on the continental
shelf off North-West-Africa. ICES, Shellfish and Benthos Comm. C.M. K30, 1-3 (1974)
Gosselck, F.: The distribution of Branchiostoma senegalense (Acrania, Branchiostomi-
dae) in the offshore shelf region off North West Africa. Int. Revue Ges. Hydrobiol.
60 (2), 199-207 (1975)
Hargrave, B.T.: Coupling carbon flow through some pelagic and benthic communities.
J. Fish. Res. Bd. Canada 30, 1317-1326 (1972)
Hargrave, B.T., Peer, D.L.: Comparison of benthic biomass with depth and primary pro-
duction in some Canadian east coast inshore waters. ICES, Shellfish and Benthos Comm.
C.M. E, 1-16 (1973)
Hart, T.J., Currie, R.I.: The benguela current. Discovery Rep. 11, 123-298 (1960)
Horn, W., Mein9ke, J.: Note on the tidal current field in the continental slope area
off Northwest Africa. Mem. Soc. Roy. Sci. Liege, 6. Ser. 10, 31-42 (1976)
Kirkegaard, T.B.: The polychaeta of West Africa. Part 1. Sedentary species. Atlantide
Rep. 5, 7-117 (1959)
Khusid,-T .A.: Distribution of benthic foraminifers off the West Coast of South
America. Oceanology !i, 900-904 (1974)
Kudrass, H .-R.: Sedimentation am Kontinentalhang vor Portugal und Marokko im Spat-
pleistozan und Holozan. "Meteor" Forsch. Ergebn. C .!1., 1-63 (1973)
Lange, H.: Herkunft und Verteilung von Oberflachensedimenten des westafrikanischen
Schelf- und Kontinentalhanges. "Meteor" Forsch. Ergebn. C. 22, 61-84 (1975)
Lorenzen, C.J.: A method for the continuous measurement of in vivo chlorophyll con-
centration. Deep-Sea Res. 11, 223-228 (1966)
Marchig, V.: Zur Geochemie rezenter Sedimente des indischen Ozeans. "Meteor" Forsch.
Ergebn. C 11, 1-104 (1972)
Maurin, C.: Ecologie ichtyologique des fond chalutables Atlantiques (de la Baie
Ibero-Marocaine a la Mauritanie) et de la Mediterranee occidentale. Rev. Trav.
Inst. Pech. Marit. 32, 1-47 (1968)
Miro Orell, De M.: Sedimentos recientes del margen continental de Mauritania (expe-
dici6n Sahara II). Res. Exp. Cient •. blo Cornide 2, 1-12 (1973)
Mittelstaedt, E.: Der hydrographische Aufbau und die zeitliche variabilitat der
Schichtung und Stromung im nordwestafrikanischen Auftriebsgebiet im FrUhjahr 1968.
"Meteor" Forsch. Ergebn. A 11, 1-57 (1972)
Mittelstaedt, F.: On the currents along the Northwest African coast south of 22~.
Dt. Hydrogr. Z. 29 (3),97-117 (1976)
Murty, P.S.N., Reddy, C. V.G., Varadachari, V. V.R.: Distribution of organic matter
in the marine sediments off the west coast of India. Proceed. Natl. Inst. Sci.
India, Vol. 35, B, ~, 377-384 (1969)
Nakajima, K., Nishizawa, S.: Exponential decrease in particulate carbon concentra-
tion in a limited depth interval in the surface layer of the Bering Sea. In: Bio-
logical Oceanography of the Northern North Pacific Ocean. Takenouti, A.Y. (ed.).
Tokyo: Idemitsu Shoten, 1972, pp. 495-505
Newton, R.S., Seibold, E., Werner, F.: Facies distribution patterns on the Spanish
Sahara continental shelf mapped with side-scan sonar. "Meteor" Forsch. Ergebn.
C ~, 55-57 (1973)
Pamatmat, M.M.: Oxygen consumption by the seabed. IV. Shipboard and laboratory ex-
periments. Limnol. Oceanogr. 16 (3), 536-550 (1971)
Peterson, W.T.: Qpwelling indices and annual catches of dungeness crab, Cancer ma-
gister, along the West coast of the United States. Fish. Bull. U.S. 21, 902-910
(1972)
Peterson, W.T., Miller, C.B.: Year-to-year variations in the planktology of the Oregon
upwelling zone. Fish. Bull. 73, 642-653 (1975)
138
Pflaumann, U.: Late quaternary stratigraphy based on planktonic Foraminifera off

Senegal. "Meteor" Forsch. Ergebn. C 23, 1-46 (1975)
Rowe, G.T.: Benthic biomass and surface productivity. In: Fertility of the Sea.
Costlow, J.D. (ed.). ~, 441-454 (1971a)
Rowe, G.T.: Benthic biomass in the Pisco, Peru upwelling. Inv. Pesq. 35, 127-135
(1971b)
Rowe, G.T., Menzel, D.W.: Quantitative benthic samples from the deep Gulf of Mexico
with some comments on the measurement of deep-sea biomass. Bull. Marine Sci. 11
(1-2), 556-566 (1971)
Rowe, G.T., Polloni, P.T., Horner, S.G.: Benthic biomass estimates from the north-
western Atlantic Ocean and the northern Gulf of Mexico. Deep-Sea Res. ~, 641-650
( 1974)
Ruger, H.J.: Bakteriensporen in marinen Sedimenten (Nordatlantik, Skagerak, Biscaya
und Auftriebsgebiet vor Nordwestafrika) - quantitative Untersuchungen. Veroff. Inst.
Meeresforsch. Bremerh. 12, 227-236 (1975)
Ryther, J.H.: Photosynthesis and fish production in the sea. Science .166, 72-76
( 1969)
Sanders, H.L.: Marine benthic diversity: A comparative study. Am. Naturalist 102,
243-282 (1968)
Sanders, H.L.: Benthic marine diversity and the stability-time hypothesis. Brookhaven
Symp. Biol. 22, 71-80 (1969)
Sanders, H.L., Hessler, R.R.: Ecology of the deep-sea benthos. Science 163, 1419-1424
(1969)
Sankaranarayanan, V.N., Qasim, S.Z.: The influence of some hydrographical factors
on the fisheries of Cochin area. Bull. Natl. Inst. Sci. India~, 846-853 (1968)
Schulz, S., Kaiser, W.: Estimation of the productivity of the NW African upwelling
region. ICES, Plankton Comm. C.M. L~, 1-5 (1974)
Shuman, F.R., Lorenzen, C.J.: Quantitative degradation of chlorophyll by a marine
herbivore. Limnol.Oceanogr.1Q. (4),580-586 (1975)
Siedler, G., Seibold, E.: Currents related to sediment transport at the Ibero-Maroc-
can contineptal shelf. "Meteor" Forsch. Ergebn. A H, 1-12 (1974)
Smith, K.L., Jr., Rowe, G.T., Clifford, C.H.: Sediment oxygen demand in an outwelling
and upwelling area. Tethys ~ (1-2), 223-230 (1974)
Thiede, J.: Planktonische Foraminiferen in Sedimenten vom ibero-marokkanischen Kon-
tinentalrand. "Meteor" Forsch. Ergebn. C 7, 15-69 (1971)
Thiel, H.: Ein Fotoschlitten fur biologische und geologische Kartierungen des Meeres-
bodens. Marine Biol. 2, 223-229 (1970)
Thiel, H.: The size structure of the deep-sea benthos. Int. Revue Ges. Hydrobiol.
60, 575-606 (1975)
Tlirkay, M.: Decapoda Reptantia vom Kap Blanco. Auswertung der FAhrt 26 (1972) von
F.S. "Meteor". "Meteor" Forsch. Ergebn. D 20, 71-75 (1975)
Walsh, J.J.: A biological sketchbook for an eastern boundary current. In: The Sea.
Steele, J.H. et. al. (eds.). wiley-Interscience, 1977, ~, 923-968
Weichart, G.: Meereschemische Untersuchungen im nordwestafrikanischen Auftriebsge-
biet 1968. "Meteor" Forsch. Ergebn. A ii, 33-70 (1974)
Role of Bacteria in an Upwelling Ecosystem
S.WWATSON
The role that bacteria play in the carbon cycle in an upwelling eco-
system or in any other marine ecosystem is poorly defined. To estimate
the role that bacteria might play in the oceans one must first examine
the source of substrates available to them. Many investigators have
studied primary production in the world's oceans (Steemann-Nielsen and
Jensen, 1957; Ryther, 1963). In general these studies have shown that
25-75 g of Care fixed/m 2 /year in the open ocean. In coastal regions
usually 100 g of Care fixed/m 2 /year, but in upwelling areas, such as
off the coast of Africa, Peru, and in the Arabian Sea, up to 300 g or
more of C may be fixed/m 2 /year (Ryther, 1969)
The questiori is what percentage of this 300 g of carbon is mineralized

by bacteria and what percentage is utilized by other trophic levels.
Ecologi~ts in general seem to believe that everything in the ocean is
eaten and that nothing over dies (Cushing, 1959). Indeed, the work of
Menzel and Ryther (1961) in a nutrient poor area such as the Sargasso
Sea would tend to verify this hypothesis. However, in coastal waters
this hypothesis must be questioned. Riley (1956) found that zooplank-
ton utilized no more than 25% of the carbon fixed by plants in Long
Island Sound. He postulated that the remaining 75% of the phytoplankton
cells were either decomposed in situ in the water column or sank to
the bottom where they were buried or were decomposed by bacteria. If
a similar situation occurs on the continental shelf in upwelling areas,
100-200 g of carbon/m 2 would be directly available for bacterial min-
eralization.
Another source of soluble organics available to bacteria is that ex-

creted by phytoplankton cells. Fogg (1966) indicated that 50% or more
of the photoassimilated carbon could be released by phytoplankton into
the water as dissolved organic compounds. Hellebust (1965) studied 22
species of unicellular algae in culture and found that most of them
excreted 3-6% of their photoassimilated carbon during the logarithmic
phase of growth, but a few species excreted 10-25% of their photoas-
similated carbon during this growth period. Hellebust also concluded
that natural populations excreted 4-16% of their photoassimilated carbon
during the peak of the bloom and 17-38% of the carbon at the end of the
bloom. Parsons and Seiki (1970) suggested that a maximum of 30% and
an average of 15% of total carbon fixed by phytoplankton is released
as soluble organic compounds. From this estimate i t appears that 30 g
of organic carbon might be excreted/m 2 /year in an upwelling area.
From studies by Ryther .( 1969) and others we know that the energy cap-
tured by photosynthetic processes and stored in organic compounds is
utilized by 3-5 additional trophic levels. We also know that this trans-
fer of energy is not an efficient process. Slobodkin (1961) estimated
that only 10% of the organics assimilated at each trophic level was
transferred and utilized for cellular growth by the organisms of a
higher trophic level. Ryther (1969) suggests that the efficiency varies
from 10-20% depending upon the geographic location in the ocean.
140
While organisms of each trophic level rarely utilize more than 10-20%
of the assimilated carbon for cellular growth, they probably utilize
on the average another 30-40% of the assimilated carbon for maintenance
energy. If only 50-60% of the organics assimilated are utilized for
cellular growth and maintenance energy, then 40-50% of the organics
must be accounted for. It seems obvious, a priori, that such organics
are either buried in the sediment or that they are decomposed by hetero-
trophic bacteria. We believe that 40-50% of the organics ingested at
each trophic level are excreted as soluble or particulate organic mat-
ter and that bacteria subsequently decompose such compounds. Thus even
if we assume that members of the first trophic level eat 300 g of C/m2/
year, they probably excrete 120-150 g C/m 2 /year, and this amount should
be available for bacterial mineralization.
If we consider the three sources of organics just mentioned, it seems

likely that in an upwelling area 150-200 g of organics would be min-
eralized by bacteria/m 2 /year. If this amount of carbon is mineralized
by bacteria, then one should find a significant concentration of dis-
solved organics in the water. According to Wangersky (1965) most of
the organic matter in the ocean exists in soluble form. Duursma (1961)
found an increase of 20 g of soluble C/m2 after the spring bloom and
an increase of 6 g C/m2 after the fall bloom of phytoplankton. From
these observations Duursma calculated that 52 g of soluble carbon would
be produced m2 /year. Studies by Andrews and Williams (1971) suggested
that bacteria in the English Channel oxidize 2.6 g of glucose/m2 and
29 g of amino aCids/m 2 each year. These authors also suggested that
heterotrophic processes may utilize 100 g of C/m 2 /year which would be
50% of the carbon fixed annually by photosynthetic processes in the
English Channel.
These few literature references seem to verify the hypothesis that

over 50% of the organics produced by photosynthetic processes are uti-
lized by bacteria at least in shallow waters of the ocean. The findings
of Andrews and Williams (1971) also suggest that a large percentage
of these soluble organics is composed of small molecular weight com-
pounds that are readily mineralized by bacteria.
If bacteria really mineralize 150-200 g of C/m 2 /year one can calculate

the number of bacteria produced annually in an upwelling area. This
calculation is based on the following assumptions: (1) that the average
volume of a marine bacterium is 0.1 ~3 (snhere with radius of 0.29 ~);
(2) that each bacterium contains 10 x 10-15 g of C; (3) that each bac-
terium would utilize about 50% of the organic carbon for cellular
synthesis and the remaining 50% for maintenance energy; (4) that the
water depth is approximately 100 m and (5) that all mineralization
takes place within the water column .. Based on these assumptions 0.75-
1.0 x 10 14 bacteria would be produced/m 3 /year and this would be equi-
valent to 0.75-1.0 x 10 8 bacteria/ml. If the bacteria had an average
generation time of 36.5 days the standing crop would be 0.75-1.0 x 10 7
bacteria/ml, or if the generation time were 3.65 days a standing crop
of 0.75-1.0 x 10 6 bacteria/ml should be found. However, if we assume
that 90% of the mineralization takes place in the sediment and that the
bacteria have a 3.65-day generation time, then we should find about
0.75-1.0 x 10 5 bacteria/ml in the water column in a rich upwelling
area, assuming that all of the mineralization takes place in the im-
mediate area of high productivity.
Now that we have formulated a working hypothesis on the role and con-
centration of bacteria in an upwelling area, let us examine the evi-
dence to see if this hypothesis can be experimentally verified.
141
Numerous investigators have attempted to enumerate by cultural and

direct counting techniques the concentration and distribution of bac-
teria in the ocean. In general, surveys employing cultural techniques
have revealed only 5-1000 bacteria/ml (Sieburth, 1971; Gunderson et
al., 1972), but it has long been known that such techniques reveal
only a small fraction of bacteria present (ZoBell, 1946). However,
Kriss (1963) reported finding 2.9 x 10 3 to 2.3 x 10 4 bacteria/ml in
the upper 100 m of Pacific waters. Unfortunately, most other investiga-
tors have not been able to experimentally verify Kriss's findings and
his results are thus regarded with some degree of skepticism.
If sea-water contains 10 5-10 6 bacteria/ml, it would seem that it should

be a comparatively easy task to enumerate them by direct counting tech-
niques. Direct microscopic observations, such as conducted by Jannasch
and Jones (1959) and by Kriss (1963), are greatly subject to error. Un-
fortunately the concentration of detrital material of similar size and
density to bacteria exceeds the concentrations of bacteria. Jannasch
and Jones (1959) compared the efficiency of five cultural and two di-
rect counting methods and found 13-9700 times more bacteria with direct
methods than with cultural methods. The authors admitted difficulty in
distinguishing small microbial forms from inorganic particles. Jannasch
and Jones co,unted up to 10 4 bacteria/ml in their sea-water samples
using a 1% solution of erythrosine in 5% phenol for individual cell
counts. These are some of the highest direct counts that have been re-
ported from oceanic environments.
Fluorescent techniques have also been employed to enumerate the concen-

tration of bacteria in samples of sea water. Using such techniques the
water sample is either stained and then filtered onto membrane filters
(Jannasch, 1958), or the cells are stained on the filters (Deufel,
1959; Schwantes, 1971). However, many p:r;oblems have been encountered
using fluorescent techniques (Madsen, 1972; Wiebe and Pomeroy, 1972).
Recently a new method for fluorescence staining of bacterial popula-

tions on membrane filters has been described that overcomes most of
the difficulties encountered using previous techniques (Zimmerman and
Meyer-Reil, 1974). With this technique, bacteria are concentrated on
Nulcepore membranes, stained with acridine orange, destained, and ex-
amined with an epifluorescent microscope. We recently tried out this
technique with good success. Unfortunately this technique was not em-
ployed in our upwelling studies but it certainly will be used in future
studies.
Few investigators thus far have employed fluorescence techniques with

any degree of success in oceanic studies. Hobbie et al. (1972), employ-
ing such techniques in open ocean studies, reported finding only 1 bac
terium/ml. Fluorescence techniques have also been employed to count
bacteria in sediments (Dale, 1974). Dale reported counts as high as
9.97 x 10 9 bacteria per gram of dry sediment in sediments from estuaries,
and this biomass is equivalent to macrofaunal biomass.
Indirect techniques have also been employed to estimate microbial bio-

masses in the marine environment. The best known of these indirect
methods is the adeno9ine triphosphate (ATP) method first employed by
Holm-Hansen and Booth (1966). Since ATP is rapidly destroyed after the
death of an organism (Hamilton and Holm-Hansen, 1967), the ATP method
is accurate for estimation of viable biomass. The only shortcoming of
this test is that it does not allow the investigator to determine the
amount of biomass contributed by photosynthetic organisms versus other
microbial populations. It is generally assumed that the ratio of cell
carbon to ATP is 250 but this can vary by a factor of three (Hamilton
and Holm-Hansen, 1967).
142
When the ATP technique is used on samples collected from below the
euphotic zone, this method may provide a fairly accurate estimate of
the bacterial biomass existing in such samples. At these depths Hobbie
et al. (1972) found from 2-39 x 10- 12 g of ATP/ml. If all of this ATP
were derived from bacteria, this concentration of ATP would indicate
a concentration of 2-39 x 10 3 bacterial/ml equivalent in size to Esche-
richia coH.
Until recently it has not been possible to use the ATP method success-
fully on sediments. However, in the last year Christian et al. (1975)
described a method to extract ATP using hot bicarbonate. Using this
method in our laboratory we were able to obtain 100% recovery of ATP
from samples spiked either with ATP or bacterial cells. This is the
method we have utilized on sediments. Others (Hobson et al., personal
communication; Karl and LaRock, 1975) extract with cold acid and re-
port that this method is reliable.
In recent years isotope tracer techniques have been employed to study

the metabolic activity of heterotrophic microorganisms. Parsons and
Strickland (1961) were the first to modify isotope tracer techniques
to measure the heterotrophic potential of microorganisms in sea-water.
This technique was subsequently employed by several investigators to
study substrate uptake kinetics of fresh-water and marine microorganisms
(Wright and Hobbie, 1966; Vaccaro and Jannasch, 1966, 1967; Wetzel,
1967; Allen, 1967; Hobbie et al., 1968). Vaccaro and Jannasch (1966,
1967) used isotope kinetic studies for determining rates of assimila-
tion, efficiency, turnover time of substrates, and for assessing the
microbial heterotrophic potential of the sea. Wright and Hobbie (1966)
showed through kinetic studies that the heterotrophic potential of mixed
bacterial and algal populations could be separated. Further studies
compared uptake kinetics of natural populations and pure culture (Hobbie
et al., 1968). Parsons and Strickland (1961) assumed that mixed popula-
tions of cells would obey the laws of enzyme kinetics, but Vaccaro and
Jannasch (1967) and Williams (1973) have shown that the primary assump-
tion that bacteria obey the Michelis-Menton type reaction holds true
only when there is a dominant population. As previously indicated,
Andrews and Williams (1971) used the 14C technique to estimate the
rates at which amino acids and glucose were mineralized in the English
Channel. To date, most of the isotope work has concentrated on uptake
kinetics of heterotrophic potential without attempting to correlate
this activity to the concentration of bacteria present at various depths.
Nor has the isotope work done on mineralization rates in the sea been
equated with the existing bacterial populations.
Upon perusal of the literature it became immediately obvious that the

techniques used were inadequate to estimate bacterial biomass in the
ocean and yielded l~ttle information concerning the distribution of
bacteria in the marine environment. During the last three years we have
been trying to develop a new assay, referred to as the Limulus amebocyte
lysate (LAL) technique or lipopolysaccharide test (LPS test) to deter-
mine the concentration and distribution of bacterial biomass in marine
environments.
The LPS test is based on a test described by Levin and Bang (1964).
LAL, an essential ingredient used in this test, is an aqueous extract
made from the blood cells of the horseshoe crab, Limu"lus po"lyphemus. When
LAL is reacted with lipopolysaccharides, which are components of the
cell wall of all gram-negative bacteria, a firm clot is formed. The
lipopolysaccharides activate an enzyme in the LAL (Young et al., 1972).
Once activated, this enzyme triggers a low-molecular-weight protein
to polymerize and clot. This test is specific for lipopolysaccharides
from gram-negative bacteria and can detect picogram quantities/ml of
143
these compounds. The test is so sensitive that 100 bacteria/ml can be

detected. We believe that this test can be used for bacterial biomass
determinations in sea-water since most bacteria in this environment
are gram-negative, and since LPS comprises more or less a constant
percent of the dry weight of each cell.
There are still obstacles that must be overcome in this test. For ex-
ample, this test does not distinguish between the LPS derived from
living and dead bacteria nor from free LPS existing in seawater. LPS
still associated with bacterial cells is referred to as bound LPS and
LPS existing free in seawater as free LPS. By centrifugation bound LPS
can be separated from free LPS.
We recognized early in these studies that the LPS test had to be used
in conjunction with standard techniques commonly used by other inves-
tigators. Thus in all of our upwelling studies we combined this test
with the ATP, chlorophyll a, and 14C techniques.
The Limulus test was performed in our studies by mixing 0.1 ml of the
LAL with 0.1 ml of an LPS standard or seawater sample to be tested.
This mixture was incubated in a 10 x 75 mm test tube for 1 h at 37 0 C.
After 1 h the tubes were inverted and scored positive if a firm gel
was formed that did not break when the tube was inverted.
To make this a quantitative test twofold dilutions were made of an LPS

standard. The standard we used was LPS from Klebsiella pnewnoniae obtained
from the United States Food and Drug Administration. Serial dilutions
were made of this standard from 1 ng/ml down to 0.01 ng/ml. Each dilu-
tion was reacted with the lysate and the minimum concentration of LPS
needed for clot formation was determined.
To determine the concentration of LPS in seawater, a sample was serially

diluted and the highest dilution causing clot formation was recorded
and this dilution factor was multiplied by the minimum concentration
of endotoxin standard needed for clot formation. The resulting answer
gave the ng/ml of LPS in the seawater sample.
Using the clot method of reading the test, the absolute concentration
of LPS in a seawater sample cannot be determined. Using twofold dilu-
tions one can underestimate the LPS concentration in seawater by nearly
a factor of two. In the last six months we have developed a spectro-
photometric assay that will provide a more accurate measurement of LPS
in seawater. When the test is read in a spectrophotometer, one can
detect as little as 0.02 pg/ml of LPS.
To determine if bacterial biomass could be quantitated using the LPS

test, the growth of E. coli was followed in culture using the LPS tech-
nique. Simultaneously, the growth of this bacterium was followed using
plate and direct counting methods as well as the ATP technique. Also,
the heterotrophic potential of the cells was followed using 14C-Iabeled
glycine as a substrate (Fig. 1). The growth of this bacterium in cul-
ture was followed for 9 h and samples were taken hourly for the anal-
yses. The re$ults were compared as indices for determining the number
of bacteria present at each time period and factors were derived for
use in field studies.
During growth, the bacteria excreted into the medium approximately

5-20% of the LPS produced. This fraction, called free LPS, was sep-
arated from the cellular LPS by centrifugation.
As shown in Figure 1 the growth of E. coli could be followed with all

techniques employed to estimate bacterial biomass through all phases
144
---.-- Fig. 1. Growth curve of Escherichia

I ~ ~ coli determined by various techniques
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Table 1. Cellular concentrations of ATP and LPS in cultures of Escherichia coli
Lag phase Logarithmic phase Stationary phase
fg LPS/cell 31.6 46.7 28.3

fg ATP/cell 0.8 0.42 0.85
LPS/ATP ratio 39.5 111.2 33.3
of growth. Table 1 presents the average concentration of LPS and ATP

per cell during the lag, log, and early stationary phases of growth.
As shown in this Table, both the LPS and ATP/cell varied during the
various growth phases. We thought initially that most bacteria in the
ocean would have LPS and ATP concentrations similar to those found in
the stationary phase of E. coli. Thus we concentrated on the LPS, ATP,
and carbon content of cells in this phase of growth. The results of
these studies are presented in Table 2.
However, in recent studies we found that the LPS and ATP concentrations
in marine bacteria differed markedly from those found in E. coli. The
average E. coli cell during the early stationary growth phase was
145
Table 2. ATP, LPS, and carbon concentrations of E. coZi cells in stationary phase
ATP fg
LPS 20 fg
Wet weight 2.5 pg
Dry weight 625 fg
Carbon/ATP 250
Carbon/LPS 12.5
LPS/ATP 20
2.0 x 1.26 with a volume of 2.5 ~3. The average cell volume of marine
bacteria growing in situ was 0.1 ~3. Thus the average volume of a marine
bacterium is 1/25th that of an E. coli cell in the early stationary
phase of growth. One would expect the marine bacteria to contain far
less carbon, LPS, and ATP per cell than do E. coli cells.
Although it is impossible to measure directly the carbon content of

marine bacteria growing in situ we can estimate how much carbon each
cell contains. Escherichia coli cells during the stationary phase of
growth contain 250 fg of C/cell and since the volume of a mari.ne bac-
terium is 1/2,?th that of an E. coli cell we estimate that each marine
bacterium would contain 10.4 fg of carbon.
To obtain a useful C/LPS ratio, the LPS content of marine bacteria

growing in situ must be measured, and this has been done on a recent
cruise (Watson et al., unpublished data). Direct counts of bacteria
were obtained using epifluorescence techniques (Hobbie, unpublished
data) and the LPS was measured with Limulus amebocyte lysate employing
a spectrophotometric technique. These studies indicated that the aver-
age marine bacterium growing in situ contained 2.5 fg/cell of LPS.
Thus the C/LPS ratio would be 4.16, and we have multiplied all our
LPS values by this number to obtain the bacterial carbon present in
each sample in the present investigation.
One of the criticisms of the LPS technique is that i t does not dis-
tinguish between living and dead bacteria. While this is a valid criti-
cism, we believe that dead bacteria do not remain intact for prolonged
periods of time in the natural environment, thus most of the bacteria
detected with the LPS technique are believed to be living. However,
additional studies are needed to clarify how much of the LPS measured
in the ocean is derived from living versus dead cells.
On the fourth leg of the Coastal Upwelling Ecosystems Analysis cruise

we attempted to measure the concentration and distribution of micro-
organisms utilizing the LPS, ATP, chlorophyll a, and 1 4 C techniques.
The station locations on this cruise are shown in Figure 2.
The bacterial, total microbial, and phytoplankton biomasses measured

in shallow and deep waters are summarized in Table 3. In the water
column of the shallow shelf stations the total microbial biomass, as
measured by the ATP method, averaged 6.25 g C/m2 while the bacterial
biomass averaged 0.59 g C/m2. In the shelf stations bacteria comprised
9.4% of the total microbial biomass. In the slope stations the average
total microbial biomass was 7.68 g c/m2 while the average bacterial
biomass was 4.03 g C/m2 indicating that 52.5% of the microbial biomass
was composed of bacteria. In the one offshore station the total micro-
bial biomass was 8.8 g and the bacterial biomass 8.3. These data sug-
gest that about 94% of the microbial biomass was composed of bacteria.
The total microbial, bacterial, and phytoplankton biomasses measured

in the top 20 meters of the water column of shelf, slope, and offshore
146
ATLANTIS .II 177-0

22"
CRUISE #82 LEG 4
••
167-C •
165-8
50'
176 169-H
4O' (LO '8"092') •
175 • 168-G
174'" ••
170 •
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• • ~
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173-L 162-1 ~
'0'
P 171-M
2'"
'S" 50' -'1O' '7" 30' 20'
•
'0' '7" 50'
Fig. 2. Location of stations on Cruise No 82, leg 4 of Atlantis I I
Table 3. Comparison of biomasses in the water column between shallow and deep sta-
tions mg C/m2
Area Station No_ Depth Bacterial Total Phytoplankton

biomass biomass biomass
Shelf 161 30 399 3277 1377

171 35 982 7510 6119
162 39 370 3457 3575
166 35 592 6628 4016
165 57 602 3665 3896
163 63 596 11497 10881
164 86 604 7734 4871
Slope 167 130 815 6973 4170

168 245 1506 10100 7065
174 365 3311 5584 2358
173 469 9160 4127 4265
177 366 3569 13114 1308
169 800 5791 6211 2948
Offshore 176 1800 8320 8754 630

147
Table 4. Comparison of biomasses in the top 20 m between shallow and deep stations
mg clrrJ2
Area Station No. Depth (m) Total Bacterial Phytoplankton

biomass biomass biomass
Shelf 161 30 2427 266 1138

171 35 3850 682 3735
162 39 2218 191 2431
166 35 3525 399 2460
165 57 1575 300 1659
163 63 4833 266 7445
164 86 3353 266 2798
Slope 167 130 2601 300 1976

168 245 1425 166 3013
174 365 1840 266 868
173 469 1078 799 1170
177 366 3340 333 443
169 800 954 732 1580
Offshore 176 1800 461 399 140
Table 5. Biomass averages per depth in slope and offshore stations
Depth (m) Bacterial biomass Total biomass Phytopl. biomass

mg c/m 3 mg C/m 3 mg c/m 3
0 29.0 71.9 61.0

10 32.0 50.5 57.5
20 10.0 110.1 64.1
30 23.3 28.5 106.5
40 15.0 63.4 44.2
50 13.0 52.4 66.5
60 20.0 38.7 14.0
70 6.7 32.1
80 22.0 9.9 9.0
90 5.0 7.4
100 10.0 9.2 8.7
200 7.7 7.6
300 9.0 3.9
400 9.0 10.1
500 3.3 3.1
600 3.3 1.0
700 3.3 0.5
800 1.7 0.8
1000 0.8 0.8
1200 0.8 1.0
1400 1.7 0.5
1600 1.7 0.8
1800 0.8 0.3
waters were compared (Table 4). The average total microbial biomass/m2
(expressed as grams of carbon) in the top 20 meters of the water column
in shelf, slope, and offshore waters was 3.1, 1.8, and 0.46 g C/m2. The
average bacterial biomass for these same three areas was 0.34, 0.43,
and 0.39 g C/m2. The average phytoplankton biomass for these areas was
3.1, 1.5, and 0.14 g C/m2.
148
These studies imply that the phytoplankton biomass concentrations,

while similar in shelf and slope waters, decreased markedly in offshore
waters but that the bacterial biomass concentrations remained relatively
constant in all three water masses. This suggests that the concentration
qf bacteria in different water masses is not directly proportional to
the concentration of phytoplankton present.
However, the data in Table 5 show that the vertical distribution of

bacteria observed within a given water mass would tend to indicate that
there is a relationship between bacterial and phytoplankton populations.
Bacteria were generally more abundant within the euphotic zone or just
beneath the euphotic zone than at deeper depths. However, at certain
depths below the euphotic zone the concentration of bacteria nearly
equalled that found within the euphotic zone. For example, a high con-
centration of bacteria was found at 400 meters (Table 6), which was
also the site of the oxygen minimum layer.
One of the questions asked in this investigation was whether most of

the mineralization of organics was occurring in the upwelling area or
whether the organics produced in the upwelling area were being trans-
ported and oxidized far out to sea. Table 6 presents the data from one
Table 6. Station 176, Depth 1800 m - biomasses at various depths in an offshore sta-
tion
Depth Bacterial Total Phytopl. LPS

(m) biomass microbial biomass biomass ATP
mg C/m 3 mg C/m 3 mg C/m 3 Ratio
0 20.0 27.0 6.0 44

10 20.0 21.75 7.5 55
20 20.0 21. 75 7.0 55
40 20.0 34.5 10.5 35
60 20.0 25.75 5.5 47
80 20.0 8.25 3.5 145
100 20.0 4.25 3.0 282
200 5.0 3.5 86
400 13.3 24.0 33
600 3.3 1. 75 114
800 1.7 0.75 133
1000 0.8 0.75 67
1200 0.8 1.0 50
1400 0.8 0.5 100
1600 1.7 0.75 133
1800 0.8 0.25 200
Integrated 8320 mg c/m2 8753.75 mg C/m2 630 mg C/m2

total
offshore station. The bacterial biomass in the upper 100 meters at this
deep station far exceeded the bacterial biomass measured in the upper
100 meters at most other stations. Note that the bacteria comprised a
large percentage of the total biomass measured and the bacterial bio-
mass was greater than the phytoplankton biomass. We believe that active
mineralization of the organics produced in slope and shelf waters was
occurring at this station, especially in the top 100 meters of the
water column.
On occasion in these studies the phytoplankton biomass and/or the bac-

terial biomass, exceeded the total biomass measured by the ATP tech-
149
niques (Table 5 and 6). We cannot explain this discrepancy with cer-
tainty. A factor of 50 was used to convert the chlorophyll a measure-
ments to phytoplankton biomass. This factor varies depending upon the
physiologic state of the phytoplankton, and on some occasions we have
overestimated the phytoplankton biomass. We know that in local Woods
Hole waters the C/chlorophyll a ratio is around 30 during winter months
when light intensity is low, but in summer months this ratio may rise
to nearly 100. Thus one can easily be off by a factor of three when
estimating phytoplankton biomass from chlorophyll a concentrations.
Similarly our ATP measurements on occasion may have been in error. The
ATP technique is a difficult technique and subject to many sources of
error.
Because of the small cell volume of marine bacteria, called minibac-

teria, growing in situ, the ATP/cell in these bacteria would be far
less than that found in cultured cells. During the early stationary
phase of growth, E. coli cells in our laboratory were found on the
average to contain 0.85 fg of ATP/cell. Since the volume of a minibac-
terium is 1/25th that of an E. coli cell, each minibacterium should
contain 0.034 fg of ATP/cell. In a recent cruise (Watson et al., un-
published dat~) we found that each minibacterium growing in situ below
the euphotic zone contained 0.05 fg of ATP/cell. The LPS/ATP ratio of
viable minibacteria growing in situ thus appears to be around 50.
An estimate of what percent of the minibacteria are viable can be ob-

tained from the LPS/ATP ratio. At station 176 (Table 6) the average
LPS/~TP ratio below the euphotic zone was 101. If all of the ATP mea-
sured were derived from bacteria this ratio would indicate that about
50% of the bacteria were in a viable state. However on our most recent
cruise the average LPS/ATP ratio below the euphotic zone was 50, sug-
gesting that most of the bacteria were alive.
Table 7. Comparison of total water and sediment biomasses in shallow and deep sta-
tions (mg C/m2)
Area Sta.No. Depth Total % in H2O Bacterial % in H2O

(m) biomass biomass
Shelf 161 30 40928 8.0 1251 31.9

171 35 19288 38.9 5729 17.1
162 39 11558 29.9 2783 13.3
Slope 173 469 4507 91.6 12573 72.9

169 800 13298 46.7 11071 52.3
Offshore 176 1800 9145 95.7 10159 81.9
In Table 7 we have compared the percentages of the total microbial bio-

mass and bacterial biomass that existed in the water column to those
found in the sediments. This Table indicates that a large percentage
of the total biomass as well as the bacterial biomass existed in the
sediments of shallow stations but that this percentage decreased as
the water depth increased. This suggests that most energy captured by
photosynthetic processes is funneled directly into the sediments and
is unavailable to the higher trophic levels within the water column.
This finding would agree with Riley's (1956) findings on Long Island
Sound and our own findings in local Woods Hole waters.
150
Table 8. Biomass of various components of the ecosystem
Component Grams Estimated by/from
Zooplankton 7.0 Dr. Morris Blackburn

Phytoplankton 4.3 Chlorophyll a data
Bacteria 12.6 LPS technique
Fish 0.01 Dr. Morris Blackburn
Infaunal benthos 1.6 Dr. Gilbert Rowe
(Percentage of total biomass comprised of bacteria = 49)
If bacteria are important in the carbon cycle, they should comprise a

significant percentage of the total biomass. In Table 8 we have esti-
mated the biomass of the various components of the ecosystem that exis-
ted at station 173, which was 469 meters deep. As shown in this Table,
bacteria comprised 49% of the total biomass.
We used 14C-Iabeled compounds in these studies, not to estimate the

heterotrophic potential, but rather to provide some information re-
garding relative metabolic activites in different areas. Table 9 merely
shows that the total bacterial metabolic activity increases with the
depth of the water column.
Table 9. Comparison of total microbial activity in the water column from shallow and
deep stations
~g C oxidized/h/m2 mg c/m2
Station Depth Acetate Glucose Glycine Urea Bacterial Total Phyto-
No. (m) biomass bio- plankton
mass biomass
161 30 4.5 1.8 134.8 45.0 399 3277 1377

166 35 182.0 217.9 727.8 111.9 592 6628 4016
171 35 85.9 179.3 377.0 26.0 982 7510 6119
162 39 1.8 67.2 441.0 147.3 370 3457 357:
165 57 432.9 516.1 549.7 1004.2 602 3665 38910
163 63 252.0 367.2 1201.0 728.9 596 11497 lCb'::l
164 86 273.0 760.0 654.5 645.8 604 7734 4(,"1_,
172 230 551.6 11.2 350.3 283.9
174 365 7885.6 3213.1 2041.4 105.5 3311 5584 2358
177 366 1646.5 4619.3 1092.6 682.9 3569 13114 1308
173 469 2448.7 3169.9 2483.8 434.3 9160 4127 4265
176 1800 8249.5 1199.4 6692.0 7792.6 8320 8754 630
The rates at which acetate, glucose, glycine, and urea were oxidized
in the upper and lower portions of 25-em sediment cores are shown in
Table 10. In the slope stations 50-100% of the oxidative activity mea-
sured occurred in the upper 10 cm of the cores. In shallow stations
30-70% of the microbial activity was localized in the top 10 em of the
sediment cores.
In Table 11 we have compared the relative rates of oxidation of orga-

nics in the water column to those in the sediments. These data again
suggest that a large percentage of the microbial activity as well as
a large percentage of the total microbial biomass existed in the sedi-
ments of shelf waters. In slope sediments the microbial activity and
the bacterial biomass were-far less than in shelf sediments. In off-
151
Table 10. Comparison of microbial activtiy in sediments from shallow and deep sta-
tions
mg C oxidized/h/m 2 mg C/m2
Sta.No. Depth Acetate Glucose Glycine Urea Total Bacterial
(m) biomass biomass
Total 171 35 6.946 3.020 10.250 0.110 11778 4747

activity 162 39 1.167 1.452 0.490 1.244 8102 2413
in 0-25 cm P 480 1.010 0.232 0.476 0.032 1023 1559
173 469 0.832 2.031 2.036 0.007 380 3443
169 800 0.051 0.262 0.428 0.001 7088 5280
176 1800 0 0.022 0.504 0.001 391 1839
Activity 171 35 2.706 1.313 5.381 0.053 11778 2346

in top 162 39 0.829 0.806 0.269 0.799 4778 1280
10 cm P 480 0.577 0.221 0.466 0.027 885 960
173 469 0.620 1.721 1.837 0.001 380 1813
169 800 0.051 0.138 0.376 0.001 3889 1813
176 1800 0 0.022 0.476 0.001 391 1039
% Activity 171 35 38.9 43.5 52.5 48.2 70 49.4

in top 162 39 71.0 55.5 54.9 64.2 59 53.0
10 cm P 480 57.1 95.3 97.9 84.4 87 61.6
173 469 68.8 71.6 87.7 100.0 100 53.1
169 800 100 52.7 87.9 100.0 55 34.4
176 1800 0 100.0 94.6 100.0 100 56.5
Table 11. Comparison of total water and sediment microbial activity and biomass as
a function of depth of the water column
llg C oxidaized/h/m 2 g C/m2

Station No. Acetate Glucose Glycine Urea Total Bacterial
and depths biomass biomass
171 H2O 85.9 179.3 377 25.96 7.51 0.98

(35 m) Sed. 6946 3020 10250 110 11.78 4.75
% in
Sed. 98.7 94.4 96.4 80.9 61.1 82.9
173 H2O 2449 3170 2484 434 4.13 9.16

(469 m)
Sed. 832 2031 2036 6.7 0.38 3.41
% in
Sed. 25.4 39.1 45.0 1.5 8.4 27 .1
shore water, where the water column is 5000 meters, we have found on
other cruises that less than 1% of the bacterial activity and bacterial
biomass under 1 m2 of sea surface would be localized in the sediments.
The average bacterial biomass in the water column and sediments at

shelf and slope stations shown in Table 7 was 6.68. If this standing
crop contains 10% of the yearly carbon incorporated into bacterial
cells, and if an equal amount of carbon is used for cellular mainte-
nance, bacteria would use 133.6 g of the 300 g C/m2 that phytoplankton
fix each year in this upwelling area. Certainly this percentage seems
high, but it does suggest that bacteria play an important role and prob-
152
ably utilize nearly 50% of the energy captured by photosynthetic pro-

cesses in an upwelling area.
Initially we estimated, based on several assumptions, that there should

be 0.75-1.0 x 10 5 bacteria/ml in the water column in an upwelling area,
provided that the organics were not transported horizontally to another
area prior to mineralization. In recent studies at a 180-meter station
in an upwelling area off the southwest coast of Africa we found, em-
ploying the epifluorescence direct counting technique, 8 x 10 5 -5 x 10 6
bacterial/ml.
Our evidence indicates, however, that a considerable amount of organic

matter within the water column is transported horizontally seaward.
Because of this horizontal transport the concentration of bacteria
measured within the water column in an upwelling should be less and
not more than we originally estimated, provided that all our original
assumptions were correct. However, our original estimate was based on
several assumptions that have not been experimentally verified and
need reexamination. One such assumption is that the average marine bac-
terium divides every 3.65 days, and we have no evidence to indicate
that such is the case. Our estimate was also based on the assumption
that 90% of the mineralization takes place within the sediments in the
immediate area of an upwelling system. The preliminary evidence pre-
sented here provides cursory evidence to support this assumption, but
much additional work needs to be done before any such broad generali-
zation can be considered true.
No information is available upon the fate of the bacterial biomass. If

our evidence is correct that the annual production of bacterial carbon
is as great as we have indicated, then i t follows that an equal amount
of bacteria~ carbon must similarly be removed from the environment or
else this particulate source of carbon would accumulate. We expect that
bacteria are removed by predation. If this is so, bacteria must serve
as a major food source in the sediments and to a lesser degree within
the water column. However, exactly which groups of organisms serve as
the primary predators of bacteria have not been determined.
In summary this preliminary report suggests that bacteria play an im

portant role in an upwelling area where they are major energy consumers.
Most organics appear to be mineralized within the sediments near the
site where they are produced, while the remaining fraction of organics
are mineralized within the water column near the site of upwelling or
are transported horizontally some distance seaward where they are min-
eralized. Our evidence also suggests that bacteria may serve as a major
food source for larger organisms.
Aeknowledgments. This work was supported in part by the United States Energy Research
and Development Administration E(ll-l) 3565 (Reference No. COO-3565-06), National
Science Foundation grant 20/21270, and a research grant from the Arthur Vining Davis
Foundation. Contribution number 3717 from the Woods Hole Oceanographic Institution.
References
Allen, H.L.: Acetate utilization by heterotrophic bacteria in a pond. Hildrol. Kozl.

