Lake Communities
Sarian Kosten, Department of Aquatic Ecology and Environmental Biology, Institute for
Water and Wetland Research, Radboud University Nijmegen, Nijmegen, The Netherlands
Mariana Meerhoff, Centro Universitario Regional Este, Universidad de la Republica,
Maldonado, Uruguay
Based in part on the previous version of this eLS article Lake Commu-
nities (2007) by Christer Bronmark and Lars-Anders Hansson.
As in all other ecosystems, lake community structure is
determined by processes at different spatial and temporal
scales, including biogeographical, regional and local
environmental conditions, speciation, and local biotic
interactions such as competition and predation. Lakes are
commonly classified according to: (1) productivity (e.g.
oligotrophic versus eutrophic), (2) where the major car-
bon input comes from or (3) thermal stratification pat-
terns and morphometry. Most lakes worldwide are
shallow and small. Within lakes we distinguish three dif-
ferent zones or habitats: the near-shore littoral zone, the
open-water pelagic zone and the lake bottom or benthic
zone. Each of these zones has a characteristic biological
community, although they interact in different ways. The
relative importance of each community to the whole
ecosystem functioning varies with lake morphometry and
productivity. Different anthropogenic activities influence
lake communities at a local or regional scale; global
changes such as climate change, however, represent a
new threat to lakes worldwide.
Introduction
A community is generally dened as an assemblage of
species coexisting in a specic environment at a specic
time, and community ecology deals with patterns and
processes at this scale. Accordingly, a lake community
consists of the species coexisting in a specic lake, including
shes, macroinvertebrates such as snails, shrimps and
insects, zooplankton such as watereas, vascular plants
(macrophytes), phytoplankton (typically microalgae and
cyanobacteria), periphyton, and heterotrophic micro-
organisms and sometimes aquatic mammals. To under-
stand what determines the structure of lake communities
eLS subject area: Ecology
How to cite:
Kosten, Sarian; and Meerhoff, Mariana (November 2014) Lake
Communities. In: eLS. John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0003177.pub2
eLS & 2014, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
. Introduction
. Within-lake Communities
. Local Biotic Interactions
. Current Threats to Lake Communities
Online posting date: 17th November 2014
we need to understand the processes that determine species
coexistence.
What determines community structure in
lakes?
For each lake, there is a regional pool of species that are
potential members of the local community, but any given
lake community is not just a random subset of the potential
colonists. Rather, the community structure of a specic
lake is a function of a number of processes operating at
dierent spatial and temporal scales (Figure 1). The location
on the globe (which roughly determines climate, that is,
solar radiation, temperature and precipitation patterns),
for instance, is an important determinant of community
composition and structure, but regional factors inuencing
colonisation and extinction chances as well as present-day
abiotic and biotic interactions in the local environment also
play a role (Figure 1; Bronmark and Hansson, 2005).
A useful approach to understand the interaction of pro-
cesses at dierent scales is to view the species composition
in a particular lake as the result of a series of dierent lters.
Biotic factors operating at the local scale, for example,
mutualism, competition and predation, can only aect the
species from the regional pool that have successfully
colonized the lake (Figure 1). See also: Community Ecology:
An Introduction
The size and composition of the regional species pool,
that is, the number and identity of species available for
colonisation of local lakes in a region, is determined by
factors operating at large geographic and historical scales.
At the largest scale, we can consider plate tectonics, that is,
the movement and collisions of continents that may
determine why some taxonomic groups are present or
absent on a continent. For example, Antarctic lakes lack
sh due to their biogeographic history. Most of these lakes
are melt water lakes with extremely low productivity, but
the ones closest to the seashore can be very productive due
to frequent visits by seals and birds using the lakes as water
closets and swimming pools. Hence, the lakes range in
productivity from ultraoligotrophic to highly productive,
but have only two major trophic levels (phytoplankton
and zooplankton, besides the microbial organisms)
(Hansson, 1992). See also: Biogeographical Regions;
Trophic Cascades
1
 Lake Communities

Global species pool a b c d e f g h i j

Biogeographic
processes,
history,
regional climate

Cl
im
at
e
ch
an
ge
Regional species pool b c e f g h

Abiotic frame:
local climate, Land use, water level
catchment, Anthropogenic pressures

lake characteristics

)
(x
c e f g

n
io
Biotic interactions:

ct
du
competition, predation, tro
in
p

parasitism, mutualism
Sp

Local assemblage e f x

Figure 1 A conceptual model showing how filters operating at different spatial scales determine the community structure of lakes, and how different
anthropogenic activities add new or modify natural filters. Modified from Bronmark and Hansson (2005).

