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REVIEW
Towards a functional classification of the
freshwater phytoplankton
COLIN S. REYNOLDS1, VERA HUSZAR2, CARLA KRUK3, LUIGI NASELLI-FLORES4 AND SERGIO MELO5
1CEH ALGAL MODELLING UNIT, THE FERRY HOUSE, AMBLESIDE, CUMBRIA, LA LP, UK; 2LABORATORIA DE FICOLOGIA, MUSEU NACIONAL/UFRJ, SÃO

, BRASIL; 3LIMNOLOGY SECTION, UNIVERSIDAD DE LA REPÚBLICA,  MONTEVIDEO, URUGUAY;

CRISTÓVÃO, RIO DE JANEIRO (RJ)
DIPARTIMENTO DI SCIENZE BOTANICHE, UNIVERSITA DI PALERMO, VIA ARCHIRAFI , I- PALERMO, ITALY; 5INSTITUTO DE BIOLÔGIA,
UNIVERSIDADE FEDERAL DE RIO JANEIRO, ILHA DO FUNDÃO, FIO DE JANEIRO (RJ), , BRASIL
CORRESPONDING AUTHOR: COLIN S. REYNOLDS. E-MAIL: csr@ceh.ac.uk

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This paper considers the structure of freshwater phytoplankton assemblages and promotes a scheme
of ‘vegetation recognition’, based upon the functional associations of species represented in the plank-
ton. These groups are often polyphyletic, recognizing commonly shared adaptive features, rather than
common phylogeny, to be the key ecological driver. Thirty-one such associations are outlined and the
basic pattern of their distinctive ecologies is outlined. An invitation to other plankton scientists to
assist in the development of this scheme is issued.

I N T RO D U C T I O N distinguish the differences in energy flow through the

No less than other scientific disciplines, biology is founded
on a vast amount of verified but miscellaneous infor-
mation that, necessarily, must be classified and ordered.
Linnaeus may not have been the first to name and group
organisms together but the essence of his binomial system
of species names, strictly within higher hierarchical
groupings based upon morphological affinities, has sur-
vived intact through over two centuries of continuous
expansion and refinement. The new age of molecular
biology and genetic verification has not made Linnaean
taxonomy redundant but, rather, has greatly enriched it,
at once furnishing an independent test of suspected evol-
utionary affinities and a means of verifying the mechan-
isms of gene expression. Modern molecular techniques
consolidate the strength of a sound phylogenetic base to
taxonomy and, more often than not, lend weight to many
past judgements about the interrelationships of individual
species.
Ecology is a predominantly biological discipline con-
cerned with the distributions of organisms and their inter-
relationships with each other and their environments.
Ecologists also recognize that unless they can name
species or, at least, have a close idea of their affinities, the
point of their work is very largely lost. At the same time,
however, whether they are trying to work out how a system
is organized and its functions are allocated, or to
community structures of old meadow or arable cropland,
various types of forest or coral reefs, ecologists will soon
resort to additional schemes of classification (Lavorel et al.,
1997). These will be concerned with the functional roles
and structural adaptations of the main species, whether
they are producers or consumers, whether they are
invasive or tolerant of site maturation, whether they share
tolerances to acidity or waterlogging or other environ-
mental constraints, and so on. Terrestrial ecologists recog-
nize types of system and predominant pathways of energy
flow based wholly on their ability to distinguish (say) sandy
heath from ombrogenous bog, or Carex paniculata swamp
from Salix carr or from Quercus woodland, simply on the
grounds of which kinds of species are present. Com-
munities are better, more reliable indicators of habitat
conditions than are the presence or absence of com-
ponent species. The work of the great phytosociologists
[(Tansley, 1935; Tüxen, 1955; Braun-Blanquet, 1964); see
also (Shimwell, 1971)] has bequeathed a system of diag-
nosing and naming the very distinct associations of plant
species that constitute vegetation. In essence, the associ-
ations are the basic functional units. Each is named after
one or two species that are characteristically represented
in that particular community-type, using a distinctive
binomial construction based on the name of one of them:
Lemnetum minoris is the association of free-floating mats of
duckweed, typically including Lemna minor (Haslam et al.,