47, 295-297 (1967)
Andrews, P., Williams, P.J. LeB.: Heterotrophic utilization of dissolved organic com-
pounds in the sea. J. Marine Biol. Assoc. U.K. 21, 111-125 (1971)
Christian, R.R., Bancroft, K., Wiebe, W.J.: Distribution of microbial adenosine tri-
phosphate in salt marsh sediments at Sapelo Island, Georgia. Soil Sci. 119, 89-97
(1975)
153
Cushing, D.H.: On the nature of production in the sea. Fish. Invest. London, Ser. 2,
Vol. 22, no. 6 (1959)
Dale, N.G.: Bacteria in intertidal sediments: Factors related to their distribution.
Limnol. Oceanog. ~, 509-518 (1974)
Deufel, J.: Zahlung von Wasserbakterien auf Membranfiltern mittels Fluoreszenzmikro-
skopie. Naturwissenschaften ~, 654 (1959)
Duursma, E.K.: Dissolved organic carbon, nitrogen, and phosphorous in the sea. Neth.
J. Sea Res. !, 1-148 (1961)
Fogg, G.E.: The extracellular products of algae. Oceanog. Marine BioI. Annu. Rev.
i, 195-212 (1966)
Gunderson, K., Mountain, C.W., Taylor, D., Ohye, R., Shen, J.: Some chemical and
microbiological observations in the Pacific Ocean off the Hamaiian Islands. Limnol.
Oceanog. 12, 524-531 (1972)
Hamilton, R.D., Holm-Hansen, 0.: Adenosine triphosphate content of marine bacteria.
Limnol. Oceanog. 12, 319-324 (1967)
Hellebust, J.A.: Excretion of some organic compounds by marine phytoplankton. Limnol.
Oceanog. lQ., 192-206 (1965)
Hobbie, J.E., Crawford, C.C., Webb, K.L.: Amino acid flux in an estuary. Science
159, 1463-1464 (1968)
Hobbie, J.E., Holm-Hansen, 0., Packard, T.T., Pomeroy, L.R., Sheldon, R.W., Thomas,
J.P., Wiebe, W.J.: A study of the distribution and activity of microorganisms in
ocean water. Limnol. Oceanog. 17, 544-555 (1972)
Holm-Hansen, 0., Booth, C .R.: The measurement of adenosine triphosphate in the ocean
and its ecological significance. Limnol. Oceanog. 11, 510-519 (1966)
Jannasch, H.W.: Studies of planktonic bacteria by means of a direct membrane filter
method. J. Gen. Microbiol. ~, 609-620 (1958)
Jannasch, H.W., Jones, G.E.: Bacterial populations in sea water as determined by
different methods of enumeration. Limnol. Oceanog. 4, 128-139 (1959)
Karl, D.M., La Rock, P.A.: Adenosine triphosphate measurements in soil and marine
sediments. J. Fish. Res. Bd. Can. 32, 599-607 (1975)
Kriss, A.E.: Marine Microbiology (Deep Sea). Shewan, J.M., Kabata, Z. (Transl.).
London: Oliver and Boyd Lt., 1963
Levin, J., Bang, F.B.: A description of cellular coagulation in the Limulus. Bull.
Johns Hopkins Hosp. ~, 337 (1964)
Madsen, B.L.: Detritus on stones in small streams. In: Mem. 1st. Ital. Idrobiol.
Melchiorri-Santolini, U., Hopton, J.W. (eds.). 1972, 29. Suppl., pp. 385-403
Menzel, D.W., Ryther, J.H.: Zooplankton in the Sargasso Sea off Bermuda and its re-
lation to organic production. J. Cons. Perm. Int. Explor. Mer. 26, 250-258 (1961)
Parsons, T.R., Strickland, J.D.H.: On the production of particulate organic carbon
by heterotrophic processes in sea water. Deep-Sea Res. ~, 211-222 (1961)
Parsons, T.R., Seiki, H.: Importance and general implications of organic matter in
aquatic environments. In: Organic Matter in Natural Waters. Hood, D.W. (ed.).
U. Alaska Occ. Publ. No.1, 1970, pp.1-27
Riley, G.A.: Oceanography of Long Island Sound, 1952-1954. II. Physical oceanography.
Bull. Bing. Oceanogr. ColI. 15, 15-46 (1956)
Ryther, J.H.: In: The Sea. Hill, M.N. (ed.). London: Interscience, 1963, pp. 347-380
(1969)
Schwantes, H.O.: Mikroskopischer Nachweis von Mikroorganismen auf Membranfiltern
nach Filtraten aus Flussig~eiten mit Hilfe der Akridinorgane-Fluoreszenz. Microsc.
Acta 70, 236-238 (1971)
Sieburth, J. McN.: Distribution and activity of oceanic bacteria. Deep-Sea Res. 18,
1111-1121 (1971)
Slobodkin, L.B.: Growth and regulation of animal populations. New York: Holt, Rine-
hart and Winston, 1961, Chap. 12
Steemann-Nielsen, E., Jensen, E.A.: Galathea Report, F. Bruun et al. (eds.). London:
Allen and Unwin, 1957, Vol. 1, p. 49
Vaccaro, R.F., Jannasch, H.W.: Studies on heterotrophic activity in seawater based
on glucose assimilation. Limnol. Oceanog. 11, 596-607 (1966)
Vaccaro, R.F., Jannasch, H.W.: Variations in uptake kinetics for glucose by natural
popUlations in seawater. LimnoL Oceanog. g, 540-542 (1967)
154
Wangersky, P.J.: The organic chemistry of sea water. Am. J. Sci. 53, 358-374 (1965)
Wetzel, R.G.: Dissolved organic compounds and their utilization in two marl lakes.
Hidrol. Kozi. 47, 298-303 (1967)
Wiebe, W.J., Pomeroy, L.R.: Microorganisms and their association with aggregates
and detritus in the sea: a microscopic study. In: Mem. Ist. Ital. Idrobiol.
Melchiurri-Santolini, U., Hopton, J.W. (eds.), 1972, 29. Suppl, pp. 325-352
Williams, P.J. LeB.: The validity of the application of simple kinetic analysis to
heterogenous microbial populations. Limnol. Oceanog. 18, 159-165 (1973)
Wright, R.T., Hobbie, J.E.: Use of glucose and acetate by bacteria and algae in
aquatic ecosystems. Ecology 47, 447-464 (1966)
Young, N.S., Levin, J., Prendergast, R.A.: An invertebrate coagUlation system ac-
tivatedby endotoxin: evidence for enzymatic mediation. J. Clin. Investig. 21,
1790-1797 (1972)
Zimmerman, R., Meyer-Reil, L.-A.: A new method for fluorescence staining of bacterial
populations on membrane filters. Kieler Meeres Forsch. 30, 24-27 (1974)
Zobell, C.E.: Marine Microbiology. Waltham, Mass: Chronica Botanica Co., 1946
Heterotrophic Activity in the Mauritanian Upwelling in
March 1973: Assimilation and Mineralization of Amino Acids
AHERBLAND
1. Introduction
The heterotrophic bacteria in natural waters are extremely difficult

to study because of difficulties encountered in biomass measurement
and in activity measurement.
Firstly, the bacteria are known to be the smallest living organisms in

seawater, and they cannot be fully separated by differential filtration
because the smallest phytoplankton cells and the larger bacteria are
equal in size,. Moreover, the bacteria are commonly associated with the
detrital particles, which are larger than bacteria, so they are re-
tained by membrane filters with greater porosity than the true size
of the bacteria. Also, it is not possible to measure directly the car-
bon or nitrogen of bacteria in a sample of seawater.
Secondly, the bacteria do not contain a specific compound, such as

chlorophyll in phytoplankton cells. Although the carbon-chlorophyll
relation is not clear, the chlorophyll measurement associated with
other parameters (e.g., specific determination) allows the comparison
of phytoplankton abundance in two oceanic regions. This is not possible
for bacteria.
Thirdly, in contrast to photosynthetic algae, which use only carbon

dioxide or bicarbonate, the heterotrophic bacteria may use many dif-
ferent sources of carbon that are not exactly known, extremely diver-
sified, and that vary with the ambiant populations of phytoplankton
and zooplankton. While the 14C02 added in primary production experi-
ments behaves like natural C02 (except for isotopic discrimination) ,
the simple labeled organic compounds added in the heterotrophic activ-
ity measurements do not reflect the overall organic matter present,
and the uptake -of labeled compounds does not preclude the behavior of
the organic matter. Hence, the rate of production of bacterial cellular
material and the rate of decomposition of organic matter under natural
conditions are almost impossible to measure.
However, the study of the uptake of single substrates gives valuable

information about the substrates that are being used and about the
activity of bacteria, which are certainly the main organisms responsible
for uptake of organic solutes at natural concentrations (Wright and
Hobbie, 1966), although it is not possible to distinguish this from
algal heterotrophic activity.
This paper deals with the uptake of amino acids in an upwelling area,
during a bloom of phytoplankton. Comparisons are made with glucose
uptake under the same conditions, a year before in the same area.
156
2. Methods
The satisfactory results obtained in March 1972 by following a drogue

in the Mauritanian upwelling between Cape Timiris and Nouakchott (Herb-
land et al., 1973) led us to undertake a similar study in 1973 in the
same area during the same season. The coldest water (15 0 C), the richest
in nutrients (20 ]Jgat 1 N0 3 ) , and the poorest in oxygen (60% saturated)
and chlorophyll (2 ]Jg/l) were determined by surface survey, with con-
tinuous measurements. The area was east of a little canyon south of
Cape Timiris where the depth of the bottom is 25-30 m. The drogue was
set at a depth of 3 m, and followed for six days. Three times each day
(6 h, 12 h, and 18 h) the ship was brought to the drogue, and a series
of observations was made. Each series has a separate station number.
The methods used for measuring oxygen, nitrate, ammonia, organic phos-
phorus, and chlorophyll have been given in detail elsewhere (Herbland
and Voituriez, 1974). The organic excretion of phytoplankton was mea-
sured after filtration of particulate matter, followed by the removal
of H14C0'3 by acidification (pH 2.5), bubbling (10 min), and direct
IBON IB o
r
.30
• '3 31
2
35 • • 36
.37
-3B
E
0
E
e
r
E 0
0 cv
0
J?
)
0
/
CV
/
1972 1973
Fig. 1. Position of stations as determined by the location of the drifting buoy in

1972 C\nd 1973
157
50
<I
CHL a
mg/m3
m
38 40
0
;;
Fig. 2. Chlorophyll a and oxygen evolution during the drift of the buoy in 1973
counting of the remaining labeled organic compounds by liquid scintil-

lation (Anderson and Zeutschel, 1970). The heterotrophic utilization
(assimilation and respiration) was measured by 3-H incubation experi-
ments, in the dark, with a 14C amino acid mixture. The mixture contained
glycine (11%), serine (29%), lysine (27%), aspartic (18%) and glutamic
(15%) acids, at a total concentration of 15 ~g ell. After incubation,
the filtration was made with low suction pressure (200 mmHg) on Milli-
pore membrane filters. The C02 respired was trapped directly in a scin-
tillation vial, suspended at the top of the conic incuqation flasks,
with one ml of hyamine hydroxide dropped on a glass fiber filter. The
radioactivity was counted with a liquid scintillation spectrometer.
For details, see Herbland and Bois (1974).
3. Results and Discussions
The tracks of the drogues were very similar from one year to the next
(Fig. 1). They followed a co~se parallel to the shore, but in 1973,
the drogue veered offshore on the last day.
158
38 40 42 44 46 48 50 52 St.
0'-~--~~~7-~~L-~~--~~~~~--~~
50
Z 100~
40 42 44 46 48 50 5 St.
0 38
.O~
V
:::\
50/
50
BAC.
30/
jJgC/m 3 I h
Fig. 3. Nitrate and heterotrophic activity evolution during the drift of the buoy
in 1973
3.1 Heterotrophic Activity Evolution Versus Other Biologic Parameters
Where upwelling takes place (cold water), the chlorophyll a values are
relatively low (2 ~g/l). They increases regularly and reach 10 ~g/l
the third day and are higher than 30 ~g/l the fourth and fifth days
(Fig. 2). The oxygen concentrations follow parallel pattern. Initially
the water is undersatured (60%). Saturation is reached the third day.
The biologic production of 02 is not sufficient to explain the in-
crease of oxygen in the water both years. There is evidence of atmo-
spheric enrichment (Herbland and Voituriez, 1974). During the same
period, nitrate decreased from 18 to 5 ~g at/l (Fig. 3). The hetero-
trophic activity is very low during the first days. It increases sharp-
lyon the third day and remains very high during the bloom of phyto-
plankton (Fig. 3).
159
CHLo 02
g/m2 mill
o
120
7 110
6 100
5 90
4 80
3 70
2 60
50
39 40 42 43 45 46 48 49 51 52
2 3 4 5 days
Fig. 4. Evolution of integrated values (0-20 m) during the drift of the buoy in
1973
The evolution of integrated values (Fig. 4) shows clearly the simul-

taneous increase of chlorophyll and oxygen, and later, the increase
of heterotrophic activity. The amino acid uptake bloom seems to occur
immediately after the phytoplankton bloom. Ammonia increases subsequent
to heterotrophic activity (Fig. 5). Then we have the successive evolu-
tions in the water mass: (1) oxygen increase with simultaneous phyto-
plankton increase and nutrient decrease, (2) heterotrophic activity
increase, and (3) ammonia increase.
Phytoplankton cells are certainly not yet decomposed because nutrients

are available. Some authors have recently found tha-t excretion products
of phytoplankton can be taken up by marine bacteria in culture (Wright
and Shah, 1975) or in natural populations (Tanaka et al., 1974; Herb-
land, 1975).
160
(]
WPz
Zoopk.
I
/"'\\ \
I \
ml/m 2 mg at! m 2 I \
~ " \\ +
50 \ 5
\
20 \
\
40 \ 4
\
\
o
30
20 o 2
10
St. 39 40 42 43 45 46 48 49 51 52
I 2 3 4 5 ooys
Fig. 5. Evolution of integrated values (0-20 m) during the drift of the buoy in 1973
In this drogue study, the excretion of phytoplankton was measured.

There is a simultaneous increase of heterotrophic activity and excre-
tion of phytoplankton (Fig. 6). It does not prove that there is a tro-
phic link but it is a strong indication. The vertical distribution of
heterotrophic activity shows that, at the beginning, in recently up-
welled water, it is homogeneous (stations 39 and 42 in Fig. 7). At the
end, however, during the bloom of phytoplankton, heterotrophic activ-
ity is located essentially in the upper fifteen meters, where the phy-
toplankton and organic matter (measured by organic phosphorus) are
abundant (stations 51 and 52). The same result was obtained in 1972
(Fig. 8) with glucose uptake.
3.2 Growth Yield or Growth Efficiency
A certain amount of amino acids was taken up, and a percentage of this
amount was oxidized to carbon dioxide. The growth yield is the ratio
of amino acid assimilated/assimilated + oxidized. This ratio has sharp
variations: An average value of 78% was found by Williams (1970) for
an amino acid mixture. Hobbie and Crawford (1969) reported results in
natural fresh-water populations taking up sugars, amino acids, and
acetate; most of their values lay in the range 70-80%, the overall
range being 43-92%. Despite these variations, the two amino acids leu-
cine and lysine showed the highest growth yields while aspartic and
glutamic acids had the lowest (Crawford et al., 1974; Gocke, 1976).
1 61
PHYTO
mgl4C/m 2/h EXC.
7 BAC. ACT gC/m 2
/J
6 3
5
EXCRETION PHYTO·
o
4
2
3 BAC
39 40 42 43 45 46 48 49 51 52
Fig. 6. Comparison between the phytoplankton organic excretion and heterotrophic

activity (integrated values), during the drift of the buoy in 1973
We measured simultaneously assimilation and respiration during incu-

bation. There is a direct relation between the carbon assimilated and
the carbon respired (Fig. 9). For all samples, the mean value of growth
efficiency is 50%. In 1972, the mean value with glucose as substrate
was 35%. These results are low compared with those in the literature.
This ratio was not constant over the track of the drogue (Fig. 10).
It was low, about 40%, at the beginning, and reached a maximum of 60%
the fourth day when the bacteria and phytoplankton are actively grow-
ing. It can be seen that the yield values below 18 meters are lower
than the values between 0 and 10 meters. This result cannot be fully
explained, but it is possible that oxygen concentration (oxygen is
the last electron acceptor in respiration processes) can affect the
yield: The oxygen concentrations are low near the bottom and at the
beginning of the experiment. However, many parameters other than oxygen
can influence the metabolism of bacteria.
3.3 Geographic and Nychthemeral Variations
In 1972, after the drogue study, a transect of 120 miles was done from
the coast to the open sea to try to measure the extent of the upwel-
ling area. Except for station 50, where chlorophyll and heterotrophic
162
SAC 100 200 ,IlgC/m!1 h SAC 100 200 jJgC/mS/h
02 2 3 4 5 mVI 02 2 3 4 5 mill
0
I
< J*
t:. X
/
A~ BAC )02 I
10 10
t:.
BA£
1 02
\ j
1/
20
/ j
20
Sf. 39 St. 42
30
t:. J 30
t:. -1
,g
SAC 100. 200 300 pgC/m!/h SAC 100 200 300 pgC/m!/h
Porg.O.2 0.6 1.0 at/I Porg 0.2 Q6 I. 0 }Jg at/I
02 I 2 3 4 5 6 7 mill 02 I 2 3 4 5 6 7 mill
Fig. 7. Vertical distribution of organic phosphorus (Porg), oxygen (02)' and hetero-
trophic activity (Bact) at two stations in recently upwelled water (Sts. 39 and
42), and at two stations after the phytoplankton bloom (Sts. 51 and 52) in 1973
163
\
2 2
.
02 mill I 3 4 mill I 3 4 PoIg Q2 0 .4 0 .6
14
7
TOC 14 15 16 17 2 4 6 pg C/nr/h "C
I4t-~l'
I x)1
!
.~~
10
J
\
l:J. I! TO
/ 02
I
BAC} Porg
15
TO J I 02
I
l:J. BAC
I \l:J. 0"
x
l< 0
20 I \l:J.
25
m
St· 24 St. 25
T"C 15 17 19 21 °C
BAC 4 12 20 28 36 f914C Im 3 /h
Por9. 0.2 0 .6 1. 0 L4 1.8 }l9ot II
02 23456 7 8 9 10 mill
1 0 ./ . ; ;_ _ _ =:::x~02
l:J.
ro / Porg . /
1 BAC.
i r ;;
20
~J
<oW
St. 36
30 /
1972
Fig. 8. As in Figure 7, but in 1972. Stations 24 and 25 are in recently upwelled
water, and 36 is in old upwelled water
164
50 o o
14C02 Respired
o
pgClm 3 /h
0
0
0
40 0
0 0
0 0 0
0 0
0
0
0
0
0 0
0
0
0
30 0
0 00 0
0 0
0
0
o
cf>
o 0
0
0
20
0
0 0
0 00
0
0 0
0
0
0
0
0
o 0 14C
0
Assimilated
0
10
10 20 30 40 pgClm 3 /h
Fig. 9. Relation between carbon of amino acid "assimilated" by the cells, and carbon
"respired" during the 3-h experiments
activities were high (probably a divergence area), there was a general

decrease from the shore to the open sea (Fig. 11).
Daily variations were seen. Incubations made during the morning (M) -
gave higher values than incubations made during the afternoon (AN).
It is possible that during the night, heterotrophic activity is more
important, hence, the bacteria are more numerous in the morning sample
than in the afternoon one. It could be the result of a competition
between autotrophic and heterotrophic organisms, the "autotrophs" be-
ing active during the day and perhaps inhibiting the "heterotrophs."
165
"10
R = ASSIMILATION I ASSIM.+ RESPIRATION
60
<18m
0- 10m
yr -L-_--~+
50
+ /)
(~';~/ h " " , + ______ _
,,
'+
~----:I/ /
/
/
+, //
, /
40~~--'-r-/------~~--~--------L---~--------~---L--------~--~~
39 40 42 43 45 46 48 49 51 52 St.
Fig. 10. Change of growth yield in two layers of water, during the drift of the buoy
in 1973
35
l pgc/m2/h
M
o
20
M
M
V
0 0
15
M
0
i/
M
10 0 0
AN
0
5 AN
AN
;N \ AN
42 43 48 49 50 51 52 53 54 55 56 St.
Fig. 11. Geographic and nychthemeral variations of bacterial activity: St. 42 is near
the shore, and 56 is in the open sea. The distance between Sts. 42 and 56 is approx-
imatively 120 miles. ~ morning stations; AN afternoon stations
166
4. Conclusions
Amino acids are well taken up by heterotrophic organisms (probably

bacteria) in this upwelling area.
The same results were found in 1972 with glucose and in 1973 with an
amino acid mixture:
1. Same time succession: upwelling water rich in nutrients and poor

in oxygen and chlorophyll, phytoplankton bloom, heterotrophic bloom,
and ammonia enrichment.
2. Growth yields are somewhat higher with amino acids than with glucose
(50% against 35%). This average yield seems to be affected by the en-
vironmental conditions (oxygen concentration and an undetermined fac-
tor).
3. Nychthemeral variations have been observed, but not satisfactorily

explained.
References
Anderson, G.C., Zeutschel, R.P.: Release of dissolved organic matter by marine phyto-
plankton in coastal and offshore areas of the northeast Pacific Ocean. Limnol.
Oceanog. 15, 402-407 (1970)
Crawford, C.C., Hobbie, J.E., Webb, K.L.: utilization of dissolved free amino acids
by estuarine microorganisms. Ecology 55, 551-563 (1974)
Gocke, K.: Respiration von gel6sten organischen Verbindungen durch naturliche Mikro-
organismen. Populationen. Ein Vergleich zwischen verschiedenen Biotopen. Marine
BioI. 35, 375-383 (1976)
Herbland, A.: Utilisation par la flore heterotrophe de la matiere organique natu-
relle dans l'eau de mer. J. Exp. Marine BioI. Ecol. 19, 19-32 (1975)
Herbland, A., Bois, J.F.: Assimilation et mineralisation de la matiere organique
dissoute dans la mer. Methode par scintillation liquide. Marine BioI. 24, 203-212
(1974)
Herbland, A., Voituriez, B.: La production primaire dans l'upwelling mauritanien
en mars 1973. Cah. ORSTOM, ser. Oceanog. 11, 187-201 (1974)
Herbland, A., Le Borgne, R., Voituriez, B.: Production primaire, secondaire et re-
generation des sels nutritifs dans I 'upwelling de Maruitanie. Doc. Scient. Centre
Rech. Oceanog. Abidjan 4, 1-75 (1973)
Hobbie, J.E., Crawford, C~C.: Respiration corrections for bacterial uptake of dis-
solved organic compounds in natural maters. Limnol. Oceanog. 14, 528-533 (1969)
Tanaka, N., Nakanishi, M., Katoda, H.: Nutritional interrelation-between bacteria
and phytoplankton in a pelagic ecosystem. In: Effect of the Ocean Environment on
Microbial Activities. Colwell and Morita (eds). Baltimore, Univ. Park Press, 1974,
pp. 495-509
Williams, P.J.B.: Heterotrophic utilization of dissolved organic compounds in the
sea. I - size distribution of population and relationship between respiration and
incorporation of growth substrates. J. Marine BioI. Ass. UK 50, 859-870 (1970)
Wright, R.T., Hobbie, J.E.: Use of glucose and acetate by bacteria and algue in
aquatic ecosystems. Ecology 1], 447-464 (1966)
Wright, R.T., Shah, N.H.: The trophic role of glucolic acid in coastal sea water.
I - Heterotrophic metabolism in sea water and bacterial cultures. Marine BioI. 33,
175-183 (1975)
Physical Aspects and Biological Consequences of Ghanaian
Coastli lJpvvelling
R W.HOUGHTON and M.AMENSAH
Results of recent research on the physical and biological aspects of the

Ghanaian coastal upwelling are presented. This upwelling is unusual in
that its occurrence and intensity do not correlate with variations in
the local wind. During the upwelling the strong stratification over
the shelf breaks down as cold saline water comes to the surface, in-
i tiating a period of high biological producti vi ty. Surface current re-
cords and hydrographic sections taken in 1974 both suggest an offshore
displacement in the surface layer, but this and the change in the sea
level do not correlate with the surface wind. The upwelling does not
appear to be driven by changes in the circulation on the shelf. The
observation of low frequency waves over the shelf and on the equator
in the Gulf of Guinea suggest that they may provide a nonlocal driving
mechanism for this coastal upwelling.
Biological production during the three-month major coastal upwelling

has been compared with that du~ing the nine-month period of hydrogra-
phic stability. Plankton samples were obtained by the Hardy Plankton
Indicator and the ICITA net for the periods 1969-1970 and 1972-1974.
Production of both plankton and the fish SaI'dinella aurita is several

times higher in the upwelling season than in the stable period. Also,
during the upwelling many more species of phytoplankton cells but fewer
species of zooplank~on, especially copepods, occur. The copepod popula-
tion is dominated by Calanoides caI'inatus (Kr¢yer).
1. Introduction
The coastal upwelling in the Gulf of Guinea between Nigeria and the
Ivory Coast, a seasonal event occurring between July and September,
is of great importance to the local fishing industry. During this pe-
riod, cold, saline water is found over the continental shelf, as the
thermocline rises to the surface. This cold water is local in origin
and not advected into the area from Southwest Africa by the Benguela
current. In the Gulf of Guinea there are two prominent capes: Cape Pal-
mas in western Ivory Coast, and Cape Three Points in western Ghana,
both of which demarcate regions of distinct hydrography. Since the
colder waters are found predominantly east of each of these capes
where the coastline is roughly parallel to the prevailing southwest
wind, it has been commonly assumed that the upwelling there is the
result of wind-driven divergence along the coastline. A careful exami-
nation of the data does not support such a simple interpretation. In
this paper we shall discuss briefly the physical and biological aspects
of this coastal upwelling based primarily on recent data collected
off the coast of eastern Ghana.
168
1.1 Data
The data used to study the coastal regime are derived from a number of
sources. The Ghana Meteorological Service Department maintains stations
along the coast, marked in Figure 1, where surface conditions are re-
2°W ,OW
6°
Ghana
.
-t
Aanderaa
Fishery
Mooring
stations
*
5° 5°
Gulf of Guinea
I I I
0 40 eOkm
1000 rn
2°W 'oW 0° l°E
Fig. 1. Bathymetry along the coast of Ghana showing the location of coastal stations
maintained by the Meteorological Services Department and the Fishery Research Unit,
the hydrographic transect south of Tema, and the Aanderaa mooring
corded hourly. The Survey Department maintains tide gauge stations at

Takoradi and Tema harbors where the sea level is recorded continuously.
The Fishery Research Unit maintains stations along the coast where the
sea temperatures and salinity are measured daily from surface bucket
samples. Weekly transects are taken perpendicular to the coast south
of Tema (see Fig. 1) out to station C, 30 km offshore (extended to
station E during 1974), where temperature and salinity profiles are
measured to 200 m depth from BT and Nansen bottle casts. At stations
Al and B the current profile is measured weekly with an Ekman meter.
During 1974 these data were augmented by a mooring on the shelf edge
of three Aanderaa meters recording continuously for 78 days the tem-
perature and current during the onset of the upwelling.
In 1970, the Hardy Plankton Indicator, described by Glover (1953), with

a mesh size of 60 ~, was used to collect phytoplankton at all four
stations on the transect. The Indicator was towed at a depth of 7-8 m
for 5 min at a ship speed of approximately 12 km/h. In the laboratory,
the cells were identified to genus level and counted along two diameters
of the coverslip on the disc at right angles to each other, using a
binocular microscope.
Zooplankton samples were collected at all four stations for the period
1972-1974 using the ICITA net with a mesh size of 300 ~, fitted with
a T.S.K. flow meter. It was.towed step-obliquely for 15 min. The depth
of tow ranged between 13 and 25 m to the surface. In the laboratory,
displacement volumes of the samples were measured.
169
Also, fish catches were regularly recorded at most landing stations

along the coast.
2. General Hydrography
2.1 Seasonal Variation
The beach temperature is a good indicator of the hydrographic condi-

tions over the shelf. The variability of the Tema Harbor temperature
records for the three different years is evident in Figure 2. Since
u
o
III
:;
'lii
I;;
a.
E
~
Fig. 2. Water surface temperature recorded at Tema harbor for various years
during the summer months the mid-ocean surface temperature drops to

about 2S o C, beach temperatures below 24 0 are taken to indicate a coastal
upwelling condition. In 1967 the upwelling was unusually long and in-
tense, while during 1968 i t was virtually absent.
The coastal hydrography of the Gulf of Guinea is generally divided into

four regimes (Longhurst, 1962; Morliere and Rebert, 1972): the short
cold season, December-January; the long warm season, February-June;
the main upweiling season, July-September; and the short warm season,
October-November. During the two warm seasons, the increased insolation
and coastal rains produce a very stable stratification with a well-
mixed surface layer 30-40 m deep over a very sharp pycnocline. This
pycnocline is enhanced by the coincidence of a salinity maximum just
below the thermocline. During the main cold season this stratification
vanishes as the thermocline rises to the surface, and the entire water
column over the shelf becomes homogeneously cold and saline and is
much less stable. The biological productivity is enhanced during this
period. Although the thermocline rises briefly during the short cold
170
season and sporadically during February to April, there is no break-

down in the hydroqraphic stability and no significant increase in bio-
logical activity during these periods. Therefore, in this general review
we will confine our attention to the main upwelling period.
2.2 Upwelling Correlation with Surface Wind

Given the orientation of the coastline with respect to the prevailing
southwesterly wind, it is natural to expect the local wind to be the pri-
mary driving agent of the upwelling. In Figure 3 we show monthly mean
2 ....." ___ •__ •__ .-_............... ". ______ •

--.---""'*
.§_ ,
.~
Z4°E' .-.-.-......... / ._e-,. ..... ... j (b)
/
c 200 . -•
•_§
u
nOtO'0_0_0".
_/-0 0---- (c.) j
f 180 _0
i5
6
Fig. 3a-c. (a) is-year average speed

from meteorological stations at Ada
(solid line) and Saltpond (dashed
line); (b) vector average speed and
direction from N.O.A.A. Marine Sur-
face Data for 2°C square centered at
4.60 N, O.6 0 W; (c) same as (b) but
oJ for 2° square centered at 2.SoN,
O.70 W
Months
winds on the coast and further offshore. Along the coast the winds are
weak with little seasonal variation in strength and only deviate from
the prevailing southwest direction in December and January. Further-
more, offshore winds are stronger with a pronounced peak during June
and July but with less than a 300 variation in the mean monthly direc-
tion over the entire year.
A comparison of the yearly variation in the upwelling intensity and
the mean coastal wind strength for the years 1963-1974 is shown in
Figure 4. The mean wind is based on measurements at three coastal sta-
tions for the months of July through September. For the upwelling in-
tensity the area between the beach temperatures curve (see Fig. 2) and
a line at 24 0 C for the months of June through September is measured.
There is certainly no positive correlation between the coastal wind
171
Fig. 4. Upwelling intensity (definition explained

!! 4 in text) as a function of average coastal wind
·c::I
.ci
• speed for the years 1963 to 1974
~
3
~
·iii
c: ..
.!
.s 2 • •
It
•
.. • •
01
c:
. •
~
a.
:::l
•
2
.3
Ave. coastal wind m/s.
strength and upwelling intensity. In fact, 1968, the year with the
weakest upwelling, was the year with the strongest coastal wind.
The data for the winds further offshore are too sketchy to compare
reliably the year-to-year variation. However, for the years 1967 and
1968 there does not appear to be a significant variation that is large
enough to account for the dramatic difference in the upwelling inten-
sity.
The surface ocean current is predominantly eastward, presumably driven

by the prevailing southwest wind. Below the thermocline is a westward
countercurrent whose origin is still not clear but which is possibly
a returning branch of the equatorial undercurrent (Morliere et al.,
1974). No systematic time delay in the onset of the upwelling has been
observed along the coast to suggest that the cold water is advected
by either of these currents.
2.3 Upwelling 1974
To illustrate the changes in the coastal hydrography we will examine

the data for 1974 (see Fig. 5), a year for which the record was most
complete. During May and June the thermocline and halocline are both
at 40 m depth and rise slowly. The onset of the upwelling, 10-15 July,
is the result of an abrupt rise in the thermocline, producing a sudden
drop in the beach temperature. The thermocline remains near the sur-
face and the water over tne shelf is homogeneous until October when
the stratification is reestablished.
Although the alongshore wind increases gradually during May and June,
it does not correlate with the sudden change in hydrography. The one
sharp increase in wind around the 5th of July precedes the upwelling
by a week, and the winds do not diminish in October when the upwellin~
ceases. Throughout this period there is always an onshore component,
but it never exceeds 0.5 m/s.
The adjusted sea level drops during the upwelling period. However, the
change is nearly equal to the magnitude of the change in the dynamic
height anomaly, suggesting that isostatic adjustment rather than wind-
driven divergence is the primary cause.
The hydrographic structure over the shelf during the onset of the
upwelling is shown in Figure 6. The isopycnals that are horizontal in
172
Fig. 5. Hydrographic data during 1974

upwelling season. From top to bottom:
alongshore wind at Tema harbor; Tema
sea level, adjusted for changes in air
pressure, superimpos ed on the dynamic
height anomaly calc u lated to 50 m depth
over the shelf; Tema harbor temperature,
days on which transect data were gath-
E
u e red; the depth of is othe rms, isohalines,
and isopycna ls derived from these trans-
ect data. Dotted line: depth of the
salini ty maximum
u
o
10
a..
~
:: ~::-t:f::;J(;:~
IOO~---'------------~I-so~~~--=---~
26
100'~:1~5~J~I--~15~3~0~~15--3~1~~IS~3~1--~IS~~3~0~15~
May Jun. Jul. Aug. Sep . Oct .
June rise and become inclined in July. The vertical and horizontal
displacement of the isohalines and isothermals indicates a vertical
circulation up the shelf slope and offshore on the surface. An offshore
flow of nearly the same magnitude (7 x 10- 2 m/s) can be inferred from
the near-surface current records of the moored Aanderaa meters. How-
ever, during the rest of the upwelling period the isopycnals are hori-
zontal and the inclination that is characteristic of coastal upwelling
is absent over the Ghana continental shelf. This curious feature may
be the result of the low latitude of the Ghana coast where the hori-
zontal scale length of the boundary layer, the Rossby Radius of Defor-
mation, is approximately 80 km - much larger than the shelf width -
and the expected structure may simply be beyond the range of the trans-
ect.
2.4 Conclusions
The hydrographic changes that occur during the Ghana upwelling are ab-
rupt and dramatic. The usual stability and stratification of the water
over the shelf vanish as cold saline water characteristic of 60-100 m
depths rises to the surface. The tilting of the isopycnals and the off-
shore displacement of the isohalines suggest the surface divergence
typical of a coastal upwelling driven by local wind stress. It is puzzl-
173
Fig. 6. Cross section of tem-

perature, salinity, and density
over the shelf perpendicular to
the coast south of Tema during
the onset of the 1974 upwelling.
The position of the salinity
maximum is indicated by the
dashed line
..
.c:.
Q.
t,I
a
10 20 30 10 20 30 10 20 30
Di~tance from Tema harbour (km)
ing, therefore, that upwelling events and the yearly variation in the
upwelling intensity do not correlate with the local wind.
Hsueh and O'Brien (1971) showed that coastal upwelling can also be in-
duced by an alongshore current. In the northern hemisphere onshore flow
occurs in the bottom frictional layer when the coastline is to the
left of the current, and Hsueh and O'Brien show that this can influence
the coastal upwelling as much as the surface wind stress. Continuous
current records from the moored Aanderaa meters show that the under-
current before and during the Ghana upwelling is consistently westward
and that the flow in the bottom frictional layer is offshore. In addi-
tion there is no event or sudden change in the current structure to
suggest that the circulation is driving the coastal upwelling.
The periodic fluctuation in the surface temperatures that persisted

throughout the upwelling during 1974 was unusual. In Figure 5 we see
that this is the result of a barotropic variation in the depth of the
isotherms over the entire shelf. These variations with a period of
14 1/2 days propagated westward along the coast at a speed of 55 ± 10
cm/s. Investigation of these waves (Houghton and Beer, 1976) suggests
that they are not trapped barotropic shelf waves but some other type
of wave of oceanic origin. This is of particular interest since other
wave modes have recently been observed during GATE in the Gulf of
Guinea (Dliing et al., 1975) and because such wave modes are thought
to be important in coastal upwelling (Gill and Clarke, 1974). The sud-
den changes in the coastal hydrography during the upwelling could be
the effect of a superposition of these waves on the gradual seasonal
variation in the hydrographic structure of the Gulf of Guinea. Since
the waves can be generated nonlocally, this would account for the ab-
sence of correlation between the upwelling and the local wind. Although
174
at present there is no precise model of this mechanism, the great in-

terest and active research in the Equatorial Atl antic should provide
more information about this fascinating problem in the near future.
3. Biological Effects
3.1 Phytoplankton
Counts of phytoplankton cells do not give absolute quantities of phyto-

