Table Of ContentSeasonality of Freshwater Phytoplankton
Developments in Hydrobiology 33
Series editor
H.J. Dumont
Seasonality of Freshwater Phytoplankton
A global perspective
Edited by
M. Munawar and J.F. Tailing
Reprinted from Hydrobio/ogia, vol. 138 (1986)
.11
1986 DR W. JUNK PUBLISHERS ....
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Library of Congress Cataloging in Publication Data
~easonality of freshwater phytoplankton.
(Developments in hyarooiology : 13)
1. Freshwater phytoplankton--Seasonal variations.
I. Munawar, M. II. TaIling, J. F. \Jonn rrancis)
111. Series.
<lI(Q15.S5 1986 581.5'2632 86-15342
ISBN -13 :978-94-0 10 -8635-6 e-ISBN-13 :978-94-009-4818-1
DOl: 10.1007/978-94-009-4818-1
Copyright
© 1986 by Dr W. Junk Pubhsner:" Dordrecht.
Softcover reprint of the hardcover 1s t edition 1986
All rights reserved. No part of this pubhcation may be reproduced, stored in a
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Dr W. Junk Publishers, P.O. Box 163, 3300 AD Dordrecht, The Netherlands.
v
Preface
This volume originated in a belief, shared by the two editors, that the time was ripe for a world-wide survey
- or at least sampling - of seasonality in freshwater phytoplankton. An opportunity was provided by the
International Limnological Congress (S.I.L.), held at Lyon in August 1983, to plan a one-day symposium
on the topic. From this enjoyable and successful occasion, augmented by additional written contributions,
the present volume has emerged. As convenors and editors, we are grateful to the contributors for their
cooperation in this international venture.
The seasonality of phytoplankton is widely conditioned by that of climate. Thus one may expect to find
the geographical differentiation of climatic patterns reflected in the seasonal patterns of algal occurrence.
Diversity in the global perspective is also introduced by considerations of geomorphology, geochemistry, and
genotypically determined response.
Nevertheless, the historical base of our subject is firmly rooted in the north-temperate zone. From its fresh
waters, and seas, there have evolved virtually all of the approaches and techniques now being applied to the
analysis of phytoplankton seasonal dynamics. One can instance the correlation of events in time-series of
environmental and algal observations, aided by statistical techniques or simply very long-term sampling; the
evidence from chemical composition of the crops and element-budgets; algal responses in cu1tur~ and under
bioassay procedures, as for nutrients and contaminants; activity measurements, as of photosynthetic C bal
ance; the analysis of horizontal as well as seasonal variability in large lakes; the size-fractionation of biomass,
pigments and primary productivity; and information from small lakes, or large enclosures, available for ex
perimental manipulation. These and other 'strategies' of phytoplankton study are illustrated in the following
pages.
Our present aim of a truly global perspective, with some degree of regional synthesis, is one scarcely
represented in the literature. The greatest success in unravelling causal relationships is likely to come from
intensive local studies, preferably combining several approaches. Thus many contributions here have centred
their attention on single or adjacent sites. Each is an object-lesson in itself, and in aggregate present a world
wide array with varying factor-emphasis, amplified by some wi4er regional comparisons.
The classic ground of central and western Europe is well represented by four contributions. One (Sommer)
develops some broad regional comparisons of seasonal patterns and successions; others give especial atten
tion to depth-distributions (Dokulil & Skolaut), to phytoplankter size (Bailey-Watts), and to experimental
manipulations through large enclosures (Reynolds). Problems of physical origin in more northerly latitudes
are described from the USSR (Petrova) and'the Canadian tundra (Sheath). Aspects of the seasonality of large
temperate lakes in a continental (N. American) climate are also treated, with emphasis on the behaviour of
biological components, in the North American Great Lakes including picoplankton and ultraplankton
(Munawar & Munawar) and of sediment-water exchanges in shallow Canadian lakes (Hecky, Kling, Brunskill
& Fee).