© Oxford University Press 2002

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JOURNAL OF PLANKTON RESEARCH VOLUME
1975). The system works because the overlapping require-
ments of individual species can often be satisfied simul-
taneously in particular locations, so long as the
adaptations of each allow them to tolerate the conditions
obtaining. Most interestingly, such tolerances may require
behavioural or physiological adaptations that are not
necessarily constrained by phylogenetic affinity: plant
associations can be polyphyletic.
When it comes to describing the structure of pelagic
communities however, most aquatic ecologists might be
content to know that phytoplankton was present
(perhaps judged by its content of extractable chloro-
phyll). In contrast, specialists may well delight in being
able to provide a full list of species, perhaps separated
according to criteria distinguishable only with an elec-
tron microscope or verifiable using polymerase chain
reaction. At a more general level, the distinction of a
diatom- or flagellate- or even a Cyanoprokaryote-
dominated phytoplankton may convey a suitable
summary of what is there. Yet the question as to why one
phylogenetic category of organisms does better than
another under sketchily defined environmental con-
ditions remains largely unanswered, while the ability to
explain, much less predict, which species dominate con-
tinues to puzzle (Huisman and Weissing, 2001).
In this paper, we seek to consolidate a view that plank-
ton ecology will benefit from the adoption of an alterna-
tive scheme of ‘vegetation recognition’, based upon the
representation of functional associations of species. These
may be selected or excluded on the basis of major
adaptive features but which are not specific to one or a few
phylogenetic groups. Similarly, selection or exclusion of
these very features by the conditions obtaining in given
systems will not predict species dominance but they may
well narrow down the probabilities to a particular func-
tional group well-represented in a given habitat or some
given variation in the conditions.
Before elaborating the notion that microscopic algae
might form sociological groupings much as do terrestrial
plants, it is necessary to say a word about our choice of
group terms, none of which has universal understanding
or acceptance. ‘Communities’ is rejected for, however for-
tuitous is their composition, the species present interact
and some may perhaps benefit each other in some way.
We should also recognize the community role of het-
erotrophs and phagotrophs, but this is not a feature of the
scheme we describe here. ‘Assemblages’ is better in recog-
nizing the fortuity of species presence. However, material
collected in a single sample of plankton may well com-
prise several dozens of taxa, few of them numerous, some
being there only because they have been introduced by
advection or entrainment (for example) from the benthos
or littoral. Some species will have been increasing in
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representation in conditions they had found favourable,
others will have been in decline because the same con-
ditions were not altogether suitable for them. Our scheme
seeks to distinguish these behaviours, at least among the
relatively more numerous species.
‘Associations’ is the word used by terrestrial plant ecol-
ogists. It has the merit of conveying the notion of a group
of species responding similarly to a single set of environ-
mental conditions, even when the positions in a fluid
environment are more difficult to deal with. Yet this too
suffers a drawback in failing to acknowledge the fact that
some species may show analogous adaptations to similar
conditions but have yet to be found simultaneously ‘in
association’ at the same localities—thus, Cylindrospermopsis
and Anabaena minutissima show similar antennal properties
and nitrogen-fixation capacities that suit them to turbid,
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nitrogen-deficient water columns but, to our knowledge,
they have never been found in mutual association in the
same location.
The collective term closest to encompassing our pro-
posal to bracket together species with similar morpho-
logical and physiological traits and with similar ecologies
is perhaps, ‘narrowly defined functional groups’. Accept-
ing that the expression ‘functional group’ does not have a
unique or universal interpretation either, we emphasize
that the terminology seeks to differentiate among phyto-
plankton on the basis of specialist adaptations and
requirements (such as having a high affinity for phospho-
rus or carbon dioxide at low external concentrations, or of
requiring skeletal silicon, or of being a good light
antenna). The separation then permits two useful deduc-
tions to be made. One is that a functionally well-adapted
alga will be likely to tolerate the constraining conditions of
factor deficiency more successfully than individuals of a
less well-adapted species. The other is that a habitat
shown typically to be constrained by light, or C or N or
whatever, is more likely to be populated by species with the
appropriate adaptations to be able to function there (but
NOT that they WILL be there). Thus, our application of
the term ‘functional group’ is sensitive to the sets of appro-
priate adaptive specialisms and the clusters of species that
have them. Until such time as it is possible to forecast the
presence of individual species in given locations, this
scheme seems to us to offer the clearest way into under-
standing and predicting the distributions and dynamics of
natural populations of phytoplankton.
Adopting this concept of ‘associations’ and of ‘func-
tional groups’, we set out here to demonstrate from our
individual experiences the measure of consensus we have
attained about the general applicability of the model. We
seek to encourage others to use it and, because the method
is in its infancy, we invite them to share in the means of its
development.
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C. S. REYNOLDS ET AL. FUNCTIONAL CLASSIFICATION OF FRESHWATER PHYTOPLANKTON
T H E VA L I D I T Y O F T H E
F U N C T I O N A L G RO U P I N G
The idea of developing ecological categories of phyto-
plankton and the desire to be able to use them to describe
variations in composition among the natural lakes has a
long history (Hutchinson, 1967). The inspiration for
Reynolds’ first attempt (Reynolds, 1980) to devise a system
of classification of planktonic algae to be sensitive to
environmental change (essentially to eutrophication but
also to shorter seasonal fluctuations in stratification and in
the accessibility of adequate nutrient supplies) came from
the sorting methods of Tüxen and Braun-Blanquet
(Tüxen, 1955; Braun-Blanquet, 1964): the relevé, the
small arbitrary unit of vegetation in which plant presence
is scored for abundance, was substituted by a list of the
species counted (in excess of a cut-off point) in a particu-
lar sample from a particular lake on a particular date.
Species frequently found to co-exist and to increase or
decrease in number simultaneously were delimited and
given association identities. Although these were claimed
to be analogous to plant associations, to accord them
formal phytosociological names seemed premature and
the alphanumeric devices have been retained provision-
ally.
Fourteen groups of phytoplankton were identified in
Reynolds’s original study (Reynolds, 1980). Some have
been subdivided since, although the biggest change has
been to re-label them (Reynolds, 1984). Such validity as
the sorting method carries continues to underpin these
associations. Today, the list of associations (Table I) has
more than doubled, with most of the later groups having
been added, as it were, ‘by eye’. Whilst still retaining the
ideal that the subdivisions reflect the simultaneity of
responses of individual species to environmental variabil-
ity, it has to be emphasized that the new groupings have
been accommodated on intuitive grounds, without the
benefit of even the sociological separation. This does not
make them wrong but an invalid principle gives no
grounding for robust advocacy of the functional grouping.
However, two developments enhance the validity of the
functional grouping. In the analysis of the phytoplankton
in a small urban lake in Montevideo, Kruk et al. (Kruk et
al., 2002), applied canonical variate analysis (ter’Braak
and Smilauer, 1998) to discriminate phases of differing
species composition with several classification approaches.
They found that the eigenvalue for the first CCA axis for
the associations was over 0.7: 78% of the cumulative vari-
ance was explained by the scheme as it had evolved by
1997, despite having been designed primarily for Euro-
pean lakes. This was a statistically better performance
than any of the other schemes considered. The func-
tional-group classification seems to capture much of the