plankton present; they only provide relative values for the different
samples. Figure 7 shows these phytoplankton counts at station A1 on
BOO
700
29
r'\,..,/,.. _-J'\
~~~
.
f .~ I
i i
500 \J 1 !\ I
400 \! \\ i
300 if .1
200 ~ ~I
.
..,..
....
100 ........_.--jC Temperature I:'
24
E • Phytoplankton ~
~ 23
Z
~ ) 22
21
=~O !\/\ )
~: J~ ~ 20
Fig. 7. Numbers of phyto-
I O~ ... J''-'-____ ~-- 19 plankton cells at station
AI' 1970
J
days when samples were taken for the year 1970. Similar values were
obtained at stations A 2 , B, and C. This diagram shows that, on the
average, quantities of phytoplankton cells during the period of hydro-
graphic stability (October-June) are low while the values for the up-
welling period are high. The counts during October to June were about
100 cells per sample while during July to September, a peak as high as
2700 cells per sample was obtained.
However, a number of very low values obtained during the upwelling

season calls for further comment. One characteristic feature of the
upwelling season is the occurrence of phytoplankton blooms. Between
175
the end of one bloom and the beginning of another, quantities of cells
are very low. It is considered that the samples with low cell counts
were taken during the period between one bloom and the next.
It is not possible from the data (Fig. 7) to show if there is any time
lag between the occurrence of the upwelling and the high phytoplankton
production. However, it is assumed that if there were any time lag i t
would be less than one week (the samples were taken once a week). This
might be the reason why the time lag was not discernible from the data.
Many more species of cells occur during the upwelling season than during
the stable hydrographic period. During the upwelling season a few of
these species become dOminant, sometimes forming blooms. The dominant
species during the upwelling period include: SkeZetonema sp., CosainodiscUB
sp., ThaZassiosira sp., Nitzschia sp., ThaZassiothrix sp., Chaetoceros sp., Asterio-
neUa sp., and RhizosoZenia sp.; the first four usually form blooms.
Common species that occur during the stable hydrographic period include
Ceratiwn sp.,· Dinaphysis, Peridinidae, and Cyanophyceae.
3.2 Zooplan~ton
For all three years (1972-1974) under consideration, a good correlation

was obtained between the fall in temperature and the increase in zoo-
plankton production at all four stations (Figs. 8-10). The pattern of
Station B
~ 1,400
°c
as 1,200 30
>01000 2S
58 'soo 26
ij"3- 600
ge 400
<i.e
is'- 200
O~~~~~-L~~~~~~
Station A2 Station C
"e
::I
1,400r-r-r-.-.-,-,-....=,-.,...-r-.-,-.....,·C
g'S ~OO
1,000 o~_.","',, ,.,
~
28 ...._-r......."
... 0 0-' .--,
eO 800 ..."
II1Q 6 00 ____ 0 Temperature \
,,-
, 26
24 0---_ Temperature'-\
u_ Displacement - Displacement \ /
ae 400 volume 22 volume V
<i.e
.!.- 200 20
o
° ~
Fig. 8. Mean monthly displacement volumes of zooplankton and surface temperatures
at four stations off Tema, 1972
fluctuations in zooplankton displacement volumes was similar though

the absolute values of production were different for the three years.
The values were high during the upwelling period and low during the
period of hydrographic stability.
Of the three years, highest production occurred in 1972. Displacement

volumes fluctuated between 1PO and 300 ml/1000 m3 in the stable hydro-
graphic period; these values increased with the onset of the upwelling
176
Station A I Station B
Col 1,400r-r-.-.-.--.--.-........-r-r-r-,---,°C OC~.--.-,-........-r-r-,--,--.-.-,-,oC
~ E 1,200 Temperature 30
,,~...........
0 _____ •
.............................. , 30
~ 0 1,000 - Displacement volume... B
.C; g
BOO ..,........... , / -"0 26
..' \
,
\ ,l"
~' 2B
26
~-3 600 ,.,... " ...,........ // 24
60 0 - - - - 0 Temperature
'---.../ 24
~ e 400 ~
•.,-." 22 400 - DisPlacem~nt 22
-[.: 200 20 200 ................................
volume 20
.!!l
o 0 ~ O~~~~~~~~~~-L-L~IB
/---'----/v
1400
e..,
Col
0----- Temperature °C Col
Temperaturrz
°c
" e 30 ~E 1200
e---_e 30
00 - 0 i~e.!~crzmrznt volumrz", Displacrzment volume,-"',...
>0 .............................. \ ,/' ...... 2B go 1000
.. 0 BOO
2B
c;o \ /", 26 50 26
... "
CoI-
e- 24 e- 600 24
CoI- \ \- ' Col"'::::
u e ge 40
---_
22 22
.." .-
<i.e
a
20
IB a"'.-
<i.e 200
........................
0 ~~~~~~-7-7~-=~~~IB
20 -
J F M A MJ J A S 0 N D
Fig. 9. Mean mbnthly displacement volumes of zooplankton and surface temperatures
Station AI Station B
1,400 1,400
Col
Trzmprzraturrz °C
Col
°c
~.., 1,200 .,.- 30 ;.., 1,200
.....- ......... ,, 30
"
~!~~acemrznt volumrz -e
2B gOlpoo 2B
,,
g6 1,000 # •• 0 _,
................. fI"/ •.•..

\,,
-'.0
C;O BOO 26 ~g BOO ,-' 26

0---- Trzmprzraturrz '''.,',-
\
~Q 600 \
\, 24 ~.::::. 6 / 24
Col-
ge 400 22 g"E 400 - Displacrzment
22
Q.e 200 20 <i.e 200 _ ...• volumrz 20
a ......
",.-
"--e...........
a
",.-
0 IB 0 18
,
1,400 1,400
Col
TrzmJIIZrature °c Col
30 ~ ... 1.200
...................-...........
~ ... 1,200 DispkIcrzmrznt volumrz
~6IPOO ............................" 28 gljlPoo
C; g 800 ~
" , '\
'.- .......'\ ,."

,,.--" 26 "0 BOO
50
0"
'-, ,.'
~"- 60
~e 400
'0'- 24
22
e..::::.. 60
CoI_
g e 400 -
0---... Trzmf,rzraturrz ' .. / '
Disp acemrznt volume
.2 e <i.e 200
Q..- 200 20 ",'-
.!!!
0 0 IB is 0
Fig. 10. Mean monthly displacement volumes of zooplankton and surface temperatures
(or drop in temperature) to the maxima, which ranged from 770 to 1280
ml/1000 m3 of water filtered. In other words, maximum production ob-
tained was as high as 6 to 11 times as much as in the stable hydro-
graphic period, while minimum multiples were 3 to 5. In 1973 and 1974,
though absolute values were lower than in 1972, maximum production in
the upwelling period was as high as 5 to 12 times as in the stable pe-
riod while minimum production was 2 to 5 times as much.
Whereas more species of phytoplankton cells occur during the upwelling

period than in the stable period, fewer species of zooplankton, espe-
177
cially copepods, occur in the upwelling period. The copepod population

is domina ted by the herbivorous calanid, Calanoides carinatus (Kr¢yer).
In a normal sample at the peak of the season, C. ca:I'inatus may constitute

a significant proportion by number (60-80%) of the zooplankton.
This species is not found on the continental shelf during the period
of hydrographic stability (Mensah, 1974a). It has been found as
stage V copepodites in very low concentrations (less than 500 specimens
per 50000 1 of water) at depths of 500 m and below beyond the con-
tinental shelf.
A detailed study of its life cycle in relation to the upwelling has

shown that as soon as the drop in temperature begins, this copepod at
stage V copepodite initiates a shoreward migration to the shelf along
the bottom. As they migrate down the shelf, they move up toward the
surface waters and moult into the adults, having fed well on the abun-
dant phytoplankton and bacteria populations available. The life span
of the copepod (i.e., generation time) is about 28-32 days (Mensah,
1974b). The adults die after one spawning. Depending upon the duration
of the upwelling (2 1/2-4 months) there may be 3 or 4 generations,
respectively. At the end of the upwelling season when the temperature
begins to rise, the offspring of the last generation initiate offshore
migration toward the continental slope. This last generation suffers
from high mortality leaving a very low concentration of stage V cope-
podites that stay in the deep waters until the next major upwelling
(Mensah,1974a).
3.3 Fish Production
Fish production is also very high during the upwelling. This is espe-
Cially so with the temperature-dependent zooplankton-feeding clupeid
v
"
~-
c- .-.!:;C
.. E
Ec
bI":':;' lI----.,. Monthly beech temperatures
.. 0
v .. :::!:~
.EE
"'-
Q.:::J
.- 0
- ......... Plankton displacement
45,000 volumes JL ·C
0> - - - Monthly lendings of 0
1,400 35,000 S. aurita (me tric tons) 29
1,300
""-----.... --_..----_...... . . ..... "" 28
1,200 30,000 " 27
""
\
\
1,1 00 ~ \\ 26
1.000 25,000 25
900 24
800 20000 23
700 22
600 15000 21
500 20
400 10000 19
300 18
200 5,000 ............-----.---. ~.-. 17
100 16
0 0 15
J F M A M J J A S
Fig. 11. Relationship between the upwelling and production of zooplankton and Sa:I'-
dinella aurita in 1972
178
Sardine lZa au:ti. ta . S . auri ta has been economically the most important pe-
lagic fish in Ghana, and i t constitutes approximately 45% by weight of
the total marine landings annually. However , . it is seasonal, appearing
on the traditional fishing grounds mainly during the upwelling season,
i.e., from approximately late June to the end of September or the first
half of October. Figure 11 contains a summary of the relationship be-
tween the upwelling and the production of zooplankton and S. au:ti.ta.
It is evident from this diagram that the maximum occurrence of this
fish on the shelf takes place during the upwelling season.
Also, Figures 12 and 13 show that even though the species carries out
minor spawning for the greater part of the year, maximum spawning takes
place during the upwelling season.
12,0001
1
.. 5,000
..
C>
C>
a ~ooo
Oi
c Station Al Station AZ
'0
;; 3.000
If)
'0
2,000
~
Of
.c
E
z"
1,000
o
o 10 20 30 40 SO o
.--
10 20 30 40 SO
W te k 5
Fig. 12. Numbers of Sardi nel la eggs per 1000 m3 of water in 1969
3.4 Ev aluation of the Biological Conclusions
It is significant that the high production of phytoplankton and zoo-

plankton during the three-month period of the upwelling coincides with
the period of the major spawning of the S. au:ti. ta. Thus the availability
of sufficient plankton as food for the larvae as well as for the adult
S. au:ti.ta is assured: Kwei (1964) showed that the adult S . au:ti.ta feeds
on zooplankton, principally, the calanid CalanoidEs carinatus. It is also
important that the zooplankton population is dominated by C. carinatus ,
which has been found to be a phytoplankton feeder (Mensah, 1974b).
This copepod feeds on the abundant phytoplankton and undergoes 3 or
4 generations during the upwelling season, thus providing sufficient
numbers of this copepod as food for the adult sardine.
Acknowledgments. We are grateful to the Ghana Me teorological Services Department

and the Survey Department for their c ooperation in providing important data for this
work. M.A. Mensah is grateful to Miss E . R. Ana ng of the Fishe ry Research Un i t for
providing the phytoplankton data, and to Dr. E.A. Kwe i of the volta Lake Research
Proj e ct, for his criticisms of the manuscript.
179
SPOOr
2,000 2POO
Station A1 Station A 2
1,500 1,500
__1 .---
1.000 1,000
..a
~
>
.5!
500 500
I
'0 2,000
0
10 20 30 40 50
0
2,000
.
III
~
.a 1,500 Station B 1,500 Station C
E
:J
Z
Ji
1,000 1,000
500 500
--JV
0 0
I
0 10 20 30 40 50 0 10 20 30 40 50
Weeks
Fig. 13. Numbers of SardineZZa larvae per 1000 m3 of water in 1969
References
Diiing, W., Hisard, P., Katz, E., Meincke, J., Miller, L., Moroshkin, K.V., Philander,
G., Ribinikov, A.A., Voigt, K., Weisbert, R.: Meanders and long waves in the Equa-
torial Atlantic. Nature (London) 257, 280-284 (1975)
Gill, A.E., Clarke, A.J.: Wind induced upwelling, coastal currents and sea level
changes. Deep-Sea Res. 11, 325-345 (1974)
Glover, R.S.: The Hardy Plankton Indicator and Sampler: a description of the various
models in use. Bull. Marine Ecol. 4, 7-20 (1953)
Houghton, R.W., Beer, T.: Wave propagation during the Ghana Upwelling.J. Geophys. Res.
81, 24, 4423-4429 (1976)
Hsueh, Y., O'Brien, J.J.: Steady coastal upwelling induced by an alongshore current.
J. Phys. Oceanog. 1, 180-186 (1971)
.Kwei, E.A.: Migration of SardineUa aurita (Val. et CUv.). Ghana J. Sci. .! (1), 34-43
(1964)
Longhurst, A.R.: A review of the oceanography of the Gulf of Guinea. Bull. IFAN A24,
633-663 (1962)
Mensah, M.A.: The occurrence of the marine copepod CaZanoides carinatus (Kr¢yer) in
Ghanaian waters. Ghana J. Sci. ~ (2), 147-166 (1974a)
180
Mensah, M.A.: The reproduction and feeding of the marine copepod Calanoides carinatus
(Krpyer) in Ghanaian waters. Ghana J. Sci. 14 (2), 167-192 (1974b)
Morliere, A., Rebert, J.P.: Etude hydrologiq~du plateau continental ivoirien. Doc.
Scient. Centre Rech. Oceanog. Abidjan 1 (2), 1-30 (1972)
Morliere, A., Hisard, Ph., Citeau, J.: Le courant de Lomonosov dans le fond du Gulf
de Guinee Moi 1973. Doc. Scient. Centre Rech. Oceanog. Abidjan 5 (1-2), 85-102
(1974)
Oceanographic Conditions in the Galapagos Archipelago and
Their Relationships with Life on the Islands
G. T. HOUVENAGHEL
1. Intr.oduction
The Galapagos Islands, formed by the emerging tops of huge submarine

volcenoes rising from the sea floor, are located on the equator about
900 km west of the South American mainland.
In this part of the eastern tropical Pacific Ocean the distribution

and circulation of the water masses are intricate, as outlined by Wyrtki
(1966) •
The oceanographic information and data published for the Galapagos area
are scarce and sporadic; oceanographic cruises in the eastern tropical
Pacific included just a few stations close to or within the archipelago.
Wooster and Hedgpeth (1966) reviewed the existing information to de-

scribe the oceanographic setting of the Galapagos. Goering and Dugdale
(1966), Barber and Norton (1968), and Wiebe et al. (1968) focussed
their interest on the submerged crater that forms an isolated basin
(Bahia Darwin on Genovesa Island). More recently, Sibert (1971) de-
scribed briefly the hydrologic profile in Bahia James, Santiago Island
and Pak and Zaneveld (1973) analyzed the Cromwell Curr~nt flowing to
the east of the Islands.
During a Belgian zoologic expedition to the Galapagos Islands in 1967-

1968, we made hydrologic, chemical, and biologic observations. This
gave us the opportunity to study the oceanographic conditions in the
archipelago and more specifically its seasonal fluctuations.
2. Methods
One part of our work was carried out in Bahia Academia on the southern
coast of Santa Cruz, a central island of the archipelago (Fig. 1).
Samplings were made from April to September 1968. Three stations located
in different environments were visited about every ten days: Station 1
in coastal waters at the entrance of the bay, Station 2 in sheltered
inshore waters at the southern end of Bahia Academia, and Station 3
in the exposed waters at its northern end. All the samplings were car-
ried out at the surface from an outboard engine dinghy belonging to
the Charles Darwin Research Station, where we analyzed them within
two hours after collecting.
We measured the following parameters: temperature, salinity, oxygen,

phosphate, silicate, nitrite, nitrate, pH, alkalinity, primary produc-
tion (oxygen method). They were determined using the methods given by
Strickland and Parsons (1960). The plankton was sampled at the surface
with a biconical net with an,entrance of 25 cm diameter and a 75-cm
long filtration gauze of 140 ~m mesh size. The open area ratio and the
182
....
" .
.. '.
.. . .
• .
, .,..
'
.
."
':
. ..
o
.. ... . .'
• I·
.' • •
....
..
.,..
o ~. e•••••
•
. -:·6Crist6~·'!.I: .....
~
...... J •
.. .
tv....
-...
• "..
:
,:.. "..
... '- .... '\
S
Floreana
.. ... .- .
;. :
..
.. . ..
•f
'
Fig. 1. Chart of the stations considered in this study. 0, samplings made during the
Belgian Expedition in 1967-1968, and e, A, respectively, the BT and Station data on
file at the NOAA. (a) Bahia Wreck, (b) Bahia Academia
fitration efficiency (at 100 cm/cm/s) calculated for this gear accord-
ing to Tranter and Smith (1968) are 5.77 and 0.996, respectively.
Another set of samplings concerned the entire archipelago and covered

the period from September 1967 to September 1968. The stations in off-
shore waters are represented by circlets on Figure 1. For logistic
reasons and boat limitations, these samplings were limited to tempera-
ture, salinity, and plankton. In offshore waters a Tregouboff type
net was used. The sizes are 50 cm diameter at the entrance and 220 cm
length and the 4 parts of network have mesh sizes of 400, 347, 123,
and 68 ~m, giving a global open area ratio of 4.52 and a filtration
efficiency of 0.927. The plankton biomass values are expressed as dis-
placement volume (ml/100 m 3 ) and were evaluated according to the method
of Yentsch and Hebard (1956).
To achieve a better understanding of the hydrology within and especial-

ly around the islands we also took advantage of bathythermographic and
station data in archives at the National Oceanic and Atmospheric Ad-
ministration (NOAA, Washington D.C., USA). Station data from the cruises
S.O.E. 17 and 19 of the R.V. Te Vega in 1968 supplemented this study.
183
3. Temperature
In the Galapagos surface water temperatures are neither uniform in

space nor stable in time although the islands are on the equator. The
annual temperature cycles as observed both at Bahia Wreck, San Crist6-
bal Island, (Abbott, 1966) and at Bahia Academia (Houvenaghel, 1973)
are shown in Figure 2. The annual cycle includes two seasons: a warm
a b
28 28
26 26
24 24
22 22
20 20
18 18
16 16
JAN MAR MAY JUL SEP NOV JAN MAR MAY JUL SEP NOV
Fig. 2a and b. Monthly variation of the sea temperature in surface waters at Bahia
Wreck (1958-1963) (a) and at Bahia Academia (1965-1971) (b). The mean and extreme
values are shown
one from February till April and a cold one during the rest of the year.
Year to year fluctuations in the intensity and the duration of both
seasons are noticed. These cycles correspond roughly to those calculated
according to the harmonic analysis of sea surface temperature in the
Pacific Ocean by Wyrtki (1965a) and Panfilova (1968). When comparing
the observed and the theoretical curves, discrepancies appear in the
amplitudes and the length of the seasons.
The most numerous data by far for describing hydrology in the Galapagos
area are temperature data. Besides the daily routine measurements at
Bahia Academia and Bahia Wreck some 1248 surface temperatures from all
over the archipelago have been used in the present study. The extreme
surface values are 15.0o C and 29.3 0 C with a mean of 22.6 o C.
In a first step, surface temperatures have been plotted against time

for all quadrates of 1° longitude bg 1° latitude covering the Galapagos
region from 880W to 93 0 W and from 2 N to 20S. By smoothing out a curve
through the pOints in each diagram, we determined the mean temperature
cycle experienced in the area. Another set of similar plots and curves
was constructed for 15 coastal stations (embayments and anchorages)
frequently visited. All these curves show that the lowest surface tem-
peratures are encountered to the west and the southwest of the archi-
pelago and that there is a decreasing gradient in coolness when going
toward the north and the northeast.
184
Fig. 3a and b. Monthly

charts of surface tem-
peratures. The isotherms
for March (a) are plot-
ted from 56 values and
those for September (b)
from 60 values
21
26 25
r
24
23
22
ZI
&:7
<::I
8 w b
Monthly charts of surface temperature have also been drawn. When data
were lacking for a particular region the values predicted by the annual
curve for that region were used.
The charts clearly reveal the distribution and the extension of the
cold waters (Fig. 3). A cold-water patch is always present west of
Isabela Island even during the warm season (chart a, Fig. 3). This
patch gets colder and grows in area, extending mainly to the south and
the east of the archipelago during the cold season (chart b, Fig. 3).
The maximum extension of the cold waters occurs from June to August-
September.
185
Analysis of the data from the bathythermographic stations, shown as

dots on the chart in Figure 1, provided the link between the vertical
structure of temperature distribution in the waters surrounding the
islands and the surface patterns described by the monthly charts.
The basic feature of the thermal structure is the presence of a per-

manent thermocline that is nearer to the surface at the equator than
away from it. This corresponds to the general pattern described in the
eastern tropical Pacific by Cromwell (1958) and Wyrtki (1965b). To the
north but mostly to the northeast of the archipelago an upper mixed
layer about 25 m deep, increasing in thickness to the north, is limited
in depth by a sharp thermocline with gradients of O.3 0 Cjm to 1.8 0 Cjm.
To the west and south of the archipelago such a highly stratified struc-
ture disappears during the cold season or may even be lacking through-
out the year. The thermocline here is weak, with gradients from O.05 0 Cj
m to O.18 0 Cjm and frequently reaches the surface, revealing an upwelling
pattern. The bathythermographic stations where the thermocline inter-
sects the surface are shown on the chart of Figure 4; they appear to
10N 91 0 W 89 BW
~
.
• <:> ..
00 0 A •
~. ) , ... )~O
• 0,
•
~ ··.U
• .. ,.:.'
.. . •. 0
•
Fig. 4. Chart showing

• .<:1 -C ~ the bathythermographic
• • • • • stations where the
• 00
0
2BS • • • 0 thermocline intersects

the surface
the west and especially to the south of the islands where cold waters
are cornmon according to the monthly surface charts. Many of these sta-
tions where an upwelling occurs lie above the southern slopes of the
underwater relief of the archipelago and especially where the bathy-
metric chart indicates shallow regions or island pedestals (see Fig.
14). The relation between topography and upwelling may also be proved
by the bathythermographic tansects from series of stations across the
archipelago (Fig. 5). In the transition region between the well-stra-
tified waters in the north and northeast and the upwelling regions in
the south and west, the thermocline is irregular and thick and has
relatively weak gradients. This transition zone is a characteristic
feature of the archipelago waters east of Isabela Island.
From series of station data along tracks similar to those presented

for BT sections in Figure 5, further information is deduced concerning
water distribution, reduction of stratification in shallow areas, etc.
186
o
1 o
--
m 20
l-
I 2Q m
I --!
100
15~ 1 .-1.15
100
I
J
15
T-----j
~
200 200
15~
300 300
100
200
3OO ~----------------~~~~~~~~----~
t'
5
100
200
Fig. 5. Examples of isotherm profiles determined from serial bathythermographic

stations corresponding to the tracks sho~m on the map
The distribution pattern of water stability (10 8 E'), described, e.g.,

by the transects in Figure 6 (west-east track across the southern part
of the archipelago), indicates that the uppermost layer, where stability
reaches high values, is sharply limited by the waters from below, where
stability is reduced and may even be negative.
The low and negative values are observed for the waters reaching shal-
low parts of the archipelago and, in the left part of these sections,
correspond to stations located south and south-west of and close to
Isabela Island.
113 173 172 171
I drt_ , 'u
~P:,UtI '" F~
50
lDO
150
200
400
600
800
lDOD
Fig. 6. stability transect (108E ,) constructed for stations along a west to east track across the southern part of the
archipelago. Mean depth of the maximum salinity is the dotted line. Shaded areas represent negative stability
ex:>
-..J
188
These waters belong to extensions of the Equatorial Undercurrent around

and across the archipelago as shown by the close relation between the
stability pattern and the depth of the maximum salinity distribution;
the deflections in their flow by bottom topography seem to be respon-
sible for the presence of unstable water bodies.
The reduction of the water stratification, observed as a broadening and

a weakening of the thermocline in the BT sections, should result from
the turbulence induced by the topography upon water flow.
The role of tidal mixing in reducing stratification in the water column

does not extend below the upper layer since a comparison of the pro-
files for hydrologic parameters (S%o, oxygen, phosphate, silicate,
nitrate, nitrite) with the temperature profile reveals that the upper
layert which is stable, is also accompanied by noticeable nutrient con-
sumption and oxygen production. Within the thick thermocline found in
these waters a staircase structure is frequently observed, and a dis-
tinct secondary superficial thermocline may also be present. This stair-
case structure indicates the presence of complicated circulation with
antagonistic components consisting of the superficial waters flowing
to the west and the subsuperficial waters flowitig to the east. The
secondary superficial thermocline is the consequence of local heating
and is linked with an increase in stability of the superficial layers.
Such a phenomenon occurs especially in protected regions in the central
part of the archipelago or in bays, where surface temperature charts
indicate wedged areas of maximum temperature, and also in the super-
ficial waters downstream from an upwelling area. This thermocline pat-
tern with a secondary superficial thermocline may be related to the.
seasonal shallow thermocline described by Wyrtki (1965b) for the Peru-
vian waters; however, in the Galapagos, this feature is not restricted
to the warm season, but may be found throughout the year.
The seasonal, horizontal, and vertical distribution of temperature,

and the patterns and locations of upwellings contribute to the evidence
that the waters involved in the upwellings and influencing the hydro-
logy of the entire archipelago, are subsuperficial waters belonging to
the Equatorial Undercurrent.
4. Salinity
During our seasonal survey at Bahia Academia, the salinity fluctuated

from 33.88 to 35.51%0 . As a general rule, cold waters are characterized
by high salinities while warm waters exhibit the lowest salinities.
The analysis of the surface salinity distribution in the archipelago

was based on the data from our expedition in 1968 (128 values) and on
those from the NOAA archives (174 values). The upwelling waters can
be recognized by their high salinity while the waters to the north and
the north-east are always less saline. These tropical surface waters,
cornmon in the warm season over most of the archipelago, disappear from
this area by the end of April when a concomitant and progressive ex-
tension of the cool waters with high salinity takes place. The maximal
extension of the 35%. waters is reached at the surface in May-June.
Later on, the 34%0 isoline, first appearing to the north and the east
of the islands, spreads over the archipelago in such a way that it may
cross the 10 N latitude as early as September.
The salinity data help to describe the extension of the upwelling waters
and their further mixing and heating when flowing downstream. Such
189
heating is a cornmon feature especially in the warm season, as indicated

by the temperature/salinity diagram of Fig.ure 7. The salinity distribu-
tion pattern also indicates that at that time the flow of the tropical
33 34 35 3& S ~D
'e
-•• ...,
0 0
u •
•• •
00
• 0 · • . @•• ~
. 0
00
0
0
2. 0
~ :. j
:.;
0
~ cfo-;t• 00 .. . 0
• . . .
':*1'*\.....
....
~
22
. . 0 'i0 •
*,.
I*' ...... ..;.. :
lan o ,." ••• : e
0 eb
•• •
..
mar .. @* .... :
20 apr • II-. ••• •
!'lay
• ,ul
jUn
••
•
e•
* o ••
(!) aug
II ~ up
33 3~ 3S 38
Fig. 7. Temperature/salinity diagram for the surface samples collected in 19 68

throughout the archipelago
33 34 3S 3S SX.
25
20
15
10
Fig. 8. Temperature/salinity
diagram for the Te Vega sta-
tions during summer, 1968
5
(~February-March) and
winter, 1968 (~August)
33 34 3S 34 3S
surface waters toward the archipelago does not reach further south than
north of Isabela Island on the western edge of the archipelago and the
southern part of Santa Cruz Island in the central area . This, in turn,
explains the striking difference noticed between the temperature re-
gimes in the waters of Bahia Academia and Bahia Wreck (Fig. 2).
190
Figure 8 contains the temperature/salinity diagrams constructed from

data collected from Te Vega stations during the summer cruise (Fig. 8,
A February-March, 1968) and the winter cruise (Fig. 8, B August, 1968).
The temperature/salinity curves for the winter are more scattered than
those for the summer.
During both seasons some curves indicate upwelling while others de-
termined from stations to the north and the east of the archipelago
represent mixing patterns between the Equatorial Undercurrent core
water and the tropical surface waters.
Some transects and profiles describing the vertical distribution of

the salinity show a distinct, localized core of high salinity in the
subsurface layers. This feature is a property of the Undercurrent as
demonstrated since Montgomery and Stroup (1962). The salinity maximum
observed in 1968 by the R.V. Te Vega appeared between the levels of
thermosteric anomaly of 180 and 300 cl ton- 1 • This is consistent with
the Piquero and Eastropac data collected west of the Galapagos, where
at 10 S 93°5 W the high salinity and high velocity cores are found
virtually at the same depth (Taft and Jones, 1973).
The analysis of salinity distribution corroborates the role played by

the Equatorial Undercurrent in determining the upwellings in the Gala-
pagos Islands.
5. Nutrients
The seasonal cycle of the thermohaline properties in Bahia Academia

indicates sudden changes in the water quality: They result from ad-
vections or from locally induced inshore upwellings.
Phosphates at the three stations are in phase with the thermohaline

fluctuations. The nitrate and nitrite concentrations are irregular;
however, they exhibit a tendency to be higher in the cold season. The
silicates, also irregular, behave differently at each station: At Sta-
tion 1, in open coastal waters, the silicates remain low - around 1 pg-
at/l during the entire survey, while at the other two stations, in in-
shore waters, peaks reached 6 to 7 pg-at/l at times. These peaks could
be linked with inshore upwellings, which brought waters from the bottom
of the bay into the surface layer.
At the time of our observations the R.V. Te Vega visited the Galapagos
twice and established eight hydrologic stations within or close to the
archipelago. The data provided by these cruises gave us the opportunity
to compare the nutrient concentrations in coastal and inshore waters
with those from the surrounding oceanic waters. For each sample the
nutrient concentration was plotted against the density of the water
(Figs. 9-12).
For all the nutrients examined the scattering of points is divided
into two parts: one that is relatively homogeneous at crt < 26 and an-
other one, much more scattered, in the lighter waters above this value.
The limit of crt = 26 corresponds to the salinity maximum observed in
the temperature/salinity diagram constructed from the Te Vega data
(Fig. 8).
An enrichment of phosphates (Fig. 9) is noticed in the coastal waters;

this could be explained by ,regeneration processes taking place in those
waters. At one Te Vega station close to San Crist6bal Island the phos-
phate values are similar to those measured in Bahia Academia. Such
191
~ <::!.
z
OJ PHOSPHATE OJ NITRITE
0> 0>
::l. ::l.
3.0 3.0
2.5 2.5
2.0 2.0
1.5 1.5
1.0 .!' 1.0

o
0.5 0.5
0·°22
i S-
en
OJ NITRATE OJ
0> 0>
:l. :l.
15
10
0
Bahia Academia
5
0
stal. 1
stal. 2
0 stat. 3
Te Vega stal.
... ' 0
-surf.
S 'i'oo max .
°22 6t
Fig. 9. Phosphate/density (at) diagram for the Te Vega hydrologic stations (shown
as joined dots) from S.O.E. 17 in February-March 1968 and S.O.E. 19 in August-Sep-
tember 1968, and for the data measured at the surface frcm April till September 1968
at 3 stations in Bahia Academia
Fig. 10. Nitrite/density (at) diagram. For details see Figure 9
Fig. 11. Nitrate/density (at) diagram. For details see Figure 9
Fig. 12. Silicate/density (at) diagram. For details see Figure 9

192
features also suggest a local enrichment in phosphates above the shal-

lower parts of the archipelago.
The nitrite distribution (Fig. 10) is characterized by maxima above

the pycnocline. This could explain the appearance of peaks of nitrite
concentration observed at the surface in Bahia Academia. All the distri-
bution patterns in the nutrient/density diagrams constructed for phos-
phates, nitrites, and nitrates also (Fig. 11) indicate that the waters
on the southern coast of Santa Cruz Island are not undercurrent waters
reaching directly the central part of the archipelago; they are recently
upwelled waters, which, owing to solar heating, became lighter and in
which primary production and regeneration is going on. The silicate
diagram (Fig. 12) distinctly shows a co~sumption pattern leading, as
we mentioned in the seasonal analysis in Bahia Academia, to extremely
low values in superficial coastal waters. This is evidence for the im-
portant development of diatoms in those waters. The fact that silicate
concentrations may reach such low levels also indicates that this nu-
trient could be a limiting factor.
Some of the station data from NOAA enabled the construction of trans-
ects describing the vertical distribution of nutrients in selected
regions of ~e archipelago. In upwelling areas and further away down-
stream in the upper layer, above the pycnocline, the nutrient concen-
trations are lower than those in the maximum salinity core of the Under-
current. For nitrites, however, highest concentrations occur at the
bottom of the upper layer when the stability at this level is high.
For the transects crossing shallow parts of the archipelago, enrich-
ment in phosphates is also noticeable above the sea-bottom.
The range of the nutrient concentrations found in different types of

waters in the galapagos area is summarized in Table 1, which clearly
demonstrates that the eutrophication of the waters of the Galapagos
archipelago is the result of the input into this area of the nutrient-
rich waters belonging to the core of the Undercurrent.
Table 1
Nutrient concentration in ~g-at/l
Source: NOg
Archipelago
Offshore sur- NOAA 0.04- 1.47 0.0-11.8 0.00- 0.62 0.0-15.0
face waters archives n =96 n =32 n =56 n =49
Te Vega 0.27- 0.85 1.1- 8.5 0.03- 0.11 7.5-16.1

1968 n = 8 n = 8 n = 8 n = 6
Maximum salin- Te Vega 1.20- 2.00 14.7-26.5 0.00- 0.35 8.3-20.0

ity core n = 8 n = 8 n = 8 n = 6
Bahia Academia present

study 0.45- 2.90 2.3-28.8 0.00- 0.70 0.5- 8.0
Coastal and
1968 n =51 n =51 n =51 n =51
inshore sur-
face waters
193
6. Oxygen
The common occurrence of waters recently upwelled and belonging to the

Undercurrent is also attested by the distribution of dissolved oxygen.
From the station data, the amounts of dissolved oxygen at the surface
range from values as low as 2.25 ml/l up to 6.20 ml/l corresponding
to 43 to 119% saturation (Fig. 13). The most prominent fact is that
6 ••
S· ..
...
5
r-. . ." -tIC.·· '-: ~.. .. ., ~ I . •
7t"~ ~:
.: .-. . . . ~~ • • • • • -. • ••
.,. .:...:...\. ., .
• I • ••fI'. •• .....
• . ....
3
: .
2
17 19 21 23 25
Fig. 13. Oxygen/temperature diagram for the surface waters in the Galapagos area
from the data in file at the NOAA. The saturation levels indicated on this diagram
are determined from oxygen solubility calculated for waters with Cl = 18%0 (upper
curve) and Cl = 19%0 (lower curve) • Values above these limits correspond to super-
saturation, those below represent undersaturation
77.3% of the samples (132 in total) collected at the surface through-

out the archipelago were undersaturated although primary production
rates are relatively high. In the coastal and inshore waters of Bahia
Academia the dissolved oxygen contents measured were at and above the
saturation level, indicating that the waters in this region were drift-
ing in the upper layers for a time long enough to allow the biologic
production to increase the oxygen content up to saturation and even
supersaturation levels. The oxygen contents in Galapagos waters are
very similar to those mentioned by Zuta and Guillen (1970) for the
Peruvian waters where, in upwelling zones, values as low as 2 ml/l may
also be found at the surface.
Owing to the oxygen distributi.on discussed above, the Galapagos area

appears to be much more undersaturated in oxygen than it is decribed
in the Eastropac Atlas (Love, 1970-1972), where saturation values of
90-105% are mentioned for February-March and 80-100% for August-Septem-
ber. This discrepancy also indicates that the upwelling is a regional
phenomenon that falls beyond the spatial resolution power of large
oceanwide expeditions.
7. Biologic Parameters
7.1 Primary Production
The analysis of dissolved oxygen distribution pointed out the impor-

tance of the primary production processes in these waters.
194
In Bahia Academia, the rates .of grass praductian that we measured by

the .oxygen methad averaged 16.7 mg C/m 3 /h but have reached 51.7 mg C/
m3 /h. The mean chlaraphyll a cancentratian was 2.18 mg/m 3 with values
ranging fram 0.62 ta 6.30 mg/m 3 • The mean praductivity index deduced
from these measures is about 8 mg C/mg ChI a/h with a minimum .of 2.2
and a maximum of 30.3.
No distinct seasanal cycle could be deduced. The peaks occur when the
water quality has changed. During the warm seasan the increase in pra-
duction corresponds, according ta the ather parameters, ta the horizan-
tal advection of trapical surface waters, while during the cald season,
the peaks only show up when the presence of cold, nutrient-rich waters
is naticed.
The relation between primary production and nutrients is indicated by

the scattering pattern in a diagram depicting the distribution of
gross production versus phasphate concentratian using all the data from
the stations in Bahia Academia. The .observed carrelation is negative
and equals -0.41 (with r = 0.37 for a p = 0.01 and n = 46). As a general
rule, it may be deduced that primary production is higher in waters in
which phytoplankton have already been developing over some days.
7.2 Plankton Biomass
The plankton biomass, expressed as biovolumes (plankton displacement),

has been studied from 250 surface samples collected throughout the ar-
chipelago (Tregouboff net) .
No seasonal cycle or trend was discernible as for primary production. The

samples from upwelling regions such as the ones to the west and south
of Isabela Island were low - about or under 5 ml/100 m3 • In coastal
and inshore waters, and especially next to upwelling zones, the bio-
volumes are much more important, and values as high as 82 ml/100 m 3
have been found to the north of Isabela Island.
From the distribution of the biovolume values in a temperature/salinity

diagram, we may also notice that the highest plankton volumes are more
common in waters of high salinity and high temperature. This means
that biologic production is enhanced by upwellings but that the plank-
ton develops highly only in these upwelled waters after they have been
drifting for some time in the upper layer.
In Bahia Academia, in the coastal waters of Station 1, when a peak of

primary production is noticed, a succeeding peak of plankton biovolume
is found from the samplings taken about 10 days later. At the two in-
shore stations such a concordance is less obvious, perhaps owing to
the greater heterogeneity of factors affecting these waters.
An explanation for the delayed start of biologic production in the

nutrient-rich upwelling waters has been offered by Barber and Ryther
(1969), who demonstrated that in the Undercurrent waters a better phyto-
plankton development was ob,tained when organic chela tors were added
to the medi um.
195
8. Conclusions
8.1 Evidence of Upwelling
Physical, chemical, and biologic parameters examined in this study

demonstrate the presence of upwelled waters in the Galapagos area and
depict their distribution within and around the archipelago. All the
oceanographic properties in the Galapagos are determined or influenced
directly or indirectly by the waters flowing with the Undercurrent.
arw
~--~--------------~------------~~~~ ____________~~____________~2 ' s
Fig. 14. Bathymetric chart of the archipelago showing the main flows of the east-
going Undercurrent waters (schematized by arrows) and the location of the most prom-
inent upwellings (shaded areas)
The Undercurrent flow patterns through the archipelago as deduced from

the distribution of the studied parameters are summarized on the chart
of Figure 14 showing the bathymetry of the archipelago. The strongest
and most regular upwelling zones are also indicated (shaded areas).
The general patterns of these flow pathways and upwelling sites suggest
that both features are linked directly to the submarine topography of
the archipelago, counteracting the west-east Undercurrent. This circula-
tion is complicated by the presence at the surface of the west drift,
196
the South Equatorial Current, which carries the superficial layer along.
All of the physical, chemical, and biologic parameters describing the
surface properties suggest that the waters driven by this superficial
current are upwelled waters that are eventually mixed and progressively
heated up while flowing downstream to the west.
In Galapagos, local upwelling should be due to the bottom topography,

which introduces an obstacle along the path of the Equatorial Under-
current. It results in deflecting the waters to the north, the south,
or on both sides of the archipelago and in the piling up and rising
of the waters upstream from the obstacles with upwelling at the surface.
An alternative explanation for the rise to the surface of the Equatorial

Undercurrent is that a divergence produced leeward of the islands in
the surface current would induce the subsuperficial water to upwell.
Such a mechanism is the result of the west-flowing surface current and,
behind it, of the trade winds.
As a result of this hypothesis the divergence zone should be restricted

to the area between the west and north-west of the westernmost island.
This of course explains the main upwelling area west of Isabela, but
it is more difficult to understand how such a mechanism could produce
the small local upwellings observed inside of the archipelago, wind-
ward of the huge Isabela wall.
When taking into account a divergence leeward of this island, one should
not expect that the rise of sub superficial water would cause a con-
comitant rise in the sea level. Examination of the tide levels for
the seven tidal stations in the Galapagos Islands (Fig. 15) shows that
there is a noticeable difference between the stations in the center
and to the east of the archipelago and the two stations to the west
of Isabela. There, the mean tidal amplitude is lower than further east.
This would indicate that piling up of water is going on against the
wall of Isabela.
The seasonal temperature patterns of the surface waters and upwelling

occurrences are not exactly in phase with trade winds; there is still
evidence of small-scale, localized upwelling west of Isabela during
the summer when winds are irregular and surface current is much weaker.
The dependence of the upwelling in Galapagos upon Equatorial Undercur-

rent activity is furthermore exemplified by the correlation found be-
tween the annual sea surface temperature cycles (8 years) in Galapagos
and those from Chimbote [7 0 41'S, Peruvian. coast, data in Zuta and Guil-
len (1970)]. This correlation gives r = 0.867 (with r = 0.443 for P =
0.001). -
Further work is necessary to establish the contribution of each mech-

anism to upwelling in Galapagos. The present data provide strong evi-
dence for the mechanical action of the topography on the Countercurrent
flow.
8.2 Seasonal Patterns
The relative importance of the different elements determining the cir-

culation in the Galapagos waters flutuates seasonally.
In the summer, when the intertropical convergence zone migrates to the

southern hemisphere, the dominant feature in the surface circulation
is the presence of the southeast flow of warm tropical surface waters
(with temperatures of 27 0 C and more and salinities of 33%0 or less)
197
N
A
I
o'
2' S 88 '
--
-- -
~~
--
;::-
-
~~
~
- -
- - .-..
=
oSW IE
1000
III 0
34
, 2 5 II 1 7
:
t ~ ~ i ~
Hp
5
3~---------------------------------- ______________
Fig. 15A and B. (A) Chart showing tide levels for 7 tidal stations in the archipelago.
Amplitudes are in feet. (B) Evolution of the mean tidal amplitude along a southwest
to northwest axis (1 to 7 = stations of tidal predictions)
coming from the north . At that time of year, a local patch of colder
waters is still present to the west of Isabela Island contrasting with
the general warming up. From the salinity distribution we may deduce
198
that the warm tropical surface waters do not normally reach the southern
edge of the archipelago. Later in the year, from March-April on, the
upwelling zone originating west of the islands extends south and east
of the Galapagos, bringing more and more cold, saline waters into the
superficial layers.
From June to August, but especially in August, the east-flowing waters

driven by the Undercurrent counteract with the west-going surface waters
drifting with the South Equatorial Current. This leads to a circulation
pattern suggesting the presence at the level of the archipelago of a
broad cyclonic movement.
During the last part of the year, the east-going flow passing to the
north of the archipelago increases, as demonstrated also by the local
shift of the surface isotherms to the north. A sudden decline of the
extension of the cold waters at the surface occurs at the end of the
year (December-January) when the summer season brings tropical surface
waters to the south.
The annual cycle of the surface circulation pattern coincides with the
meteorologic seasons, especially the annual shift of the intertropical
convergence zone. However, upwellings in the Galapagos waters are pre-
sent throughout the year.
8.3 Ecologic and Biogeographic Consequences of Hydrology
Surface currents have been proposed as the main vector for colonization
of oceanic islands like Galapagos since the islands are of volcanic
origin. Nevertheless, the presence, most of the year, of a crown of
upwelling waters isolates the Galapagos and prevents them from landings
and settlings of most plants and animals. Such a water barrier, acting
against potential immigrants from outside the archipelago or from other
islands, is the dominant factor promoting isolation and endemism.
The combined action of the hydrologic, climatologic, and edaphical

properties at the shore, and especially temperature and dryness, local-
ly introduces another type of selection for littoral and landing or-
ganisms. As a whole, Galapagos shores are not a fully suitable environ-
ment either for tropical or subtropical species or for temperate ones.
Examples are found in the intertidal zonation, which responds to a sharp
thermal and dessication stress. On rocky shores (Houvenaghel and Hou-
venaghel, 1974) there is a vertical gradient in the composition of the
fauna and algal flora going from temperate to tropical when walking up
the intertidal zonation area. Above the mean sea level, the rocks are
almost barren. There the species exhibit highly specialized adaptation
to dessication. Most of the intertidal life is restricted to the lower
levels of the zonation and is sheltered mainly in cr'evices and under
boulders. The infralittoral fringe is covered by a coralline with al-
gal turf community similar to those described in warm seas without
corals and in temperate seas. At this level as well, in the infralit-
toral fringe, a gradient in biofacies related to water agitation and
temperature is found. For example the Blossevillea fringe on open coasts
corresponds to kelps and laminarians in temperate waters while Caulerpa
mats and mangroves in sheltered areas are tropical characters.
All this demonstrates that the water temperature plays a fundamental

role in niche diversity and isolation within the littoral environment.
Another striking character in the infralittoral zone is the absence
of coral reefs and the scarcity of hermatypic species to the west of
Isabela Island. Coral colonies and beach accumulation of coral sand
are found in the shallow areas of the central part of the archipelago
199
(Houvenaghel, 1973). In such an environment, as measured at Bahia Aca-

demia, carbonate activity values are low, close to, or even less than
those predicted from the aragonite and calcite solubility products al-
though photosynthesis is reducing pC0 2 • Away from sheltered areas where
CO 2 is consumed in large quantities, carbonate dissolution is favored,
and carbonate deposition and coral growth would be prevented for thermo-
dynamic reasons.
The terrestrial life conditions, determined by climate, are also direct-

ly related to surrounding hydrology. We have presented (Houvenaghel,
1974) the correlation between rainfall and local sea temperature and
the year to year influence of the sea temperature in determining the
dryness of the climate. The existence of a broad transition zone laying
between the desert coastal belt and the humid vege,tation zones above
200 m in altitude is due to the yearly shifts of the boundary between
dry and humid climates. All biologic cycles influenced by dryness will
also depend on these fluctuations.
As for the marine environment, the cold waters driven into the Galapa-
gos region by the Equatorial Undercurrent and taking part in upwelling
also directly determine the life conditions in the terrestrial environ-
ment.
The sharp isolation of the Galapgos Islands provided by the upwellings,

and the severe hydrologic, ecologic, and climatologic conditions pro-
duced by these waters, are the leading factors promoting species isola-
tion, endemism, and evolution to such an extent that nowadays Galapagos
remains one of the clearest showcases for biologic evolution.
Acknowledgments. This expedition to the Galapagos Islands was financially and logis-
tically supported by the Belgian Ministery of Education, the Charles Darwin Founda-
tion, and the Free University of Brussels; we gratefully acknowledge their help.