The scene then shifts to the sub-tropics. Here no lake has been more extensively studied than L. Kinneret,
where the resting stage of a dominant phytoplankter copes with the problem of 'over-summering' rather than
VI
of over-wintering (PoIlingher). The diversity of seasonal patterns in the continent of Africa is surveyed (TaIl
ing), and experience from Kenyan lakes used to examine some general issues of geographical variation (Kalff
& Watson). From South India, patterns of seasonal development under a monsoonal climate are described
(Zafar). From South America there are unusually long-period records for the lowland L. Valencia (Lewis)
and the elevated andine L. Titicaca (Richerson, Neale, Wurtbaugh, Alfarc & Vincent); these are used to
statistically and objectively define successional sequences and categories of periodic variation.
Lastly, one of the very few records of phytoplankton seasonality in temperate latitudes of the Southern
Hemisphere is presented (Duthie & Stout).
We hope that this assemblage - the first of its kind - will appeal to the interests and imagination of
a wide variety of readers. We would like to thank the numerous referrees for meticulously reviewing the
manuscripts in addition to the reviews performed by both the editors which indeed ensured the highest level
of scientific quality. We wish to express our gratitude to Wil Peters of Dr W. Junk Publishers and Henri
Dumont, Editor-in-Chief of Hydrobiologia for their interest, encouragement and assistance in the publica
tion of the proceedings.
M. Munawar, Fisheries & Oceans Canada, Canada Centre For Inland Waters, Burlington, Ontario, Canada
J. F. TaIling, Freshwater Biological Association, Ambleside, Cumbria, England
The global distribution of numbered contributions.
Contents
Preface................................................................................ V
CLASSIC GROUND: CENTRAL AND WESTERN EUROPE
The periodicity of phytoplankton in Lake Constance (Bodensee) in comparison to other deep lakes of
central Europe
by U. Sommer ........................................................................ .
Succession of phytoplankton in a deep stratifying lake: Mondsee, Austria
by M. Dokulil and C. Skolaut. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Seasonal variation in size spectra of phytoplankton assemblages in Loch Leven, Scotland
by A. E. Bailey-Watts.................................................................. 43
Experimental manipulations of phytoplankton periodicity in large limnetic enclosures in Blelham Tarn,
English Lake District
by C. S. Reynolds.................................................................... 43
HIGH NORTHERN LATITUDES
Seasonality of Melosira-plankton of the great northern lakes
by N. A. Petrova..................................................................... 65
Seasonality of phytoplankton in northern tundra ponds
by R. G. Sheath........................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
GREAT LAKES OF NORTH AMERICA
The seasonality of phytoplankton in the North American Great Lakes, a comparative synthesis
. by M. Munawar and I. F. Munawar. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Seasonality of phytoplankton in relation to silicon cycling and interstitial water circulation in large,
shallow lakes of central Canada
by R. E. Hecky, H. J. Kling and G. J. Brunskill ...... , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
THE SUB-TROPICS AND TROPICS
Phytoplankton periodicity in a subtropical lake (Lake Kinneret, Israel)
by U. Pollingher...................................................................... 127
The seasonality of phytoplankton in African lakes
by J. F. Talling....................................................................... 139
VIII
Phytoplankton and its dynamics in two tropical lakes: a tropical and temperate zone comparison
by J. Kalff and S. Watson.... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Seasonality of phytoplankton in some South Indian lakes
by A. R. Zafar. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Phytoplankton succession in Lake Valencia, Venezuela
by W~ M. Lewis, jr.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Patterns of temporal variation in Lake Titicaca. A high altitude tropical lake. I. Background, physical
and chemical processes, and primary production
by P. J. Richerson, P. Neale, W. Wurtsbaugh, R. Alfaro andW. Vincent. . . . . . . . . . . . . . . . . . . .. 205
THE SOUTH TEMPERATE ZONE
Phytoplankton periodicity of the Waitaki lakes, New Zealand
by H. C. Duthie and V. M. Stout.............. ...... ...... ... . ...... ... .... .......... .. 221
Hydrobiologia 138: 1-7, (1986).