statistical variation. Analogous findings have been
reported by Fabbro and Duivenvoorden (Fabbro and
Duivenvoorden, 2000), although they used a different
sorting method.
The second enhancement comes from the finding that
algae forming a single functional group also have similar
morphologies, as quantified by the dimensions of the algal
‘units’ (cells or colonies, as appropriate, together with any
peripheral mucilage): surface area (s), volume (v) and
maximum linear dimension (m) are powerful predictors of
optimum dynamic performance (Reynolds and Irish,
1997). Typical representatives of each main association
strike mutually similar co-ordinates on a plot of ms/v
against sv. The plot pulls apart rounded and stout cylin-
drical units of differing sizes but separates those larger
units that preserve a high s/v by being attenuated in one
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plane (sometimes two, but not three). Separation of the
algae on the basis of their morphologies coincides sub-
stantially with the distributions of the same species among
different types of habitat distinguished on the basis of
accessibility to light and all nutrient resources (Reynolds,
1984, 1997).
The converse of this is that it is now easier to fit hitherto
functionally unclassified species into existing functional
categories, at least on a provisional basis. What to do
about those that do not readily fit the classification is to
become one of the recommendations of our essay.
THE RECOGNITION AND
N O M E N C L AT U R E O F T H E
F U N C T I O N A L G RO U P S
The latest list of trait-differentiated functional groups
(Table I) has 31 entries. Alphanumeric terms are retained
but some groups that have stood robustly for the last 20
years are named provisionally for the first time. In this
section, their distinguishing properties and ecologies are
briefly reviewed. At the beginning, the alphanumeric coda
were allocated in blocks to reflect seasonal shifts (A–D for
vernal blooms, E–H for associations at the start of
summer stratification, and so on) and within each block,
the trophic ‘preferences’ were also distinguished (so B →
E → L → N might represent the seasonal progression in
a mesotrophic temperate lake and C → G → M → P
would summarize a more eutrophic system). As the system
has become adapted to accommodate other types of
water and especially waters at other, non-temperate, lati-
tudes, so the intrinsic significance has been lost. In the text
below and in the Tables, we have found it unhelpful to run
through the coda in alphabetical order, preferring instead
to develop the distinctions among near-related functional
groups.
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Table I: Trait-separated functional groups of phytoplankton (updated from Reynolds, 1997)

Codon Habitat Typical representatives Tolerances Sensitivities

A Clear, often well-mixed, Urosolenia, Nutrient pH rise

base poor, lakes Cyclotella comensis deficiency
B Vertically mixed, mesotrophic Aulacoseira subarctica Light pH rise,
small-medium lakes Aulacoseira islandica deficiency Si depletion
stratification
C Mixed, eutrophic small- Asterionella formosa Light, C Si exhaustion
medium lakes Aulacoseira ambigua deficiencies stratification
Stephanodiscus rotula
D Shallow, enriched turbid Synedra acus Flushing nutrient
waters, including rivers Nitzschia spp depletion
Stephanodiscus hantzschii
N mesotrophic epilimnia Tabellaria Nutrient stratification

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Cosmarium deficiency pH rise
Staurodesmus
P eutrophic epilimnia Fragilaria crotonensis Mild light and stratification
Aulacoseira granulata C deficiency Si depletion
Closterium aciculare
Staurastrum pingue
T deep, well-mixed epilimnia Geminella Light deficiency Nutrient
Mougeotia deficiency
Tribonema
S1 turbid mixed layers Planktothrix agardhii highly light flushing
Limnothrix redekei deficient
Pseudanabaena conditions
S2 shallow, turbid mixed layers Spirulina light flushing
Arthrospira deficient
Raphidiopsis conditions
SN warm mixed layers Cylindrospermopsis light-,nitrogen- flushing
Anabaena minutissima deficient
conditions
Z clear, mixed layers Synechococcus low nutrient light deficiency
prokaryote picoplankton grazing
X3 shallow, clear, mixed Koliella low base mixing,
layers Chrysococcus status grazing
eukaryote picoplankton
X2 shallow, clear mixed layers Plagioselmis stratification mixing,
in meso-eutrophic lakes Chrysochromulina filter feeding
X1 shallow mixed layers in Chlorella, Ankyra stratification nutrient deficiency
enriched conditions Monoraphidium filter feeding
Y usually, small, enriched Cryptomonas low light phagotrophs!
lakes
E usually small, oligotrophic, Dinobryon low nutrients CO2 deficiency
base poor lakes or Mallomonas (resort to
heterotrophic ponds (Synura) mixotrophy)
F Clear epilimnia colonial Chlorophytes low nutrients ?CO2 deficiency
e.g. Botryococcus high turbidity
Pseudosphaerocystis
Coenochloris
Oocystis lacustris

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C. S. REYNOLDS ET AL. FUNCTIONAL CLASSIFICATION OF FRESHWATER PHYTOPLANKTON

Table I: continued

Codon Habitat Typical representatives Tolerances Sensitivities

G Short, nutrient- Eudorina high light nutrient deficiency

rich water columns Volvox
J shallow, enriched lakes Pediastrum, Coelastrum settling into low
ponds and rivers Scenedesmus light
Golenkinia
K short, nutrient-rich Aphanothece deep mixing
columns Aphanocapsa
H1 dinitrogen-fixing Anabaena flos-aquae low nitrogen mixing, poor light,
Nostocaleans Aphanizomenon low carbon, low phosphorus
H2 dinitrogen-fixing Anabaena lemmermanni low nitrogen mixing, poor light,
Nostocaleans of Gloeotrichia echinulata
larger mesotrophic lakes