References
Abbott, D.P.: Factors influencing the zoogeographic affinities of Galapagos inshore

marine fauna. In: The Galapagos. Bowman, R. (ed.). Berkely and Los Angeles: Univ.
California Press, 1966, pp. 108-122
Barber, R.T., Norton, J.: Hydrography of Bahia Darwin, Isla Genovesa. stanford Oce-
anog. Exped • .!.2., 78-84 (1968) (Unpubl. manuscript)
Barber, R.T., Ryther, J.H.: Organic chelators: factors affecting primary production
in the Cromwell Current upwelling. J. Exp. Marine BioI. Ecol. 3, 191-199 (1969)
Cromwell, T.: Thermocline topography, horizontal currents and "ridging" in the eastern
tropical Pacific. Bull. Int. Am. Trop. Tuna Comm. 3, 135-144 (1958)
Goering, J.J., Dugdale, R.C.: Denitrification rates in an island bay in the equatorial
Pacific Ocean. Science 154, 505-506 (1966)
Houvenaghel, G.T.: Contribution a l'etude de l'ecologie marine des lIes Galapagos.
Mem. Acad. r. Sci. d'outre mer Cl. Sci. Natl. Med. (N.S.) 14, 1-102 (1973)
Houvenaghel, G.T.: Equatorial Undercurrent and climate in the-Galapagos Islands.
Nature (London) 250, 565-566 (1974)
Houvenaghel, G.T., Houvenaghel, N.: Aspects ecologiques de la zonation intertidale
sur les cotes rocheuses des lIes Galapagos. Marine BioI. 26, 135-152 (1974)
Love, C.M. (ed.): Eastropac Atlas, Circular 330, Washington-n.C.: U.S. Dep. Commerce,
Natl. Marine Fisheries Service, 1970-1972
Montgomery, R.B., Stroup, E.D.: Equatorial Waters and Currents at 1500 West, in July-
August 1952. Baltimore: John Hopkins Press, 1962
Pak, H., Zaneveld, J.R.: The Cromwell Current on the East side of the Galapagos Is-
lands. J. Geophys. Res. 78, 7845-7859 (1973)
200
Panfilova, S.G.: Seasonal variations of the surface temperature in the Pacific Ocean.
Okeanologiia ~I 801-806 (1968)
Sibert, J.: Some oceanographic observations in the Galapagos Islands. Am. zool. 11,
405-408 (1971)
Strickland, J.D.H., Parsons, T.R.: A manual of sea water analysis. Bull. Fish. Res.
Bd. Canada 125, 1-185 (1960)
Taft, B.A., Jones, J.H.: Measurements of the Equatorial Undercurrent in the Eastern
Pacific. In: Progress in Oceanography. Warren, B.A. (ed.). Oxford: Pergamon Press,
1973, Vol. 6, pp. 47-110
Tranter, D.J., Smith, P.E.: Filtration performance. In: Zooplankton sampling. Tranter,
D.J. (ed.). Paris: Unesco, 1968, pp. 27-56
Wiebe, P., Cox, J., Malone, T.: A hydrographic survey of Darwin Bay. Stanford Oceanog.
Exped. 19, 44-56 (1968) (Unpubl. manuscript)
Wooster, W.S., Hedgpeth, J.W.: The oceano~aphic setting of the Galapagos. In: The
Galapagos. Bowman, R.I. (ed.). Berkeley and Los Angeles: Univ. California Press,
1966, pp. 100-107
Wyrtki, K.: The annual and semiannual variation of sea surface temperature in the
North Pacific Ocean. Limnol. Oceanog. 10, 307-313 (1965a)
Wyrtki, K.: The thermal structure of the eastern Pacific Ocean. Deutsche Hydrogr. Z.
Erganzungsheft, 1-84 (1965b)
Wyrtki, K.: Oceanography of the Eastern Equatorial Pacific Ocean. In: Oceanography
and Marine Biology. Barnes, H. (ed.). London: Allen & Unwin, 1966, Vol. !, pp.
33-68
Yentsch, C.S., Hebard, J.F.: A gauge for determining plankton volume by the mercury
immersion method. J. Cons. Perm. Int. Explor. Mer 22, 184-190 (1956)
Zuta, Sr, Guillen, 0.: Oceanographia de las aguas costeras del Peru. Bol. Inst. Mar.
Peru~, 157-324 (1970)
Hydrographical Aspects of Upwelling Regions
Role of Mixing in the Dynamics of Upwelling Systems
J.D. THOMPSON
Observational data and theoretical models of coastal and equatorial

upwelling have improved dramatically in recent years. Nevertheless,
parameterization of turbulent mixing processes in upwelling regimes
remains a serious, unsolved problem. In this contribution the role of
mixing in the dynamics of upwelling systems is examined. An historical
perspective is provided via a review of early parameterizations of
horizontal and vertical mixing in upwelling models, beginning with
Ekman (1905). Recent ananlytical and numerical models of upwelling
systems are examined in view of their inadequate mixing recipes. Some
attempts to quantify the role of mixing in upwelling regimes by ob-
servational ptudies are described.
Numerical model simulations of coastal upwelling off Oregon during

August, 1973 are compared with a portion of the Van Leer Cyclosonde
data for the same period. Results suggest that greater interaction be-
tween theoreticians and observationalists is essential for improving
simulations of actual upwelling systems.
1. Introduction
My lecture topic for the Symposium is "The Role of Mixing in the Dyna-
mics of Upwelling Systems." In preparing this presentation I found it
difficult to settle on a working definition for the term "mixing" or
to set the spatial and temporal scales over which i t should apply. Un-
fortunately, there is some truth to the cliche that "one scientist's
signal is another scientist's noise."
Let us instead use the term "turbulent mixing" and recognize that tur-
bulent motions transfer (or mix) momentum, kinetic energy, vorticity,
or a scalar tracer at rates generally several orders of magnitude great-
er than the rates due to molecular diffusion. Let us further recognize
that oceanic turbulence is anisotropic due to the imposed geometry of
ocean basins and the effects of stratification. The effects of turbu-
lent mixing having horizontal scales less than the baroclinic radius
of deformation and vertical scales as large as the depth of the water
column, will be the focus of this lecture. Turbulent time scales will
extend to the seasonal. Formally, let us use Hinze's (1959) definition
of turbulence as "an irregular condition of flow in which the various
quantities show a random variation with time and space coordinates, so
that statistically distinct average values can be discerned."
Despite the emphasis on smaller horizontal eddy scales, I wish to avoid

the implication that geostrophic turbulence or eddy motions on scales
larger than the baroclinic radius of deformation are dynamically un-
important. Certainly, mixing produced by alongshore topographic varia-
tions (capes or canyons), spatial variations in the wind field, or
baroclinic instability can be important in the horizontal and vertical
exchange of heat, salt, momentum, and nutrients in upwelling regimes.
204
Almost certainly these scales are an energy source for higher wavenumber
eddies. Some mention of these larger-scale mixing processes will be
made during the lecture.
2. Coastal Upwelling Theories: An Overview
Time and/or space averaging of the Navier-Stokes equations gives rise

to an eddy (or Reynolds) stress tensor which represents the effect of
turbulent velocity correlations on the mean flow. Unfortunately, the
resulting double correlation (covariance) functions must be expressed
in terms of triple correlation functions, which give rise to quadruple
correlation functions, and so on. This so-called closure problem is
often "solved" by a priori relating the correlation functions to the
mean flow. The simplest (and most tempting) approach is to relate the
velocity covariance functions to the mean flow gradient using postulated
"eddy diffusivities," in analogy to the simple Fickian form for molec-
ular viscosity. The value of the "diffusion coefficient" may vary in
space and time and may be substantially different for different oceanic
variables. Unfortunately, in using this analogy one may begin to think
of turbulence as a property of the fluid, rather than a property of
the flow. Nevertheless, the temptation is great, since i t is mathemat-
ically much simpler to parameterize these eddy stresses through pos-
tulated diffusivities than to introduce more complicated closure schemes.
The molecular analogy is then easily extended to passive scalar equa-
tions for heat, salt, nutrients, etc.
In theoretical meteorology and oceanography the "eddy diffusion" clo-

sure hypothesis has been used extensively - and with some success. How-
ever, as theoretical models have been pushed to account more accurately
for observed features of the ocean and atmosphere, the limitations and
errors of these simple parameterizations have become clearer. A sig-
nificant portion of this lecture will be addressed to how turbulent
mixing processes have been parameterized in theoretical upwelling
models, what dynamic effect they exert, and to what extent they are
valid representations of the physics, as best can be determined from
observations. A more extensive review of dynamical upwelling models
can be found in O'Brien (1975).
The first parameterization of vertical turbulence in an analytical up-

welling model is well known. We need not dwell on the Ekman-Sverdrup
model, but it is most useful to employ i t as a point of reference. If
one considers the simplest possible case of a steady, infinite, homo-
geneous ocean driven by a steady uniform wind and postulates that the
pincipal momentum balance is between Coriolis accelerations and vertical
"friction," then the simple Ekman (1905) model obtains:
fv -F x ' (1)
fu Fy '
where u and v are the eastward and northward-directed currents re-

spectively in a right-handed Cartesian coordinate system, f is the
Coriolis parameter, and Fx and Fy are turbulent shear stresses. In Ek-
man's model the turbulent stresses are parameterized in terms of a ver-
tical eddy viscosity and the velocity gradient:
fv = -1 d (A d u) (2)
P dZ v az '
205
fu = J...p ~
3z (A v ~)
az . (2 )
For constant eddy viscosity this is exactly the molecular analogy. Ob-
servational values for Av vary by several orders of magnitude, but
generally range from 100 cm 2 sec- 1 in the upper ocean to 1 cm 2 sec- 1
in the deep ocean. The "Ekman spiral" for the horizontal velocity vec-
tor resulting from Equation (2) and appropriate boundary conditions
is perhaps the best known diagram in meteorology and oceanography.
The Ekman theory provides one important length scale to the mixing
problem, the depth of frictional influence Dv = (Av/ f ) 1/2. Often this
depth is confused with the mixed layer depth. On specifying the equi-
valence of the wind stress and the Reynolds shear stress at the sea
surface and the existence of an ocean depth where velocity and shear
stress vanish, then the x- and y-directed mass transports are imme-
diately determined.
The beauty and utility of transport theory is the independence of the

transports on the detailed specification of the vertical eddy viscosity.
Ekman theory as modified by Sverdrup (1938) remains the cornerstone
of wind-driven coastal upwelling theory. The offshore mass transport
away from the coast, determined by the ratio of longshore wind stress
to the Coriolis acceleration, is one of the most widely cited parameters
in coastal upwelling theory.
The principal advantage of a mass transport upwelling theory, being

independent of details of the vertical structure of velocity, density,
and turbulence, is also a distinct limitation. The theory is by defini-
tion useless for predicting velocity profiles with depth, upwelling
distributions, or details of an upwelling circulation. While mass tran-
sport theory has contributed significantly to studies of large-scale
ocean circulations, it has been of limited use to the modeler of coastal
upwelling ecosystems.
In addressing this limitation Hidaka (1954) attempted to extend the

Ekman-Sverdrup model. He introduced a horizontal friction term in the
momentum equations having a form identical to Ekman's vertical turbu-
lence term. Hidaka's steady, homogeneous, infinitely deep coastal up-
welling model produced an offshore transport in the familiar vertical
Ekman layer fed by upwelling in a zone of horizontal extent determined
by the horizontal eddy viscosity, analogous to the vertical Ekman depth.
For horizontal eddy viscosities of order 10 8 cm 2 s-l the upwelling
width scale is of order 10 km at mid-latitudes. Hidaka also showed that
the ratio of vertical motion to horizontal motion in a frictionally
dominated homogeneous coastal upwelling regime was equal to the square
root of the ratio of vertical to horizontal eddy viscosity.
It is sobering to note that the state-of-the-art in upwelling dynamics

just 20 years ago was a steady homogeneous model, totally dependent
on a parameterization of turbulent mixing by constant eddy viscosities.
In Hidaka's model, assuming a known wind stress, all length and veloc-
ity scales are determined once the eddy viscosities are selected.
A number of substantial papers on homogeneous coastal upwelling circula-

tions have appeared since publication of Hidaka's original work. While
details very considerably between papers, a common feature is a strong
bottom Ekman layer required to feed the offshore Ekman flux. This
strong bottom Ekman layer is rarely found in coastal upwelling observa-
tions, though admittedly the data are sparse. Garvine (1971) clearly
showed that in a steady, homogeneous model of coastal upwelling on an
f-plane the bottom Ekman layer could be eliminated by introduction of
a north-south barotropic pressure gradient. Unfortunately, the vertical
206
and horizontal lengths scales of the upwelling circulation are still

determined by the horizontal and vertical eddy viscosities. It appears
that the unsatisfying necessity of relying on uncertain turbulent mix-
ing processes in steady homogeneous upwelling models is unavoidable.
If that be true, then we must be very pessimistic about using such
models to progress beyond a qualitative upwelling theory.
Clearly there are alternatives to steady homogeneous upwelling models.

Observations indicate that coastal upwelling is a time-dependent pro-
cess characterized by stratification. One logically asks if the role
of turbulent mixing processes in upwelling systems might be reduced in
highly stratified regimes. Yoshida's 1955 and 1967 papers were the first
to present an analysis of upwelling for the simplest possible strati-
fied model that retains a portion of the transient dynamics. Perhaps
the most important result of Yoshida's investigations was the intro-
duction of a new length scale for the width of the upwelling zone -
the baroclinic radius of deformation. This scale does not depend in
any crucial manner on the value of horizontal eddy viscosity. Physical-
ly, the baroclinic deformation scale is the distance an internal gravity
wave propagates in an inertial period. In an upwelling region at mid-
latitudes, say for the Oregon case, that length scale is 0 (10 km) .
Yoshida introduced the possibility that the upwelling width scale,

during spin-up at least, does not depend on turbulent mixing. The ratio
of the width scales of the horizontal diffusive boundary layer to the
baroclinic radius then becomes quite important. A small value for the
ratio lessens our dependence on turbulent mixing recipes. At the moment
we choose to ignore the obvious complications of forced topographic
scales and the time dependence of the stratification. Recall that the
baroclinic radius is proportional to the square root of the stratifica-
tion.
Yoshida demonstrated the usefulness of hydrostatic, layered models in

upwelling studies. Briefly, their advantages are related to their sim-
plicity and ease of interpretation. A host of researchers (for example,
O'Brien and Hurlburt, 1972; Hurlburt and Thompson, 1973; Peffley and
O'Brien, 1976) have significantly extended Yoshida's two-layered hydro-
dynamic upwelling models to obtain new information about upwelling cir-
culations. The equatorward surface jet, the poleward undercurrent,
shelf-induced upwelling, and the effects of coastline irregularities
and bottom topography has been investigated using the layered approach.
Significant progress has been made in understanding the spin-up pro-
cesses associated with coastal upwelling using these models. However,
since these hydrodynamic models allow no mixing or entrainment between
layers, they become invalid when the internal interface, which models
the seasonal pycnocline, breaks the surface. Physically, the limitation
of these models involves the assumption that the time scale for thermo-
dynamic mixing is long compared to the upwelling time scale of a few
days. There is no interesting steady state in these models. The time
scale for interface surfacing with reasonable wind stress values and
upper layer depth is a few days.
One major contribution of the layered hydrodynamic models is the pre-

diction of an equatorward surface jet occurring within the baroclinic
radius of deformation. This feature, now commonly reported in upwelling
observations, can be explained via a potential vorticity argument for
a two-layer inviscid fluid (Charney, 1955; O'Brien and Hurlburt, 1972).
However, if horizontal turbulent mixing is large, with eddy viscosities
0(10 8 cm 2 s-l) or larger, the inviscid potential vorticity approximation
is invalid, and the surface jet is swallowed up in the horizontal dif-
fusive boundary layer.
207
Yoshida attempted to extend the time scale of validity of his layered

model by introducing a vertical mixing process in which mass transfer
through the interface was proportional to the deviation of the inter-
face from its initial state. Using this approach, Yoshida arrived at
a steady state where the dynamic balance was between vertical advection
of heat and turbulent mixing. While the physical rationale for Yoshida's
parameterization of vertical mixing is not entirely clear, the neces-
sity for doing so is. The correct magnitudes of the vertical and hori-
zontal velocities cannot be obtained in the absence of a vertical tur-
bulent mixing effect in Yoshida's model!
The layered models appear well adapted for the study of transient up-
welling dynamics on short time scales. Their lack of detail in the
vertical dimension is a significant deficiency. When one attempts to
extend their time scales, turbulence mixing must be parameterized as
in the homogenous models, and we are back to the original dilemma -
how to avoid such parameterizations in the face of our gross ignorance
about turbulent processes.
Unfortunately, the continuously stratified models of coastal upwelling

have met with the same fate as the layered models in dealing with tur-
bulent mixing', Although considerable difficulties have arisen in model-
ing the coastal corner region, models of coastal upwelling in a con-
tinuously stratified ocean have been constructed. In several of these
papers (Allen, 1973; Pietrafesa, 1973; Pedlosky, 1974) significant
qualitative agreements with some recent upwelling observations have
been presented. For example, in Allen's 1973 study of spin-up to steady
state, a reasonable baroclinic coastal surface jet was obtained. Ped-
losky found a poleward undercurrent in his steady model. Pietrafesa
demonstrated how one- or two-celled upwelling circulations are induced
depending upon the relative importance of stratification, diffusion,
and density advection.
Allen's f-plane analysis uncovered three relevant time scales for coast-
al upwelling. Each time scale involves a horizontal or vertical eddy
viscosity. The formation of the coastal jet is initiated on the shortest
time scale, while the diffusive adjustment of the coastal current to
a steady state occurs on a time scale comparable to the length of the
upwelling season. Only for time scales shorter than approximately five
days can inviscid dynamics by employed.
From these papers and others (Durance and Johnson, 1970; Hsueh and
Kenney, 1972; Blumsack, 1972) one concludes that for time scales longer
than a few inertial periods, theoreticians have found it absolutely
necessary to include some simple parameterization of turbulent mixing
processes in their models. Ominously, Tomczak (1973) has shown that
the type of upwelling circulation obtained in these models depends
strongly on the ratio of horizontal to vertical Prandtl numbers as-
sociated with turbulent exchanges of properties. Tomczak concludes
that since all eddy or diffusion coefficients are probably not con-
stan ts, "... and since only little is known about their order of mag-
nitude and their dependence on density, current shear, etc., a theory,
the results of which depend critically on these coefficients, can only
be a qualitative one. The more coefficients are involved, the more this
is true."
The impasse we face in coastal upwelling theory also exists in equato-

rial upwelling theory. As Yoshida (1959) and Gill (1971) have noted,
there exists a close relationship between many problems associated with
the equatorial boundary layer and problems associated with coastal
dynamics. The historical development of equatorial upwelling models
closely parallels that of coastal upwelling models in their dependence
208
on turbulent exchange processes. Gill suggests that equatorial circula-

tion can only be accurately simulated when a suitable parameterization
of turbulent processes is introduced. Philander (1973) in an excellent
review article on theoretical models of equatorial flows, noted that
appropriate scaling for equatorial flow is dependent on the choice of
vertical eddy viscosity as well as the vertical distribution of the
parameter. In this regard, Robinson (1966) developed a model in which
the eddy viscosity was a function of Richardson number and found that
the model produced more realistic velocity profiles than did the con-
stant eddy viscosity model. Observations by Jones (1973) indicating
that the Richardson number is a key dynamic parameter in equatorial
flows suggests that field observations could be well employed to find
a suitable empirical relation for relating vertical turbulent mixing
to Richardson number. We will return to this point later.
3. Recent Observations: More Questions Than Answers
Theoretically, turbulent mixing processes must be included in upwelling

models, MatQematically, the qualitative and quantitative nature of the
modeled circulations crucially depends on how such mixing processes
are parameterized. Can one attack these seemingly insurmountable prob-
lems by using clues from the observations? Is it possible, for example,
to describe the upwelling circulation as a system with relatively small
"background" mixing, but with limited regions of intense mixing such as
in frontal zones or very near the coastal boundary? Can we consider
the alongshore flow to be essentially inviscid and geostrophic, and
only concern ourselves with mixing in the onshore-offshore component?
Are our present parameterization schemes using constant "eddy diffusion"
terms hopelessly invalid? How can we improve the "realism" of our models
without making them analytically intractable at best, and at worst,
impractical for economic computer integrations?
Considering the dependence of theoretically predicted upwelling circu-
lations on turbulent mixing processes, it is frustrating to discover
the paucity of observational papers on the problem. Reports from recent
large-scale field programs [CUE-I (1972) and CUE-II (1973) off Oregon;
JOINT-I (1974) off Northwest Africa; JOINT-II (1976) off Peru] are im-
proving our description of coastal upwelling dynamics, but so far we
can offer only tentative answers to the questions raised above. New
instrumentation and improved observing techniques give us hope that
more definitive answers can be expected in this decade.
Vertical mixing of momentum in upwelling models has generally been
characterized by a constant vertical eddy viscosity term or, in the
case of layered models, an interfacial stress as a function of veloc-
ity shear. The observations suggest a much more complicated vertical
mixing process, more intermittent and spatially inhomogeneous.
Jones (1973) studied the vertical turbulent mixing process in the
equatorial upwelling area along the equator about 400 miles west of
the Galapagos Islands. On April 1, 1968, during the EASTROPAC Expedi-
tion, velocity shear was measured to a depth of 450 m using current
meters separated by a 10-m vertical interval. The Vaisala frequency
was determined from the values of density averaged over the distance
separating the two current meters. The Richardson number was then cal-
culated from the square of the ratio of Vaisala frequency and the shear
determined from the two current meters. The entire up-down sampling
process required almost six hours.
209
Q Fig. 1. Shear, Vaisala frequency

and Richardson number calculated
at 10 m spacing from the observa-
co
1:E tions. Data obtained on the way
down represented by solid circles,
::I
c <0 open circles on the way up. The
r::
0
dashed line on the right-hand
'" panel indicates the value Ri =
"E .".
'"
.r::.
<.)
1/4 (from Jones, 1973)
[}'
N
'7
~
7Q
~ ..,
~
'" "0
-
~O~__~__~____L -__~__~____~__- L__~~__~
-0 ~ ~ g §
(w) 41daO
Figure 1 is a presentation of shear, Vaisala frequency, and Richardson
number calculated from Jones' observations. Near the 35-me depth, where
the horizontal velocity maximum occurs, there is a minimum in shear
and a corresponding maximum in Richardson number. Note that below 125 m
to 300 m the Richardson number was less than 1. Between 190 m and 270 m
it was less than 1/4, the customary value below which transition from
laminar to turbulent flow is thought to occur.
Based on laboratory evidence vertical eddy viscosity can be estimated

from rather simple recipes involving Richardson number and friction
velocity. These recipes are derived from empirical studies of turbulence
in a thermally stratified fluid. Jones employed the Monin and Yaglom
(1971) formula:
ku*z
(3)
Av = 1+aRi'
where k is Von Karman's constant, z is the separation distance between

the two current meters, a is an empirical constant, and Ri is Richardson
number. The friction velocity is given by
u = kllu (4)
* ln (z/L)
210
Table 1. Observed values of shear, depth, and stability as calculated by Jones for
STD station 552 and current meter station I (00 07'S, 97 0 40'W) on April 1, 1968.
Calculations of Richardson number, U*, and K are described in the text (from Jones,
1973)
Depth Shear Stability Richardson U* K

(m) (s-l x lO- 2 ) (s-l x lO- 2 ) number (cm s-l) (cm Z s-l)
25.6 3.57 2.24 0.39 1.55 210.09

35.0 0.36 1.98 30.88 0.16 0.40
40.1 1.14 1. 35 1.41 0.49 24.58
50.6 1.65 1.53 0.86 0.72 54.05
55.4 1.67 1.05 0.40 0.72 96.64
65.3 1.13 1.22 1.17 0.49 28.64
75.2 0.75 0.59 0.61 0.33 32.15
84.6 0.25 0.46 3.30 0.11 2.48
94.4 0.81 0.75 0.86 0.35 26.53
104.3 0.65 0.65 1.02 0.28 18.50
113 .9 0.70 0.64 0.83 0.30 23.60
123.5 0.70 0.67 0.90 0.30 22.09
132.9 0.64 0.39 0.38 0.28 38.31
141.9 0.69 0.44 0.40 0.30 39.93
150.8 0.65 0.44 0.47 0.28 33.68
161.0 0.94 0.43 0.21 0.41 79.60
169.7 0.70 0.37 0.28 0.30 50.63
178.7 0.76 0.46 0.37 0.33 46.29
186.8 0.37 0.37 1.00 0.16 10.71
196.4 0.65 0.18 0.08 0.28 80.60
203.9 0.94 0.20 0.05 0.41 130.55
210.8 0.67 0.25 0.14 0.29 68.42
218.8 0.69 0.15 0.05 0.30 95.83
229.3 0.85 0.27 0.10 0.37 98.37
237.2 1.00 0.31 0.10 0.43 115.73
245.3 1.61 0.43 0.07 0.70 207.03
252.2 2.10 0.65 0.10 0.91 243.04
257.6 2.30 0.53 0.05 1.00 319.42
263.5 1.87 0.57 0.09 0.81 223.88
271.6 1.28 0.62 0.24 0.56 101.00
284.0 1.27 0.82 0.42 0.55 71 .12
295.7 1.89 0.91 0.23 0.82 152.61
where L is the Kolmogorov length scale and ~u is the observed shear.

Table 1, from Jones' article, provides estimates of the vertical eddy
viscosity at 10-m depth intervals using Equations (3) and (4). The
values are remarkable in several respects: (1) They vary over two orders
of magnitude between 25 m and 300 m. (2) They do not have an exponential
or even monotonic decrease with depth. (3) The largest eddy viscosity
is found near 250 m. The inadequacy of a constant vertical eddy viscos-
ity for this problem is obvious. This pOint was emphasized by Robinson
(1966) in his attempt to model the Euqatorial Undercurrent.
Clearly, even at depth, beneath the core of the Undercurrent, vigorous

vertical mixing can occur in conjunction with equatorial upwelling, as
evidenced by the uniformity of temperature, salinity, and oxygen in
this dynamically unstable layer. Is this local, persistent lack of
dynamic stability common in other upwelling systems, particularly coast-
al upwelling? Does intense vertical mixing often occur in those regions?
Perhaps the best answer to date comes from the work of Van Leer and
colleagues of the University of Miami. Van Leer's automatic profiling
current meter, termed a Cyc~esonde, provides very detailed vertical
profiles of the current and temperature structure in an upwelling zone
211
on time scales from hours to weeks (see Van Leer et al., 1974 for de-
tails of the instrumentation) .
Using such an instrument Johnson et al. (1976) have produced depth-time

sections of north-south and east-west velocity components and temper-
ature during active coastal upwelling. The data from the NE Station
was obtained in August, 1973, during CUE-II 12 km off the Oregon coast
in 100 m of water. The Mark I Cyclesonde had vertical resolution of
about 5 m amd sampled the water column roughly once every 40 min. Using
the instantaneous velocity and temperature measurements and mean salin-
ity profiles obtained from hydrographic sections, the vertical shear,
Vaisala frequency, and Richardson number were calculated. Figure 2 is
CUE-n NE CYCLESONDE STATION

LAT ' 45 0 18.5' N LONG ' 1240 07.5'W
o LOG RICHARDSON NUMBER VALUES <: -0.5 SHADED CONTOUR INTERVAL ' .5(looRI)
20
~40
J:
l-
n.
:g 60
80
100 i I I
o o o
28 AUG 29AUG 30 AUG
Fig. 2. Time-height section of the logarithm of Richardson number calculated from
data obtained by the Van Leer Cyclesonde during August 28-31, 1973, off Oregon (from
Johnson et al., 1976)
the resulting depth-time section for the logarithm of the Richardson

number. The shaded area corresponds to values of Richardson number less
than about 0.3, or near the so-called critical Richardson number of
0.25. Note that subcritical Richardson numberS occurred in a depth zone
of 40-60 m, corresponding to the base of the permanent pycnocline. The
temporal variation in the fraction of the water column having subcriti-
cal Richardson number appears to be inertial/semi-diurnal.
Were we to use Jones' technique for estimating vertical eddy viscosity

based on the Cyclesonde data from Oregon, we would find, as in the
equatorial case, that vertical eddy viscosities varied by orders of
magnitude through the column. We would also find that Av has no simple
functional form with depth, although maxima tend to occur at the base
of the permanent pycnocline. It is interesting to note that there oc-
curred a rapid thickening of the subcritical Richardson number layer
with the onset of strong winds after 1200 GMT, August 28. While other
Cyclesonde stations did not show such clear-cut low stability modula-
tion, it does appear from these observations thcit vertical mixing is
very important in mid-latitude coastal upwelling. Additional observa-
212
tional evidence (Halpern, 1976; Mooers et al., 1976) tends to support

the Cyclesonde results. The observations indicate that vertical mixing
in upwelling zones is highly intermittent, spatially inhomogeneous,
and dynamically important on time scales from days to seasons. Any up-
welling model which attempts to parameterize vertical mixing processes
must account for these facts.
Although vertical mixing processes appear to be complicated, perhaps
we can still consider the longshore flow to be inviscid and geostrophic,
and only concern ourselves with vertical mixing in the onshore-off-
shore direction. For example, Mooers et al. (1976), using current and
hydrographic data obtained off Oregon during 1965 and 1966, concluded
that between 20 and 60 m, in water depth of 80 m, 10 kID offshore, the
alongshore current was essentially in geostrophic balance during a two-
week averaging period. However, they found relatively high vertical
shear existed across the frontal layer in both the mean and fluctuating
component of the alongshore flow. They suggested that the frontal layer
may be the site of shear instability and may have characteristics of
an internal, turbulent boundary layer.
Using the Cyclesonde data discussed earlier, Johnson et al. (1976)
concluded that relative to the crossisobath flow the alongisobath flow
was strong, persistent, and nearly geostrophic. They found that the
corssisobath flow was quite weak, variable, and ageostrophic. However,
as in Mooers et al. (1976) they discovered low dynamic stability in
the frontal layer, due to large tidal and near-inertial shear, suggest-
ing the existance of an internal boundary layer and important ageostro-
phic mean flow components.
With the exception of the narrow frontal zones and perhaps thin boundary
layers at the top and bottom of the water column, sparse observations
to date support the hypothesis that the alongshore flow is essentially
geostrophic during active coastal upwelling. Vertical mixing is far
more important to the onshore-offshore momentum balance.
Do observations also indicate that horizontal mixing is intermittent,
and spatially inhomogeneous in upwelling zones? Are horizontal mixing
processes small compared to vertical mixing processes? Can one parame-
terize horizontal momentum mixing in terms of Laplacian friction with
a constant eddy viscosity? Can one consistently estimate the value of
the horizontal eddy viscosity from observations?
Stevenson et al. (1974) attempted to estimate horizontal eddy viscosity
by tracking surface drogues from aircraft during an upwelling event
off Oregon during August, 1972, in the Coastal Upwelling Experiment
(CUE-I). Eighteen surface drogues were deployed on 23 August, along a
line normal to the coast at intervals of 1/3 nautical miles. These
drogues were tracked for roughly 20 h. The main result of the drogue
study was the smooth, nearly monotonic shape of the longshore velocity
component. A longshore equatorward component of the flow clearly domi-
nated the pattern with speeds of 30 cm s-l at the seaward edge. The
velocity profile decayed toward the shore with longshore speeds de-
creasing from 30 cm s-l to 2 cm s-l from seaward to shoreward edge,
respectively. Using a uniform eddy viscosity hypothesis, Stevenson
et al. (1976) made a tentative estimate of the horizontal eddy vis-
cosity of 2 x 10 7 cm2 s-l. The corrresponding boundary layer thickness
was roughly 19 km. Unfortunately, this experiment has not been repeated
for various phases of an upwelling event off Oregon. We would require
a great number of these experiments, off Oregon and in other upwelling
regions, to feel confident about using such an estimate.
213
Mooers et al. (1976) have made a few exploratory estimates of horizontal

eddy viscosity coefficients based on direct current observations. They
examined current data taken at 10-min intervals for two current meter
time series at 60 m depth, 10 and 30 km offshore in water 80 and 200 m
deep. They calculated the uv cospectrum from this data set in two ways:
(1) using 30 days of low passed data (data filtered with a low pass
filter with a half power point of 40 h) and (2) 40 days of hourly aver-
ages of the 10 min data. In the former case, contributions to the Rey-
nolds stress spectrum came from time scales longer than two days. The
latter case incorporates those contributions on time scales longer than
one hour.
The low passed results indicated a value for the horizontal eddy vis-
cosity of 2 to 5 x 10 5 cm 2 s-l between the coast and the shelf edge.
Thus the energy flux was down the mean momentum gradient. However, they
found that for the hourly results, AH was negative (-10 6 to -10 7 cm 2
s-l) between coast and shelf edge, with energy flux up the mean momentum
gradient. Their preliminary results seem to indicate that inertial
internal waves having periods between one hour and two days contribute
to the mean flow. The authors caution against drawing any firm con-
clusions from these results considering the limited number of current
records available and the difficulties in defining the mean motion in
a regime where the time-variable signal is as large as the mean motion.
On larger space and time scales baroclinic instability may be important
for horizontal mixing in an upwelling zone. Evidence presented by Mooers
et al. (1976) shows that, off Oregon, although the winds continue to be
favorable for upwelling for more than a week, the inclined frontal layer
does not continue to propagate further offshore. This suggests that a
strong dissipative process such as baroclinic instability might be im-
portant. Holladay and O'Brien (1975), using sea surface temperature
variance spectra, also find evidence for strong baroclinic excitation
in the Oregon upwelling region. It is not yet clear how significant
baroclinic instability might be to horizontal mixing processes in coast-
al upwelling areas, or whether geostrophic turbulence theory might be
useful in parameterizing the process.
In summary, the limited observations of upwelling systems indicate that
turbulent mixing is dynamically important to the onshore-offshore circu-
lation on time scales as short as a few hours. The alongshore flow may
be considered largely geostrophic and relatively inviscid except per-
haps in frontal layers, very near the coastal boundary in a viscous
boundary layer, and in thin Ekman layers at the top and bottom of the
water column. Given the intermittency and spatial inhomogeneity of mix-
ing in upwelling systems, our present parameterizations using constant
eddy diffusion terms must be deemed inadequate. Vertical mixing proces-
ses in upwelling models should be related to the dynamic stability of
the water column, with the possible inclusion of a small background
eddy diffusion. Horizontal mixing processes are not well observed and
are perhaps less important than vertical mixing. Background horizontal
eddy viscosity is likely to be less than 10 8 cm 2 s-l on time scales of
a few days to a week. On shorter time scales the inertial-internal mo-
tions may actually contribute to the mean current and completely in-
validate the eddy viscosity formalism. Observationally the difficulty
in determining the mass balances in the onshore-offshore and longshore
directions leads to large uncertainties in the details of the upwelling
circulation. Further, the observational uncertainty in current and
hydrographic measurement makes accurate determination of mixing pro-
cesses extremely difficult. Unfortunately, the present theoretical cir-
culation models are highly s'ensi ti ve to small changes in the mixing
parameters.
214
4. A Simple Mixing Model: Formulation and Comparison with Observations
Scientists have generally been unable to make or verify quantitative

predictions of actual upwelling circulations using theoretical models.
Observationalists tend to suspect a model's practical value and the-
oreticians comfort themselves with· even gross agreement between model
and observations. With few exceptions, theoretical models of upwelling
regimes have not adequately guided design of field experiments nor
have the observations significantly improved model design. This is
hardly the climate for further understanding of upwelling dynamics in
general, or the effects of mixing on dynamics in particular.
What is needed is unprecedented cooperation between observationalist

and theoretician. As the observational data base improves in both quan-
tity and quality, it may become possible to verify a numerical simula-
tion of an upwelling circulation, driven by observed local and/or remote
forcing, using observed currents and hydrography for verification. This
approach, recently applied to the Simpler simulation of lake circula-
tions (Simons, 1976), has long been used in numerical weather predic-
tion, though we are far from having sufficient observational coverage
of either the upwelling circulation or the wind forcing to overly con-
cern ourselves with predictibility limits such as those encountered
in weather forecasting. Nevertheless, we can begin to hindcast our mo-
dels, forcing and verifying them with the increasingly dense observa-
tions. Such hindcasts should prove useful in hypothesis testing as well
as in experimental design. The idea of testing model predictions using
a wide range of turbulence closure schemes and verifying those pre-
dictions against a standard data set is extremely exciting. Defant
(1961) may be right that "a totally satisfying explanation of upwelling
is probably not possible at all since the total process is composed
of a number of substages each of which is controlled by other factors."
Yet one wonders if observations and simulation could be employed itera-
tively to attack the parameterization problem Defant alludes to.
A few primitive attempts to follow the procedure outlined above have

recently been made. In the remaining paragraphs of this section I wish
to describe a relatively simple upwelling model that has been applied
to an actual upwelling regime and the results compared with observa-
tions. My purpose is to emphasize the potential for interaction between
theoretician and observationalist. Certainly more detailed upwelling
models could be used in the simulations, but presently little could
be gained from additional complexity.
The simple hydrodynamic two-layer model was extended to include vertical

mixing across the fluid interface. Explicit reconstructions of Ekman
layers at the surface, bottom, and interface were developed using per-
turbations of velocity from the vertical average. High (1 km) grid re-
solution was used with proper ocean interior dynamics (Hurlburt and
Thompson, 1973; Pedlosky, 1974; Garvine, 1974) matched to the upwelling
circulation. For the simulations considered, longshore variations in
velocity and density fields were considered negligible compared to off-
shore variations. The verification period was selected as August, 1973
off Oregon during CUE-II. Model results are compared with the Van Leer
Cyclesonde data for that period.
First consider the model. Assume a stable stratified, rotating, hydro-

static fluid on a continental shelf-slope cross section. Assume the
interface is permeable to heat, mass, and momentum through turbulent
mixing. The vertically integrated momentum equations are (Thompson,
1974):
215
[v • (p IhlAH~) ]~1 gh l +
+ - 2pO Vp 1 + 81 , (5)
POhl
aV2
+
+ ++
+
'r - +
'B
+ V2 • VV2 + k X f~2 g~(hl + h2 + D) + g' ~hl +
at Poh2
[~ • (P2 h 2A ~)] ~2 gh l ~
+ POh2 --po- PI (6)
where
g' g(P2 -Pl)/p2'

P2Ql (~2 - ~Il
(7)
P Ihl
P lQ2 (~2 -~l)
P2 h 2
Vp (P2 -pd/pl.
-; r Cr I~1 - ~2 I (~1 - ~2)

-;B CBI~2 1~2
Density and are predicted by
(8)
(9 )
(P2) t + V2
+
. ~P2 = ( 10)
(h 2 ) t + ~ . (~2h2) ( 11)
where
PI Po - yTl
P2 Po - y T 2·
The subscripts 1 and 2 refer to upper and lower layers. The usual nota-
tion and right-handed coordinate system is used. The stresses are at
the surface (8), interface (I), and bottom (B). The height at the bot-
tom topography above a reverence level is given by D. Ql and Q2 repre-
sent turbulent entrainment rates, where Ql represents entrainment of
lower layer fluid upward into the turbulent upper layer. The effective
thermal expansion coefficient for seawater is represented by y, and
Po is a reference density. The eddy diffusion coefficient for heat is
KH and is AH for momentum. Diabatic effects (solar insolation, back
radiation, etc.) are represented by H.
There are two main reasons for including the details of the model here.
First, it emphasizes that even a "simple" model of upwelling dynamics
is rather complex and difficult to formulate. Second, mixing processes
are now explicitly set down, with horizontal diffusion and vertical
216
entrainment clearly indicated in the equations. Note that subgrid scales

now refer to horizontal space scales less than 1 km, vertical scales
less than the layer thickness, and times scales less than one hour.
We have chosen KH and AH as 10 6 cm 2 s-l for our integrations. From the
laboratory work of Rouse and Dodu (1955), Kato and Phillips (1969),
and others we have specified that the entrainment of non turbulent fluid
into turbulent fluid is related to the bulk Richardson number and can
be expressed as
P ¢ u 3
o
Ql = gllph 1
*
where ¢ is the fraction of wind energy available for turbulent mixing
(Denman, 1973) and u* is simply the friction velocity, (::;-s/po) 1/2. Q2
has a similar form except that hI is replaced by h2 and u* is replaced
by the bottom friction velocity. Tidal mixing has not been included
in the model. This model is distinguished from earlier upwelling models
because it includes a vertical turbulent mixing whose intensity is
tied to the dynamic stability of the water column, and which is self-
determined during model evolution. Further the vertical mixing has been
divided into a "large-scale" entrainment and a "small-scale" diffusion.
Essentially we have identified two dominant scales of turbulent eddies
which accomplish the mixing. The "small-scale" diffusion allows us to
calculate departures from the vertically-averaged velocity field in
each layer under the assumption of hydrostatic balance, small Rossby
number, and quasi-equilibrium conditions. Perturbation equations are
then:
fa~~ a3~'
k x _ _J = LV p. + Av ~z" (12 )
az Po J 0
j=1,2.
The primed quantities are departures from the vertical average within
a layer and the subscript refers to the upper or lower layer. We have
chosen Av to be 50 cm 2 s-l for each layer. The boundary conditions are
->- ->-
av _
A
V az - T
s
, at the sea surface
at the interface (13 )
at the interface
az az
at the bottom
with the integral constraints
upper lower
layer layer
5 ~l dz 5~2 dz = O.
Note that the perturbation velocities can be calculated only if the

averaged flow and density fields are known. In practice these values
are obtained during the course of the integration.
During the CUE~II experiment off Oregon in August, 1973, sufficient

observational data were obtained to attempt a preliminary test of the
preceding numerical model. The wind driving was supplied by the hourly
217
values of Newport winds and wind stress was calculated from the usual
bulk aerodynamic formula. The model was driven from rest on August 1,
1973, with the initial stratification provided from observed hydro-
graphic data. The upper layer thickness was taken to be the undisturbed
depth of the seasonal pycnocline, about 30 m. The total water depth
far offshore was 2500 m. The east-west model extent was 3000 km, with
large-scale wind-stress curl estimated from weather maps. The bottom
topography was smoothed to correspond to the offshore section coincident
with the CUE-II buoy line.
_ -- --i 50
E
100 ..c
! Fig. 3. Longshore current speed

contours off the Oregon coast to
25 km at day 28 of the model in-
tegrations. Dashed lines represent
negative (southward) values. Con-
150
tour intervals are 2 em s-2. The
wind speed and direction are in-
dicated by the ~circle dia-
Radius 1/2 dyne gram. An arrow pointing downward
represents a wind from the north.
25 20 15 10 5 0200 Length of the arrow indicates
Km wind speed
The verification period corresponded to the time of extensive Cycle-

sonde observations from 28 August to 31 August. Figure 3 shows the con-
tour field of longshore current on August 28 as shown by the model.
The wind had just begun to blow from the north at this time. Note the
equatorward surface jet of minimum value <-16 cm s-l 10 km offshore
and the weak poleward undercurrent <2 cm s-l within 15 km of the coast.
The longshore flow was found to be largely geostrophic. The u-w flow
for the same time is depicted by vector plots in Figure 4. Note the
difference in horizontal and vertical scale in the diagram. In this
plot the maximum onshore flow (6 cm s-l) occurs about 5 km offshore
near 40 m depth. Maximum instantaneous vertical flow occurs within
4 km off the coast below 20 m. Note the offshore flow in the surface
Ekman layer and at the base of the pycnocline. There is no simple one-
cell or two-cell circulation evident in this instantaneous picture.
Figure 5 gives the more classical one-cell picture of coastal upwelling

at day 29. However, during other times much more complicated one- and
two-cell circulation were predicted. The cellular nature of the up-
welling circulation depends upon wind stress, stratification, current
shear, and bottom relief. Our results indicate that it is quite un-
reasonable to expect unambiguous evidence for one or more cells in ob-
served current data. It is not surprising that various researchers have
seen one or more cells in their data under varying conditions.
218
u-w Vector plots Elapsed time = 28.00 day Fig. 4. Plot of u-w vector-tra-
" ,.
. . . . 4' . . . . . . . . . . . . . . . . _
. .....
" ,.
........ ,
......... .
-,r'!-...........,...,....,................,-.-.-.-......,........ -.-...,....... .. --.-.- 0
....~ ........... ,
jectory field at day 28 of the
integrations. Note the difference
_-.- _ _ . . . . . . ....... _.".....,. .......... ~.......... lilt' f
in horizontal and vertical scales.
... . . . . . . . . -...... . - . . _ . if if I .,~ .... ~,
Vector lengths are scaled to the
,~ ........... ~,. ... III' fI' , " , II ...... ~ .............................- . ... maximum in the field
•• • ~~~~,.."-,,-,,,..A ... 50
E
100
a
L
o'"
150
25 20 15 10 5
Km
u- W Vector plots
50
E
100 L
Q.
<I>
a
. 150
25 20 15 10 5 Fig. 5. Same as Figure 4 but for

Km day 29 of the integrations
Figure 6 provides a comparison of the model simulation of the current

at the NE Station compared to the observed values. These are averaged
values over the Cyclesonde observation period of about 64 h. Note that
the zonal flow predicted by the model near the surface underestimates
the actual value. One suspects that the underestimate is due to a low
estimate for the wind stress since the wind data were obtained from a
coastal anemometer 50 km away. Effects of wave action on the Cyclesonde
might also be important. The model does rather well in zonal flow pre-
219
,
NE 0 NE V
-3 -2 -1 0 2 -40 -30 -20 -10 0
0 0
"- \ ,,
o~ ) \, ,
.",..,.,~O-
""