© Dr W. Junk Publishers, Dordrecht
The periodicity of phytoplankton in Lake Constance (Bodensee) in comparison to
other deep lakes of central Europe
Ulrich Sommer
University of Constance, Institute of Limnology, P.D. Box 5560, D-7750 Constance, FRO
New address: Max Planck Institute of Limnology, P.O. Box 165, D-2320 Pion, FRO
Keywords: phytoplankton succession, inter-lake comparison, oligotrophic-eutrophic gradient, central Eu
ropean lakes
Abstract
Phytoplankton periodicity has been fairly regular during the years 1979 to 1982 in Lake Constance. Algal
mass growth starts with the vernal onset of stratification; Cryptophyceae and small centric diatoms are the
dominant algae of the spring bloom. In June grazing by zooplankton leads to a 'clear-water phase' dominated
by Cryptophyceae. Algal summer growth starts under nutrient-saturated conditions with a dominance of
Cryptomonas spp. and Pandorina morum. Depletion of soluble reactive phosphorus is followed by a domi
nance of pennate and filamentous centric diatoms, which are replaced by Ceratium hirundinella when dis
solved silicate becomes depleted. Under calm conditions there is a diverse late-summer plankton dominated
by Cyanophyceae and Dinobryon spp.; more turbulent conditions and silicon resupply enable a second sum
mer diatom growth phase in August. The autumnal development leads from a Mougeotia - desmid assem
blage to a diatom plankton in late autumn and winter.
Inter-lake comparison of algal seasonality includes in ascending order of P-richness K6nigsee, Attersee,
Walensee, Lake Lucerne, Lago Maggiore, Ammersee, Lake Zurich, Lake Geneva, Lake Constance. The oligo
trophic lakes have one or two annual maxima of biomass; after the vernal maximum there is a slowly develop
ing summer depression and sometimes a second maximum in autumn. The more eutrophic lakes have an
additional maximum in summer. The number of floristically determined successional stages increases with
increasing eutrophy, from three in K6nigsee and Attersee to eight in Lake Geneva and Lake Constance.
Introduction Lake Constance' (Leader: M. Tilzer). The most im
portant physical and chemical environmental varia
It is the aim of this contribution to present a bles (Stabel & Tilzer, 1981), the primary productivi
comparison of the phytoplankton seasonality in ty (Tilzer, 1984), the seasonality of the
deep central European lakes of considerable size. phytoplankton (Sommer 1981a, b), of the
All are characterized by basins with steep sides, zooplankton (W. Geller, in prep.) and sedimenta
small littoral zones, and a development of mainly tion (H. H. Stabel, in prep.) have been studied at
pelagic communities. Unfortunately heterogeneity weekly intervals. In the second section a compari
and incompleteness of available data make an son with other central European lakes is attempted.
equal treatment of all lakes impossible. Therefore, The data are taken from literature and from unpub
in the first section the periodicity of phytoplankton lished lists (Zurichsee, Walensee, by courtesy of U.
in Lake Constance is given in greater detail. This Zimmermann). Less frequent sampling and ab
lake has been thoroughly studieq from 1979 to 1982 sence of background data on physical and chemical
by the working group of the Deutsche Forschungs conditions and zooplankton make it inevitable in
gemeinschaft - sponsored project 'Carbon cycle in some cases to explain developments in those lakes
2
by analogy with Lake Constance. For example, the (2) Mass growth of algae starts with the onset of
June minimum of algal biomass in Lake of Geneva thermal stratification in April/May. High nutrient
has been interpreted as a grazing-induced clear availability and low grazing pressure gives rise to an
water phase in spite of absence of zooplankton assemblage of rapidly growing, nanoplankton spe
data, because phytoplankton data are nearly identi cies dominated by Rhodomonas lens, R. minuta,
cal to those from Lake Constance for the respective Cryptomonas ovata and small centric diatoms,
phase. mainly Stephanodiscus hantzschii. All these algae
Inter-lake comparison needs some degree of ab are heavily grazed by Cladoceran filter-feeders.