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U summer epilimnia Uroglena low nutrients CO2 deficiency
LO summer epilimnia in Peridinium segregated prolonged or deep
mesotrophic lakes Woronichinia nutrients mixing
Merismopedia
LM summer epilimnia in Ceratium very low C, mixing, poor
eutrophic lakes Microcystis stratification light
M dielly mixed layers of small Microcystis high insolation flushing, low total
eutrophic, low latitude lakes Sphaerocavum light
R metalimnia of mesotrophic P. rubescens low light, strong instability
stratified lakes P. mougeotii segregation
V metalimnia of eutrophic Chromatium, very low light, instability
stratified lakes Chlorobium strong
segregation
W1 small organic ponds Euglenoids, Synura high BOD grazing
Gonium
W2 shallow mesotrophic lakes bottom-dwelling ? ?
Trachelomonas
Q small humic lakes Gonyostomum high colour ?

As indicated, the groups A to C were devised originally
to apply to the vernally growing diatoms in temperate
lakes of differing trophic levels. Group A is characterized
by a number of centric diatoms of the genus Cyclotella
(C. glomerata, C. comensis) and, especially, of Urosolenia (for-
merly Rhizosolenia) that are prominent in the plankton of
many medium-to-large high-latitude lakes that are typi-
cally clear, dilute in solutes and deficient in phosphorus.
The algae in this group are mostly unicellular (103–104
µm3 in volume), apparently have high affinity for nutri-
ents, though not, apparently, for carbon. Its distinctiveness
may earn the title ‘Urosolenietum’. The A association is also
represented in the clear, low-phosphorus Amazonian
flood-plain lake, Lago Batata, where it occurs together
with Merismopedia and Peridinium umbonatum (Group LO,
q.v.) during the filling and high-water stages (Melo and
Huszar, 2000). Group B diatoms belong to more enriched
lakes, merging into Group C without precise boundaries.
The large-celled Stephanodiscus (rotula, neoastraea) are indica-
tive of calcareous, often very phosphorus-rich, systems
where they may co-dominate with Asterionella and such
species of Aulacoseira as A. ambigua. Asterionella and other
Aulacoseira spp. (A. subarctica, A. islandica) and Cyclotella (C.
meneghiniana, C. stelligera) co-exist in softer-water lakes.
Diatoms of groups B and C form larger units (104–105
µm3 in volume) but shape contributes to the maintenance
of high s/v and efficient light-harvesting. Population
development is often subject to the availability of silicon