<:: ---... 0 ____
_ _0
~ ""\
~
................
~
CI.>
.......
.... 0
....
<F)
CI.> \:; \
a; a; 'n I
'b___ "-
0
E \ E I
..s=
50
\ ..s=
50 ~'n I
I
D.. 0
I
oI
0.. \ I
1----
CI.> CI.>
0 ' 0 t:t I
0 L(\
\ /'0 D.\
r/ / d~
~l,
100 / 100
Fig. 6. Comparison of model simulation and observations at the NE station for the
CUE-II period 2300 GMT to 1300 GMT, August, 1973. Profiles represents averages of
zonal (left) and meridional (right) velocity components over the period. Solid lines
represent observed values, dashed lines represents model prediction
diction down to about 30 m, but it completely misses the strong off-

shore flow at 50 m. There are a number of reasons why this discrepancy
might appear. It is not clear whether it is due to the three-dimen-
sionality of the actual flow, poor vertical resolution of the model,
inadequate turbulence parameterization, or something else. Below 70 m
the model and observations are in fair agreement, even to the maxima
in onshore flow at 80 m in the observations and at 90 m in the model.
The meridional flow is perhaps more useful for verification due to its
large signal as compared to the zonal flow. Once again, near the sur-
face the model underestimates the current strength - indicating too
weak forcing. It is important to note that the vertical shear in the
longshore current is nearly the same for model and observation down to
about 30 m. The model tends to weaken the current at too shallow a
depth. It appears that an additional layer is required by the model
for better simulation. Perhaps of most importance in the comparison
is the absence of a strong subsurface flow. If the model did not have
the proper ocean interior it would have predicted a much larger baro-
tropic mode.
Our comparison may be entirely fortuitous. Perhaps we cannot seriously

begin yet to compare coastal upwelling models with what is actually
observed. Perhaps the observations and the models are still too inade-
quate for quantitative comparisons. Nevertheless, we continue to study
this simulation and the Cyclesonde data. We plan to rerun the model
to undertake additional simulation when the data become available.
Whatever the outcome, the interaction of the theoretical modeller and
the observationalist in this particular research has proved to be both
intructive and exciting.
220
5. Conclusions
Theoretical models of upwelling circulations have advanced at a remark-

able pace during this decade. Unfortunately, simplistic descriptions
of turbulent mixing processes remain a crucial element in the model
formulations. Observational knowledge of upwelling circulations has
markedly increased with the advent of extensive new field programs and
advances in instrumentation technology. The new observations underscore
the inadequacy of our mixing recipes. Although we are able to qualita-
tively understand and reproduce many observed features of upwelling
circulations, a major obstacle to further theoretical advances is the
proper parameterization of the observed intermittent and inhomogeneous
turbulent mixing. Some suggest that in general this may be an impossible
task (Ramage, 1976). Surely it is a difficult one (Kraichnan, 1976).
We have reached a plateau in modeling upwelling systems - to extend

our models beyond qualitative agreement with observations will prove
extremely difficult. If we have any hope for improving our understanding
of upwelling systems using theoretical models it must be based on un-
-precedented cooperation between theoretician and observationalist. Ob-
servations are becoming sufficiently dense in space and time so that
the theoretician can no longer insulate his model from quantitative
verification. While on the short term we may circumvent the turbulence
closure problem by model "tuning," on the long term we must be skeptical
of any theory of upwelling dynamics that does not realistically account
for turbulent mixing in the dynamics of upwelling systems.
Acknowledgments. A portion of this research was undertaken while the author was a
Postdoctoral Fellow in the Advanced Study Program of the National center for Atmo-
spheric Research, Boulder, Colorado. NCAR is sponsored by the National Science Founda-
tion. Support for completion of the study and travel expenses to Kiel were provided
by the Office of Naval Research through a subcontract wi th the Naval Research Lab-
oratory. Appreciation is extended to the Symposium Organizing Committee and to SCOR
for financial assistance during the author's stay in Kiel. This paper is a theoretical
contribution to the Coastal Upwelling Ecosystems Analysis (CUEA) Program, an IDOE-NSF
sponsored project. Computer time was provided on NCAR's Control Data Cooperation
6600 and 7600. The Naval Research Laboratory, Washington, DC, provided technical
assistance in preparation of the manuscript. Appreciation is extended to Mr. Richard
Grotjahn for programming assistance, to Ms. Karon Christensen, Ms. Margaret Mikota,
and Mrs. Cathy Turesko for typing the manuscript, to Dr. Harley Hurlburt for reading
the manuscript, and to Dr. J.J. O'Brien for several helpful suggestions.
References
Allen, J.S.: Upwelling and coastal jets in a continuously stratified ocean. J. Phys.
Oceanogr. 1, 245-257 (1973)
Blumsack, S.L.: The transverse circulation near a coast. J. Phys. Oceanogr. ~, 34-40
(1972)
Charney, J.G.: The generation of oceanic currents by wind. J. Marine Res. li, 477-498
(1955)
Defant, A.: Physical Oceanography. New York: Pergamon Press, 1961, Vol. I, 729 pp.
Denman, K.L.: A time-dependent model of the upper ocean. J. Phys. Oceanogr. 1, 173-184
( 1973)
Durance, J.A., Johnson, J.A.: East coast ocean currents. J. Fluid Mech. 44, 161-172
(1970)
Ekman, V.W.: On the influence of the earth's rotation on ocean currents. Arkiv. Mat.
Astron. Fysik. ~, 1-52 (1905)
221
Garvine, R.W.: A simple model of coastal upwelling dynamics. J. Phys. Oceanogr. l'
169-179 (1971)
Garvine, R.W.: Ocean interiors and coastal upwelling models. J. Phys. Oceanogr. !,
121-125 (1974)
Gill, A.E.: The equatorial current in a homogeneous ocean. Deep-Sea Res. ~, 421-431
(1971)
Halpern, D.: Structure of a coastal upwelling event observed off Oregon during July
1973. Deep-Sea Res. ll, 495-508 (1976)
Hidaka, K.: A contribution to the theory of upwelling and coastal current. Trans.
Am. Geophys. Union ~, 431-444 (1954)
Hinze, J.O.: Turbulence. New York: McGraw-Hill, 1959, 586 pp.
Hsueh, Y., Kenney, R.N.: Steady coastal upwelling in a continuously stratified ocean.
J. Phys. Oceanogr. 2, 27-33 (1972)
Holladay, C.G., O'Brien, J.J.: Mesoscale variability of sea surface temperatures.
J. Phys. Oceanogr. ~, 761-772 (1975)
Hurlburt, H.E., Thompson, J.D.: Coastal upwelling on as-plane. J .. Phys. oceanogr.
1, 16-32 (1973)
Johnson, W.R., Van Leer, J.C., Mooers, C.N.K.: A cyclesonde view of coastal upwelling.
J. Phys. Oceanogr. ~, 556-574 (1976)
Jones, J.H.: Vertical mixing in the euqatorial undercurrent. J. Phys. Oceanogr. 1,
286-296 (1973)
Kato, H., Phillips, O.M.: On the penetration of a turbulent layer into a stratified
fluid. J. Fluid Mech. ]2, 643-655 (1969)
Kraichnan, R.H.: Eddy viscosity in two and three dimensions. J. Atmos. Sci. ll,
1521-1536 (1976)
Monin, A.S., Yaglom, A.M.: Statistical Fluid Mechanics. Cambridge: The MIT Press,
1971, Vol. I, 769 pp.
Mooers, C.N.K., Collins, C.A., Smith, R.L.: The dynamic structure of the frontal
zone in the coastal upwelling region off Oregon. J. Phys. Oceanogr. 6, 3-21 (1976)
O'Brien, J.J.: Numerical models of the ocean. Nat. Acad. Sci. 6, 204-215 (1975)
O'Brien, J.J., Hurlburt, H.E.: A numerical model of coastal upwelling. J. Phys.
Oceanogr. 2, 14-26 (1972)
Pedlosky, J.~ On coastal jets and upwelling in bounded basins. J. Phys. Oceanogr.
!, 3-18 (1974)
Peffley, M.B., O'Brien, J.J.: A three-dimensional simulation of coastal upwelling
off Oregon. J. Phys. Oceanogr. ~, 164-180 (1976)
Philander, D.G.H.: Equatorial undercurrent: Measurements and theories. Rev. Geophys.
Space Phys. 11, 513-570 (1973)
Pietrafesa, L.: Steady Baroclinic Circulation on a Continental Shelf. Ph.D. Disser-
tation. Univ. Washington, Seattle, 1973
Ramage, C.S.: Prognosis for weather forecasting. Bull Am. Meteor. Soc. 57, 4-10
(1976)
Robinson, A.R.: An investigation into the wind as the cause of the equatorial under-
current. J. Mar. Res. 24, 179-204 (1966)
Rouse, H., Dodu, J.: Turbulent diffusion across a density discontinuity. La Houille
Blanche 10, 530-532 (1955)
Simons, T.J.: Verification of numerical models of Lake Ontario III. Long-term heat
transports. J. Phys. Oceanogr. ~, 372-378 (1976)
Stevenson, M.R., Garvine, R.W., Wyatt, B.: Lagrangian measurements in a coastal up-
welling zone off Oregon. J. Phys. Oceanogr. 4, 321-336 (1974)
Sverdrup, H.U.: On the process of upwelling. J~ Marine Res. l' 155-164 (1938)
Thompson, J.D.: The coastal upwelling cycle on a S-plane: Hydrodynamics and thermo-
dynamics. Ph.D. Thesis, the Florida State Univ., 1974
Tomczak, M.: Note on diffusion in coastal upwelling. J. Phys. Oceanogr. 1, 162-165
( 1973)
Van Leer, J., Diiing, W., Erath, R., Kennelly, E., Speidel, A.: The cyclesonde: An
unattended vertical profiler for scaler and vector quantities in the upper ocean.
Deep-Sea Res. 11, 385-400 (1974)
Yoshida, K.: Coastal upwelling off the California coast. Rec. Ocn. Works in Japan
2, 8-20 (1955)
222
Yoshida, K.: A theory of the Cromwell current and of the equatorial upwelling. J.
Oceanogr. Soc. Japan 15, 154-170 (1959)
Yoshida, K.: Circulation in the eastern tropical oceans with special reference to
upwelling and undercurrent. Japan J. Geophys. i, 1-75 (1967)
The Circulation of Large Lakes
IRBENNETI
Highlights of the last two decades of research on the circulation of

large lakes and its implications for the study of oceanic coastal up-
welling are reviewed. First, progress in understanding the effects of
rotation and nonlinearity on the internal seiches of lakes is traced
from the detection of small geostrophic effects to evidence for non-
linear effects on internal Kelvin waves. Second, models of thermal
circulation of lakes are discussed. Third, Birchfield's recent theory
of wind-driven flow in a homogeneous lake is described and numerically
evaluated and an attempt is made to assess the strength of the Taylor-
Proudman constraint in confining the deep waters of the Great Lakes.
1. Introduction
The analogy between currents on continental shelves and in large lakes

is intriguing because many observations which are not understood appear
to be very similar. For example, in summer the mean current near the
north shore of Lake Ontario is towards the west - opposite the wind
(Blanton, 1974, 1975; Pickett and Richards, 1975). There are no models
at present which explain this. A very similar discrepancy between
theory and observation exists on both the East and west continental
shelves of the United States. On the West Coast there is a poleward
undercurrent during the upwelling season (Mooers et al., 1976) and on
the East Coast both the surface and bottom drift are against the mean
wind (Bumpus, 1973).
Because lakes are physically simpler than oceans many attempts to use
simulated lake currents to predict biological and chemical parameters
are now being made. This ambitious task, given the present pressure
for answers to environmental questions, will undoubtedly be repeated
for continental shelves. However, the main applications of environ-
mental modeling have been to rivers, reservoirs, and harbors. These
small scale water bodies do not have the complicated vertical circu-
lations of large lakes and continental shelves.
Thus this essay has two purposes. I want to assess both our ability
to quantitatively explain vertical motion in the Great Lakes and our
ability to simulate i t using numerical models. This is not a trivial
distinction; there are many observations which we think we understand
but which we cannot duplicate by numerical simulation because of math-
ematical difficulties. To a large extent, however, numerical modeling
is the medium used to transfer our understanding of lake circulation
to other water bodies. This is particularly true for basins of com-
plicated geometry such as the Baltic Sea (Kielmann, pers. corom., 1975)
and where observations are sparse or difficult to obtain such as in
Tomczak jr. and Diaz (1975). Although even crude models may help plan
field work by suggesting which measurements are most crucial to make,
i t is clear that it is timely to critically review their validity.
224
Anyone interested in a more thorough review of Great Lakes physical

limnology should read Boyce (1974) or Mortimer (1974).
I have chosen to divide the review into three sections corresponding

to the three most common idealizations of lakes - the summer, spring,
and winter models. In the summer the Great Lakes are usually stratified
everywhere and the main energy source is the wind. In the spring the
iakes are divided into two regions, the deep region where the tempera-
ture is below the temperature maximum densi ty and surface heating causes con-
vective overturning, and the stratified shore zone. The main energy source is
the densi ty contrast between the two regions. The fall is similar to spring
wi th the shore water less than the temperature of maximum densi ty and will
not be separately discussed. In the winter all density variations are
ignored and the only energy source is the wind. These three idealiza-
tions have been very useful in interpreting observations. More compli-
cated observed flow patterns can often be understood as the combination
of these basic flows.
2. Summer Circulations
It has long been recognized that in small stratified lakes the first
uninodal internal seiche is often generated by the wind. As a first
approximation the observed period can be accounted for by neglecting
rotation and nonlinearity (Mortimer, 1953). In some lakes, however,
it has been possible to detect thermocline slopes consistent with geo-
strophic flow parallel to the long axis of the lake (Mortimer, 1955).
In addition the internal seiches are sometimes found to have a surge
behavior (Thorpe, 1971; Ahrnsbrak, 1974). These phenomena can be under-
stood for long thin lakes by assuming the flow to be parallel to the
axis of the lake but with a slight tilt of the thermocline across the
lake. The resulting equations are the same as for nonrotating flow in
a long channel. However, in wider lakes such as Leman and Michigan the
internal seiches must take the form either of long internal Kelvin
waves trapped near the boundary or near intertial frequency Poincare
modes. Both types have been observed but the Kelvin wave is the most
relevant to coastal upwelling and will therefore be discussed in detail.
The classical work on observations of internal Kelvin waves is Mortimer

(1973). He first inferred them from water level observations around Lac
Leman, first showing that long period water level fluctuations were
strongly correlated with oscillations of the thermocline. Later he
obtained many long time series of temperature from municipal water in-
takes on Lake Michigan which showed a similar counterclockwise progres-
sion of temperature. A particularly well defined case of a large up-
welling-downwelling episode followed by wave propagation is shown in
Figures 1 and 2. Figure 1 (from Ayers et al., 1958) shows the surface
temperature after three days of winds from the north have caused an
upwelling of the seasonal thermocline along the east shore. Mortimer's
data, Figure 2, show a sharp rise of temperature at the water intakes
on the west shore and a sharp fallon the east. In the following week
of weak winds, the relaxation of the thermocline on the east coast shows
a clear phase propagation towards the north. The phase speed is roughly
45 cm/s, consistent with the internal wave speed.
Csanady and Scott (1974) observed a very similar propagation of the

thermocline displacement in Lake Ontario; but they had current measure-
ments also. The currents showed the vertical shear across the thermo-
cline consistent with an internal Kelvin wave, but they also had a net
vertically integrated flow. Clearly, therefore, the Kelvin wave model
225
Km.
400
300
TEMPERATURE
·C
SYNOPTI C 'lI
200
9 AUGUST 1955
100
HAVEN
Fig. 1. Lake Michigan, 9 August 1955. Distribution of surface temperature from six
transections (dashed lines) by Ayers et al. (1958). The corresponding temperatures
at coastal water intakes are indicated by vertical arrows in Figure 2
226
20
10 10
5 5
10 10
20 20
20 20
15 15
iO 10
5 5
25
2 20
15 \
,'\~,,II 15
5
25
20 20
15..,. 15
10
- - - - . r t- - - - - + 15
10 10
5 JULy ----+--------1 - - - --+5

·c 20 25 30 25 ·c
Fig. 2. Lake Michigan 1955. Temperatures, °C, at municipal water intakes (positions
shown in Fig. 1, depths as indicated), six hourly means at Milwaukee, Six-hourly
readings at Racine, otherwise daily readings. Also illustrated are daily means (and
"fastest mile") of wind speed and direction at Mi.l waukee. Black bars above the line
represent the prevailing directi'on S to W; black bars below the line represent winds
from the quadrant N to E; and shaded bars above and below the line represent winds
from the quadrants W to Nand E to S, respectively
227
is not complete; any wave interpretation must allow for bottom topo-
graphy and the shelf wave propagation mechanism associated with it.
In addition, one cannot neglect inertial effects since current speeds
are comparable to the phase speed. A simple model of Bennett's (1973),
combining Kelvin wave dynamics and inertial effects predicts steepening
similar to that of the thin lake case and observed in Loch Ness (Thorpe,
1971) .
These thin regions of large vertical and horizontal motion propagating

for long distances around the perimeter of a lake make the summer cir-
culation very difficult to model. There are no numerical models at
present that can reproduce the wave propagation accurately, though all
give the initial upwelling. Figure 3 summarizes my own feeble attempt
to model the Lake Ontario case studied by Csanady and Scott (1974) with
a 7-level, 5-km grid model. The observed data (from the Canada Centre
for Inland Waters) in the upper half show a strong depression of the
thermocline near the north shore. Csanady and Scott's data clearly show
that this thermocline depression originated on the south shore during
strong winds from the west about a week earlier and propagated counter-
LAKE ONTARIO
TEMPERATURE C 20 - 40 METRES
AUGUST 1 - 3 1972
Fig. 3. Temperature observed in Lake Ontario Aug 1-3, 1972 and simulated by a three-
dimensional numerical model
228
clockwise to the north shore. Both the current records and the large
temperature gradient indicate that frictional dissipation must be small.
However, the response of the numerical model is strongly damped. In
the predicted temperature field in the lower half of Figure 3 the thermo-
cline depression has reached only the east end of the lake and has de-
creased in amplitude. A very similar episode that occurred the follow-
ing week has been simulated by Simons (1975) with similar results.
While Simons was able to reasonably simulate the current meter records,
his model predicts a cooling of the north shore water over ten days
when the observations show a warming. I interpret this to mean that
the model predicts upwelling but misses the wave-like rebound of the
thermocline.
The depression of the thermocline on the north shore is associated with
a flow toward the west strong enough to make the monthly mean current
in the same direction - opposite the mean wind (Pickett and Richards,
1975) and opposite the mean current predicted by the numerical model.
Since the mean current is only 1 cm/s and the wave motion has an
amplitude of 20-50 cm/s it is natural to suspect rectification effects
of the cause of the mean flow. At present, however, there are no con-
vincing mechanisms for this rectification. I believe the reason for
the model's sluggish respons.e is the poor resolution of the shore
zones combined with the lateral friction needed to keep the model
stable. A new version of the model with more resolution near the
shore and a lateral friction acting only on the divergent component
of horizontal motion seems to work better but has not been fully
tested yet.
3. Spring Circulation
Because of the nonlinear equation of state of fresh water the spring

warm-up period is complicated. In early spring the water is below the
temperature of maximum density (4 o C) and very weakly stratified. Then
surface heating causes convective overturning until a given column
reaches 40 C and begins to stratify. This point is reached first in the
shallow water; the stratified region gradually expands until it covers
the entire lake in about six weeks, only occasionally being distorted
by the wind.
There was a flurry of research on this process in the 1960s and early
1970s. There were observations (Tikhomirov, 1963; Rodgers, 1966, 1968;
Rodgers and Sato, 1970), theoretical studies (Bennett, 1971; Csanady,
1971; Huang, 1972; Brooks and Lick, 1972) and a laboratory experiment
(Elliott and Elliott, 1969). The consensus was that flow was a weak,
time-dependent Hadley type circulation with a cyclonic circulation of
the ring of stratified water and a secondary flow of. upwelling near
the shore and sinking at the outer edge of the stratified zone. Advec-
tion of heat by the secondary flow tends to warm the deep water and
cool the shallow water '(Rodgers, 1968). A numerical estimate of this
advection (Bennett, 1971) confirmed that it was important, but since
the estimate was not very sensitive to the eddy viscosity it gives
little insight into how to parameterize turbulence for more complex
flows. Compared to the summer flows with thin jets and large transient
effects the spring circulation is easy to model. Further research in
spring will probably concentrate on the details of the front at the
edge of the stratified zone, and on the coupling between the wind and
the water in the presence of the large surface temperature gradients.
229
4. Winter Circulation Models
In the winter density gradients are much smaller than in spring or

summer and winds are stronger. It is therefore reasonable to expect
that the restoring force of gravity will not be important and that
vertical motion will be limited only by friction and the earth's ro-
tation. In the deep ocean and on the continental shelves it is often
assumed that rotation is the dominant constraint; in shallow lakes it
is assumed that friction dominates. This division between the rotation
and frictional constraints is an old idea that is rarely made more
quantitative. A major reason is the mathematical difficulty of even
the simplest model of the upwelling zone. It is generally agreed that
for a closed basin with a flat bottom, upwelling would occur at the
upwind shore for a shallow basin and at the shore to the left of the
wind (in the northern hemisphere) in a deep basin. These ideas follow
directly from Ekman's (1905) classical paper. He inferred these results
from the flow directions at various depths in steady flow profiles,
which assumed that the current is horizontally uniform; he did not
complete the circulation pattern by trying to solve for the structure
of the upwelling zone. The current profiles were determined by choosing
a horizontal pressure gradient so that there is no vertically inte-
grated flow. 'Since, in these profiles, there is no vertical average to
the current, flow in the surface layer must be compensated for in the
deep water. For the shallow water limit, the return flow is opposite
the wind and for the deep water limit the flow is to the left of the
wind.
It is true that, in agreement with Ekman's theory, upwelling is observed
at the shore to the left of the wind in deep lakes. It does not fallow,
however, that the flow in the deep water is to the left of the wind.
I have found that in my numerical model the flow of the deep central
water is to the right of the wind - implying that, since it is nearly
geostrophic, the free surface is actually depressed in the down-
wind direction. (This free surface slope is, of course, compensated
by set-up in the shore zones so that all the forces on the lake are
balanced when integrated over the whole volume.) This fact can be under-
stood by adding topography to the Ekman problem.
It has been known that topography radically alters the nature of the
problem even without rotation. For example, We1ander (1957) showed that
with variable depth the pressure gradient is not simply a constant but
varies in space and must be determined as the solution of an elliptic
equation with a mixed boundary condition at the shore. This implies
that the pressure gradient at one point in principle depends on the
topography, wind, and shoreline shape everywhere in the basin. The
solutions of this problem have not been systematically studied. Recent-
ly, however, Birchfield (1973) developed an interesting approximate
solution.
Birchfield showed that in the limit of small Ekman number the flow con-
sists of three regions. In the deep region the flow is stagnant at
large depths and in the surface Ekman layer there is a transport to
the right of the wind. Near the shore there is a double boundary layer
structure. The inner layer is from the shore to the point where the
water is as deep as the Ekman depth. The outer layer, matching with
the shore layer and the deep water, has a width equal to the geometric
mean of the radius of the basin and the width of the inner layer. In
this region, since the water is deeper than the Ekman depth, the flow
consists of a geostrophic flow with bottom and surface Ekman layers of
the classical type. It therefore follows that if the wind has no curl
the vertical motion due to geostrophic flow up the slope must be com-
230
pensated by a curl of the bottom stress. In the thinner shore layer

this does not apply and upwelling into the surface layer can occur to
complete the circulation. The important difference between this theory
and Ekman's is that, due to the Taylor-Proudman constraint, there is
no geostrophic current to the left of the wind in the center to com-
pensate for the surface drift to the right. Instead, the surface drift
is recirculated around the shoreline in a three-dimensional boundary
layer.
This theory immediately leads to two questions:
1. How small must friction be for Birchfield's solutions to apply?
2. How can one recognize the flow predicted by this theory from the
observed circulations of lakes or from numerical models?
To answer these questions it is necessary to solve the Ekman problem
for finite Ekman number and to numerically evaluate the solutions for
realistic physical parameters. I have done this for the case of a uni-
form wind blowing over a circular lake with a parabolic bottom. By
separation of variables this can be reduced to a complex second order
ordinary differential equation. The first step is separation of the
vertical structure as in Welander (1957), giving the mass transport
at a horizontal location in terms of the local surface pressure gradient
and wind. Welander continued by using the continuity constraint to
eliminate the mass transport and to find an equation for the surface
pressure alone. This equation, however, is not well posed since where
the depth is small the solution has a logarithmic singularity. I have,
therefore, chosen instead to eliminate the surface pressure to derive
an analogous equation for the stream f~nction, which is well behaved
even where the depth is small. The numerical solutions presented here
used 1000 equally spaced grid pOints in the radial direction with stan-
dard second order central difference approximations.
I have computed the solutions for a wide range of Ekman numbers but the
main results can be summarized in three figures. In the first the stream
function has been nondimensionalized by the wind drift transport:
where T is the wind stress, R is the radius, p is the water density

and f is the Coriolis parameter. In terms of the stream function,
radial and azimuthal volume transports are defined as:
0
- 1 aljl
J
-H(r)
urdz r aa
0
aljl
J ue dz
-H(r) ar
The radial coordinate is defined as positive outward and the azimuthal
angle increases counterclockwise and is zero where the wind is long-
shore (see Fig. 4). The Ekman number is defined as
E __2_A_
where A is the vertical eddy viscosity and H is the maximum depth. The
bottom has a parabolic shape rising to zero at the shore.
231
180 Fig. 4. Nondimensional stream func-

tion for steady flow in a circular
paraboloid driven by a uniform wind
o 90
-04 I~
-OU
o
STREAM FUNCTION
E=0.0016
N=IOOO
WIND
10 STREAM FUNCTION - - AMPL ITUDE
P: =0.2 cm 2 sec-2 - - - - PHASE

_~180
8 --~,.
H= 141m /-
It)
Q) f = 10-4
til
"'E 6 150
t)
C1>
Q 4 w
w (f)
a
::;)
120 <t
I
t:: 2 Q.
....J
Q.
~
----------
L -______~--~--~------~~----~~----~60
50
r(km)-
Fig. 5. stream function for a circular paraboloid lake model of the dimensions of
Lake Ontario for two values of the Ekman number
In Figures 5 and 6 dimensional variables are used. The depth of 141 m

and radius of 50 km correspond to a circular paraboloid of approximately
the same volume and surface area of Lake Ontario or the southern basin
of Lake Michigan. The wind stress of 0.2 dyn cm- 2 is a typical monthly
mean and the eddy viscosities 100 and 16 cm- 2 s-l are in a realistic
range. The corresponding Ekman numbers are 0.01 and 0.0016.
232
VERTICAL AVERAGE OF AZIMUTHAL VELOCITY

4 N= 1000
/'
/' -- /
/
Tu
H =141m /'
/'
/ "
"
'"
til
f;=0.2 cm 2 sec- 2 /'
/'
/'
/
300
/
E 3 f = 10-4 /' " /
/
~
w /
/ " /
/
240
00// "
/ W
0 /
=> (f)
t:: 2 ~'/'
/'\", /
/ <l:
I
-.J /
a.. /'
/
/ 180 a..
2 /' \, 10,/
<l:
./
/'
-- .,. "
\>-/'
---
-" /'
-" /'
..- -" --
------- --- 60
I I
00 10 20 30 50
r(km)-
Fig. 6. Depth averaged longshore current in a circular paraboloid for two Ekman num-
bers
The stream function for the smaller Ekman number, Figure 4, clearly
shows the Ekman drift in the center; however, there is no complicated
boundary layer structure. The reason is that, as the Birchfield theory
predicts, the horizontal mass balance requires only the Ekman drift
and the relatively thick El/4 boundary layer. Thus, within a fairly
wide range of eddy viscosities the stream function looks similar.
If you look closely at the streamlines in the center of Figure 4, you

will note that the flow has a small component in the direction of the
wind. This is opposite to what you would expect for a shallow lake.
The transition occurs at an Ekman number between 0.0016 and 0.01, as
can be seen in Figure 5 where the solutions are plotted in pola~ form.
Again, a striking feature of the solutions is that since the El/4 bound-
ary layer is thick, there is not much fine structure. Phase goes mono-
tonically from close to 90 0 in the center to 180 0 at the edge. Note,
however, that the inner El/2 boundary layer shows up as a region near
the edge where the phase is very close to 180 0 indicating that maximum
longhsore flow occurs at the coastline positions where the longshore
component of the wind is largest.
This inner boundary layer shows up better in Figure 6, a polar repre-

sentation of the vertically averaged longshore component. Within this
region the current has a jet structure. For smaller Ekman number, the
jet is stronger and thinner. However, the region where the current is
large enough to be observable with present technology is small for both
Ekman number s .
Thus the answers to the two questions posed earlier are:
1. The Birchfield theory's main conclusion - that the Taylor-Proudman

constraint causes the deep water to be stagnant, is true for Ekman
numbers much smaller than 0.01. This is probably true for deep lakes
such as Lakes Ontario and Michigan, particularly in summer when light
winds and stratification limit turbulence. This conclusion assumes that
the effective eddy viscosity for mean motions is much less than 100 cm 2
233
The model simulations of Simons (1975) showed that to reproduce

5- 1 •
the current speeds, values higher than this were needed only during
high winds. In addition, the evidence for internal Kelvin wave propa-
gation is clear evidence that dissipation is low most of the time. In
any case, because of topography, the deep water is not deflected to
the left of the wind as in Ekman's flat basin model.
2. Despite the strong distortion of the transport streamlines, the

maximum current speeds still occur where the longshore wind is strong-
est. It would be very difficult to detect rotational effects on mean
circulation by direct measurement of currents. Indirect evidence could
be provided by measuring the residence time of the deep water.
Acknowledgments. This work was supported by the Great Lakes Environmental Research
Laboratory of the National Oceanic and Atmospheric Administration, under Contract
No. 03-5-022-57.
References
Ahrnsbrak, W.F.: Some additional light shed on surges. J. Geophys. Res. 79, 3482-3483
(1974)
Ayers, J.C., Chandler, D.C., Lauff, G.H., Powers, C.F., Henson, E.B.: Currents and
water maSses of Lake Michigan. Univ. Michigan, Great Lakes Res. Div. Pub. 2, 169 pp
(1958)
Bennett, J.R.: Thermally driven lake currents during the spring and fall transition
periods. Proc. 14th Conf. Great Lakes Res., Intern. Assoc. Great Lakes Res.
Michigan: Ann Arbor, 535-544 (1971)
Bennett, J.R.: A theory of large-amplitude Kelvin waves. J. Phys. Oceanogr. 2,
57-60 (1973)
Birchfield, G.E.: An Ekman model of coastal currents in a lake or shallow sea. J.
Phys. Oceanogr. 2, 419-428 (1973)
Blanton, J.O.: Some characteristics of nearshore currents along the north shore of
Lake Ontario. J. Phys. Oceanogr. i, 415-424 (1974)
Blanton, J.O.: Nearshore lake currents measured during upwelling and downwelling of
the thermocline in Lake Ontario. J. Phys. Oceanogr. ~, 111-124 (1975)
Boyce, F.M.: Some aspects of Great Lakes physics of importance to biological and
chemical processes. J. Fisheries Res. Board Canada 11, 689-730 (1974)
Brooks, I., Lick, W.: Lake currents associated with the thermal bar. J. Geophys.
Res. 77, 6000-6013 (1972)
Bumpus, D.F.: A description of the circulation on the continental shelf of the east
coast of the United States. In: Progress in Oceanography. New York: Pergamon,
1973, Vol. VI
Csanady, G.T.: On the equilibrium shape of the thermocline in a shore zone. J. Phys.
Oceanogr. l, 263- 2 70 (1971)
Csanady, G.T., Scott, J.T.: Baroclinic coastal jets in Lake Ontario during IFYGL.
J. Phys. Oceanogr. !, 524-541 (1974)
Ekman, V.W.: On the influence of the earth's rotation on ocean currents. Ark. Mat.
Astron. Fys. ~, 1-53 (1905)
Elliott, G.H., Elliott, J .A.: Small-scale model of the "thermal bar". Proc. 12th
Conf. Great Lakes Res., Intern. Assoc. Great Lakes Res. Michigan: Ann Arbor 553-557
(1969)
Huang, J.C.K.: The thermal bar. Geophys. Fluid Dynamics 2, 1-25 (1972)
Mooers, C.N.K., Collins, C.A., Smith, R.L.: The dynamic structure of the frontal
zone in the coastal upwelling region off Oregon. J. Phys. Oceanogr. ~, 3-21 (1976)
Mortimer, C.H.: The resonant response of stratified lakes to wind. Schweiz. Z.
Hydrol. Basel ~, 94-151 (1953)
Mortimer, C.H.: Some effects of the earth's rotation on water movement in stratified
lakes. Verh. Inter. Ver. Limnol. 11, 66-77 (1955)
234
Mortimer, C.H.: Frontiers in physical limnology, with particular reference to long

waves in rotating basins. In: Publ. 10, Great Lakes Res. Div., University of Michi-
gan, 9-42 (1973)
Mortimer, C.H.: Lake hydrodynamics. Mitt. Intern. Ver. Limn~l. 20, 124-197 (1974)
Pickett, R.L., Richards, F.P.: Lake Ontario mean temperatures and currents in July
1972. J. Phys. Oceanogr. 5, 775-781 (1975)
Rodgers, G.K.: The thermal bar in Lake Ontario, spring 1965 and winter 1965-1966.
In: Publ. 15, Great Lakes Res. Div., University of Michigan, 369-374 (1966)
Rodgers, G.K.: Heat advection within Lake Ontario in spring and surface water trans-
parency associated with the thermal bar. Prod. 11th Conf. Great Lakes Res., Intern.
Assoc. Great Lakes Res. 480-486 (1968)
Rodgers, G.K., Sato, G.K.: Factors affecting the progress of the thermal bar of spring
i~ Lake Ontario. Proc. 13th Conf. Great Lakes Res., Intern. Assoc. Great Lakes Res.
Michigan: Ann Arbor 942-950 (1970)
Simons, T.J.: Verification of numerical models of Lake ontario. II. Stratified cir-
culations and temperature Changes. J. Phys. Oceanogr. 5, 98-110 (1975)
Thorpe, S.A.: Asymmetry of the internal seiche in Loch N;ss. Nature (London) 231,
306-308 (1971)
Tikhomirov, A.I.: The thermal bar of Lake Ladoga. Bull (Izvestiya) All-Union Geogr.
Soc. 95, 134-142. Amer. Geophys. Union Trans., Soviet Hydrology: Selected Papers
2 (1963)
Tomczak, M., jr., Diaz, C.G.: A numerical model of the circulation in Cienfuegos
Bay, Cuba. Estuarine and Coastal Marine Sci. 3, 391-412 (1975)
Welander, P.: Wind action on a shallow sea: some generalizations of Ekman's theory.
Tellus~, 45-52 (1957)
Hydrologic Aspects of the Main Upwelling Areas off Peru
s. ZUTA, T. RIVERA, and A BUSTAMAN1E
Field observations are described, using historical data from the main
coastal upwelling areas off Peru: one between 40 S and 60 S (northern
zone) and the other between 14 0 S and 16 0 S (southern zone). Vertical
cross sections and time sections of temperature, salinity, density,
oxygen, phosphate, and geosbrophic motions are presented. The data are
not the most adequate for a good description of the time variation.
In the northern zone the hydrographic structure shows evidence of strong
coastal upwelling in May and September, above 70 m, associated with the
southern extension of the Equatorial Undercurrent. The surface layer
exhibits the greatest changes of salinity during December-March and
September-November, being homosaline from March to September.
In the southern zone, the main upwelling takes place in June (Autumn)
and August (Winter) above 70 m, bringing to the surface waters of two
sources, depending on the time of the year. In the weak upwelling of
spring and summer the source water comes mainly from the south (sub-
antarctic water) 1 in autumn and winter the source is mostly from the
north. The hydrographic distribution shows evidence of great variations
in the surface layer, mainly above 20 m, and weak changes below 100 m.
Both in the northern and southern zones, tongues and patches, associated
with cyclonic and anticyclonic eddies, seem to govern the dynamics.
The tongues are typical along the Peruvian coast, apparently as a
result of the interaction of the inflow of the open ocean and an out-
flow of the upwelled water. These features establish a strong front,
especially in summer, and a meander-like distribution of the isotherms
and the surface flow, typical in May and June. The tongues extend 70
to 130 miles from the coast, especially in the southern hemisphere
autumn 1 the patches are 10 to 30 miles in diameter and are more common
in the southern zone.
In the northern zone the winds are predominantly from the south and
southwest, being strongest in May, July, and September. In the southern
zone the predominant winds are from the southeast and strongest in June,
August, and September. The latter are more parallel to the coastline
and facilitate the net upwelling, the strongest along the whole coast.
During an EI Nino period upwelling persists in the southern zone,
south of 140S, and seems to become stronger than normal in summer time.
1. Introduction
One of the world's main coastal upwelling areas occurs along the Peru-
vian coast where the biggest single fishery of the world, the anchoveta
fishery, develops.
The Peruvian coast has been. explored intenSively since 1961, through
national and foreign cruises as well as the so-called Eureka operations1
236
these employ about 20 fishing boats for a quasi-synoptic observation

of the whole coast and contribute to the fishery stock assessment. For-
eign cruises, mainly from the United States (1966 and 1969) and U.S.S.R.
(1972-1974), obtained helpful information about the coastal upwelling
off Peru, permitting theoretic considerations and conceptual models
usable for the CUEA (Coastal Upwelling Ecosystems Analysis) program.
For this presentation use is also made of the periodic cruises of the
Instituto del Mar del Peru (IMARPE).
Many questions require more field observations to obtain a good or

better description of the upwelling ecosystem. The present paper tries
to contribute some information on the hydrologic aspects of two main
areas of upwelling off Peru, with emphasis on the time variation.
2. Data
We used all available data on temperature, salinity, oxygen content,

and phosphate for the two shaded areas and dotted lines of Figures 1
and 2. The summary of the data sources given in Appendix 1 indicates
that we cannot speak properly of long-term averages.
8r 81 0
\
2200
50
Fig. 1. Topographic chart

(contours in fathoms) of
the band between 4-6 0 8.
The dotted line and the
shaded square show the
locations of hydrographic
sections and the coastal
6°~~~~~-t~~~~~~~~~--~--~~--~
8~lo~--~ 6°S area off Paita, respectively
237
Fig. 2. Topographic chart (contours in fathoms) of the coastal band: 140 30'-16 0 00'S.
The dotted line and the shaded square show the locations of hydrographic sections
and the coastal area off San Juan, respectively
The meteorologic data of Table 1 are related to coastal stations, and

in this respect we have good long-term averages of air and sea surface
temperature, atmospheric pressure, and wind speed but not wind direc-
tion; in this case we refer to published papers.
The vertical sections presented are not the most adequate to show the
features of the strong upwelling period for the lines shown in Figures
1 and 2. For instance, the sections off Paita belong to June and August
instead of May and September; those off San Juan are of September and
November, instead of August and February, and the other one is for
early June. For the two places we do not have observations deep enough
to choose a reference level better than that used in the geostrophic
calculations.
3. Coastline and Bottom Features
Mooers and Allen (1973) and Hurlburt (1974) emphasized the influence
of bottom topography and the coastline geometry on the dynamics of
coastal upwelling. With this idea in mind we prepared Figures 1 and
Table 1. Monthly means of sea level atmospheric pressure (P a ), wind speed (Ws), wind direction (Wd), surface air temperature N
LV
(T a ), sea surface temperature (Ts), and difference Ta-Ts-(~T). The atmospheriC pressure is given as P -1000 (X)
Variables Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec A.M. Period Source
a) Talara
Pa (mb) 10.9 10.6 10.5 10.9 11.5 12.5 12.7 12.2 12.0 12.1 11.9 10.8 11.5 1943-72 Senamhi a
Ws (kt) 14.6 12.4 11.7 14.9 17.5 19.4 18.4 17.9 18.4 18.0 17.2 16.8 16.4 1960-73
Wd (p) s s s s s s s s s s s s s 1960-73
Ta (OC) 24.0 25.3 25.3 24.3 22.5 20.7 19.3 18.9 18.8 19.2 20.0 21.8 21.7 1949-73
Ts (oC) 20.9 22.0 21.4 20.1 19.2 18.6 17.9 17.7 17.5 17.8 18.2 18.5 19.2 1956-73 DHNMb
~T (OC) 3.1 3.3 3.9 4.2 3.3 2.1 1.4 1.2 1.3 1.4 1.8 3.3 2.5
b) Chicama (17 0 41'S, 79 0 26'W)

Pa (mb) 9.6 8.1 7.8 9.0 10.0 10.8 11.5 12.0 12.3 12.8 11.7 10.7 1925-60 Senamhi
Ws . (kt) 4.5 4.5 4.6 4.6 4.5 4.3 4.2 4.4 4.4 4.5 4.4 4.4 4.4
Wd (p) SE SE SE SE SE SE S SE S SE S SE SE
Ta (oC) 21.8 22.8 22.6 21.3 20.3 19.2 18.8 18.6 18.5 18.5 19.4 20.4 20.2 1949-68 Gildemeister C
Ts (OC) 17 .4 19.3 19.0 17.8 17.6 17.1 16.8 16.4 15.9 15.7 15.8 16.4 17.1 1925-72
~T (OC) 4.4 3.5 3.6 3.5 2.7 2.1 2.0 2.2 2.6 2.8 3.6 4.0 3.1
c) Callao/Lima (Ta)/Chancay (Ts)

Pa (mb) 11.5 10.9 10.9 11.8 13.0 14.2 14.2 14.3 14.1 13.6 13.6 12.4 12.9 1949-72 Senamhi
Ta (oC) 21.4 22.2 21.9 20.2 17.8 16.1 15.3 15.1 15.3 16.1 17.6 19.3 18.2 1929-74 Min. pesqueriad
Ta (oC) 17 .0 17.2 17.5 17.2 16.8 16.0 15.8 15.0 14.8 15.3 15.7 16.3 16.2 1952-74 Senafere
~T (oC) 4.4 5.0 4.4 3.0 1.0 0.1 -0.5 0.1 0.5 0.8 2.1 3.0 2.0
d) San Juan
Pa (mb) 10.1 9.6 10.7 11.7 12.9 14.3 14.4 14.6 14.1 13.7 13.3 11.8 12.6 1959-72 Senamhi
Ws (kt) 10.1 10.7 11.2 11.8 11. 1 12.1 12.1 12.6 13.4 13 .1 11.8 10.4 11. 7 1960-73
Ta (oC) 21.9 22.4 22.0 20.2 18.5 16.6 15.5 15.3 15.7 16.8 18.4 20.3 18.6 1949-73
Ts (oC) 15.5 16.0 16.1 15.8 15.0 14.8 14.1 13.8 13.2 13.3 14.2 14.7 14.7 1958-74 DHNM
~T (oC) 6.4 6.4 5.9 4.4 3.5 1.8 1.4 1.5 2.5 3.5 4.2 5.6 3.9
e) Pta. Coles
Pa (mb)
Ws (kt) 11.8 12.2 12.8 11.7 11. 3 10.3 9.9 10.9 9.9 9.9 10.3 11. 3 11. 1 1965-73 Senamhi
Wd (p) SE SE SE SE SE SE SE SE SE SE SE SE SE 1965-73
Ta (OC) 21.8 21.8 20.9 19.6 18.2 17.0 15.8 15.7 16.1 16.4 19.0 20.5 18.6 1965-73
Ts (OC) 16.6 16.8 16.6 16.1 16.1 15.3 14.6 14.4 14.1 14.6 15.2 16.1 15.5 1954-73(15) n
~T (OC) 5.2 5.0 4.3 3.5 2.1 1.7 1.2 1.3 2.0 1.8 3.8 4.4 3.0
239
I and 2, based on navigation charts of the Peruvian Hydro-

.fl ..; graphic Service from 1970 •
~~ .....
0 (l'I
Figure 1 shows a coastline oriented meridionally in
I-< 0 .....
$:2: 0 general. The continental shelf is 3-12 miles wide, ex-
<<: cept for the parts at the Paita and Sechura Bights,
~:> III
0 where i t becomes 18 and 27 miles wide, respectively.
0'<: .i To some degree the isobaths parallel the coastline be-
OM ...: S
u OM
tween Talara and Pta. Falsa. The maximum depths, related
OM ..:I
>M to the northern side of the Peru trench, in some places
1-<\0 Q)
Q)(l'I
00 .....
'0 exceeding 2900 fathoms, appear about 35-60 miles from
the coast.
Figure 2 shows a coastline orientated mainly 135 0 (SE)

from Pta. Dona Maria to Pta. Lomas. The continental
shelf is 2 to 13 miles wide, and the isobaths tend to
parallel the coastline and make a canyon-like distribu-
tion off Pta. Lomas and farther south. The profile for
the section off Pta. Caballas (Cabo Nazca) shows a shelf
13 miles wide with a slight slope, and a steep continen-
tal slope (12-15 0 above 1000 fathoms and 21 0 down to
2000 fathoms). This profile is representative for the
hydrographic sections off San Juan.
4. Meteorologic Variables
In this section only the gross features of the mean

distribution of sea level atmospheric pressure, wind,
and surface air temperature along the Peruvian coast
are given. The long-term averages are given in Table 1,
which also includes sea surface temperature and its dif-
ference from the air temperature.
4.1 Sea Level Atmospheric Pressure
The monthly values show minima in March, except that of

San Juan in February. The high values prevail from June
to September, with maxima in July-August; the exception
is Chicama, which has high values in August-October
with the maximum in October. Both the annual and geo-
graphic amplitudes for Talara, Chicama, Callao, and San
Juan are of the order of 2.0 to 5.0 mb.
4.2 Surface Winds
Table 1 indicates that the prevailing winds are from

the south at Talara and from the southeast farther south.
The intensity also changes with latitude and time of
the year. At Talara the stronger winds appear in June
and the weaker ones in March; at San Juan the stronger
winds are in September and the weaker in January, and
at Pta. Coles the high wind speeds are in March and the
low ones in September-October.
The mean distribution given by zuta and Guillen (1970)

for the coastal areas off Talara (MS 308-41), Chimbote
(MS 307-89), Callao (MS 343-17), and San Juan (MS 343-55),
240
shows the following pattern. At Talara the dominant winds from the S
and SW are stronger in May, July, and September, and weaker in February,
August, and November. At Chimbote the winds are generally weak and
mainly from the SE. At Callao the winds are mostly from the SE, with
high speeds in June, September, and November, and low ones in January
and October. At San Juan the winds are from the SE throughout the whole
year with the highest speeds in June and September and the lowest in
March and t1ay. Evidently Talara and San Juan are two places of strong
winds most of the year.
The strongest winds do not coincide with the highest atmospheric pres-
sure; they appear one month earlier at Talara and Chicama, and one
month later at Callao and San Juan. The lowest wind speeds coincide
with the lowest pressure at Talara, and lag the pressure minimum by
one month at San Juan.
Burt et al. (1973) made the first study of surface winds during upwel-
ling, off San Juan, from a buoy near shore, and found a mean direction
of 163 0 ± 16 0 and a mean speed of 4.6 m s-l, values close to those given
by Zuta and Guillen (1970). There is a need to make detailed studies
of the wind field off the Peruvian coast as was emphasized at the EI
Nino Workshop in Guayaquil (Zuta et al., 1976).
4.3 Surface Air Temperature
Data for Talara, Chicama, Callao, San Juan, and Pta. Coles are shown
in Table 1. For their locations see Figures 1 and 2.
At Talara and Chicama the maximum air temperature values appear in

February, and the minimum values in September, with an annual range
of 4.40C. The maximum values occur in February at Callao, San Juan,
and Pta. Coles, with minima in August and an annual range of 7.10C.
Air temperatures are 1.0-6.5 0 C higher than sea-surface temperatures,

the greatest differences (3.0-6.5 0 C) occurring from December to April
and the lesser (0-2.5 0 C) from June to August. The maximum and minimum
values of these two variables coincide at Talara and Chicamai but at
Callao, San Juan and Pta. Coles sea-surface temperatures lag behind
air temperatures by one month. In general, the air temperature is de-
termined by the sea-surface temperature. P. Lagos and H. Montes (per-
sonal communication) made a statistical annalysis of temperatures from
Talara and Pisco and found that similar changes at these places occur
almost simultaneously with predominant periods of 12 and 6 months.
5. Hydrographic Conditions
The main features of the sea-surface temperature and the vertical dis-
tribution of temperature, salinity, density, oxygen, phosphate, and
geostrophic velocities in the upper 300 m will be discussed, giving