straction from actual data. Here, the main abstrac (3) Increased zooplankton grazing leads to a
tion is the deduction of a 'main sequence' in the dramatic decrease of phytoplankton densities in
succession of phytoplankton (Margalef, 1978). This early June. The scarce phytoplankton of this 'clear
concept is based on the experience that in a strati water phase' (Lampert & Schober, 1978) is domi
fied body of water nutrients are successively deplet nated by Cryptomonas ovata and Rhodomonas
ed, interactions with higher trophic levels increase minuta.
and non-motile algae replaced by motile or buoy (4) Towards the end of the clear-water phase
ant ones. Partial destratification or erosion of the shortage of food leads to a decline of zooplankton
thermocline by weather-induced mixing may lead abundance. Algal biomass rises again. As long as
to re-capitulation of earlier successional stages nutrients are not yet depleted, Cryptomonas ovata,
('reversions' sensu Reynolds, 1980). In contrast to C. rostratiformis and Pandorina morum are the
temporary reversions the autumnal erosion of dominant algae.
stratification is a regular event and belongs to the (5) When soluble reactive ·phosphorus becomes
'main sequence'. An experimental demonstration depleted in July algal dominance shifts to the large
for the distinction between the 'main sequence' and and poorly grazed diatoms Asterionella formosa,
'reversions' has been provided by destratification Fragilaria crotonensis, Stephanodiscus binderanus,
experiments (Reynolds, Wiseman, Godfrey & But Melosira granulata. Current competition research
terwick, 1983). has shown by continuous culture experiments that
several species of net-planktonic diatoms tend to
outcompete all other taxa if P is in short supply
Lake Constance (Bodensee) and rich silicate supply prevents Si-limitation (Til
man, 1977; Tilman, Kilham & Kilham, 1982; Som
Lake Constance (500 km2 surface area, maxi mer, 1983).
mum depth 250 m, mean depth 100 m) is a warm (6) When silicate is depleted (Sommer & Stabel,
monomictic lake. The process of rapid eutrophica 1983) the diatoms are replaced by a dominance of
tion was stopped in the late 70's (Elster, 1982). At Ceratium hirundinella (mid-August).
present the overturn concentrations of total phos (7) In some years the Ceratium-stage is followed
phorus are about 100 p.g .1-1 • Seasonality and by a diverse phytoplankton dominated by the heter
stratification of nutrients and temperatures are ocystous blue-green algae Anabaena f/os-aquae, A.
available from Stabel & Tilzer (1981), and details on planctonica, A. spiroides, Aphanizomenon f/os
the counting method and further information on aquae and the Chrysophyceae Dinobryon sociale
phytoplankton seasonality from Sommer (1981a, and D. divergens. There were some deviations from
b). Phytoplankton species composition throughout this scheme of summer succession. Upwelling of
the whole text refers to biomass, measured as cell phosphate gave rise to short growth pulses of Cryp
volume. tophyceae and upwelling of silicate led to a second
The seasonal periodicity of phytoplankton spe sommer diatom stage, which then replaced the
cies composition followed with slight deviations the Cyanophyceae-Dinobryon assemblage. The de
same general pattern (Fig. 1) during the whole peri velopment of the Cyanophyceae-Dinobryon assem
od of study (1979-1982). blage is considered the 'main sequence' of algal
(1) During winter a long period of deep circula succession, whereas the late summer pulses of dia
tion and low light intensities lead to extremely low toms and Cryptophyceae are classified as rever
algal densities. sions.