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JOURNAL OF PLANKTON RESEARCH VOLUME
and dependence upon turbulence for suspension leaves
them relatively sensitive to mixed depth and the seasonal
onset of near-surface density stratification.
Diatoms of Group D are mostly found in shallow, nutri-
ent-enriched, well-ventilated waters, liable to be turbid.
Small-celled (≤ 103 µm3 in volume) and fast growing, such
diatoms include planktonic Nitzschia spp., Synedra acus,
Stephanodiscus hantzschii, Cyclotella ocellata and, possibly,
Cyclotella pseudostelligera.
Groups N and P also involve diatoms but both are
associated either with lower latitudes or with the summer
period in temperate lakes. The dependence upon physical
mixing is strongly apparent, requiring a continuous or
semi-continuous mixed layer of 2–3 m in thickness. Both
associations can be represented in shallow lakes where the
mean depth is of this order or greater, as well as in the
epilimnia of stratified lakes when the mixing criterion is
satisfied. Species of Group P (which include Aulacoseira
granulata and Fragilaria crotonensis) are apparently more tol-
erant of carbon dioxide depletion than those of Group N
(Tabellaria spp.) and tend to be present in the more
eutrophic waters, subject to their other requirements
being satisfied. Both groups are characteristically associ-
ated with a presence of desmids, though these associations
have still to be investigated in detail. Some of the slender-
celled Closterium spp. and hard-water desmids (Staurastum
pingue) are P members, whereas the affinities of some
common planktonic Cosmarium, Staurodesmus, Pleurotaenium
and Xanthidium spp. are with N. Desmids may be more or
less abundant than diatoms in established assemblages
dominated by Groups N or P. Strong dominance by
desmids has also been noted in some tropical lakes as a
consequence of atelomixis, where the surface mixed layer
is alternately attenuated and deepened on a diel basis
(Barbosa and Padisák, 2002).
In more persistently mixed layers, in which light is
increasingly the limiting constraint, filamentous algae
show preferential adaptations. Some of these (Binuclearia,
Geminella, Mougeotia, Tribonema) were grouped relatively
recently (Reynolds, 1997), in spite of their disparate phylo-
genetic affinities, into Association T. This is one of the
groups that merges into another, the S group of photoad-
apting solitary filamentous Cyanoprokaryotes (Cyanobac-
teria). The most familiar of these is Planktothrix agardhii,
which is common among enriched, exposed and generally
shallow lakes at most latitudes, where it can aspire to
monocultural populations persisting throughout the year,
constituting ‘the third stable state’ to which shallow lakes
may gravitate (Scheffer, 1998). Other potentially dominant
species from the group include Limnothrix redekei, Pseudan-
abaena limnetica and Planktolyngbya contorta. The original,
temperate-lake Group (S1) reasonably aspires to a ‘Plank-
totrichetum’. The subdivision S2, comprising the group of
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genera, Spirulina, Arthrospira and Raphidiopsis that inhabit
warm, shallow and often very alkaline waters, was pro-
posed by Reynolds (Reynolds, 2000). The solitary habits of
Cylindrospermopsis and Anabaena minutissima often in rela-
tively phosphorus-rich lakes and their tolerance of vertical
mixing set them apart from other dinitrogen-fixing
members of the Nostocales. They had been allocated
earlier to Group SN (Padisák and Reynolds, 1998).
At the other extreme of warm-water systems are the
clear, low-biomass oligotrophic and ultra-oligotrophic
water columns in which relatively the largest biomass
resides in the picophytoplankton. Group Z was an ad hoc
creation to accommodate the organisms and what, at first,
seemed to be their habitats. The significance of
autotrophic picoplankton [photoautotrophs in the size
range 0.2–2 µm: (Sieburth et al., 1979)] to the carbon flow
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in the sea and in large lakes has been recognized for some
time (Fogg, 1986; Stockner, 1988; Pick, 2000). Picophyto-
plankton were originally understood to include single-
celled synechococcoid and Cyanobacteria and, certainly
in lakes, a number of chlorococcal species of Chlorella and
Choricystis. Prochlorophyte Cyanobacteria also figure in
the oceanic picoplankton (Chisholm et al., 1988). The
emphasis in the designation is the small unit sizes (unicells,
< 10 µm3 in volume), which have a tendency not to sink
out rapidly but to be persistent, at least for so long as they
remain substantially ungrazed. Colonies comprising small
cells of Cyanobacteria (such as Aphanocapsa) or of chloro-
phytes (such as Dictyosphaerium or Coenochloris) belong in
other groups (K and F respectively). Neither is it the case
that picophytoplankton is confined to oligotrophic
systems. The high biomasses of Synechocystis aquatalis,
Synechococcus and Chlorella minutissima observed by Huszar et
al. in certain coastal lagoons suggested to them a more
eutrophic assignation (X1) than is implied by the use of Z
for the synechococcoid picoplankton (Huszar et al., 2000).
Provided that the energy-, carbon- and nutrient-require-
ments of these algae can be met, they are not obliged to
live at low population densities (Vörös et al., 1991). Some
of the highest recorded densities of synechococcoid
picoplankton come from shallow, eutrophic ponds in
Antarctica where grazing on the autotrophs was also
apparently restricted (Izaguirre et al., 2001). In addition,
there is now recognized a functional difference between
the deep-water prokaryotes that contain phycoerythrin
pigment and those living in surface waters and do not (or
contain rather less; Padisák et al., 1997). Pending further
investigation, this distinction has not been formalized.
It is apparent that the Z association requires further
consideration. Huszar et al. ((Huszar et al., 2000) have gone
as far as a proposal to transfer all eukaryotic members
(such as Chloromonas and Chlorella minutissima) to X, on the
grounds that, functionally, these are simply the smallest
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C. S. REYNOLDS ET AL. FUNCTIONAL CLASSIFICATION OF FRESHWATER PHYTOPLANKTON
members of the nanoplankton. For the present, we retain
the Z category to refer to the picocyanobacteria that
dominate in the photic layers of oligotrophic lakes, either
in the upper mixed layers or forming deep chlorophyll
maxima in the upper hypolimnion, as they do in the
Stechlinsee (Padisák et al., 1997). Elsewhere, Olrik’s (Olrik,
1997) discrimination among the smaller nanoplankters
(mostly unicells, < 103 µm3 in volume) is sufficient to justify
the ascription of oligotrophic (X3), meso-eutrophic (X2)
and eutrophic-hypertrophic (X1) associations.
The Y association was set aside for larger nanoplanktic
flagellates (103–104 µm3 in volume), although it has been
used more or less synonymously with the larger Crypto-
monad species. Well-adapted to live in a wide-range of
habitats, their dynamics are, nevertheless, vulnerable to
the feeding activities of a wide range of crustacean zoo-
plankton and even of some rotifers and protistans.
In the earliest attempt to separate phytoplankton
associations, the sparse, oligotrophic–mesotrophic early
summer plankton, often dominated by chrysophytes and
mucilaginous colonial green algae, provided difficulties.
With more information, however, the groups E and F are
known to be represented in the plankton of a wide spec-
trum of lakes but not for the same reasons. Some of the
chrysophyte genera are demonstrably capable of strong
growth in the well-insolated water, where they can sup-
plement nutrient uptake from dilute solution by the
phagotrophic ingestion of bacteria. They are, however,
obligate consumers of carbon dioxide gas (i.e. they cannot
resort to bicarbonate as many eutrophic algae do). The
non-motile but near-neutrally-buoyant colonial green
algae of Group F seem to have an elevated light thresh-
old: they function best in clear water and are otherwise
tolerant of deep mixing. Thus, algae of both groups E and
F have a strong representation among mesotrophic lakes
but both are sensitive to nutrient enrichment and the
additional demands that high biomass may place on the
carbon and light fluxes.
In contrast, the strongly motile colonies of Volvox, Eudo-
rina and their allies are able to regulate their position in
relation to the light field. They respond to nutrient-rich
conditions in stagnating water columns and are, therefore,
most familiar in small eutrophic lakes and during very
stable phases in larger river-fed basins and storage reser-
voirs. The distinctive Group G could be proposed as a
‘Volvocetum’.
Another distinctive group ( J) of mainly non-gelatinous,
non-motile Chlorococcales is prominent in shallow, highly
enriched systems (including many low-gradient rivers),
represented by Scenedesmus, Pediastrum and Coelastrum. At
one end, that of barely colonial, mucilage-free chlorococ-
cals (such as Golenkinia, Treubaria), it merges into X1; at the
other, where light becomes critical, it merges into S.