emphasis to time variations.
5.1 Synoptic Sea-surface Temperature
Figures 3 and 4 represent a quasi-synoptic sea-surface temperature

distribution in late May 1974. They show, as main features, tongues,
patches, and fronts along most of the Peruvian coast.
241
83° 81° 79° 77°

4°r-------,-------r-------,-------,-------.-------.-------,4°
1918-- PAITA
20 -C20
22 ~21
6° ~UNTAFALSA 60
,
23
I
/
/
I
I
8° 80
100 10°
12° ALLA 12°S

83°W 77°
Fig. 3. Synoptic chart of sea-surface temperature (OC), 28-30 May 1974. Northern
part of Eureka 29 (with 51-vessel fishing fleet), lMARPE
Six prominent tongues of cold water extend off Pimentel, Chimbote-

Huarmey, Callao, Pisco, San Juan, and Atico, apparently as a result
of advection of cold water offshore (outflow) and the inflow of warm
oceanic water. These tongues extend 70 to 130 miles from the coast,
and the result is a meander-like distribution of the isotherms. Data
from the Eureka and seasonal cruises tell us that this distribution
is typical in May and June, with a similar distribution of the surface
flow (Zuta and Santander, 1974). The tongue off Pisco parallels the
coastline and is an even more prominent feature in Figure 5 (late
February, 1974), in which a warm nearshore cell at Pisco is also ap-
parent.
The warm and cold patches are of different sizes and are more cornmon
south of 12 0 S, being 10-30 miles in diameter. They seem to be the re-
sult of cyclonic and anticyclonic eddies of a meander-like surface
circulation, which is evident in the topographies of the 1S o C isotherm
and the dynamic depths (e.g., Zuta and Santander, 1974; Zuta et al.,
1974). It is argued that the patches are elements of intensified up-
welling within an area of large-scale coastal upwelling (Tomczak, 1973).
Evidence for this idea seems 'to be clearer for Peru than for the up-
welling area off Northwest Africa as mentioned by this author.
242
,---r------~-------.------~~-----.------~~-----,12°
16° 16°
lS0 lS0S
Fig. 4. Synoptic chart of sea-surface temperature (Oe) , 28-30 May 1974. Southern
part of Eureka 29 (with 51-vessel fishing fleet), lMARPE
The fronts, which are very important for the fishery too, appear in
some places near the shore and in others offshore (Figs. 3 and 4) and
become strongest in the southern hemisphere summer (Fig. S). In some
cases there is an SoC temperature difference within a 30-mile wide
band. Also, in Figure S there is a branching of the main front off
Pta. Dona Maria, with one band 20 miles offshore and the other about
40-60 miles from the coast.
S.2 Mean Seasonal Distribution
Figure 6 shows monthly long-term mean sea-surface temperatures from

coastal stations located at the northern (Talara and Paita), central
(Chicama and I. Don Martin), and southern (San Juan and Pta. Coles)
parts of the Peruvian coast. The southern region has moderate changes
in the annual cycle, while the other two regions have quite clear
peaks in February-March.
The more or less continual cooling period is from February to Septem-

ber (7 months) at Talara and Pta. Coles, from March to August (S months)
at Paita, from March to September (6 months) at I. Don Martin and
San Juan, and from February to October (S months) at Chicama. The an-
nual amplitude is 3-S o C, increasing northward.
Figure 7 shows the variations of four variables at the standard levels

- 0, 20, SO, and 100 m. The temperature, salinity, and density in the
243
Fig. 5. Synoptic chart of sea-surface temperature (oC), 26-28 February 1974. A por-
tion of Cateo No.4 (with 22-vessel fishing fleet), IMARPE
area off San Juan have large fluctuations at 0 and 20 m, but little
variation at 50 and 100 m; the reverse is true for oxygen, which ex-
hibits notable changes at 50 and 100 m related to the intensity of
the subsurface southward flow. The changes in salinity are high from
December to February, and very low for the remainder of the year. There
is no constant cooling or warming, and oxygen shows no clear seasonal
variations. A similar picture exists for the area off Paita (not shown
here), but with marked peaks of salinity at all levels in July of the
same magnitude as the January peak at San Juan.
Figures 8 and 9 show the monthly mean variations of the vertical dis-
tribution of four variables in the upper 250 m for the coastal areas
off Paita and San Juan, respectively (see shaded squares in Figs. 1
and 2).
In the area off Paita (Fig. 8) the temperature and density display
similar patterns with large changes in the upper 150 m, where the
thermocline, pycnocline, and a weak oxycline (transition layer) de-
velop. The significant changes in salinity, especially above 50 m,
take place from October to March, with the presence of equatorial sur-
face water above 50 m, and subtropical water underneath. Appreciable
changes in oxygen distribution occur from May to September, possibly
due to the surfacing of the extension of the Equatorial Undercurrent
244
22
r-' Talara (1956-73)
I \ (4°34'S 81°15'W)
I \ / '
20 / \ ./
I \
,, /
/
I
\ I
18 f"'", //
Paita (1963-75) " I
(5°05'S, 81°07'W) '-,'--. ,//
16 ............ -----~
18 Chicama (1925-72)
(7"41'S 79°26'W)
f '
15
16 San Juan (1958-74)

(15°20' S, 75°1O'W)
7
14
16 Pta. Coles (1954-73)

FV (17"30'S, 71°20'W)
14
Jan Mar Jut Sep Nov Jan

Fig. 6. Monthly variations of mean sea-surface temperature at six Peruvian coastal
stations (for location see Figs. 3 and 4)
during this time of upwelling. Waters of high temperature and low den-
sities appear in February and June, and waters of low temperature and
high density appear in May and September, the months of main upwelling.
The upwelling takes place above 50 m in May and above 150 m in Septem-
ber; in the latter case of maximum upwelling the 25.8-26.1 Gt band
reaches the sea surface.
Displacements of the 25.3, 25.5, and 26.0 isopycnals indicate a mean

upwelling velocity of 12 m/month (4.6 x 10- 4 cm s-l) for autumn, and
of 58 m/month (2.2 x 10- 3 cm s-l) for winter. These speeds are evident-
ly much lower than those calculated by Halpern (1974) from measure-
ments of the displacement of the 25.5-26.0 G t band, conventionally used
in the Oregon upwelling regime (Pillsbury, 1972), but not appropriate
for these two areas of the Peruvian coast.
245
Jan Mar Jul 26.5 Jan Mar May Jul Nov Jan
24r-~'-'--r-r~-,-,--r-,-.-.-,-,
...............:;;~:~...........,;....... /· .......... 100m

22
26.0 ........ ,/ "',,'
........ J ~ 'J
t
20
! '--
,/' \
.,;\
t25.5 I
~ 18 20m'" \
\ .,Q " I
" I
f-
16
\
V
-'
,'\
25.0 ".J
5om------, " '.
.... ". \ "
• \ I \'
14 100m .. ···· ....~~;.............. . 24.5
12
24.0
35.4r-----------------, 7
1.3 6 @
~.2 5 ~ ~om
- ............. '"---~;-.""'""\ -
(f) .1 :;4
35.0 "::::"3 /,. ....... --~, " \~20m
.9
EO
~2 .............. ,' ........ ',,' ",\
0 ... ,J .: -.. " '50m

.8 1 '---' : .
.................... / ......... ·· .. 10 m
.7L-~~~~_L_L_L~~_ _L_~~~~
0
Nov Jan Jan Mar May Jul Sep Nov Jan
Fig. 7a-d. Monthly variations of mean temperature (a), salinity (b), density (c),
and oxygen (d) at four standard depths (0-100 m) for the coastal area off San Juan
In the area off San Juan (Fig. 9), similar patterns of temperature and
densi ty show marked changes in the upper, 100 m, where the thermocline
(mostly above 50 m), the pycnocline (strongest above 50 m), and the
isolines 1-5 ml/l of the oxycline (above 75 m) develop. Fluctuations of
isotherms and isopycnals are moderate with respect to those in the area
off Paita. The appearance at the surface of warm low-density waters
takes place in January, April, and July, and the coldest and most dense
water appears in June and August, the months of strong upwelli~g. In
August the 25.8-26.1 0t band reaches the surface. Evidently, in Decem-
ber and February, the weak upwelling brings to the surface subantarctic
waters, while in June and August strong upwelling brings waters to the
surface whose sources are mostly in the north. This is also evident in
the spreading and deepening of the isolines of oxygen during these
months.
An estimate of the movements of the 25.5 and 26.0 0t isopycnals gives

upwelling speeds of 54 m/month (2.1 x 10- 3 cm s-l) for autumn, and
103 m/month (4.0 x 10- 3 cm s-l) for winter, in agreement with the
values found by Zuta and Guillen (1970) for the area off Callao.
Significant changes in salinity take place from December to February,

with the appearance of subtropical surface water above 50 m and sub-
antarctic water below. During the remainder of the year the whole column
appears highly homogeneous, similar to the area off Paita for the
period March to October.
The general features of the areas off Paita and San Juan are mostly in
agreement with those off Pta. Falsa and Callao as presented by Zuta and
Guillen (1970).
246
100 100
15r 14
200
13@)\ Q \
\
\
\
200
,
\
\ \
I,' \
Om Om
3
2
100
Jon Mor Moy Jul Sep Nov Jon Jon Mor Moy Jul Sep Nov Jon
Fig. 8a-d. Monthly mean distribution of temperature (a), salinity (b), density (c),
and oxygen (d) for the coastal area off Paita
The main seasonal distribution of the four variables in the upper

300 m shows clearly the large variations in the 0-50 m layer, especial-
ly above 20 m, and homogeneity in the 100-300 m layer. Remarkable dif-
ferences for the two areas occur in the surface layer. There are great
variations in salinity and a strong pycnocline off Paita, in contrast
to the features off San Juan, where a strong oxycline and a conspicuous
oxygen minimum develops in the layer below 100 m depth. This minimum
is dominant from October to March (Fig. 9) and is probably associated
with the strong southward flow of the Peru-Chile Undercurrent (Wooster
and Gilmartin, 1961). On the other hand, off San Juan the maximum
values of phosphate are closer to the surfa ce than off Paita (Fig. 10).
The vertical sections of temperature, salinity, density, and oxygen

given in Figures 11 through 15 complement the preceding descriptions.
For their locations see Figures 1 and 2.
Figure 11 is a section made off Paita in early June, 1973, and in this,
as in all others, temperature and density have the same pattern with
a close correspondence between the 15 and 12 0 C isotherms and the 26.0
and 26.5 isopycnals, respectively. The divergence of the isopycnals
247
III V IX XI VII IX
I I I I I I
JOO JOO
34.8
3485
....
....
200 200
1
05 ....
100 100
.:" ".
<025
.... ,.' i \.....
200
>05 >05 @) <025 200
Jon Mar May Jul Sep Nov Jon Jon Mo r Moy Jul Sep Nov Jon
Fig. 9a-d. Monthly mean distribution of t e mperature (a), salinity (b), density (c),
and oxygen (d) for the c oastal area off San Juan
takes place at 70 m within 40 miles of the coast, which means that the
upwelling occurs above the 70-m depth. In late August, 1963 (Fig. 12),
the upwelling was above the same level but within 20 miles of the coast,
with the 25.8-26.1 isopycnal band much deeper than in June.
Figures 13 to 15 show three stages of the hydrographic conditions off

San Juan (Cabo Nazca). The location is almost the same as section II
of R/V Thompson Cruise 26 on 31 March-1 April, 1969 (Smith et al.,
1971). In early June (Fig. 13) the divergence of the isotherms takes
place at 70 m within 25 miles of the coast, and below 100 m and within
100 miles from the coast there is a notable deepening of the isotherms
and isopycnals. In mid-September, 1968 (Fig. 14), the picture was dif-
ferent, even though the upwelling took place above 70 m as in the other
cases and within 30 miles of the coast. In late November, 1966 (Fig.
15), the distribution was similar to that for June (Fig. 13), but with
weaker upwelling. In the three cases or stages, the subtropical sur-
face water was dominant in the surface layer farther than 100 miles
from the coast; the subantarctic water appeared as a subsurface minimum
in June and November with its axis at about 80 m depth and farther
than 100 miles from the coast, being notable in June; the oxygen min-
T (OCl_ S("Ioo)- - at - Oz(ml/L) .... P04 (j.(g-ollL) N
12 14 16 18 20 22 34.6.7 .8 9 35.0 .1 .8 24.0 2 .4 .6 .8 25.0 .2 .4 .6 .8 26.0 2 .4 .6 0 2 3 4 5 0 1 2 3 ~
CD
Om. : : .',' ' _ : Om.
./~/:/" .... - ----------- ---------- ------\,.~.~.---~.~~.....•..•
100 /7 \«:::. 100
~!\\:\\
I . \
SUMMER
AUTUMN ",
200 200
WINTER
...\,,,i·, ... '1
SPRING :
:
,,
, :
@ @ @) o
Jf ,~I (C) ~ I~ \, 1'00
T (OC )- S("Ioo)-- at - Oz(ml/L) P0 4 (J.<g-ot/L)-
10 12 14 16 IS 20 34.7.8 .9 35.0 .1 24.6 .8 25.0 .2 .4 .6 .8 260 .2 .4 .6 0 2 3 4 5 0 1 2 3 4
Om. ''- • .. _ • Om.
f:;>;>;> --- l' - - ---"""\\ ",;;:>:;,;:::>/:- """'<~

100 100
i='
/i/ (/)
~;
\\ ';
(:
El/
i(~\
\~~!
:
V 1i \ ::: ":
tff !!
:': \ Ii~: ' :\
I ::f \ ;: :;
:'i ~ I: :'
200 ( ". ~ l' ., 200
:': .os t= .
, #! I
ti ~': !,01 !~
l
: \
IQ\: J , ' nfb\ ~
0V \.i
') ~ @) o
300 300
Fig. lOa-e. Vertical profiles of seasonal mean temperature (a), salinity (b), density (c), oxygen (d), and phosphate (e) for
the coastal areas off Paita (above) and off San Juan (below)
249
n·c) 5 ("!..)
S45
i
544
,
543
i
Om
100 100
~ 14
200 200
~ 13
.~
34.9
300
\ 300
Om Om
100 100
~ 263"
200 200
~ .....
'-~~" 05
' ........ _.. .-- ...
_______ 26.5•.
300 83"W 8ZO

Fig. 11a-d. Vertical sections of temperature (a), salinity (b), density (c), and
oxygen (d) off Paita, 9-10 June 1973, (cr. 7305 BEe Mesyatsev)
imum layer is a dominant feature below 100 m and within 100 miles of
the coast, especially in November.
Evidently the density field depends on the temperature regime, even

in the area off Paita. The divergence of the isopycnals at 70 m is in
agreement with the view of Wyrtki (1963) that upwelling near the Peru-
vian coast is limited to depths less than 100 m.
250
noel
0~t;lP~;=~~==~14~i====jl~~1~7~1~~
16
100 100
200
~1 4 ~ 200
~ '3
300 300
Om Om
100 100
200 262 200
. 05
26.4
82"
oxygen (d) off Paita, 19-20 August 1963, (Cr. 6308 BAP Bondy)
6. Geostrophic Flows
The theoretic (e.g., Allen, 1973) and numeric (e.g., O'Brien and Hurl-
burt, 1972) models and the field observations (Smith et al., 1971;
Smith, 1974; Huyer and Smith, 1974) show that the along-shore geostro-
phic flow is an important feature in coastal upwelling events.
Figure 16 shows geostrophic flows in June (a) and August (b) off Paita
and in June (c) and November (d) off San Juan; these correspond to
Figures 11, 12, 13, and 15, respectively, and are consistent with the
density field of these sections.
251
T ('C)
51 66 65 64 63 r,
62 61 60
I j
Om
~20
:=:::-=
100
.---34.6:--J ~~
_ _ _ _-_34 7 - J 1 . 100
34.8
34.9
200 200
300~------------~--~----~----~u L-----------~~--~----------~ 300

omr-------~~r_._~~~--~--~nr~_, r-----------~~~~~--~----._,__,Om
5
,..-/ 4 ._----,
.,
:.-- 2 Ilr" i
II I
100 -1
:- - - 05 " II ,
I
100
I
I
/
I
\
,
I
, ,- I
I
I
I
/
I
200 I 200
..... , /~
/
""-0.25 /
/
"- ,_ ..... /
/ @
300 150 L-~------~~------~~------~300

150 100 :il 0
Miles
oxygen (d) off San Juan, 2-3 June 1967, (Cr. 6705 BAP Unanue)
The southward flow off Paita (Fig. 16a, b) with maximum speeds up to
10-15 cm s-l at about 50 m depth nearshore, is related to the south-
ward extension of the Equatorial Undercurrent east of the Galapagos
Islands (Cochrane and Zuta, 1968, unpubl. manuscript; Stevenson et
al., 1971); however, the offshore flow, with the axis near the surface,
seems to be related to a surface flow carrying equatorial surface wa-
ters. The northward flowing currents (one between the above-mentioned
southward flows and one further west) with speeds up to 15-20 cm s-l,
must belong to the northern side of the Peru Coastal Current. The dis-
tribution of these flows tells us that off Paita the upwelled waters
corne from that southern branch of the Equatorial Undercurrent or Crom-
well Current that brings waters of high oxygen content.
252
T ('C) 5 ('1 •• )
SI 52
I
51I 50
I
1.9
I
1.8I ,
1.7 ,
52 51I 510 1.9I 1.8
I ,
1.7
Om Om
/
~.
, ,.
:0-
17 ..... I
I
-_/
\
100 349 100
; 12,5
.,- /
.,-
.,-
..-
200
-- ----- -- . 12
® @
11
300
(milL)
Om Om
200
' 26.5 ./ /".,/.' -0.25
~© 266
/'
/'
/'
"-
\
@)
"-
300
Miles 100 50 100 50
Fig, 14a-d. Vertical sections of temperature (a), salinity (b) .. density (c), i?Ild
oxygen (d) off San Juan, 13-14 September 1968, (Cr. 6809 BAP Unanue)
The basic pattern of circulation suggested by SCOR-WG 36 (SCOR Pro-

ceedings, Vol. 10, No.2, 1975), for upwelling off Peru, seems to be
consistent with the hydrologic features off San Juan, and even more
so with those off Chirnbote (see Figs. 25, 34-36 of Zuta and Guillen,
1970) .
We have described normal conditions, but during abnormal years, when

the circulation pattern is greatly modified for a period of 4-14 months,
the southward flow seems to become stronger than normal, and the up-
welling does not disappear south of 14 0 S, being stronger than normal
in the southern summer (Zuta et al., 1974).
T(°C) 5 (" •• )
51. 39 40 41 42 43 44 45 39 40 41 42 43 44 45
r I i I I i I i I ...... <: I I <: i I 1
O m. }:/ 7 10m
- - - 35 .1
100 100
200 200
34.8
~:: /
) ,), '\ ), .
300' <' at , " " '" 300
Om I ,""" ~-.?LL ~~
,
100
" 100
" "\
\
.\\
1
I
200 I
\
26.5
/
26.7_
I , >" ,.'.... r)' '\ 1-
300 1 . <' 2&0 c 150 ,-- IV
Miles 200 150 100 Ul
W
Fig. 15a- d. Vertical sections of temperature (al, salinity (b), density (c), and oxygen (d) off San Juan, 24-25 November
1966, (Cr. 611 BAP Unanue)
254
8l'W 1''30' 8l'W

Om ;-r "al I!IO.J
100
am
00 100
200
1
300 ISO
200 100 SO
Fig. 16a-d. Vertical sections of geostrophic velocity (em s-l) relative to 300 m;
(a) 9-10 June 1973 off Paita (Cr. 7305 BEC Mesyatsev); (b) 19-20 August 1963 off
Paita (Cr. 6308 BAP Bondy); (c) 2-3 June 1967 off San Juan (Cr. 6705 BAP Unanue);
and (d) 24-25 November 1966 off San Juan (Cr. 6611 BAP Unanue) . Case (a) is relative
to 200 m
7, Summary and Conclusions
Analysis of the available data for the two main areas of upwelling off
the Peruvian coast yields the following features or characteristics:
1. The coastline is oriented almost parallel to the direction of the

dominant winds, i.e., meridional between Talara and Pta. Falsa, and
135 0 (SE) between Pta. Dona Maria and Pta, Lomas, with a narrow con-
tinental shelf 3-28 miles wide in the first case, and 2-13 miles wide
in the latter, the shelf being a bit flatter in the former case,
255
2. The sea surface temperature displays prominent tongues of cold water

extending 70 to 130 miles from the coast. Cold and warm patches of
10-30 miles in diameter are more common south of 120S, especially in
autumn; nearshore and offshore fronts are strong in the southern hemi-
sphere summer. All these result from the outflow and inflow of cold
upwelled water and the inflow of warm oceanic water, which give a
meander-like surface circulation.
3. The large seasonal changes in the hydrographic properties take place

in the 0-50 m layer, especially above 20 m; the 100-300 m layer has
low variability.
4. The temperature and density fields display a similar pattern: The

15 and 12 0 C isotherms nearly coincide with the 26.0 and 26.5 at iso-
pycnals, respectively. The 25.8-26.1 at band is a good one to show the
region of intense upwelling, instead of the 25.5-26.0 at band commonly
used for the Oregon coast.
5. The main periods of upwelling are May and September off Paita, and
June and August off San Juan. In both places the upwelling takes place
above 70 m and within 20-40 miles of the coast. Off Paita the only
source of water is the southward extension of the Equatorial Undercur-
rent. Off San Juan the source water is mostly from the north in June
and August, and from the south in the weak upwelling of December and
February.
6. Displacements of selected isopycnals give mean upward speeds of

12 m/month (4.6 x 10- 4 cm s-l) for autumn and 58 m/month (2.2 x 10- 3
cm s-l) for winter off Paita. The speeds off San Juan for the same
season are 54 m/month (2.1 x 10- 3 cm s-l) and 103 m/month (4.0 x 10- 3
em s-l), respectively.
7. Geostrophic speeds give southward flows of up to 15-45 cm s-l off

Paita, and up to 10-15 cm s-l off San Juan, as well as northward flows
with speeds up to 15-35 cm s-l off Paita and up to 5-15 cm s-l off San
Juan.
8. Taking into account the geostrophic motion and the hydrologic struc-
ture, we can say that the basic pattern of circulation for Peru sug-
gested by SCOR seems consistent with the observed features off San Juan
and even more so off Chimbote.
AaknOlJ)ledgments. We wish to express appreciation and thanks for the help given us
by Mr. R. Gomero (data collecting and drafting) and Mr. P. Torres (drafting). We
appreciate the valuable suggestions of R. Jordan, A. Landa, and J. Csirke on our
original manuscript. Mr. o. Guillen put at our disposal the tabulated meteorologic
data for the airports of Talara, Callao, and San Juan. J. Valdivia (SENAMHI) gave
us the meteorologic data of Chicama and Punta Coles. This work was supported by the
Instituto del Mar del Peru (IMARPE). Typing was done by Sonia Salazar. We wish to
thank Drs. J. O'Brien, M.B. Peffley, M. Tomczack for their valuable suggestions and
review of the manuscript.
References
Allen, J.S.: Upwelling and coastal jets in a continuously stratified ocean. J. Phys.
Oceanog. 1, 245-257 (1973)
Burt, W.V., Enfield, D.B., Smith, R.L., Crew, H.: The surface wind over an upwelling
area near Pisco, Peru. Boundary~Layer Meteor. l' 385-391 (1973)
Halpern, D.: Variations in the density field during coastal upwelling. Tethys. 6
(1-2), 363-374 (1974)
256
Hurlburt, H.E.: The influence of coastline geometry and bottom topography on the
eastern ocean circulation. In: Technical Report, Mesoscale Air-Sea Interaction
Group, Dept. of Meteorology. Tallahassee: Florida State Univ., 1974
Huyer, A.: Smith, R.L.: A subsurface ribbon of cool water over the continental shelf
off Oregon. J. Phys. Oceanog. i, 381-391 (1974)
Lagos, P., Montes, H.: Spectral analysis of the oceanicatmospheric data from the
Peru coast. Instituto Geofisico del Peru (1975) (to be publ.)
Mooers, C.N.K., Allen, J.S.: Final Report of the Coastal Upwelling Ecosystems Anal-
ysis Summer 1973 Theoretical Workshop, School of Oceanography. Corvallis: Oregon
State Univ., 1973
O'Brien, J.J., FUrlburt, H.E.: A numerical model of coastal upwelling. J. Phys.
Oceanog. 2, 14-26 (1972)
Pillsbury, R.D.: A description of hydrography, winds, and currents during the up-
welling season near Newport, Oregon. Ph.D. Dissertation. Corvallis, Oregon: Oregon
State Univ., 1972, p. 163
SCOR: Proceedings, International Gouncil of Scientific Unions 10 (2) (1975)
Smith, R.L.: Upwelling. Oceanog. Marine Biol. Ann. Rev. 6, 11-47 (1968)
Smith, R.L.: A description of current, wind, and sea level variations during coastal
upwelling off the Oregon coast. July-August, 1972. J. Geophys. Res. 79, 435-443
(1974)
Smith, R.L., Enfield, D.B., Hopkins, T.S., Pillsbury, R.D.: The circulation in an
upwelling eC9system: the PISCO cruise. Investigacion Pesquera 35 (1),9-24 (1971)
Stevenson, M.R., Taft, B.A.: New evidence of the Equatorial Undercurrent east of
the Galapagos Islands. J. Marine Res. 29, 103-115 (1971)
Tomczak, M., Jr.: Problems of physical oceanography in coastal upwelling investiga-
tions. Geoforum 11, 23-34 (1972)
Tomczak, M., Jr.: An investigation into the occurrence and development of cold water
patches in the upwelling region off NW Africa. In: "Meteor" Forsch-Ergebnisse.
Berlin-Stuttgart: 1973, Reihe A, No. 13, pp. 1-42
Wooster, W.S.: Gilmartin, M.: The Peru-Chile Undercurrent. J. Marine es. 19, 97-122
(1961)
Wyrtki, K.: The horizontal and vertical field of motion in the Peru Current. Bull.
Scripps Inst. Oceanog. 8, 313-346 (1963)
Zuta, S., Guillen, 0.: Oceanography of the coastal waters of Peru. Bol. Inst. Mar.
Peru ~ (5), 157-324 (1970)
zuta, S., E~field, D., Valdivia, J., Lagos, P., Blandin, C.: Physical aspects of the
"El Nino" phenomenon. El Nino Workshop, Guayaquil, Ecuador, December 4-12. Rome
FAO 185, 3-61 (1976)
Zuta, S., Santander, H.:. The marine environment and its relationship with the bio-
logical resources (1974) (to be publ.)
257
Appendix 1. Available hydrographic observations used for the two coastal areas
Year Ship Cruise No. Area off Paita Area off San Juan
J F M A M J J A SON D JFMAMJJASOND
1961 Bondy 6109 2 -

1962 Bondy 6201 2 -
1963 Bondy 6302 - 2
1963 Bondy 6308 2 - -
1963 Bondy 6311 2 -
1964 Unanue 6402 1 -
1964 Unanue 6405 - 1 -
1964 Unanue 6408
1964 Unanue 6411 1 - 2
1965 Unanue 6504 - - - 2 -
1965 Unanue 6507-08 ------2-
1965 Unanue 6510-11 - 4 -
1966 Unanue 6602 2 -
1966 Unanue 6611 3 -
1967 Ecorrastreo VIII
1967 Unanue 6705 1 -
1967 Unanue 6708
1967 Unanue 6711 1 -
1968 Unanue 6802 - 1
1968 Ecorrastreo IX 5
1968 Unanue 6809 1 -
1968 Ecorrastreo X 2
1968 Unanue 6811 1 -
1969 SNP-1 6901-02 - 1
1969 SNP-1 6905-06 - 4 - 2
1969 SNP-1 6907 - - - - - - 1
1969 Ecorrastreo XIV - 9 -
1969 SNP-1 6910
1969 SNP-1 6912
1970 SNP-1 7005 - - - - 1 -
1970 Unanue 7009 1 -
1970 Unanue 7011-12 2 -
1971 Unanue 7105 2
1971 Unanue 7108 1 "- -
1971 Eureka XIX - - - - - - - 4 -
1971 Chatyr Dag 7109 - 2
1971 Unanue 7111 - 2 -
1972 Unanue 7202 - 2 - -
1972 Unanue 7204 1 -
1972 Eureka XX - - -6 -
1972 SNP-1 7205
1973 Eureka XXVI
1973 Unanue 7302-03 - 1
1974 SNP-1 7402 - 1
1974 SNP-1 7405
1974 Eureka XXIX ----20-
1974 Eureka XXX - - - - 9 - - - -
Total observations: 322 6 251-34 6 5 9 23 4 226 5 3 15 2

Geological Aspects of Upwelling Regions
Sediments as Indicators of Upwelling
L. DIESTER-HAASS
1. Introduction
The investigation of recent sediments from upwelling regions is con-

cerned with three main questions: (1) the possible mechanisms of forma-
tion and typical parameters that allow the classification of a sediment
as being influenced by upwelling; (2) whether i t is possible to apply
these parameters to fossil sediments to reconstruct past oceanographic
conditions, a question that becomes especially interesting within the
Deep-Sea Drilling Program; and (3) whether the upwelling-influenced
sediments have any economic significance.
There are two major kinds of upwelling: coastal and equatorial upwel-
lings (the arctic and antarctic divergence zones will not be considered
here). The coastal upwelling off N and S Africa and Nand S America
is situated in the trade wind belt in the eastern boundary current
region (Smith, 196B). Off Somali, S Arabia, and off the west coast of
India it is in the monsoon region (Smith, 196B).
The main characteristics of the surface water masses in upwelling re-

gions are lowered temperat.ures (up to 9 0 lower than the normal values
for the latitude); lowered salinity; lowered oxygen contents; high
nutrient contents and thus increased productivity.
Two of these characteristics, lowered temperatures and increased bio-

logic production, are important for geologic observations, because they
may be reflected in the sediments and thus lead to characteristic dif-
ferences between sediments underlying upwelled water masses and sedi-
ments from the same latitudes that have no upwelling influence.
These characteristics, lowered surface temperatures and high biologic

productivity, are only reflected in the sediments if the upwelled water
masses influence the bottom sediments for sufficiently long periods
of time. A 2-cm thick surface layer of the bottom sediments reflects
the conditions during a period of between 200 and 1000 years on the
upper and lower continental slopes, respectively, in the Northwest
African upwelling region (Diester-Haass, 1976). On the shelf, the time
period included in a 2-cm thick surface layer is highly variable in
different upwelling regions. Bottom currents may lead temporarily to
nonsedimentation, topographic features with basins -and hills may lead
to high sedimentation rates directly adjecent to regions with reduced
or no sedimentation. Terrigenous input can locally increase sedimenta-
tion rates. In the Southwest African upwelling region a maximum sedi-
mentation rate of 150 cm/1000 years in basins on the shelf has been
observed (Meyer, 1973). Here a 2-cm thick surface layer reflects 13
years.
So surface sediments do not reflect the contemporaneous hydrologic

conditions like plankton, nor do they reflect a one-year average of
hydrologic conditions like benthos (Thiel, this vol.), but they do re-
flect average hydrologic conditions during past decades or centuries.
262
Even if upwelling occurred during past centuries or decades, i t is not

certain whether i t is clearly reflected in the surface sediments. First,
there is the problem of constancy of upwelling during the year. It is
known that upwelling can change in position and intensity during a year,
due to changes in wind and currents (Hoflich, 1972). For example, off
Northwest Africa, upwelling phenomena are observed throughout the year
off Cape Blanc (20ON). Off Senegal, however, upwelling was measured
only during 5.5 months per year (Schemainda et al., 1975). This leads
to differences in the underlying sediments. In the first case we find
the influence of the cool and fertile water masses throughout the year,
and in the second case we find a mixture of "normal" sedimentation and
upwelling-influenced sedimentation.
Secondly, there is the problem of currents. On the shelf and upper

slope, currents may be strong enough to erode bottom sediments (Newton
et al., 1973; Diester-Haass, 1975), so that typical upwelling influences
in the sediments disappear. On the continental slope off Sahara current
velocities up to 40 cmls have been measured 10 m above the sea-floor
in 500 m water depth (Meincke et al., 1975).
Thirdly, there is the problem of a continuous downwelling front (Ri-

chert, 1975). Richert believes that planktonic diatoms, a main con-
stituent of qpwelling-influenced sediments, do not reach the bottom
fast enough in regions where a continuous downwelling front is absent.
Thus, dissolution of the opal skeletons occurs in the water column and
the sediments do not show an enrichment of opal tests.
Lowered surface-water temperatures and high organic production are the

main features that produce differences in the sediments from upwelling
regions 90mpared to adjacent regions not effected by upwelling. These
two parameters are reflected in textural, geochemical, and biologic
characteristics of the sediments.
2. Textural Indicator of Upwelling
Sediments from well-oxygenated regions are normally burrowed by benthic

organisms except where sedimentation rates are very high. Benthic or-
ganisms destroy the original bedding. When, however, bottom water lacks
02 or if H2S is present due to high organic matter supply, bottom life
is impossible and sediments show undisturbed layering. Finely laminated
sediments have been observed in diatomites from the center of the South-
west African upwelling region (Meyer, 1973; Diester-Haass, in prep. b),
in the Gulf of California (Calvert, 1966), and off Peru (Ssaidova,
1971) as well as in black-dark-green muds from Santa Barbara Basin off
California (Berger and Soutar, 1970; Soutar, 1971) and off the west
coast of India (Stackelberg, 1972). Moving away from the reducing zone
benthic life will reappear and sediments are burrowed. Laminated sedi-
ments, however, are not only typical for the 02-minimum zone in highly
fertile regions, but also for deep basins where O2 exchange is inhibited.
3. Geochemical Indicators of Upwelling
3.1 Organic Matter
The high fertility of the surface water masses in upwelling regions

allows high organic producti'on in the surface waters so that large
263
amounts of organic matter can sink to the bottom and be included in

the sediments. This is reflected in contents of organic carbon, which
are distinctly higher than in surrounding regions without upwelling
influence.
"
20'
Fig. 1. Organic carbon contents in sur-

face sediments from the NW African up-
welling region. Dots, Cores from Meteor
25 expedition .. Organic carbon val~
from Hartmann et al. (1975); crosses,
profile I of Meteor 36 expedition;
black squares, profile 2 of Meteor 36
expedition; triangles, organic carbon
values from Miro de Orell (1973); white
squares, organic carbon values from
RUhl in Diester-Haass et al. (1974)
In the Northwest African upwelling region (Fig. 1) organic carbon con-

tents are low on the shelf and lower continental slope (less than 2%).
On the upper continental slope, however, values up to 7% off Cape Ti-
miris and up to 5% off Cape Blanc indicate high organic production.
Calvert and Price (1971) measured organic carbon values up to 26% off
Southwest Africa . Maximum organic carbon values given by Bremner
(1974a), Birch and Rogers (1973), and Rogers (1974) for the same re-
264
gion are between 8 and 17% (the lower values are from the region to
the south, and the higher ones to the north of Walvis Bay). Davey and
Rogers (1975), however, give one value of 26% organic carbon at a lati-
tude of 25 0 S. The organic carbon-rich muds begin close to the coast
and extend seaward as far as about 150 m water depth (Marchand, 1928;
Copenhagen, 1953; Surnrnerhayes, 1972).
Off Peru organic carbon contents are "very high" (Ssaidova, 1971) in
his zone II (30-250 m water depth). Bandy and Rodolfo (1964) describe
negligible amounts up to 5.6%. Rhyther et al. (1971) record a percentage
of organic carbon of 2-7% off Peru. They believe that this percentage
would be several factors higher if all plant material produced in the
surface water was deposited in the sediments. Most of the organic ma-
terial is taken up by herbivorous animals in the water column.
In the Gulf of California organic carbon values can be greater than

10% (Calvert, 1966). In the basins off California contents increase
up to 11 % (Emery, 1960).
Contents of organic carbon up to 11% have also been found off the west
coast of India and attributed to seasonal upwelling (Murty et al.,
1969) .
Percent values of organic carbon do not reflect only surface produc-

tion but also dilution and oxidation. So it is better to give sedimen-
tation rates of organic matter than percent values (see Mliller, 1975).
3.2 Minor Metals
Planktonic organisms can enrich certain elements compared to the sea-

water (Seibold, 1970; Bostrom et al., 1974) and after their burial in
the sediments these elements can be enriched in the sediments. In high-
ly fertile regions supply of organic matter is so high that enrichment
of metals can be important (Brongersma-Sanders, 1965). In the Southwest
African muds, which are extremely rich in organic carbon, Ni (35-455
ppm) and Zn (18-337 ppm) are enriched (Calvert and Price, 1971). Cu
and Pb, however, do not show an enrichment compared to other marine
sediments.
Hartmann et al. (1976), who studied sediments from the Northwest African
upwelling region with Corg contents up to 4% did not find increased
contents of Cu and Zn, nor was there any correlation between organic
carbon and content in these metals. They concluded that these elements
are lost by remineralization soon after deposition of the organic de-
tritus. They believe that only in extremely Corg-rich sediments, such
as off Southwest Africa, is a correlation between some minor metals
and organic matter possible.
This interpretation agrees with results of Murty et al. (1970), who

did not find a correlation between Ni and organic carbon contents in
sediments from the shelf off the west coast of India, where upwelling
occurs and where organic carbon contents are high (up to 6% in sedi-
ments investigated for Ni content) .
3.3 Barium
Barium content in equatorial Pacific sediments is greater in the high

fertility zone of equatorial upwelling, than in nonupwelling-influenced
regions (Wolgemuth and Broecker, 1970; Hanor, 1971). Goldberg and Ar-
rhenius (1958) found barium:clay ratios up to 20 times higher in sedi-
265
ments underlying the equatorial Pacific upwelling than in neighboring,

less fertile regions.
Barium is enriched up to 700 ppm in organic matter of planktonic for-

aminifers and up to 5400 ppm in radiolaria Acantharia (Arrhenius, 1963).
3.4 Phosphorite and Uranium
Phosphorite formation is believed to occur in coastal upwelling regions

(e.g., see Kolodny and Kaplan, 1970, for further literature). Phospho-
rite deposits are described from several coastal upwelling regions.
However, it turned out that those off California (Emery, 1960), off
Northwest Mexico (D'Anglejean, 1967) and those off Northwest Africa
(Tooms et al., 1971, Summerhays et al., 1972) are fossil: Cretaceous-
Eocene off Marocco, Lower Pliocene off northern Sahara, Middle Miocene
and Pliocene/upper Pleistocene off California. Their presence in the
actual sediments is linked to local outcrops and reworkins processes.
In the Peruvian and Southwest African upwelling regions actual forma-

tion of phosphorites occurs. Off Peru phosphate nodules were formed
in the region of the upper and lower boundaries of the oxygen minimum
zone, in about 100 and 400 m water depth. The major component is apatite
rich in fluorine. P 20 S contents range from 17 to 35%. The ages corre-
sponding to the 234U:238U ratios are about 300,000 years to recent
(Veeh et al., 1973).
Late Quaternary to recent phosphorite nodules h",ve been found by Batu-

rin et al. (1972) off Chile. Ssaidova (1971) described abundant phos-
phorite nodules off Peru in water depths between 200 and 2000 m, with-
out giving indications as to their age.
Another upwelling region where recent phosphorite formation is known

is the Southwest African shelf (Bezrukov and Senin, 1971; Baturin et
al., 1972; Birch, 1973). Here two generations of phosphorites are de-
scribed by Veeh et al. (1974): lithified black phosphorite pellets and
cream to gray nodules of an age greater than 200,000 years, and soft,
unconsolidated phosphorite blebs and laminae of recent age, found in
the diatomaceous muds. P20S contents of the soft and friable nodules
are 24-28% (Baturin et al., 1972). Bremner (1974a) gives detailed maps
of phosphorite concentrations on the Southwest African shelf. However,
no distinction between recent and fossil phosphorite has been made.
In the recently formed phosphorites off Southwest Africa uranium con-

tents are increased compared to normal pelagic clays and river muds
(Veeh et al., 1974). The 23 4U: 238 U ratios are similar to the seawater
values and thus prove that the f'ixation of uranium occurred simulta-
neously with the sediment depositon. Apatite is probably the mineral
that contains most of the uranium. U contents are 79-158 ppm (Veeh et
al., 1974) and 5-93% (Meyer, 1973) on a salt-free, dry weight basis.
They are significantly higher than in black muds from the Black Sea
or Baltic Sea (12-20 ppm, Meyer, 1973).
Results from Southwest Africa, Gulf of California, and the continental

slope off Peru, where upwelling produces seasonally high organic pro-
duction and thus high sedimentation rates of organic matter, indicate
the importance of upwelling regions for uranium concentration in the
sediments (Veeh, 1967).
The described high contents in organic matter, metals, and phosphorite

make these sediments from upwelling regions interesting for economic
exploration in some areas.
266
4. Biological Indicators of Upwelling
The influence of upwelling on phyto- and zooplankton and on benthos,

on their production and preservation in the sediments, is studied in
this section. Only organisms with skeletons can be preserved in the
sediments and are therefore considered here. Organic remains are studied
here in terms of the chemical composition of their hard parts and not
of their biology (plank~on-benthos, plant-animal), because opal skele-
tons react quite differently than calcium carbonate skeletons during
descent in the water column and in the sediment during diagenesis.
4.1 Opal Skeletons
Three main groups of planktonic organisms build up opal skeletons.

Planktonic diatoms are the most important contributors of opal to the
sediments. They are primary producers and their productivity increases
greatly in fertile regions (e.g., Hart and Currie, 1960; Margaleff,
1973). Radiolarians are less abundant than diatoms. Silicoflagellates
occur together with diatoms in smaller amounts. They are not discussed
separately. (Sponges, benthonic organisms with opal skeletons, will be
mentioned in Sect. 4.2).
The doubling time of diatoms is about one day if growth conditions are
suitable (Berger, 1975); thus, large amounts of opal are contributed
to the bottom. After Heath (pers. corom) 90% of the biogenic opal is
fixed by diatoms.
Biogenous opal produced in the surface water of the oceans is dissolved

in the water column (Berger, 1968; Lisitzin, 1972; Hurd, 1972}, at the
sediment/water interface, and in the sediments (Riedel, 1959; Nigrini,
1968; Berger, 1970a; Schrader, 1972), due to undersaturation of sea-
and interstitial water with respect to opal. So in regions with normal
fertility in the surface waters, all or nearly all opal is dissolved
before being buried in the sediments. If there are some opal skeletons
they are generally badly preserved. When, however, supply of opal to
the bottom becomes sufficient to saturate the interstitial water with
respect to opal, preservation of opaline skeletons in the sediments
is possible (Johnson, 1974; Berger, 1975). This large supply of opal
is possible in upwelling regions, where nutrient-rich deep waters enter
the photic zone. That is why the sea bottom underlying upwelling regions
is covered with opal-rich sediments that exhibit well-preserved tests.
The inner shelf off Southwest Africa between 25 and 18 0 S is covered

by an up to 15-m thick layer of Holocene diatomites (Meyer, 1973).
Diatomites are also described by Bezrukov and Senin (1971); Calvert
and Price (1971); Bremner (1974a) and Diester-Haas (in prep.). Opal
contents are between 53 and 73% (Meyer, 1973) and up to 85% (Calvert
and Price, 1971) of dry sediment. Rogers (1974) gives a list of the
Southwest African diatom assemblages and states that only robust diatom
tests are preserved in the sediments. This agrees with the results of
Hart and Currie (1960) who found a striking difference between bottom
deposits and the plankton.
Off Peru Neaverson (1934), Ssaidova (1971) and Jouse (1972) found dia-
tomites. Ssaidova (1971) also describes considerable amounts of radio-
larians in the sediments from the upwelling region.
In the Gulf of California more than 50 weight percent of total sedi-

ment consists of opal, with considerable amounts of diatoms and radio-
larians (up to more than 50 and 25% respectively in the 62-250 ~m frac-
267
tion). The mean sedimentation rate of opal in this upwelling region

is 200 cm/1000 years (Calvert, 1966). In regions with high terrigenous
input, however, opal contents in weight percent of the sediment do not
indicate the quantity of opal input. Off Oregon (Pacific coast of USA)
opal is accumulating at 50 g/cm 2 /1000 years in this region with strong
upwelling and high organic production. Terrigenous material, however,
dilutes biogenous opal, so that opal concentrations are less than 5%
in general (Heath, 1974).
In the Northwest African upwelling region no quantitative opal content

determinations have been carried out. Most investigated surface sedi-
ments contain only very small amounts of opal skeletons or none at all
(Diester-Haass et al., 1973; Schrader, pers. corom.) , except for a pro-
file off Cape Blanc 20 o N) (Fig. 9) where up to 5% diatoms> 40 ~m were
found, and off Cape Timiris where Miro de Orell (1973) reports up to
9% diatoms and up to 7% radiolarians in the fractions > 44 ~m. The
amount of diatoms in the coarse fractions correlates well with the
amount of biogenous opal in the total sediment (Calvert, 1966).
The highest Holocene sedimentation rates of radiolarians have been

found in the region north of Cape Blanc, thus indicating high opal
supply in this region (Fig. 2; for position of cores see Fig. 3). On
y.,
the upper continental slope at about 23 0 N sedimentation rates are higher
0,1
1000
/;
2000
892
3000
009 .,. Fig. 2. Holocene accumulation rates of sand-sized
radiolarians as a function of water depth of cores
• J off NW Africa. Black line links cores from profil B
829 off Cape Barbas, dotted line those from profil C off
m radiolarians
836 Senegal (for position of cores see Fig. 3)
than in the cores off Senegal. Cores 03 and 28, situated in the center
of the Northwest African upwelling region, have the highest values. The
lower values off Senegal are explained by the fact that upwelling occurs
here only during 5.5 months of the year compared with 12 months off
Cape Blanc (Schemainda et al., 1975).
In the Atlantic (Ellis and Moore, 1973; Goll and Bj¢rklund, 1974) as
well as in the Eastern Pacific (e.g., Nigrini, 1968; Berger, 1970a,
1973 - refer to him for further literature; Moore et al., 1973; Johnson,
1975) the equatorial upwelling is nicely reflected in opal-rich bottom
sediments. However, this is only true when comparing nutrient-rich
waters and bottom sediments on a large scale. There are local variations
due to bottom currents, which easily transport the light diatoms (Moore
et al., 1973), thus disturbing the correspondence between nutrient-ricp
surface waters and opal content in the sediments.
Species distribution of diatoms in upwelling regions does not show

specific assemblages. In coastal upwelling regions neretic species that
support large variations in conditions (temperature, salinity etc.) are
predominant. All subtropical species are present, although surface water
268
,.. Fig. 3. Position of cores taken

during Meteor cruise 25 . C ores
1 and 3 were taken duri ng the
Trans-Atlantic Geotraverse b y
P. Rona and kindly provided by
G. Keller and P. Rona, Miami
... I------j~,____~-......-__f__,'_r_-b"''--------___l ,.
1 •
'.
• 36
..1--HI!~t_V'_::_---_t_--~II;:_-;,,'f_ r-t----
,.. '0'
temperatures are low in coastal upwelling regions (P. Richert, H. -J.

Schrader, Kiel, pers. corom.).
The distribution of biogenous opal showed that high opal content with
large amounts of organic carbon is one of the predominant features of
sediments influenced by upwelling.
4.2 Calcium Carbonate Skeletons
The main contributors of biogenous calcium carbonate to the seafloor

are planktonic foraminifers (marine protozoa), coccolithophores (marine
algae), and benthonic organisms. In upwelling regions some changes as
to species distribution and production could be observed, compared to
surrounding regions without upwelling influence.
The cool water planktonic foraminifer Globigerina bulloides is present in

the upwelling region of the west coast of India (Zobel, 1973), off
Somali (Be and Tolderlund, 1971), off Morocco (Thiede, 1973), and off
Southwest Africa (Be and Tolderlund, 1971 ; Rogers, 1974). Normally this
species is absent in these subtropical regions. It lives predominantly
in subarctic and arctic waters. In the Californian upwelling region
the cold-adapted foraminifer species Globigerina quinqueloba occurs in
large quantities during bloom situations produced by upwelling. Thus
the foraminiferal assemblages in the sediment have a cold water aspect
269
(Berger, 1975). On the Southwest African shelf (30-25 0 S) G. quinqUBZoba

is the most abundant, up to 33%, of total foraminifer species (Rogers,
1974) .
100 1000· 10-3 cml10 3yr

,
., 0
r
1000 0 0/
I
I
2000
·· ~/.
·•
I
09 028
·
3000
I
I Fig. 4. Holocene accumulation rates of sand-sized planktonic
foraminifers as a function of water depth of cores off NW
o.
029
Africa. Black line links cores from profil B, dotted line
m 036 plankt. foram. those from profil C (for position of cores see Fig. 3)
With increasing fertility the abundance of foraminifers (in numbers

per m3 water) increases (Berger, 1975). This is reflected in Holocene
sedimentation rates of foraminifers on the upper Northwest African con-
tinental slope (Diester-Haass, 1976) (Figs. 4 and 3 for position of
cores). Off Senegal accumulation rates are an order of magnitude smaller
on the upper slope than off Cape Barbas. As for radiolarians this dif-
ference is attributed to different duration of upwelling influence.
No data were available on coccolithophore species and productivity re-
flected in sediments from upwelling regions. These algae are known to
reach high growth potentials at lower nutrient concentrations. With in-
creasing fertility their numbers (in m3 water) increase only slightly
and to a lesser degree than those of planktonic foraminifers (Berger,
1975). Plankton tows by Margaleff (1973) revealed that coccolithophores
are slightly increased in the waters north of the center of the North-
west African upwelling region. However, technical problems are important
due to small size and dissolution of the calcareous tests.
Life conditions for benthos are improved in upwelling areas compared
to surrounding nonupwelling influenced regions, because the food supply
to the bottom increases as a consequence of high plankton production
(see Thiel, this vol.). In the oxygen minimum zone, however, described
for some parts of the upwelling area off Southwest Africa, Peru, Cali-
fornia, and the west coast of India (see Sect. 2), benthic life is im-
possible, and this is the zone of laminated sediments.
Some analyses of species and density of benthos in upwelling regions
have been carried out (see Thiel, this vol.). However, for geologists
only those organisms that have preservable skeletons can give indica-
tions as to upwelling influence.
In the Northwest African upwelling region the accumulation rates of
four groups of benthonic organisms during Holocene have been calculated.
It is presumed that during this period of about 10,000 years, hydrologic
conditions did not differ too much from the present ones. It can be
seen in Figure 5 that benthic foraminifers, sponges (opal skeletons),
ostracods, and echinoids have higher accumulation rates on the upper
continental slope above about 1500 m off Cape Barbas than off Senegal.
This corresponds to the results obtained for planktonic foraminifers
and radiolarians and is interpreted in the same way: different duration
270
j:;/
10
1000
/"j
2000
92 0
·09 ,-28
3000 .0'
029 29
.1
m 036 benth. foram. m .36 sponges
0,01 0,1 10'11)-3 cml103 yr

~~~~~~~~~~~
1000 1000 .I
03
2000
092
o 09
3000
829
'1
m 036
Fig. 5. Holocene accumulation rates of sand-sized benthonic foraminifers, sponges,

echinoids, and ostracods as a function of water depth of cores. Black line links
cores from profil B, dotted line those from profil C (for position of cores see Fig.3)
of upwelling influence off Cape Barbas and off Senegal (Diester-Haass,

1976 - here accumulation rates are also given in g/cm 2 /1000 years) .
This agrees with the results of Thiel (this vol.), who describes higher
macrobenthos standing stock in upwelling regions compared to less fer-
tile regions.
Only little is known of the influence of high fertility in upwelling

regions on species composition, diversity, and size of skeletons of
benthic organisms. Rogers (1974) lists molluscan species on the South-
west African shelf (25-30 0 S).
4.3 Plankton-Benthos Ratios of Foraminifers
Planktonic and benthic foraminifers are sedimented more rapidly in

strongly upwelling-influenced regions than in regions with less upwel-
ling influence, as can be seen in Figures 4 and 5. However, when form-
ing the ratio of both groups, one can see that benthic foraminifers
increase more than planktonic ones in fertile regions. In Figure 7D
the plankton:benthos ratios of foraminifers in the Northwest African
upwelling area are plotted against water depth of the surface samples.
The profile off cape Blanc, where strongest upwelling is indicated by
the highest organic carbon.and opal contents, the ratios are much smal-
ler, i.e., richer in benthos than in samples from the same water depth
in the northern profile.
271
4.4 Radiolarian:Planktonic Foraminiferal Ratio
Planktonic foraminifer production and thus their accumulation increase

in fertile regions, as shown in Figure 4 and in Section 4.2. The in-
crease of radiolarians in upwelling-influenced sediments is much more
important since they are nearly absent in normal sediments because of
dissolution. Thus, the ratio of radiolarians to planktonic foraminifers
should increase in highly fertile regions above the CaC03 lysocline.
Figure 6 shows this ratio in the surface samples from the Northwest
African upwelling region. In the most fertile region off Cape Blanc
and off Cape Timiris the highest ratios occur on the upper continental
slope, and to the north the ratios decrease. Off Morocco (33 0 N) no
radiolarians were found in the fractions > 63 vm in the surface samples
(Kudrass,1973).
( ()
()
()
..,
Fig. 6. Map of radiolarian:planktonic

foraminiferal ratios in surface sedi-
ments from the NW African upwelling
region. See legend to Figure 2. White:
no radiolarians in surface sediments
272
The formation of a ratio of the two components avoids errors in inter-

pretation due to dilution by other components. When plotting only per-
cent values of radiolarians or foraminifers there may be considerable
variations dependent on the other components.
4.5 Fish Debris
The high production of phyto- and zooplankton in upwelling regions

leads to high fish production. Their remains - otolithes, scales, bones
- are found in the sediments from upwelling regions in much higher con-
centrations than in regions not influenced by upwelling. Frankenberg
and Menzies (1968) describe sediments from the Peru upwelling region
in 126 m water depth, which were composed largely of fish bones and
cycloid scales. Ssaidova (1971) found off Peru south of 22 0 S a fish-
bone-foraminiferan facies in water depth between 200 and 400 m. At
depths between 30 and 250 m he found mainly diatoms and fish remains.
Down to 600-m water depth he found fish debris in decreasing amounts.
Also off Southwest Africa .fish remains form substantial parts of the
upwelling-influenced sediments, mainly of the diatomites (Calvert and
Price, 1971; Meyer, 1973; Diester-Haass, in prep.).
Off Northwest Africa no significant increase in fish remains could be

observed except off Cape Timiris, where Miro de Ore 11 (1973) describes
2-3% fish debris in fractions coarser than 40 ~m in those sediments
where organic carbon, diatom, and radiolarian contents are high.
4.6 Calcium Carbonate Preservation
The calcite compensation depth, that is, the depth where dissolution
of calcite is as high as supply of calcite, rises in pericontinental
regions where fertility is greater than in oceanic regions (Berger,
1970a; Lisitzin, 1971, 1972). This can be explained by higher rates
of supply of organic matter to the sediments. This causes increased
benthic activity and development of C02-rich interstitial waters that
attack calcium carbonate skeletons (Berger, 1970b, 1974). Moore et ale
(1973) found a rise in the calcite compensation depth of 650 m in the
highly fertile near-coast region of the Panama Basin compared to the
basins underlying less fertile water masses. Berger (1970a) indicates
that in general foraminiferal assemblages from highly fertile regions
are enriched in forms resistant to solution.
These observations indicate thclt in upwelling regions preservation of

calcium carbonate should be poor. However, in anaerobic sediments under-
lying highly fertile water masses, excellent preservation of calcite
shells has been found: on the continental slope off the west coast of
India in the 02 minimum zone (Stackelberg, 1972) and in the Santa Bar~
bara Basin off California (Berger and Soutar, 1970). Berner et ale
(1970) and Hartmann et ale (1973, 1976) found that in reducing sedi-
ments bacterial sulfate reduction leads to high carbonate alkalinity,
which allows even carbonate precipitation in the interstitial water.
It seems that the presence or absence of an oxidizing layer with

benthos activity above the sulfate reduction zone is essentially re-
sponsible for carbonate preservation. If the reduction zone is in con-
tact with the bottom water, calcite preservation will be good. If an
oxidizing layer with benthic activity is present, calcite will be less
well preserved.
273
In the equatorial Pacific the high fertility leads to a depression of

the calcium carbonate compensation depth. This apparent contradiction
can be explained by the much lower supply of reactive organic matter
to the bottom and smaller amounts of benthos at the great depths (Ber-
ger, 1973).
5. Two Examples of Holocene Upwelling-Influenced Sediments
5.1 Continental Slope off Northwest Africa
Two examples have been chosen to show the good correlation between some
of the parameters presented here as indicators of upwelling.
or~C 100 98 96 ./. pl for. diatoms plankt.l benth. for am.

2 3 4 'I. o 2 4'10 rad . 2 ''10 10 30 50 70
......~
.~
1000
~ .
•
A B c o
Fig. 7A-D. Some indicators of upwelling influence in surface sediments from the NW
African continental margin plotted versus water depth of samples. Dots: samples from
Meteor 25 expedition; Crosses: profile 1 from Meteor 36 expedition; Squares: profile
2 from Meteor 36 expedition; Triangles in Figure 9: profile 3 from Meteor 36 expedi-
tion situated 33 0 N off Morocco. For position of profiles see Figure 10. (A) Percent
organic carbon of total sediment; (B) Radiolarian:planktonic foraminiferan ratios;
(C) Percent diatoms in fractions > 40 ')lm; (D) Plankton:benthos ratio of foraminifers.
The four dots without numbers are from the profile off Cape Barbas
Figure 7 shows the results obtained on surface samples (for position

of the samples see Fig. 8). In Figure 7A the amount of organic carbon
is plotted against water depth. The profile off Cape Blanc has its
highest values in water depths between 1000 and 3000 m. At the same
depth radiolarian:planktonic foraminiferal ratios (Fig. 7B) are highest
off Cape Blanc, whereas most of the other samples in the northern pro-
files have no radiolarians at all, except those samples that are situ-
ated closest to the center of the upwelling region off Cape Blanc
(samples 3, 25, 26). Diatoms are absent in all samples in the coarse
fractions, except for the profile off Cape Blanc, where they occur in
amounts of up to 5% in fractions > 40 )lm between 1000- and 3000-m water
depth (Fig. 7C). Plankton:benthos ratios of foraminifers also show the
maximum fertility off Cape Blanc, where the ratios are much smaller
(i.e., richer in benthos) than in the northern region.
5.2 Shelf off Southwest Africa
The second example concerns a sediment core from the Southwest African
shelf off Walvis Bay, taken in 118-m water depth, in 22 0 30'S, 13 0 45'E,
274
Fig. 8. Position and numbers of the in-

vestigated samples off NW Africa. Dots,
cores from Meteor 25 expedition; Crosses,
profile 1 from Meteor 36 expedition;
Squares, profile 2 from Meteor 36 expe-
dition
24'
20' 20'
IS'
16
20' 16'
during "Campagne Walda" of the Centre Oceanologique de Bretagne, Brest.
The core has kindly been provided by L. Pastouret, Brest. The core
was taken in the region where Calvert and Price (1971) reported highest
organic carbon values (up to 26%). The upper meter of the core consists
of coarse skeletal debris (less than 10% < 40 ~m; calcium carbonate
contents 90%; organic carbon less than 1%; no opal skeletons; no fish
debris) (Fig. 9).
In 120-cm core depth there is a strikingly sharp boundary. Black mud

with some white molluscan shells, horizontally arranged, underlies the
coarse molluscan shell sand. Material < 40 ~m increases up to 80%,
CaC03 decreases to about 30%, organic carbon values are 4-8%, vertebrate
debris forms 2-5% of the sand fraction. No diatoms, however, have been
found in the fractions > 40 ~m. Smear slides reveal diatoms in the fine
fractions. Diatoms coarser than 40 ~m appear only in the section between
275
CaC~ ~1.lom 5 100 92"1.,,110<

20 60 100"1. 20 60 'I. 0 8'1. l ad
r---~~~
J.'----,__
> 63 ~m
---- 40-63)Jm
c d e 9
Fig. 9a-g. Core Walda 07 from the SW african upwelling region, 22 0 30'S, 130 45'E,
11S-m water depth. (a) Type of sediment: 1, coarse, yellow molluscan sand; 2, black
mud with white, thin-walled, horizontally arranged bivalve shells; 2, olive-green
diatomaceous mud; ~, sand-sized phosphorite nodules and coarse broken molluscan
shells; (b) material < 40 ~m in percent of total sediment; (c) calcium carbonate
content of total sediment; (d) percent organic carbon of total sediment; (e) percent
diatoms in 40-63 and> 63 ~m fractions; (f) radiolarian:planktonic foraminiferal ratio;
(g) percent fish debris in fractions > 63 ~m
200 and 290 cm, an olive-green finely laminated diatomite. Material

< 40 ~m forms 95% and smear slides showed that it consists mainly of
diatoms. The CaC03 content is 20%, organic carbon values are 6-10%;
diatoms from up to 70% of the sand, and 90% of the 40-63 ~m fractions.
Radiolarians, absent in the rest of the core, appear in this section.
The ratio of radiolarians to planktonic foraminifers increases up to
13% (vs. 87% planktonic foraminifers), indicating that this ratio can
be an indicator of high opal production and thus also of higb fertility
in shelf environments. Fish debris, mainly scales and bones and some
otolithes, form up to 12% of the sand fraction. This section is inter-
preted as being a typical upwelling-influenced sediment.
Between 290 and 340 cm there is again black mud and olive-green laminat-
ed sediment. The lower part of the core consists of black-brown, well-
rounded, polished, and well-sorted phosphorite grains and molluscan
shell debris. The age of the phosphorite grains has been determined
by U isotopes to be older than 500,000 years (A. Mangini, Heidelberg,
pers. corom.).
The numerous facies changes in this core, which occur abruptly without
transition zones, point to high variability in sedimentation processes
on the Southwest African shelf, which have also been stressed by Brem-
ner (1974b). It is not yet known whether these facies changes are due
to changes in the position of the upwelled water masses or to bottom
currents and rapidly varying topography.
276
6. Position of Upwelling Regions in Past Geologic Periods
The knowledge on fossil upwelling-influenced sediments is sparse. The

following examples are tentative interpretations.
6.1 Quaternary off Northwest Africa
The Northwest African upwelling, which is reflected most distinctly in

surface sediments from the upper continental slope between Cape Blanc
and Cape Timiris (19-21 o N), was intensified during the Wlirm and Riss
glacial periods and extended as far north as 27 o N. Its effect was ob-
served in cores from the upper continental slope (for position of cores
see Fig. 3), in which organic carbon contents were increased by a fac-
tor of 4-9 compared to the Holocene (MUller, 1975) (see Fig. 10). Di-
atoms, nearly absent in the Holocene core sections, increase by up to
0.001 0.D1
79 Fig. 10. Accumulation rates of organic car-
bon in three cores from the NW African con-
2500 92 tinental slope during Holocene (dots), Wurm
eo x (crosses), and Riss/Wurm Interglacial
(squares), plotted as a function of water
3000 10 o e X depth of cores. Values from Muller (1975).
m org. C For position of cores see Figure 3
250 ml wet sediment (Schrader in Diester-Haass et al., 1973); sedimenta-

tion rates of radiolarians and benthonic organisms are greatly increased
in cores 09, 10, 92 (Figs. 11, 12); radiolarian:planktonic foraminiferal
ratios are high (Fig. 13) (Diester-Haass, 1977).
0,1 10 100· '0-3 cml10 3yr

Fig. li. Accumulation rates of radio-
2500 92 • 0 x larians in five cores from the NW
09· 0 x African continental slope during
Holocene (dots), Wurm (crosses), and
10 x
"" Riss/Wurm Interglacial (squares),
29 0 • x plotted as a function of water depth
3500 1 oe of cores. For position of cores see
m radiolarians Figure 3
10 10010-3 cmll0 3yr

2500 92 x
09 • 0
""
l( Fig. 12. Accumulation rates of total benthos in
five cores from the NW African continental slope
10 De X during Holocene (dots), Wurm (crosses), and Riss/
29 Dol( Wurm Interglacial (squares), plotted as a func-
3500 01( D tion of water depth of cores. For position of
m total benthos cores see Figure 3
Mangini (1975) found increased sedimentation rates of authigenic ura-

nium in the WUrm section of core 10, which he attributes to increased
organic production.
277
100 96 92 88 100 96 92 88.,. pi. for.

0 4 8 12 0 4 8 12.,. rad.
25
1000
26
3
2000
79
92
28 09
.....
t 3000 10
29
!J m 36
HOLOCENE core no. WORM
Fig. 13. Holocene and warm radiolarian , planktonic foraminiferal ratios plotted versus
water depth of cores off NW Africa. For position of cores see Figure 3
6.2 Tertiar¥ off Northwest Africa

The early-to-middle Miocene sediments from the continental slope off
Northwest Africa (deep-Sea Dilling Site 139, 140 from leg 14) contain
abundant, well-preserved siliceous fossils. Flora of diatoms and other
siliceous algae remains were varied (Berger and von Rad, 1972). Perhaps
these Miocene siliceous-rich sediments, which were deposited at a paleo-
latitude of about 15 0 N (Berger and von Rad, 1977), are due to upwelling
influence.
6.3 Tertiary in ~he Equatorial Pacific

The equatorial upwelling in the Eastern Pacific is reflected by a typ-
ical facies distribution: calcareous-siliceous facies and lowered cal-
cium carbonate compensation depth (see Sect. 4.6) due to high organic
production below the equator. Siliceous facies is found below the cal-
cium carbonate compensation depth. North and south of this fertile zone
dissolution or nondeposition occurs (see Berger, 1973, for further
literature) •
Berger (1973) traced the position of this fertile equatorial zone back
through the Tertiary. The same facies distribution has existed for the
past 40 million years, but the position of the sediments reflecting
high fertility has shifted from about 100 S during Late Eocene, to about
5 0 S in Early Miocene, up to 0 0 today.
6.4 Cretaceous in Northwest Africa

In the upper Cretaceous coastal basin of southern Morocco, sediments
were found that indicate upwelling influence during their deposition.
Thinly laminated marls contain a cool, boreal ammonite fauna, fish re-
mains, and chert. PzOs and other heavy metals are present in large
amounts and organic carbon constitutes up to 8% (Einsele and Wiedmann,
1975) •
278
7. Conclusions (Fig. 14)
The high plankton production in upwelling regions leads to large amounts

of organic carbon buried in the sediments. Minor metals may perhaps be
UPWELLED WATE R
I
WATER
1
cool
1
high plankton production
1
rich in P04
m,"", L1 ol~Ll Phl~ ~I-.----11

cold-water
assemblages
of planktonic
foraminifers.
Fig. 14. Scheme showing pro-
perties of water masses and
influence on sediments in up-
welling regions
(metals) uraJium
enriched in sediments with high organic carbon contents. Diatoms, the

most important primary producers with preservable (opal) skeletons, are
abundant in upwelling-influenced sediments. Thus, opal content is an
important upwelling indicator. Benthos production, and thus sedimenta-
tion rates of skeletons of benthic organisms, is increased except in
the oxygen minimum zone. Plankton:benthos ratios of foraminifers can
be decreased compared to sediments from the same water depth without
upwelling influence. Fish remains are abundant. Phosphorite formation,
mainly in diatomaceous muds rich in organic carbon, occurs. The cool
temperatures of the surface waters are reflected in cold-water assem-
blages of planktonic foraminifers.
All these parameters have to be studied in sediments from upwelling

regions and compared with neighboring sediments deposi~ed without up-
welling influence, in comparable water depths and in approximately the
same latitude and the same climatic regime. Typical properties of up-
welling-influenced sediments can only be found by this comparison.
Acknowledgments. I wish to express my sincere thanks to M. Hartmann, P. Muller, P.

Richert, H.-J. Schrader, E. Seibold, E. Suess (Kiel), H. Thiel (Hamburg), and G.
Einsele (TUbingen) for fruitful discussions and to J. Rogers (Cape Town) for pro-
viding information on unpublished investigations . .1 thank G. Keller and P. Rona
(Miami), L. Pastouret (Brest) and N.B. Price (Edingburgh) for providing core material
from upwelling regions. The financial support of the Deutsche Forschungsgemeinschaft
is gratefully acknowledged.
Note: This paper has been submitted in November 1975. No literature and results ob-
tained after this date could be included.
References
Arrhenius, G.: Pelagic sediments. In: The Sea. Ideas and Observations on Progress
in the Study of the Seas. Hill, M.N. (ed.). New York-London-Sydney: John Wiley,
1963, Vol. III
Bandy, O.L., Rodolfo, K.S.: Distribution of foraminifera and sediments, Peru-Chile
Trench area. Deep-Sea Res. 11, 817-837 (1964)
Baturin, G.N., Merkulova, K.I., Chalov, P.I.: Radiometric evidence for recent forma-
tion of phosphatic nodules in marine shelf sediments. l'Iar. Geol. !l, M37-M41 (1972)
Be, A.W.H., Tolderlund, D.S.: Distribution and ecology of living planktonic fora-
minifera in surface waters of the Atlantic and Indian Ocean. In: Micropaleontology
279
of Oceans. Funnell, B.M., Riedel, W.R. (eds.). London: Cambridge Univ., 197£, pp.
105-149
Berger, W.H.: Radiolarian skeletons: solution at depth. Science 122, 1237-1238
(1968)
Berger, W.H.: Biogenous deep-sea sediemnts: fractionation by deep-sea circulation.
Geol. Soc. Am. Bull. ~, 1385-1402 (1970a)
Berger, W.H.: Planktonic foraminifera: selective solution and the lysocline. Mar.
Geol. ~, 111-138 (1970b)
Berger, W.H.: Cenozoic sedimentation in the eastern tropical Pacific. Geol. Soc.
Am. Bull. 84, 1941-1954 (1973)
Berger, W.H.: Deep-sea sedimentation. In: The Geology of Continental Margins. Burk,
C.A., Drake, D.L. (eds.). New York-Heidelberg-Berlin: Springer, 1974, pp. 213-241
Berger, W.H.: Biogenous deep-sea sediments. In: Treatise on Chemical Oceanography.
Riley, J.P., Chester, R. (eds.). London: Academic Press, 1975
Berger, W.H., von Rad, U.: Cretaceous and cenozoic sediments from the Atlantic Ocean.
In: Initial Reports of the Deep-Sea Drilling Project. Hayes, D.E., Pimm, A.C.
(eds.). Washington: 1972, Vol. XIV, pp. 787-954
Berger, W.H., Soutar, A.: Preservation of plankton shells in an anaerobic basin off
California. Geol. Soc. Am. Bull. ~, 275-282 (1970)
Berner, R.A., Scott, M.R., Thomlinson, C.: Carbonate alkalinity in the pore waters
of anoxic marine sediments. Limnol. Oceanogr. 15/4, 544-549 (1970)
Bezrukov, P.L., Senin, K.M.: Sedimentation on the west African shelf. -ICSU/SCOR
Working Party 31, Symp. Cambridge, 1970. Inst. Geol. Sc. 70/16, 3-7 (1971)
Birch, G.F.: Unconsolidated sediments off the Cape West coast. - Joint Geol. Surv./
Uni v. Cape Town. Mar. Geol. Progr., Tech. Rep. ~ (Cape Town), 48-66 (1973)
Birch, G.F., Rogers, J.: Nature of the sea floor between Luderitz and Port Elizabeth.
S. African Shipping News Fish. Industry Rev. ~, (1973)
Bost~om, K., Joensun, 0., Brohm, J.: Plankton: its chemical composition and its sig-
nific'ance as a source of pelagic sediments. Chern. Geol • .!!' 255-271 (1974)
Bremner, J.M.: Texture and composition of surficial continental margin sediments be-
tween the Kunene River and Sylvia Hill. SWA.-Joint Geol. Surv., Univ. Cape Town.
Mar. Geol. Progr., Techn. Rep. 6, 39-43 (1974a)
Bremner, J.M.: Further analyses o~ the Sardinops gravity cores·collected from the
shelf near Walvis Bay, South West Africa.-Joint Geol. Surv., Univ. of Cape Town,
Mar. Geol. Progr., Techn Rep. ~, 55-59 (1974b)
Brongersma-Saunders, M.: Metals of Kupferschiefer supplied by normal sea water. Geol.
Rdsch. E, 365-375 (1965)
Calvert, S.E.: Accumulation of diatomaceous silica in the sediments of the Gulf of
California. Geol. Soc. Am. Bull. 77, 569-596 (1966)
Calvert, S.E., Price, N.B.: Recent sediments of the South-West African shelf. - ICSU/
SCOR Working Party 31, Syrup. Cambridge, 1970. Inst. of Geol. Sc. Rep.>70/16, 175-
185 (1971)
Copenhagen, W.J.: The periodic mortality of fish in the Walvi'S region: a phenomenon
within the Benguela current. Invest. Rep. Div. Fish. U.S.A. 14, 1-35 (1953)
d'Anglejan, B.F.: Origin of marine phosphorites off Baja California, Mexico. Mar.
Geol. ~, 15-44 (1967)
Davey, R.J., Rogers, J.: Palynomorph distribution in recent offshore sediments along
two traverses off South West Africa. Mar. Geol. ~, 213-225 (1975)
Diester-Haass, L.: Sedimentation and climate in the Late Quaternary between Senegal
and the Cape Verde Islands. Meteor Forsch. Ergebn. 20C, 1-32 (1975)
Diester-Haass, L.: Quaternary accumulation rates of biogenous and terrigenous com-
ponents on the East Atlantic continental slope off NW Africa. Mar. Geol. ~, 1-24
(1976)
Diester-Haass, L.: Radiolarian/planktonic foraminiferal ratios in a coastal upwelling
region. J. Foram. Res. 2, 26-33 (1977)
Diester-Haass, L.: Sediments from the south West African upwelling region. (in prep.)
Diester-Haass, L., Schrader, H.-J., Pflaumann, U., RUbl, N., Thiede, J.: Auftriebs-
einfluB in Sedimenten. Nachrichten Deut. Geol. Ges. 9, 39-44 (1974)
Dieser-Haass, L., Schrader, H.J., Thiede, J.: Sediment;logical and paleoclimatological
investigations of two pelagic ooze cores off Cape Barbas, North West Africa. Meteor
Forsch. Ergebn. 16C, 19-66 (1973)
280
Einsele, G., Wiedmann, J.: Faunal and sedimentological evidence for upwelling in the
upper cretaceous coastal basin of Tarfaya/Morocco. IX ieme Congres International
de Sedimentologie, Nice, 1975. Vol. des Rapports, No. 1 (1975)
Ellis, D.B., Moore, T.C.: Calcium carbonate, opal, and quartz in Holocene pelagic
sediments and the calcite compensation level in the South Atlantic Ocean. J. Marine
Res. 11-Jll., 210-227 (1973)
Emery, K.O.: The Sea off Southern California. New York-London: John Wiley, 1960,
366 pp
Frankenberg, D., Menzies, R.J.: Some quantitative analyses of deep-sea benthos off
Peru. Deep-Sea Res. ~, 623-626 (1968)
Goll, R.M., Bjorklund, K.R.: Radiolaria in surface sediments of the South Atlantic.
Micropaleontology 20 (1), 38-75 (1974)
Goldberg, E.D., Arrhenius, G.: Geochemistry of Pacific pelagic sediments. Geochim.
Cosmochim. Acta 13, 153-212 (1958)
Hanor, J.S.: Rates-C>f barium accumUlation in the equatorial Pacific. Abstracts with
programs 1972, Annual Meetings of the Geol. Soc. of Amer.: 2526
Hart, T.J., Currie, R.J.: The Benguela current. Discovery Rep. ]1, 123-298 (1960)
Hartmann, M., Miiller, P., Suess, E., van der Weijden, C.H.: Oxidation of organic
matter in recent marine sediments. Meteor Forsch. Ergebn. 12 C, 74-86 (1973)
Hartmann, M., Miiller, P., Suess, E., van der Weijden, C.H.: Chemistry of Late Qua-
ternary sediments and their interstitial waters from the NW African continental
margin. Meteor Forsch. Ergebn. 24 C, 1-67 (1976)
Heath, G.R.: Dissolved silica and deep-sea sediments. In: Studies in Paleo-Oceano-
graphy. Hay, W.W. (ed.). Soc. Econ. Pal. and Min., Spec. Publ., 20, 77-93 (1974)
Hoflich, 0.: Die meteorologischen Wirkungen kalter Auftriebswassergebiete. Geoforum
11/72, 35 (1972)
Hurd, D.C.: Interactions of biogenic opal, sediment, and sea water in the central
Equatorial Pacific. PhD. Thesis, Hawaii Inst. of Geophys., Univ. of Hawaii, HIG-
72-22, 1972, 81 pp
Johnson, T.C.: The dissolution of siliceous microfossils in surface sediments of
the eastern tropical Pacific. Deep-Sea Res. 21, 851-864 (1974)
Johnson, T.C.: The dissolution of siliceous mi~ofossils in deep-sea sediments. PhD
Thesis, Univ. of California, San Diego 1975, 163 pp
Jouse, A.P.: Diatoms in the surface sediment layer of the Chilian-Peruvian region
of the Pacific Ocean. Oceanologiia 12 (5), 831-841 (1972) (Moscow, in russ.)
Kolodny, Y., Kaplan, J .R.: Uranium isotopes in sea-floor phosphorites. Geochim.
Cosmochim. Acta 34, 3-24 (1970)
Kudrass, H.-R.: Sedimentation am Kontinentalhang vor Portugal und Marokko im Spat-
pleistozan und Holozan. Meteor Forsch. Ergebn. 13 C, 1-63 (1973)
Lisitzin, A.P.: Distribution of siliceous microfossils in suspension and in bottom
sediments. In: Micropaleontology of Oceans. Funnell, B.M., Riedel, W.R. (eds.).
London: Cambridge, Univ., 1971, pp. 173-195
Lisitzin, A.P.: Sedimentation in the world ocean. Soc. Econ. Pal. Min., Spec. Publ.
17, 218 pp (1972)
Mangini, A.: Thorium- und Uran-Isotopenanalysen an Tiefseesedimenten. Unpubl. manu-
script (1975) 16 pp
Marchand, J.M.: The nature of the sea-floor deposits in certain regions off the South
African coast. S. Afr. J. Sci. 61, 444-448 (1928)
Margaleff, R.: Fitoplancton marino de la region de affloramiento del NW de Africa.
Res. Exp. Cient. B/O Cornide l, 65-94 (1973)
Meincke, J., Mittelstaedt, E., Huber, K., Koltermann, K.P.: Currents and stratifica-
tion in the upwelling area off NW Africa. Deut. Hydr. Inst. Meereskundl. Beob.
Ergebn. ±l, 1-117 (1975)
Meyer, K.: Uran-Prospektion vor Siidwestafrika. Erzmetall 26 (7), 313-317 (1973)
Miro Orell, M. de: Sedimentos recientes del margen continental de Mauretania (expe-
dicion Sahara II). Res. Exp. Cient. B/O Cornide 2, 1-12 (1973)
Moore, T.C., Ross Heath, G., Kowsmann, R.O.: Bioge-;:;-ic sediments of the Panama Basin.
J. Geol. g, 458-472 (1973)
Muller, P.: Zur Diagenese stickstoffhaltiger Substanzen in marinen Sedimenten unter
oxydierenden und reduzierenden Bedingungen. Diss. Univ. Kiel, 1975, 1/9 pp
Murty, P.S., Reddy, C.V.G., Varadachari, V.V.R.: Distribution of organic matter in
the marine sediments off the West coast of India. Proc. Natl. Inst. Sci. India
35 B (5), 377-384 (1969): Reprinted in UNESCO Ind. Ocean Exped. ~, 741-748 (1969)
281
Murty, P.S.N., Rao, Ch.M., Reddy, C.V.G.: Distribution of Nickel in the marine sedi-
ments off the West coast of India. Curro Sci. 39 (2), 30-32 (1970). Reprinted in
UNESCO Ind. Ocean. Exped. ~, 737-740 (1970)
Neaverson, E.: The sea-floor deposits. 1) General characters and distribution. Dis-
covery Rep. ~, 295-350 (1934)
Newton, R.S., Seibold, E., Werner, F.: Facies distribution patterns on the Spanish
Sahara continental shelf mapped with side-scan sonar. Meteor Forsch. Ergebn. ~,
55-77 (1973)
Nigrini, C.A.: Radiolaria from Eastern tropical Pacific sediments. Micropaleontology
l±, 51-63 (1968)
Rhyther, J.H., Menzel, D.W., Hulburt, E.M., Lorenzen, C.J., Corvin, N.: The produc-
tion and utilization of organic matter in the Peru coastal current. Working Con-
ference IIP/IBP/PM, Barcelona, 1970. Invest. Pesquera ~, 43-59 (1971)
Richert, P.: Die raumliche Verteilung und zeitliche Entwicklung des Phytoplanktons,
mit besonderer Berucksichtigung der Diatomeen im N.W.-Afrikanischen Auftriebswas-
sergebiet. Dissertation Univ. Kiel, 1975, p. 140
Riedel, W.R.: Siliceous organic remains in pelagic sediments. Silica in Sediments.
Soc. Econ. Pal. Min., Spec. Publ. 2, 80-91 (1959)
Rogers, J.: Surficial sediments and tertiary limestones from the Orange-Luderitz
shelf. Joint Geol. Surv., Univ . Cape Town, Mar. Geol. Progr., Techn. Rep. §.' 24-38
(1974)
Schemainda, R., Nehring, D., Schulz, S.: Ozeanologische Untersuchungen zum Produk-
tionspotential der nordwestafrikanischen Wasserauftriebsregion 1970-1973. Geod.
Geoph. Veroff. IV, ~, 1-88 (1975)
Schrader, H.-J.: Kieselsaureskelette in Sedimenten des ibero-marokkanischen Kontinen-
talrandes und angrenzender Tiefsee-Ebenen. Meteor Forsch. Ergebn. ~ 10-36
( 1972)
Seibold, E.: Der Meeresboden als Rohstoffquelle und die Konzentrierungsverfahren der
Natur. Chemie-Ing.-Technik 42 (23), A 2091-2103 (1970)
Smith, R.L.: Upwelling. Oceanogr. Mar. Bioi. Ann. Rev. §.' 11-46 (1968)
Soutar, A.: Micropaleontology of anaerobic sediments and the California current. In:
The Micropaleontology of Oceans. Funnel, B.M., Riedel, W.R. (eds.). London: Cam-
bridge Univ., 1971, pp. 223-230
Ssaidova, H.M.: Recent sediments off the Pacific coast of South America. Ac. of
Science USSR. Invest. P.P. Shirschov-Inst. of Oceanologie 89, 139-145 (1971) (in
russ.)
Stack~lberg, U. von: Faziesverteilung in Sedimenten des indisch-pakistanischen Kon-
tinentalrandes (Arab. Meer). Meteor Forsch. Ergebn. 9 C, 1-73 (1972)
Summerhayes, C.P.: South West African shelf sediments.~uth African Nat. Comm. for
Oceanogr. Res. Mar. Geol. Progr., Techn. Rep. !, 94-102 (1972)
Summerhayes, C.P., Nutter, A.H., Tooms, J.S.: The distribution and origin of phos-
phate in sediments off northwest Africa. Sed. Geol. ~, 3-28 (1972)
Thiede, J.: Sedimentation rates of planktonic and benthonic foraminifera in sediments
from the Atlantic continental margin of Portugal and Morocco. Meteor Forsch. Ergebn.
16 C, 1-18 (1973)
Thiel, H.: Benthos in upwelling regions. This volume, pp. 124-138
Tooms, J.S., Summerhayes, C.P., McMaster, R.L.: Marine geological studies on the
north-west African margin: Rabat-Dakar. ICSU/SCOR Working Party 31, Symp. Cambridge,
1970. Inst. of Geol. Sc., Rep. 70 (16), 9-25 (1971)
Veeh, H.H.: Deposition of uranium from the ocean. Earth Planetary Sci. Lett. ~,
145-150 (1967)
Veeh, H.H., Burnett, W.C., Soutar, A.: Contemporary phosphorites on the continental
margin of Peru. Science lQl (4102), 844-845 (1973)
Veeh, H.H., Calvert, S.E., Price, N.B.: Accumulation of uranium in sediments and
phosphorites on the South ~est African shelf. Marine Chern. 2, 189-202 (1974)
Wolgemuth, K., Broecker, W.: Barium in sea-water. Earth Planetary Sci. Lett. ~,
372-378 (1970)
Zobel, B .. : Biostratigraphische Untersuchungen an Sedimenten des indisch-pakistanischen
Kontinentalrandes (Arabisches Meer). Meteor Forsch. Ergebn. 12 C, 9-73 (1973)
Sedimentation Influenced by Upwelling in the Subtropical Baie
Du Levrier (West Africa)
B. KOOPMANN, M. SARNTIIEIN, and H.-J. SCHRADER
Marine sediments rich in opal are believed to reflect prolific primary

production in the photic zone. This assumption can best be demonstrated
in offshore upwelling areas such as in the eastern subtropical Atlantic
(Fig. 1).
Z
<t
W
U
0
....='
Z
<t
....
-I
<t . .<Sf'
Fig. 1. Generalized bathymetry off Northwest Africa

283
Despite the fact that high primary production was observed in the up-
welling area off Cape Blanc (e.g., Shaffer, 1974; Richert, 1975),
recent sediments do not show a considerable opal accumulation.
On the other hand, the sediments in the Baie du Levrier, behind the
spit of Cape Blanc, display a strikingly high opal content, generating
the following problems:
1. Is the opal content due to the extension of the influence of the

upwelling waters off Cape Blanc into the adjacent shallow bay region?
2. How does the upwelling influence take place: i.e. by the actual
generation of high biologic production inside the bay, or by lateral
supply of skeletal particles from outside the bay, where the upwelling
activity is centered?
3. Can analogous regimes of upwelling be recognized in fossil sediments?
The Baie du Levrier covers an area of about 900 km 2 • The average depth
is less than 15-20 m (Fig . 2). The bay is exposed to the northeast trade
winds from the Sahara with almost diurnal dust storms. It is subject
Fig. 2. Baie du Levrier: Bathymetry

and sample positions
to strong tidal currents (up to several knots) entering from the south-
east and south. Another salient feature is the complete absence of
fresh-water influx.
The temperatures of the upwelling water at Cape Blanc are low - approx.
16 0 C - as shown by data of the Laboratoire des Peches at Nouadhibou
(Fig. 3). The cool water enters the bay through relatively narrow chan-
nels. It leaves the area southward to the Banc d'Arguin. During this
284
Fig. 3. Surface temperatures off

• •
16,5° 16.4 TE MPERATURE
Cape Blanc during March-April 1971
(data from the Laboratoire des
°C Peches , Nouadhibou)
transport, the temperatures rise steadily from 18-20 o C in the bay to

more than 21-22 o C over the Banc d'Arguin.
A diagonal ridge divides the bay into a shallower, open Eastern Basin
and a slightly deeper and sheltered Western Basin. Because of the
sheltering effect of the "diagonal ridge" the Western Basin acts as
a trap for sediments and, temporarily, for the waters as well.
The maximum water depths of the Western Basin increase gently from 5 m
in the north to 15-20 m in the south (Fig. 2). Because of this depth
range, most of the basin floor is situated just below the local effec-
tive wave base as defined by Koopmann et al. (1978). Therefore, large
areas fall under a regime of quiet water deposition (Fig. 4). Accord-
ingly, a silty marl sediment covers more than 70% of the area, where
sand-sized components form a proportion of up to 10%. Besides a major
airborne terrigenous component from the Sahara consisting of the f.ine
sand, silt, and clay fractions, diatoms and sponge spiculae are a major
constituent of the sediment (Fig. 5).
The minimum diatom values of less than 1% of the total sediment are
restricted to the shallower areas such as banks and nearshore areas.
Maximum diatom values (up to 10% of the total sediment) occur in the
center of the Western Basin, which is mostly below the local effective
wave base .
so· Fig. 4. Locally effective surf
\-_--1-_ _-I~:_---+---___fl0 · and wave base in the Baie du
- 5 FETCH Levrier (from Koopmann et al.,
LENGTHlkm) 1978)
so·
so·
17°W SO ·
IO · n==+==~===t====! 10 '
0 ,. 7.0 ",
[3J 30-7 0".
e 0.5'/,
21DN
50' so·
Olst"butlon of diatoms
1'/, 01 tola l samp le I
Fig. 5. Distribution of diatoms in
o km 10 surface sediments of the Baie du
Levrier
so·
286
Fig. 6. Plankton-benthos ratio of

17°W 50'
Foraminifera
10' n;::::=+===ti:;;:===t====110 '
1M >0.20
0.10 - 0.20
< 0.10
50'
PIa Ilk! Oil - be Mhos - ra Ii 0