Close to J is another group of (mostly) shallow-water
small-celled colonial Cyanoprokaryotes of the genera
Aphanocapsa, Aphanothece and other non-vacuolate types.
We are uncertain of the precise significance of abundant
populations of these Group-K species, save that they seem
to survive high pH particularly well and may thus repre-
sent an association in transition towards LM or M.
The Group H is another of long standing, having been
created originally for dinitrogen-fixing Cyanobacteria of
the bloom-forming Order Nostocales. However, the
group has been subject to recent reviews: the solitary
forms were recognized to be more Oscillatoria-like than the
buoyancy-regulating types in their ecology (Padisák and
Reynolds, 1998; Huszar et al., 2000) and were distin-
guished as SN (for nitrogen-fixing S types). Now, we feel
moved to separate the classical eutrophic, low nitrogen
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group (H1), built around Aphanizomenon flos-aquae, Anabaena
flos-aquae, An. circinalis and An. spiroides, from the group of
species (H2), such as An. lemmermanni, An. solitaria and
Gloeotrichia echinulata, that are tolerant of mixing conditions
in rather larger but typically less eutrophic (hence, clearer)
lakes (Reynolds and Lund, 1988; Garcia de Emiliani,
1993).
Among well-stratified lakes, the activities of planktonic
primary producers leads to further differentiation of the
habitat, so that the upper waters, though well-insolated,
become severely deficient, either or severally, in phos-
phorus or nitrogen or carbon, whereas beneath them is
water where nutrients are less depleted, or may even
accumulate through biological transport, but where there
is too little light for massive photoautotrophic exploitation
(Margalef, 1978). In this situation, one of the more advan-
tageous adaptive strategies is to be able to make substan-
tial vertical migrations between the two compartments.
The essential adaptation is the combination of efficient
motility with large size: this is evident among the algae of
the next four categories, all of which generate unit sizes
that approach 105 µm3 or are rather greater. U is for
Uroglena, possibly the unique member of the group of
large, motile, colony-forming chrysophytes that may be
observed in stratifying oligotrophic and mesotrophic
lakes. LO refers originally to the Peridinium–Woronichinia
association of stratified mesotrophic lakes, though some
usage has tried to accommodate more acidic assemblages
involving P. inconspicuum and such Cyanoprokaryotes as
Merismopedia. Ceratium hirundinella and similar species are
actually rather eurytrophic and may be associated equally
with Cyanoprokaryote associations based on Microcystis
aeruginosa as easily as with Woronichinia; indeed, all three
may co-exist within the confines of LM.
While the L assemblages may need careful review and
recasting, the near-relative M seems better characterized,
referring to the almost monocultures of large colonies of
01reynolds (58J)(ds) 29/4/02 10:15 am Page 424
JOURNAL OF PLANKTON RESEARCH VOLUME
such as Microcystis aeruginosa, with many of its overlapping
subspecies and ecotypes, Microcystis wesenbergii and Sphaero-
cavum brasiliensis. Each may be well over 106 µm3 in volume
and its buoyancy control copes not just with segregation
per se but accommodates quite striking diel fluctuations in
stratification and mixing in low-latitude lakes.
In many of the deep, clear alpine lakes, the metalimnia
are sufficiently stable to become populated by plate-like
developments of photoautotrophs that remain centred
within narrow depth ranges for weeks to months on end,
moving upwards or downwards in response to small
changes in the light field or in the availability of other
essential resources. The behaviour is typified by that of the
Cyanoprokaryote, Planktothrix (formerly Oscillatoria)
rubescens [(Zimmermann, 1969); see also (Bright and
Walsby, 2000)] which is recognized in the group R.
Besides having a remarkable capacity for sensitive buoy-
ancy regulation, the species has a powerful facility for
chromatic adaptation during stratification deep in the
light gradient. However, the alga also tolerates holomictic
entrainment and may even grow under these circum-
stances. In smaller stratifying lakes, other species of Plank-
tothrix (P. limosa, P. mougeotii) and Planktolyngbya (P. subtilis)
showing analogous physiological and behavioural adapta-
tions, may dominate. These are also understood to fit in
Group R. However, dominance of the stable, optically
deep-water layers in the stable gradients of small, density-
stratified lakes at all latitudes (tropical forest lakes, ice-
covered polar lakes, solution hollows) may be equally open
to dominance by chrysophyte (E) or cryptomonad (Y)
flagellates (Reynolds, 1997). Where there is also a pro-
nounced gradient in redox, the key autotrophs may be
species of the purple and green sulphur bacteria that
make up Group V.
Small ponds, well supplied by organic matter (either
from farm stock or vegetal decomposition, including leaf
fall), support algal flora which have not been well charac-
terized. There may well be present elements of the
D–J–X1 group of shallow enriched systems, possibly also
Y-type cryptomonads, though one or several of the
following components may also be represented: eugle-
noids (e.g. Euglena, Phacus, Lepocinclis), small colonial
volvocines (Gonium), some chrysophytes, provided the pH
is not too high (Synura spp.), and the smaller dinoflagellates
of the genera Peridinium and Glenodinium. Grouping these
in a single association (W, now W1) is provisional and
open to further separation. Euglenoids, represented
particularly by Trachelomonas, are also found in the distinc-
tive bottom-dwelling community of shallow, aerated lakes
which appears in open water on occasions. It is tentatively
distinguished as W2. Finally, a new codon is introduced
here, Q, to refer to the populations of Gonyostomum that
dominate in humic but generally productive forest lakes,

 NUMBER  PAGES ‒ 
mostly at high latitudes, usually having a low calcium
content and a pH on the acid side of neutrality [see
especially (Korneva, 2001)].
T H E A P P L I C AT I O N O F
F U N C T I O N A L G RO U P S
In this section, we seek to show some of the uses of the
functional-group model. These include the aid to under-
standing why certain species of phytoplankton should be
more favoured than others in the assembly of communi-
ties and why planktic species composition should vary in
space and time. Conversely, the similarity in plankton
assemblages in similar lakes and at similar times becomes
easier to explain, as do the differences in assemblage struc-
ture represented in lakes of different types. Such under-
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standing is the basis of making valid predictions or, at
least, of extrapolating probabilities, about the structure of
the phytoplankton. This must be of value to managers
who now, more than ever, require decision support in
determining their policies, priorities and reactions to
developing situations.
Typical problems for managers are to do with the
capacities of the waters in their charge to support phyto-
plankton and, once certain conditions are triggered, how
quickly the capacity may be filled. These are not questions
that the functional classification is designed to meet,
although appropriate simple models are now available
(Reynolds, 1992, 2002; Reynolds and Maberly, 2002).
However, the next question is about the identity of the
dominant algae and whether it is likely to cause nuisance
because of its latent toxicity, taste/odour implications, or
the ease of treatment for potable supplies (taken to be the
converse of any propensity to block filters or to penetrate
floc blankets). In the case of storage reservoirs, helixors,
circulating pumps, multiple depth inflows and draw-off
points might be operated in different ways to deliver a
more favourable raw water for treatment.
Limnologically and hydraulically, reservoirs differ
extensively from lakes but there is no difference in the
principles that eventually influence the composition of the
phytoplankton (Reynolds, 1999). In reservoirs, as in lakes,
predicting dominant species, peak populations and their
dates is almost impossible. Even with the benefit of hind-
sight, to recount the forces, their intensities and duration,
to say nothing of the stochasticity of the contributory
events, that led precisely to the observed condition, is
defied by the chaotic nature of variability.
Yet, as has often been pointed out, certain aspects of
plankton dynamics are broadly predictable. As an
example, spring blooms in temperate lakes, in response to
lengthening days more than warming waters, may be pre-
dicted with confidence. The dominant species may also be
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C. S. REYNOLDS ET AL. FUNCTIONAL CLASSIFICATION OF FRESHWATER PHYTOPLANKTON

predicted on the basis of standing inocula furnished by
past spring blooms, while the alternative dominants will
come from a robustly predictable group of species. These
groups correspond to the functional associations: there is
merit in being able to describe in these terms the sequence
of phytoplankton dominance across temporal and spatial
sequences of water bodies. One of the most comprehen-
sive of such comparisons was that of the phytoplankton of
deep alpine lakes made by Sommer (Sommer, 1986): he
demonstrated a similarity of year-to-year behaviours in
individual lakes, sequential stages (spring bloom, summer
stratification, the summer–autumn phase of mixed-layer
deepening) common to all of them, and of different kinds
of dominant association reflecting trophic status and
limiting nutrient availability. However, the patterns are
reported in terms of a few key dominants and of the
in a particular functional group because it is in a particu-
lar kind of lake, or it is in a particular kind of lake because
of its functional affinity. A much more pragmatic view is
that most species have a cosmopolitan or even ubiquitous
distribution but they may be more sensitive to certain
properties and environmental circumstances that weaken
their dynamic performances in comparison with those
that are less sensitive or positively tolerant of the same
properties or circumstances. The longer the conditions
obtain, the poorer is the survival of the first and the rela-
tively better is the survival of the second. In this way the
standing stock becomes biased towards the survivorship of
the tolerant species. In less exacting environments, more
species will, potentially operate successfully, but as the
limiting conditions become more severe and the opera-
tional criteria discount against less tolerant species, the

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phylogenetic groupings of the main species. Setting the
same information in terms of the functional associations
provides a striking summary of the patterns detected
(Table II). The classification also holds for the large lakes
south of the Alps (Salmaso, 2000). Huszar and Caraco
have made similarly convincing deductions on the basis of
their study of a series of lakes in the northeastern United
States (Huszar and Caraco, 1998). In the meantime, the
temporal and spatial variabilities in phytoplankton com-
position of several tropical and sub-tropical systems have
been readily described in terms of association represen-
tation (Huszar et al., 2000).
In order to apply the approach to the prediction of the
consequences of changed management practices—such
as the occurrence of toxic Cyanoprokaryota—it is helpful
to carry forward an understanding of the concepts of
tolerance and sensitivity shown in Table I and which
underpin the ‘+’ and ‘–’ response signs in Table III.
Without this, it is difficult to determine whether an alga is
survivors become more predictable. This is the principle
of community assembly and it underpins the generalized
application of the original model concept that we advo-
cate. In this way, we are able to assert that we have deter-
mined a system that is sensitive across latitudes,
morphometries and trophic states, by virtue only of the
algal preferences and sensitivities and not because of
where and when the algae are found. Thus, we can antici-
pate, for example, that Group S1 is not excluded from
tropical lakes (correctly so) but the warm-water preference
of Group S2 means that we should not expect it to be rep-
resented at high latitude (hypothesis not disproved). Simi-
larly, tolerance of nutrient deficiency or an inability to
operate at low concentrations of available carbon does not
prevent populations of species from the E, LO, X3 or Z
categories from attaining high concentrations in eutrophic
environments, provided that their light and carbon
requirements can be met (all are verifiable phenomena).
Hence, there is no longer any paradox about whether

Table II: Sommer’s summary of the seasonality of dominant algae in nine central European lakes re-
rendered in terms of functional-group representation (Sommer, 1986)

Spring Clear Summer Summer Summer Summer–

bloom water P↑ Si↑ P↓ Si↑ P↓ Si↓ Autumn

Köningsee A N/Y A
Attersee N Y B
Walensee N N/Y
V’wldst’see B/Y N N H R
L.Maggiore C/R P L/R H L/R
Ammersee C/R Y R/P/Y P L H R/P
Zürichsee D/Y Y Y/H P/Y R/P
Lac Léman D/Y Y Y/G P L H T/P
Bodensee D/Y Y Y/G P L /H/ E T/P