01 Foram i n i fera
o km 10
50 '
The plankton-benthos ratio of sand-sized Foraminifera (Fig. 6) shows

high proportions of planktonic individuals in the central Western Basin.
The values vary from 5% planktonic Foraminifera on the banks and in
shallow water areas influenced by wave activity, to 30% planktonic
Foraminifera in the sheltered parts of the Western Basin. The plankton-
benthos ratio of marine diatoms demonstrates a similar trend.
Fish debris (Fig. 7) and marine ostracods were found in concentrations

of up to 30% of the sand fraction. Generally their maxima coincide
with those of diatoms and planktonic Foraminifera.
High concentrations of marine opal skeletons in the bay sediment are

related to high primary production either inside or outside the bay.
Inside the bay, no nutrification is observed from fresh-water influx.
Thus, the only possible nutrient supply must originate from the up-
welling activity off Cape Blanc (Schaffer, 1974; Richert, 1975). If
the opal fraction is autochthonous, tidal currents ought to act as a
mechanism for transporting dissolved nutrients into the bay.
Such an assumption might be supported by the benthonic diatom fraction,

which, by its state of preservat.ion, is unlikely to originate from out-
side. Similar arguments apply to the large numbers of well-preserved
ostracods and fish debris.
On the other hand, several features point to an alloch.thonous, i.e.,

outside - origin of the particles under discussion (in this case, the
bay would act as a giant trap for planktonic skeletons): the plank-
tonic and the neritic species of the diatoms increase toward the south-
ern entrance of the bay (Table 1). Further, the abundance of a cool-
water, fine-sand-sized planktonic-Foraminifera assemblage (Be, pers.
comm.) points to transportation from outside the bay. This assemblage
287
17°W so· Fig. 7. Distribution of fish debris

10'n;::::~===1:::====t:===llo '
1&8 .. 0.10·/.
~ QOS -O.l0 ./.
so·
FISh deb".
(./. of loll' samp'e)
o
-
km
50 '
10
Table 1. Selected samples and their contents
Locality Corg % Number of diatom % of planktonic Fecal pellets (%

valves diatoms of sandfraction)
Central 2.00 255 x 10 4 /g 83.6 55.0

W Basin ? 187 x 10 4 /g 84.1 62.0
Northern 1. 72 271 x 10 4 /g 78.1 5.5

W Basin 1.59 121 x 10 4 /g 67.9 12.5
Southern 1.53 218 x 10 4 /g 81.5 9.2

W Basin
Head of Bay 1.14 ? ? 4.8
resembles that described by Thiede (1975) off Cape Blanc. However, the
high proportion of fecal pellets (Table 1) does not support any of the
proposed models. At present, no definite answer concerning which type
of model prevails can be presented.
The sedimentation rates are in the order of 40-50 cm/1000 years (Koop-
mann, 1975). Assuming that 10% of the total sediment consists of dia-
tom skeletons with a specific weight of 0.2 g/cm 3 (Heath, 1974), ap-
prox. 1 g/cm 2 /1000 years of diatom valves are sedimented in the central
Western Basin. Independently, values of a similar dimension (0.8-1.7
g/cm2 /1000 years) were found by using the absolute number of diatom
valves per g dry sediment and an average weight of less than 10 x 10- 9
g per diatom valve (Table 1). The average weight was determined by
weighing 400 randomly picked diatom valves, which had been properly
cleansed of attached clay mineral particles.
288
The actual opal accumulation rate will be increased (1) by a factor of

1.5 to 2.0 when the contribution of sponge spiculae is considered
(Koopmann, 1975) and (2) by an unknown factor due to opal dissolution
during sedimentation.
Comparable diatom deposition rates of 0.15-0.44 g/cm 2 /1000 years were

estimated from Antarctic deep-sea sediments by Goodell and Watkins
(1968). On the other hand, the biogenic silica deposition exceeds 50-
100 g/cm 2 /1000 years in the most intensive upwelling areas, such as
the Gulf of California and the Cascadia Basin off Oregon (Heath, 1974).
Comparable middle Miocene sediments, which indicate a depositional

paleo-environment analogous to that of the Baie du Levrier, were re-
ported from the American East Coast in the Calvert Formation (Lohmann,
1948; Abbott and Andrews, 1975).
In conclusion, the opal sedime.ntation of the bay indicates coastal up-

welling off Cape Blanc, which is not documented in the directly under-
lying sediments on the continental margin. This is consistent, regard-
less of the fact that no final decision can be made on the allochthonous
or autochthonous origin of the planktonic particles.
AaknowZedgmenvs, The sediments of the Baie du Levrier are presently being studied
in a joint Kiel~Louvain project, the major results of which will be published else-
where (Koopmann et al., in prep.). A.W.H. Be kindly helped to determine the species
of planktonic Foraminifera, which are fine-sand-sized and accordingly difficult to
indentify.
References
Abbott, W.H., Andrews, G.W.: Miocene diatomaceous deposits of South Carolina and
Georgia. Geol. Soc. Am. Abstr. ~ (1975)
Goodell, H.G., Watkins, N.D.: The paleomagnetic stratigraphy of the Southern Ocean:
200 W to 1600 E longitude. Deep-Sea Res. 15, 89-112 (1968)
Heath, G.R.: Dissolved silica and deep-sea-sediments. Hay, W.W. (ed.). SEPM Spec.
Publ. 20, 77-93 (1974)
Koopmann, B.: Siltmergel und Schillsande in der nordlichen Baie du Levrier (Maure-
tanien) • Dipl. Thesis. Kiel: Kiel Univ., 1975, 51 pp. (Unpubl.)
Koopmann, B., Lees, A., Piessens, P., Sarnthein, M.: Skeletal carbonates and wind
derived silty marls off the Sahara coast, Baie du Levrier, Arguin Platform, Mauri-
tania, Submitted to "Meteor"-Forsch.-Ergebn. C (1978)
Lohmann, K.E.: Middle Miocene diatoms from the Hammond well. - Cretaceous and Ter-
tiary subsurface geology. Maryland Dept. Geol. Min. & Water Res. Bull. 2, 151-187
(1948) -
Richert, P.: Die raumliche Verteilung und zeitliche Entwicklung des Phytoplanktons,
mit besonderer Berucksichtigung der Diatomeen im N.W.-Afrikanischen Auftriebswas-
sergebiet. Dissertation. Univ. Kiel, 1975, 140 pp.
Shaffer, G.: On the North-West African coastal upwelling system. Dissertation. Univ.
Kiel, 1974, 178 pp.
Thiede, J.: Shell and skeleton producing plankton and nekton in the eastern north
Atlantic Ocean, "Meteor"-Forsch.-Ergebn. C 20, 33.79 (1975)
General Aspects of Upwelling Research
Upwelling Research and Ocean Affairs
w.s. WOOS1ER
Summary: The study of upwelling ecosystems involves several branches of

science and takes place in many parts of the world ocean. Properly speaking,
man is a part of these ecosystems, and upwelling research interacts
with a variety of nonscientific human activities. Examples of these in-
teractions include:
1. Justification and funding for upwelling research are often based on
possible applications of such research, although there is little sys-
tematic effort to implement potential applications.
2. Variations in upwelling and in primary production appear to be close-
ly related, but how such variations affect the abundance and availabil-
ity of commercial fish populations is largely unknown.
3. Upwelling affects the production of sediments of commercial impor-
tance (phosphates, oil deposits) and has important effects on weather
and climate, although little of present upwelling research is directed
toward such practical applications.
4. Because coastal upwelling occurs largely within a few tens of kilo-
meters of the coast, upwelling research will be importantly affected
by the regime for marine scientific research to be established by the
Law of the Sea Conference.
5. To the extent that upwelling occurs off the coasts of developing
countries, upwelling research provides an opportunity for scientists
in advanced laboratories to assist in the development of a marine
science capability in those countries.
Such examples suggest that while the best scientific research is often
done without regard to application, improved mechanisms are required
to maximize the benefits that society receives from the research that
it supports.
The list of invited and contributed papers for the Third International
Symposium on Upwelling Ecosystems illustrates the broad dimensions of
upwelling research. The scientific disciplines are well represented
- physics, chemistry, biology in all its manifestations, and even a
handful of geological papers. Geographically, both principal and secondary
coastal upwelling regions are emphasized - those on the eastern sides
of the Atlantic and Pacific and those scattered elsewhere throughout
the world ocean.
Of the eighty or so titles, there are only two that concern applica-
tion of the results of upwelling research; there is no paper relating
upwelling to the production and management of stocks of commercial
fishes. Given the focus of the meeting on the state of research, it
is perhaps not surprising to find so little attention paid to the re-
lation of that research to the broader concerns of mankind. Yet it
292
could be argued that man is a significant component of the upwelling

ecosystem and that upwelling research cannot proceed entirely within
the splendid intellectual isolation suggested by the agendas common
to such symposia.
In this paper, I discuss some of the ways in which this sort of scien-
tific inquiry affects, or is affected by, nonscientific activities of
economic, legal, or political character - that tangled web of inter-
actions known as "ocean affairs." And surely, the very first interac-
tion is that between the scientist and his source of funds.
Upwelling research is commonly justified on the basis of its potential

applications. For example, in 1968, the Secretary General of the United
Nations recommended an expanded program of international cooperation
to assist in a better understanding of the marine environment through
science. In a review of the possible scientific content of such a pro-
gram, an international group of scientists had the following to say
about upwelling research (Scientific Committee on Oceanic Research,
1969) :
"Great benefit to fisheries would result from an improved understanding, leading

to prediction techniques, of the intensity of upwelling along the eastern coasts
of ocean basins'. Such regions are heavily productive of fish and the upwelling is
sensitive to changes in the winds. The fluctuations, at present unpredictable, have
a very strong effect upon the economics of fisheries.
Regional studies of the immediate mechanisms of upwelling could be correlated with

larger scale meteorological phenomena, i. e., the subtropical anti.cyclones, which
will be carefully observed in programs planned in the implementation of the World
Weather Watch. In addition, it seems probable that variations in sea surface tempera-
ture produced by changes in upwelling have important effects on the weather conditions
in the littoral areas. Studies of such effects would be a valuable by-product of
regearch on the upwelling process. There is also considerable geological interest
in the sedimentology of such regions of high organic productivity and of the accumula-
tion of organic matter and phosphorites."
So here are the three major applications for which upwelling research
is commonly justified: prediction of fishery yields, weather forecast-
ing, and the formation of nonliving resources. If the paragraph had
been written a few years later, i t might have made more specific re-
ference to the prediction of climatic changes.
In the United States, the major impetus to upwelling research has been
given by the National Science Foundation through its Office of the
International Decade of Ocean Exploration (IDOE). The objectives of
IDOE and of related international programs coordinated by the Inter-
governmental Oceanographic Commission have been stated as follows
(Scientific Committee on Oceanic Research, 1969):
"To increase knowledge of the ocean, its contents and the contents of its subsoil,
and its interfaces with the land, the atmosphere, and the ocean floor and to improve
understanding of processes operating in or affecting the marine environment, with
the goal of enhanced utilization of the ocean and its resources for the benefit of
mankind."
Projects of IDOE, rather than being concerned with science for science's
sake, are supposed to be related to objectives such as increased net
yield from ocean resources, prediction and enhanced control of natural
phenomena, and improved quality of the marine environment. In the be-
lief that upwelling research might eventually lead to more effective
293
utilization of the ocean and its resources, the National Science Founda-
tion has allocated several million dollars to its support.
I have no doubt that the potential benefits to society of upwelling

research are large. Yet it is not obvious that investigators give much
attention to these applications once research funding has been assured.
Nor do there exist mechanisms whereby the results of this or other
marine scientific research are periodically reviewed for possible ap-
plication, a matter to which I shall return.
Of all the potential applications of upwelling research, that in the

field of fisheries is of most ancient lineage. An important early study
(Sverdrup and Fleming, 1941) was the result of a cooperative program
between the California State Fisheries Laboratory and the Scripps In~
stitution of Oceanography; from 1948 to the present, upwelling studies
on the California coast have been part of the California Cooperative
Oceanic Fisheries Investigations. And the classical study of Peru Cur-
rent upwelling, that of Gunther (1936) on "William Scoresby" in 1931,
was under the auspices of the Discovery Committee, an organization
concerned with living resources of the Antarctic. Oceanographers and
fishery scientists have long recognized how useful i t would be to under-
stand the upwelling process that is responsible for the high production
on the eastern sides of oceans.
But how, in fact, does upwelling or variations therein, affect the

abundance, distribution, and availability of fish stocks of commercial
interests? In those regions where vertical circulation ensures a supply
of nutrients in the near-surface illuminated layers of the sea, it is
evident that primary production is higher than elsewhere in the ocean
and that the consequent availability of plant material is associated
with high concentrations and production of herbivores and carnivores.
Some upwelling research is concerned with the flow of energy and mate-
rials through this food web.
Variations in the intensity of upwelling, both in space and time, are

known to lead to variations in production and biomass at various tro-
phic levels. This is clearly true of the normal seasonal changes, for
example along the Peruvian coast, and most dramatically on a monsoon
coast such as that of Somalia or Vietnam. It must also be true in ab-
normal circumstances, for example as in El Nino where coastal upwelling
and coastal productivity are temporarily reduced. However, because of
the lags in the transfer of energy through the system, it is not nec-
essarily true that seasonal or irregular variations in primary and
secondary production have disastrous consequences for fish stocks. For
example, there was a major Nino event in 1957-1958 which certainly had
disastrous effects on the bird populations. However, a major decrease
in the abundance of anchoveta was not documented although there was
ample evidence for a change in their behavior. In 1972, on the other
hand, both abundance and behavior were affected, an indication that
the effects of Nino events depend inter alia on the condition of the
stock on which they are acting. In 1957-1958 the human fishery was
rudimentary; in 1972, the anchovy stock had been under heavy fishing
pressure for a number of years. It is also interesting to note that
the 1972 event was accompanied by recruitment failure of the stocks
spawned in both late 1971 and 1972, suggesting that the changed environ-
mental conditions affected the survival of fish larvae that may have
been unsuccessful in finding adequate food at the critical time of
their life or may have been unusually affected by predation. It is
also conceivable that spawning behavior itself is affected by environ-
men tal change.
294
Since a better understanding of the upwelling process is not in our

lifetime likely to be accompanied by the ability to manipulate that
process, what useful purpose can i t serve? An interesting application
is the assessment by Cushing (1971, 1973) of the fishery potential in
the Indian Ocean, based on findings of the International Indian Ocean
Expedition. Seasonal maps of primary productivity showed distributions
closely similar to those of upwelling regions. These data, together
with those of zooplankton biomass converted to secondary production,
were used to estimate tertiary production. This analysis led Cushing
to conclude "Upwelling areas and the associated offshore divergences
are the most productive; the most promising area for (fishery) develop-
ment is that off northern Somalia and southern Arabia, followed by the
Malabar Coast, and the Indonesian area." Remarkably, both the area and
the duration of the highly productive region in the western Arabian
Sea are greater than one would have predicted merely from the indica-
tions of upwelling intensity. The oceanographic data permitted crude
estimates of the potential for increased fishery yields, for example
off southern Arabia where Cushing estimates there might be an annual
catch of four to five million tons.
In the more general case, if one could learn how variations in the
marine environment affect the abundance, behavior, and distribution
of fish stocks and could predict the location, onset, duration, and
intensity of favorable and unfavorable environmental conditions, it
would then be possible to decide where, when, and how much, fishing
effort should be applied, as well as to make the other economic, social,
and political judgments required to manage the fishery and the marketing
of its product.
It is clear that the effects of upwelling are not confined to the sur-
face layers where it occurs nor to the living resources there. The high
productivity is reflected in the deposition of sediments rich in organic
matter, which may lead to formation of valuable quantities of petroleum
or phosphorite. I have not found much information on the former apart
from the classical paper of Brongersma-Sanders (1948) on "The importance
of upwelling water to vertebrate paleontology and oil geology. 11
In the case of phosphorite deposits, a general relationship with up-

welling has long been postulated (Kasakov, 1937). The necessary condi-
tions appear to include restriction of circulation by bottom topography,
an abundance of biogenic detritus sinking to the shelf, oxygen defi-
ciency of subsurface waters, and low sedimentation rates from terres-
trial sources. The present distribution of phosphorite nodules on con-
tinental shelves is closely related to the distribution of upwelling
- i.e., deposits are known off California, Peru, and Chile, and the
northwest and southwest coasts of Africa. However, attempts to date
these deposits have usually led to the conclusion that the mineral is
not presently being formed. Recently, however, measurements off south-
west Africa and Chile (Baturin et al., 1972) and off Peru (Veeh and
Soutar, 1973) have given ages from 300,000 to Recent. It is now proposed
that phosphorite nodules have been formed more or less continuously
from at least the late Pleistocene to the present, and that the pre-
Recent age bias has arisen from reworking and enrichment subsequent
to the initial deposition. The location and assessment of such deposits
may be facilitated by improved knowledge of the upwelling process.
A more dynamic application of such knowledge concerns the effect of

upwelling on weather and climate, and vice versa. The utility of im-
proved forecasting is unquestioned, and there appear to be demonstrable
economic benefits to be derived from enhanced programs of upwelling
research, particularly as they include the relevant meteorological
causes and effects.
295
The relation between coastal upwelling and weather has long been re-
cognized and is obvious to those who have experienced the cool and un-
eventful weather on the coasts of Peru or southern California. The role
of variations of upwelling in determining climate has only recently
become apparent, for example in the possible role of the warm water
off Peru during the 1972 Nino event in the formation several months
later of Hurricane Agnes (Narnias, 1973) and the possible relation with
drought in Brazil (Caviedes, 1973). Bjerknes (1966) has long argued
that variations in the intensity of equatorial upwelling in the central
and eastern Pacific arise from large-scale changes in the atmospheric
circulation and in turn have profound effects on that circulation over
wide areas of the north and south Pacific.
Another example where upwelling may have an important climatological ef-

fect is found in the Indian Ocean. Work during and since the Inter-
national Indian Ocean Expedition has shown that much of the monsoonal
rainfall over India originates in the Arabian Sea and that fluctuations
in the wind field over that area affect the amount and timing of the
precipitation (Anon., 1974) . A low-level jet is observed to flow along
the Arabian Sea into India. It has been proposed that irregularities
in the monsoon winds and rain may result from interactions between
this jet and the cold surface waters off Somalia and downstream from
the upwelling region. Variations in the intensity of coastal upwelling
and the transport of these cold waters offshore by the Somali Current
may affect the position and intensity of the low-level jet. At the same
time, these variations control the extent of warm surface and hence
the flux of water vapor; the magnitude of this influence on the Indian
monsoon will depend on the relative extent of the affected surface
area.
Upwelling is not exclusively a phenomenon of the coast, the equatorial

divergence in the Pacific being a particularly conspicuous example of
open ocean upwelling. However, regions of coastal upwelling are exten-
sive, highly productive, heavily fished - and so located as to be crit-
ical areas in the political struggle for control of marine living re-
sources.
The four classical eastern boundary current regimes are located as

follows:
- California Current: off United States (Oregon, California) and Mexico

- Peru Current: off Chile and Peru, affects Ecuador
- Canary Current: off Morocco, Spanish Sahara, Mauritania, Senegal,
affects Gambia and the Guineas
- Benguela Current: off South Africa, Southwest Africa, affects Angola
There are other significant coastal upwelling areas of at least seasonal

or local importance, including:
- Iberian Peninsula (Portugal, Spain)

- Panama Bight (Panama, Colombia)
- Caribbean (Venezuela, Colombia)
- Western South Atlantic (Brazil)
- Gulf of Guinea (Ivory Coast, Ghana, Togo, Dahomey, Nigeria)
- Arabian Sea (Somalia, various states in southern Arabia, southwest
India)
- Northwest Australia and Indonesia
- South China Sea (Vietnam)
This listing, however incomplete, suffices to illustrate that (1) many

countries are affected by coastal upwelling, and (2) the great majority
of these are developing, rather than industrialized, countries. The
296
wide distribution of coastal upwelling has important consequences for

the continuation of upwelling research.
As the laws of the sea negotiations have progressed during the last fevl
years, a new allocation of ocean space has gradually emerged. Hitherto,
the structure of maritime jurisdiction was relatively simple. Out to
some limited distance from the coast, within the so-called territorial
sea, the coastal state has exercised extensive jurisdiction; beyond,
the great bulk of the ocean has been high seas with jurisdiction limited
to that of the flag state over its own vessels.
This simple state of affairs was modified in a significant way by the

1958 Convention on thE! Continental Shelf, which granted the coastal
state sovereign rights over the continental shelf "for the purpose of
exploring it and exploiting its natural resources." In that Convention,
the continental shelf was defined as extending not just to a depth of
200 m, but "beyond that limit, to where the depth of the superjacent
waters admits to the exploitation of the natural resources of the said
areas."
For the marine scientist, extension of coastal state jurisdiction over

this ill-defined region had a serious consequence, namely that the con-
trol of scientific research fell also into the hc:mds of the coastal
state. As the Convention puts it:
"The consent of the coastal State shall be obtained in respect of any research con-
cerning the continental shelf and undertaken there. Nevertheless the coastal State
shall not normally withhold its consent if the request is submitted by a qualified
institution with a view to purely scientific research into the physical or biological
characteristics of the continental shelf, subject to the proviso that the coastal
State shall have the right, if i t so desires, to participate or to be represented
in the research, and that in any event the results shall be published."
The provision contains the seeds of problems that haunt marine science
in the current negotiations - the coastal state is given power over
scientific research in a region that is imprecisely defined; the nature
of the affected research is equally ill-defined (for example, does
"concerning the continental shelf and undertaken there" include research
by acoustic means?); the hope is expressed that consent shall not nor-
mally be withheld, but there is no check on the coastal state's power
to make that decision; neither "qualified institution" nor "purely
scientific research" is defined, although the definitions are highly
relevant to the consent decision; the nature of coastal state parti-
Cipation in the research is not specified, nor is it made clear what
constitutes publication of results.
Implementation of this provision during the past decade has on occasion

impeded marine geological and geophysical investigations but has had no
discernible impact on upwelling research. But that and most other sorts
of marine research have been affected by the seaward expansion of coast-
al state jurisdiction beyond the former limits of territorial seas,
and the extension of coastal state control over research within these
vast, newly captured areas. The current Law of the Sea Conference seems
likely to legalize and generalize these unilateral actions.
After several years of preparatory work, the so-called Third United

Nations Conference on Law of the Sea was convened in New York in late
1973, continued in Caracas in the summer of 1974, met again in Geneva
in the spring of 1975, and is scheduled for yet another session in New
York in early 1976. Progress has been painfully slow with little evi-
dence as yet of serious attempts to negotiate. However, at the conclu-
sion of the 1975 session Of the Conference, there was produced an "In-
297
formal Single Negotiating Text" that contained several provisions af-

fecting upwelling research in coastal regions (United Nat:ions, 1975).
It is proposed that the territorial sea extend out to 12 nautical miles,
and that beyond there shall be an exclusive economic zone extending
not more than 200 nautical miles from the coastal baseline. Conference
Committee II would grant the coastal state exclusive jurisdiction with
regard to scientific research within this zone. Committee III, speci-
fically charged with developing articles on marine scientific research,
proposed that within the exclusive economic zone states proposing to
undertake research provide the coastal state with full information on
the proposed research and on the means and personnel whereby i t will
be carried out, together with the opportunity to participate in the
research, and eventually with data results and samples.
More controversial is the Committee III proposal that the State or or-
ganization in~ending to undertake scientific research shall indicate
to the coastal state whether the research project is considered (1) to
be of fundamental character, or (2) is related to the resources of the
economic zone or the continental shelf. Criteria and guidelines for
this distinction are to be elaborated by competent international or-
ganizations. If in the view of the coastal state, research is directly
related to the exploration and exploitation of living or nonliving re-
sources, the project can be conducted only with the explicit consent
of the coastal state. Even if consent is granted, the research results
are not to be published or made internationally available without the
express consent of the coastal state. If, on the other hand, the coast-
al state does not dispute that the research is of fundamental nature,
the research state can proceed with the project.
This so-called "definitional" approach will be the subject of negotia-

tions in the relevant committee. The difficulties in establishing satis-
factory definitions are nowhere more evident than in the case of up-
welling research. As noted earlier, all such research is ultimately
related to the living, and at least some of the nonliving, resources
of the economic zone or the continental shelf. If this were not the
case, financial support for the investigations might be rather less
generous than at present. But is upwelling research directly related
to the exploration and exploitation of coastal resources? In all but
a few cases, I would say i t is not.
In the first place, exploration for living resources is done principal-

ly by fishermen who are well aware of coastal regions of high produc-
tivity. There are presumably few surprises left in regions accessible
to coastal fisherman. As mentioned earlier, one of the interesting
findings of the International Indian Ocean Expedition was the extent
of seasonal upwelling off northern Somalia and the south Arabian coast.
These locations were already known as regions of high fishery potential,
and the scientific findings served chiefly to delineate the affected
zones and to permit crude estimates of potential fishery yields.
Of course, studies of open ocean upwelling may lead to discovery of

new fishing grounds. The classical studies of Sette, Schaefer, Cromwell,
and others in the Pacific Oceanic Fishery Investigations in the early
fifties revealed the vigorous and widespread nature of the highly pro-
ductive equatorial upwelling - but i t is my impression that Japanese
fishermen were already exploiting the region with their longline fishery.
It is not likely either that upwelling research has any direct rela-
tionship to exploitation. As discussed earlier, the principal applica-
tion to the utilization of living resources appears to be in manage-
ment of the fishery through,prediction of the effect of environmental
variations on the abundance, behavior, and distribution of fish stocks.
298
Whether or not upwelling research can be demonstrated to be directly

related to exploration and exploitation of resources, it is likely
that the new law of the sea will require that within the exclusive
economic zone it be conducted only with permission of the coastal state.
Of course, the consent requirement does not necessarily close off the
possibility of research. However, experience with the Continental Shelf
Convention suggests that consent will often be denied, sometimes with-
out explanation, or the application will be ignored or not acted upon
in time for the proposed research to be carried out. With the coastal
state exercising total control over research within two hundred miles
of its coast, it is evident that the future conduct of marine scienti-
fic research within that area will be significantly more difficult for
a foreign scientist than before.
How is this likely to affect achievement of full understanding of the

upwelling process and the associated biological events? You should re-
call that the two hundred mile economic zones occupy about 37% of the
ocean surface, an area only slightly less than the total land mass of
the planet. All of the coastal upwelling regions fall within this area.
Of course, scientists of some countries will be perfectly free to study
upwelling off their own coasts. As experience has shown, however, the
comparative study of upwelling regions is essential to understanding
the process. High latitude upwelling off the Oregon coast is different
from low lati,tude upwelling off Peru, both because of the different
magnitude of the Coriolis force and the different wind field. It is
not enough to examine the consequences of changes in wind stress off
Oregon or Mauritania; one must also observe upwelling systems where
the winds are strong and steady (as in the winter off Peru) and where
the changes in wind stress are large (as in monsoon regions such as
the Arabian and South China Seas). It is in such regions where studies
are most likely to lead to, or permit the confirmation of, useful pre-
diction models.
More generally, comparative studies are an essential component of ocean

research. Like the more traditional sciences, oceanography includes
both theoretical and experimental studies. However, experiments in the
usual sense are difficult to conduct in the open ocean because the
system is too large and complex for variables to be controlled. As an
alternative, the investigator compares and attempts to interpret the
results of the various experiments carried out by nature in different
parts of the world. If these "experiments" cannot be observed, this
useful approach must be abandoned.
Developing countries are presumably less concerned with control of

marine scientific research than they are with control of marine re-
sources. Whatever the politics of the negotiations, it is logical that
the coastal state should share directly in the benefits of any research
carried out in areas falling under its jurisdiction. Everyone will
agree that scientists of the coastal state should be able to participate
in the research and to have full access to data, samples, and research
results. To be realistic, however, without scientific capabilities
comparable to those of the visiting investigators, those in the coastal
state are unlikely to be able fully to utilize the results. Because
of this, the issue of transfer of technology has become closely linked
to that of marine scientific research in the law of the sea negotia-
tions.
Most scientists in developed countries appear to support the develop-

ment of marine science capabilities in other coastal states, and many
of them have contributed significantly to technical assistance efforts
in this area. The only reservations I have found relate to the in-
creased costs of marine research likely to result from the new law of
299
the sea, particularly those relating to participation and technical

assistance, and the feeling that these costs should not be charged to
limited research budgets.
Significant sums have been spent on transfer of technology during the

last few decades, particularly in applied fields such as fisheries.
Yet it is not obvious that these investments have led to increased
capabilities in marine science in very many countries. Reasons for
this lack of success include the relative inadequacy of levels of fund-
ing, poor design of assistancE" programs, and inadequate national com-
mitments by recipients of this assistance. The restrictions on coastal
research ensuing from the new law of the sea are likely to stimulate
renewed demands for bigger and better programs of technologic trans-
fer. Perhaps such programs wi.ll be more successful in the future if
they are built around research programs of common interest to both
donor and recipient countries. This has been the case in one of the
more successful examples of developing a marine science capability,
in Peru, where scientists from a number of countries have for many
years been cooperating with Peruvian scientists on studies of the vig-
orous upwelling regime offshore. This precedent might be repeated in
other coastal upwelling regions.
Curiously enough, the transfer of technology is also not notably suc-

cessful within the developed countries. Thus, although most upwelling
research is justified to funding agencies because of its real or imag-
ined applications, there are no mechanisms whereby the results of such
research are systematically examined for their applicability. Of course,
this deficiency is not restricted to upwelling research but is wide-
spread through much of oceanography. A possible solution may be for
oceanographers, together with engineers, economists, and other social
scientists, to organize application review boards where the results
of their research can be periodically assessed and related to societal
concerns such as environmental forecasting and protection, energy de-
velopment, and resource management.
In summary, there are identifiable benefits to society that could re-

sult from upwelling research. These are related to management of the
exploitation of living resources, location and assessment of nonliving
resources, and the forecasting of weather and climate. However, the
mechanisms whereby research results are identified for such applica-
tions are primitive or nonexistent. Related to this inadequacy is the
unsatisfied desire of developing coastal states to share in the bene-
fits of upwelling and other kinds of oceanographic research. This
frustration, coupled with the prevailing international politics of all
aspects of ocean affairs, is likely to result in provisions in the new
law of the sea that may seriously inhibit the future conduct of coastal
upwelling studies. It is possible that well-conceived and adequately
funded programs dealing with the transfer of marine science capabilities
might serve to mitigate the problem.
References
Anon.: Index; Design of an oceanographic program in the monsoon region of the Indian
Ocean for the First GARP Global Experiment. Unpublished report, 1974
Baturin, G.N., Merkulova, K.I., Cholov, P.I.: Radiometric evidence for recent forma-
tion of phosphate nodules in marine shelf sediments. Marine Geology 13(3), M37-M41
(1972) --
Bjerknes, J.: Survey of El Nino 1957-58 in its relation to tropical Pacific meteoro-
logy. Bull. Inter-Am. Trop. Tuna Comm. 12(2), 1-62 (1966)
300
Brongersma-Sanders, M.: The importance of upwelling water to vertebrate paleontology

and oil geology. Kon. Ned. AX. Wet., Verh. Afd. Nat. (Tweede Sectie), D1.XLV !,
1-112 (1948)
Caviedes, C.N.: Secas and El Nino; two simultaneous climatic hazards in South America.
Proc. Assoc. Am. Geogr. ~, 44-49 (1973)
Cushing, D.H.: Survey of resources in the Indian Ocean and Indonesian Area. Food and
Agriculture Organization IOFC/DEV/71/2 (1971)
Cushing, D.H.: Production in the Indian Ocean and the transfer from the primary to
the secondary level. In: The Biology of the Indian Ocean. Zeitzschel, B. (ed.).
Berlin-Heidelberg-New York: Springer, 1973, pp. 475-486
Gunther, E.R.: A report on oceanographical investigations in the Peru Coastal Current.
Discovery Rep. ll, 107-276 (1936)
Kasakov, A.V.: The phosphorite facies and the genesis of phosphorites. Trans. Sci.
Inst. Fertilizers Insecto-Fungicides 142, 95-113 (1937)
Namias, J.: Birth of Hurricane Agnes - triggered by the transequatorial movement of
a mesoscale system into a favorable largescale environment. Monthly Weather Rev.
101(2), 177-179 (1973)
Scientific Committee on Oceanic Research: Global ocean research. La Jolla, California:
(mimeo) 1969
Sverdrup, H.U., Fleming, R.H.: The waters off the coast of Southern California March
to July, 1937. Bull. Scripps Instn. Oceanogr. 4(10), 261-378 (1941)
United Nations: Third Conference on the Law of the Sea. Informal Single Negotiating
Text. A/CONF~62/WP.8/Parts I-III.A/CONF.62/WP.9/Part IV, 1975
Veeh, H.H., Soutar, A.: Contemporary phosphorites on the continental margin of Peru.
Science 181(4102), 844-845 (1973)
Subject Index
ammonium EngrauZis ringens 90, 93-97, 105, 293

Baja California 79-81 ingestion 95-97
excretion by cephalopods 91 Euphausia 35, 37-42, 68
by fish 91-94 pacifica 39-41
by zooplankton 51, 92-94 euphausiids 34-42, 66-68
NW Africa 160
feeding of clupeoids 102-107

bacteria fish, pelagic at NW Africa
biomass 145-151 bioenergetic demand 116-118, 120
growth yield 160, 161 biomass 112-116
heterotrophic uptake 150, 151, 158- debris in sediments 272, 286, 287
165 . feeding relative to food production 119,
barium in sediments 264, 265 120
baroclinic radius of deformation 206 fluorescence 20, 21
Ghana 172 food chain
temperate waters 107
upwelling areas 108
CaZanus carinatus 49-51, 53, 54, 177, foraminifers
178 plankton-benthos ratio 270, 273, 286
Chaetoceros radicans 26, 29, 30 skeletons in sediments 268-271, 275,
chaetognaths 46-48, 70 286
chlorophyll frictional layer 6', 8
Baja California 79-81 friction in upwelling models 205
Galapagos 194 fronts 6, 8, 10, 85, 87, 242
NW Africa 19, 157-159
chloroplastic pigments in sediments
131-133 ~eostrophic balance
coccolithophorids 17-19 lakes 224, 229
copepods 48-54, 65-67 Peruvian upwelling 250, 254
upwelling circulation 212, 217
~ecapods 68
diatoms 17-19, 24-30, 175 halocline see stratification
skeletons in sediments 266-268,
273-275, 284-288
diffusion coefficients 204 iet, coastal upwelling 206, 217
dinoflagellates 17-19
drift experiments 50, 155-166
~ake Michigan 224, 231
Lake On tario 227, 231
~cosystem Law of the Sea, Third UN Conference 296,
analysis and size of investigated 297
area 8-10, 13 Liriope tetraphyUa 46
definition 3
eddy diffusivity 204
eddy viscosity 204 macrofauna
horizontal 212 decrease with depths 126
vertical 209, 210, 216, 231 and oxygen conditions 127
Ekman circulation 204, 229 megafauna, numbers 132-134
Ekman layer depth 5 meiofauna, numbers 127-131
El Nino 11, 293 Mesodinium rubrum 73-87
302
Mesodinium rubrum 73-87 equatorial upwelling 209

biomass 78, 83 Oregon 211, 216
distribution 73, 74, 76, 77 Rossby radius of deformation 206
metals, minor, in sedimentS 264 Ghana 172
mixed layer depth, Galapagos 185 Rossby waves 11
mixing, turbulent
definition 203
Equatorial Undercurrent 210
scales of vertical 216
~alinity
models of upwelling circulation 204
Baja California 79-81
lakes 227
Galapagos 188
layered 206, 214
Ghana 172 , 173
stratified 207
Peru 245-253
Sardina puZahClX'dW3 91, 117
NematosaeZis 34, 35, 37, 68
SardineZZa aurita 177-179
sea level changes during upwelling 171
nitrate
sea surface temperature
Cape Blanc 284
Galapagos 190-192
difference against air temperature 238,
NW Africa 20-22, 158
240
nitrite, Galapagos 190-192
Galapagos 183-185
nitrogen budget
Ghana 169
NW Africa 92
Peru 95 Lake Michigan 225
Peru 238, 240-244
North Atlantic Central Water 17
seiches 224
Nyatiphanes 34-37, 67, 68 sensitivity analysis 4, 9, 10
shelf waves, Ghana 173
silicate
~rganic matter, sediments 262-264
ostracods 70, 71
Galapagos 190-192
oxygen
siphonophores 45, 46, 69, 70
Galapagos 193
South Atlantic Central Water 17
NW Africa 157-159, 162, 163
stability, static
Peru 245-253
equatorial upwelling 210
Galapagos 186-188
stratification
E.hosphate
Baja California 85, 86
Ghana 169, 171
Galapagos 190-192
lakes 224, 228
Peru 248
Peru 245-253
phosphorite 265, 294
thermal, Galapagos 183
phytoplankton
Sverdrup circulation 6, 205
biomass, NW Africa 145-150
numbers, Baja California 78
Ghana 174, 175
NW Africa 16, 19, 20, 22
primary production temperature (see aZso sea surface temper-
Galapagos 194 - ature)
NW Africa 26-30 Baja California 79-81
principal component analysis, between Ghana 172, 173
phytoplankton species 17, 18 temperature/salinity diagram, Galapagos
pycnocline see stratification 189, 190
ThaZassiosira partheneia 17, 24-30
and other organisms 28-30
radiolarians, skeletons in sedi- cell numbers 26-30
- ments 266, 267, 271, 275, 276 primary production 26-30
research thermocline, see stratification
control by coastal states 296 time scales in upwelling models 10, 11,
goal in upwelling studies 3, 4 206, 207
respiration, zooplankton 51-53 topography, influence on upwelling 195,
Reynolds stress 204 196, 198
Richardson number tunicates 42-45, 70
303
undercurrent, coastal upwelling 130, boundaries 4-8, 13, 14

- 132, 217 characteristics 12-14
Ghana 173 in past geological periods 276, 277
undercurrent, equatorial
Galapagos 188, 190, 192, 195 !!.ater masses
mixing 211 Galapagos 184, 188-192
Peru 243, 251 NW Africa 17
upvlelling Peru 241-254
and benthos 134-136 wind
and climate 294, 295 Ghana 170, 171
correlation with local wind 170, 171 Peru 238-240
depth, off Peru 244, 255
~ooplankton
equatorial 208
displacement volumes 65, 175, 176
intensi ty 170
enzyme activities 53, 54
velocity, off Peru 244, 245, 255
numbers 66-72
upwelling regions
wet weights 118, 119
J. N. Kremer, S. WNixon
A Coastal Marine Ecosystem
Simulation and Analysis
1978. 80 figures, 13 tables. XI, 217 pages

(Ecological Studies, Volume 24)
ISBN 3-540-08365-0
This book describes the development of a

mechanistic simulation model for Narragansett
Bay (Rhode Island, U. S.A) and its use in eco-
system analysis. Such models provide a
synthesis of analytical data on the rates ofphoto-
synthesis, feeding, nutrient uptake and ex-
cretion, respiration, reproduction, growth and
other processes of many species and groups of
organisms. The Narragansett Bay model serves
as a useful case study for systems ecologists in
general and also provides a general treatment
of the dynamics of plankton-based aquatic eco-
systems. Discussions of the biological and eco-
logical bases, as well as the mathematical
concepts involved in the model and descriptions
of the development ofthe computer algorithm
are also included.
Contents:
Perspectives. - The Narragansett Bay ModeL -
Theoretical Formulations: Physical Forcing
Functions. Phytoplankton. Zooplankton. Addi-
tional Compartments. - Simulation and
Springer-Verlag Analysis: Mathematical Considerations and the
Berlin Computer Program. The Tidal Mixing ModeL
The Standard Run. The Role of Biological
Heidelberg Detail. Sensitivity and Stability. Applications
New York and Limitations. - References.
Microbial Ecology of a
Brackish Water Environment
Editor: G. Rheinheimer
1977.77 figures, 87 tables. XI, 291 pages

(Ecological Studies, Volume 25)
ISBN 3-540-08492-4
With contributions by M. BOlter, K Gocke,

H.-G. Hoppe, 1. Lenz, L.-A Meyer-Rei!, B. Probst,
G. Rheinheimer,1. Schneider, H. Szwerinski,
R Zimmermann
This book describes the results of a IS-month investi-

gation in the Kiel Bight (Baltic Sea), in which more than
50 hydrographical, chemical, and microbiological para-
meters were measured. Using methods such as scanning
elctron microscopy and autoradiography, a comparative
analysis of most of the measured data on the bacterial
colonization of detritus and the relations between
pollution, production, and remineralization led to the
construction of a model for the ecosystem, indicating the
energy fluxes.
Together with Volume 24 by Kremer and Nixon on
ecological studies conducted at Narragansett Bay, and
system-simulation by a computer model, the work
represents a comprehensive study of coastal marine
ecosystems.
Contents:
Hydrographic Conditions. - Oxygen and Inorganic
Nutrients. - Seston and Its Main Components. - Organic
Substances and Respiration Potential. - Primary Pro-
Springer-Verlag duction. - Plankton and Fungi Populations. - Biomass. -
Distribution of Special Physiological Bacteria Groups. -
Berlin Numerical Taxonomy. - The Autoradiographic
Heidelberg Method. - Heterotrophic Activity. - Biomass Produc-
tion. - Nitrification. - Desulfurication and Sulphur
New York Oxidation. - Comparative Analysis of the Data.

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Upwelling

With 132 Figures

Dr. MATTHIAS TOMCZAK

ISBN-13: 978-3-540-08822-6 e-ISBN-13: 978-3-642-66985-9

Upwelling areas are among the most fertile regions of the

The contributions to this volume were refereed by scien-

The present debate on the future of the exploitation

Especially we wish to thank the Springer-Verlag for

Kiel, August 1978 R. BOJE

Ecosystem Research in Upwelling Regions

Ecosystem Analysis and the Definition of Boundaries

What is an Upwelling Ecosystem?

Mesoscale Heterogeneities of the Phytoplankton

On the Ecological Significance of ThaZassiosira par-

Zooplankton Communities in the West African Up-

Relative Abundance of Zooplankton Groups in the

Mesodinium rubrum in the Baja California Upwelling

Regeneration of Nitrogen by Zooplankton and Fish

Upper Trophic Levels in Upwelling Areas

Food Consumption of Pelagic Fish in an Upwelling

Benthos in Upwelling Regions

Role of Bacteria in an Upwelling Ecosystem

Heterotrophic Activity in the Mauritanian Up-

Physical Aspects and Biological Consequences of

Oceanographic Conditions in the Galapagos Archi-

Hydrographical Aspects of Upwelling Regions

Role of Mixing in the Dynamics of Upwelling Sys-

The Circulation of Large Lakes

Hydrologic Aspects of the Main Upwelling Areas

Geological Aspects of Upwelling Regions

Sediments as Indicators of Upwelling

Sedimentation Influenced by Upwelling in the Sub-

General Aspects of Upwelling Research

Upwelling Research and Ocean Affairs

Subject Index .•......•...............•.•..••.... 301

Dr. R.T. BARBER, Marine Laboratory, Duke University,

Dr. J.R. BENNETT, Department of Earth and Planetary

Dr. M. BLACKBURN, c/o Roderick Mager, P.O.Box 774,

Dr. D. BLASCO, Bigelow Laboratory for Ocean Sciences,

Dr. R. BOJE, Institut fUr Meereskunde an der UniverSitat,

Dr. A. BUSTAMANTE, Instituto del Mar del Peru, Casilla

Dr. D.H. CUSHING, Fisheries Laboratory, Pakefield Rd.,

Dr. L. DIESTER-HAASS, Geologisches Institut der Univer-

Dr. M. ELBRACHTER, Biologische Anstalt Helgoland, Lito-

Dr. M. ESTRADA, Instituto de Investigaciones Pesqueras,

Dr. P.M. HARGREAVES, Institute of Oceanographic Sciences,

Dr. A. HERBLAND, Centre de Recherches Oceanographiques,

Dr. R.W. HOUGHTON, Lamont-Doherty Geological Obser-

Dr. G.T. HOUVENAGHEL, Laboratoire d'Oceanologie, Uni-

Dipl.-Geol. B. KOOPMANN, Geologisch-Palaontologisches

Dr. R. MARGALEF, Catedra de Ecologia, Facultad de

Prof. Dr. O.A. MATHISEN, Fisheries Research Institute,

Dr. M.A. MENSAH, Fishery Research Unit, P.O. Box B-62,

Dr. T.T. PACKARD, Bigelow Laboratory for Ocean

Dr. T.• RIVERA, Instituto del r-lar del Peru, Casilla

Prof. Dr. M. SARNTHEIN, Geologisch-Palaontologisches

Dr. H.-J. SCHRADER, Oregon State University, School

Dr. H. THIEL', Institut fUr Hydrobiologie und Fische-

Dr. A. THIRIOT t, Station ZOologique, 06230 Ville-