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Table III: Responses to habitat properties of functional groups of phytoplankton [updated from
(Reynolds, 2000)]

Variablea
hm I*  [P] [N] [Si] [CO2] f
Codon <3 < 1.5 <8 < 10–7 < 10–6 < 10–5 < 10–5 > 0.4

A – ? + + + + – –
B – + + + – – – –
C – + + – – – ? –
D + + + – – – + –
N – – – + – +/– – ?
P – – – – – +/– + +
T – ? – +/– – + ? +
S1 + + + – – + + +
S2 + + – – – + + +

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SN + + – – + + + +
Z + – + + + + ? –
X3 + – + + – + – –
X2 + – + ? – + ? –
X1 + – + – – + + –
Y + + + – – + ? –
E + + + + – + – –
F + – + + – + – –
G + – + – – + + +
J + ? + – – + ? –
K + ? – – – + + ?
H1, H2 + – – – + + + +
U + – ? + – + – +
LO + – – + – + – +
LM + – – – – + + +
M + – – – – + + +
R + + – – – + ? +
V + + – – – + – –
W1 + + + – – + ? –
W2 + + + – – + ? ?
Q + +? +? ? ? + ? ?

Entries in Table are to denote tolerance (+) or no positive benefit (–) of the environmental condition set.; ‘+/–’ is used to denote that some species in
the association are tolerant; ‘?’ denotes that tolerance is suspected but not proven.
a Variables signified are: depth of surface mixed layer (h , in m from surface); mean daily irradiance levels experienced (I*, in mol photons m–2 day–1);
m
water temperature (, in °C); the concentration of soluble reactive phosphorus ([P], in mol l–1); the concentration of dissolved inorganic nitrogen ([N], in
mol l–1); the concentration of soluble reactive silicon ([Si], in mol l–1); the concentration of dissolved carbon dioxide ([CO2], in mol l–1); and the proportion
of the water processed each day by rotiferan and crustacean zooplankton (f ).
b Some representative genera or species only are listed.

the algal assemblage or the kind of lake determines the
floristic group that may be favoured. Indeed, the supposed
circularity of the argument becomes a positive strength
when algae from the same functional association are
found under analogous conditions in different lakes of
similar types, as in Table II. Other examples include the
sensitivities of the various functional groups of
Cyanoprokaryota to the depth, stability and nutrient-
content of water columns, shown in a small number of
studies (Padisák and Reynolds, 1998; Beyruth, 2000;
Huszar et al., 2000). Thus, it should be possible to decide
the best way of managing a lake or reservoir to make it less


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C. S. REYNOLDS ET AL. FUNCTIONAL CLASSIFICATION OF FRESHWATER PHYTOPLANKTON
suitable for the growth of toxic Cyanoprokaryotes (reduce
nutrients? destratify? biomanipulate?). The existing know-
ledge to be able to do this is summarized in the semi-
quantitative check-list (albeit, still lacking in some details)
presented in Table III.
DEVELOPMENT OF THE
F U N C T I O N A L C L A S S I F I C AT I O N
The remaining objective of this presentation is to advo-
cate its wider adoption and to secure a wider constituency
for its development. We are certain that not all valid
groupings have been described, while the functional
significance of many algal species has not been accorded
to functional groups and it is not clear that it will be simple
to do so. The system as we have presented it has numer-
ous weaknesses, including the prediction of assemblages
of species with overlapping habitat preferences. As yet, we
have made no attempt to deal with tychoplanktonic
entries from the periphyton or from the benthos, although
to develop a scheme that included these life-forms in their
own right is likely to be a worthwhile objective. On the
other hand, the increasing availability of statistical
methods for sorting and discriminating among numerical
and graphical information (Ten Brink et al., 1991; Seip
and Reynolds, 1995; Kruk et al., 2002) invites additional
contributions to consolidating the functional-group
concept. The application of statistics based on Bayes’
theorem on the inferences of probability distributions
from data sets relating to the definition and distribution of
the functional categories has particular appeal, as does the
introduction of Artificial Neural Networks to hypothesis
analysis. What began as qualitative shorthand for com-
paring separate plankton assemblages, in space and time,
now aspires to a verifiable quantitative method of describ-
ing community structure and change.
We hope that this article will stimulate interest and the
further contributions of other plankton scientists. There
are sufficiently tempting analogies to propose that the
approach is suited to the marine phytoplankton (Smayda
and Reynolds, 2001), while the zooplankton may be
amenable to a similar form of ecological classification
(Romanovsky, 1985). We emphasize the still-provisional
element of the classification developed thus far and the
discursive network that we have put in place, so that pro-
posed developments, additions and re-classifications are
circulated (by e-mail) among the participating authors.
This is a reasonable interim arrangement that is capable
of expanding its consultative network, which can be
joined by contacting any of the authors. E-mail addresses
are appended. All will be welcomed!

AC K N O W L E D G E M E N T S
We are extremely grateful for the perceptive comments
and helpful suggestions of the referees and the improve-
ments they enabled us to make to our original manuscript.
C O N TAC T A D D R E S S E S
Colin S. Reynolds: csr@cah.ac.uk
Vera L. de M. Huszar: vhuszar@gbl.com.br
Carla Kruk: ckruk@fcien.edu.uy
Luigi Naselli-Flores: luigi.naselli@unipa.it
Sergio Melo: melo@biologia.ufrj.br
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Received on November 2, 2001; accepted on January 5, 2002