Within a continent, climatic factors, range extensions
and contractions, glaciations and dispersal barriers aect
the number of species available in the regional species pool
by enhancing speciation and extinction. High numbers of
species are found in geologically ancient and rather isolated
lakes. New species have had ample time to evolve and
continue to do so. Owing to the poor dispersal abilities
many of these species are endemic, only occurring in that
specic lake. Good examples of lakes with endemic diverse
species ocks, resulting from the evolution of many species
from a single or a few founder species (adaptive radiation)
are the (world largest) Lake Baikal in Siberia and the
African rift valley lakes (Fryer, 1991). Lake Baikal has
many endemic species, but maybe the most spectacular
example is a group of freshwater shrimps (gammarids) with
more than 250 endemic species. The rift valley lakes have
endemic species ocks of, for example, snails and cichlid
shes. Especially the cichlid sh constitute an amazingly
diverse species ock that has evolved morphological and
behavioural specialisations and now includes species living
in dierent habitats, and/or feeding on dierent things. It is
thought that the evolution of such a large number of cichlid
species was promoted by lake level uctuations that iso-
lated bays of the present basin, but also that the wide
variation in coloration is related to sexual selection leading
to sympatric speciation (evolution without geographic
isolation). See also: Adaptive Radiation
On the regional and local spatial scales, community
structure is determined by both abiotic and biotic factors.
Although early limnologists saw lakes as microcosms and,
in fact, we still often think of lakes as ecosystems with well-
dened boundaries to the surrounding environment, lakes
are open ecosystems that are an inseparable part of their
catchments. The catchment (or drainage) area of a lake is
the region that drains the rain water to the lake. The size,
geological composition, slope, vegetation and human land
use of the catchment aect, for example, nutrient and
organic matter input, pH and the water colour of the lake.

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Lake morphometry (e.g. area, depth and exposure to pre-
vailing winds) is also determined by the landscape in the
catchment. Hence, many features of the catchment to a
great extent determine the environmental conditions that
are presented to the organisms in the lake (e.g. Kosten
et al., 2009a).
Climate and lake morphometry (in particular surface
area and depth), determine the thermal stratication pat-
tern of lakes. Solar radiation is the major source of heat in
lakes and most of the incoming light energy is converted
directly into heat that is absorbed within the rst few
metres of the water column. Wind-generated currents dis-
tribute heat within the lake but only down to a limited
depth. Thus, depending on lake depth and ambient tem-
perature, three layers of water are often formed: a warm,
less dense layer at the surface (epilimnion) and an under-
lying layer of dense, cool water (hypolimnion), separated
by an intermediate transition layer (metalimnion). This
thermal stratication is crucial for the physical, chemical
and biological processes in lakes. Stratication has strong
eects on the living conditions for lake communities, as not
only temperature but also the oxygen concentration diers
between the layers. Because of changes in solar radiation
and wind turbulence over the year, the stratication is often
not permanent, but varies among seasons (Figure 2). For a
given lake, though, there is a highly predictable yearly
pattern in temperature distribution. In dimictic lakes, the
lake straties into stable layers of water during summer and
winter, whereas during spring and autumn the lake water
mixes. Oxygen concentration is high in the epilimnion due
to photosynthesis by phytoplankton and passive diusion
Winter Spring Summer Autumn
Ice
Dimictic 4 C
4 C
Warm
monomictic
Polymictic
Figure 2 Common thermal stratification patterns in lakes. Light blue
indicates warm water, dark blue indicates cold water. Dimictic lakes mix
twice a year (in spring and autumn) and stratify in winter when they are
covered by ice and in summer when the surface waters (epilimnion) warm
up. Warm monomictic lakes never freeze, and are thermally stratified
throughout much of the year. Only during winter the surface waters cool to
a temperature equal to the bottom waters resulting in a mixing of the
different water layers (In contrast: cold monomictic lakes are covered by ice
throughout much of the year and mixing only occurs during the ice-free
period). Polymictic lakes mix throughout the year and especially in warm
regions or during heat waves stratify shortly in summer when there is
little wind or in winter when ice-covered (the latter is not shown in the
figure). Drawing by S. Kosten.
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Lake Communities
from the atmosphere. In the hypolimnion, the oxygen
concentration is generally low due to the decomposition of
organic material by microorganisms at the sedimentwater
interface and in the water column. The respiration of
benthic macroinvertebrates may consume signicant
amounts of oxygen as well. As there is no signicant input
of oxygen from the epilimnion, the oxygen concentration in
the hypolimnion decreases during the stratication period.
In eutrophic lakes with high amounts of organic matter
to be decomposed oxygen is eventually completely
depleted in the hypolimnion, making it an unsuitable
habitat for most animals and plants during a large part of
the stratication period, or even all year round in warm
climates. In other lakes there are other stratication pat-
terns. Warm monomictic lakes, for example, are never ice-
covered and mix throughout the winter. In shallow, wind-
exposed lakes in warm regions the stratication may be
quite unstable, only lasting from days to a few weeks at a
time. Such lakes are called polymictic. Lakes may also
dier in the extent of mixing during the mixing cycle. In
holomictic lakes, the whole water column is mixed during
circulation, whereas in meromictic lakes, typically lakes
with a large depth, the bottom layers are not involved in the
mixing. Most lakes worldwide are shallow and therefore
experience frequent mixing of the whole water column,
resulting in a strong water columnsediment interaction.
Climate change may strongly impact mixing regimes in
these shallow lakes, with considerable eects on oxygen
conditions, nutrient cycles and hence also community
composition (e.g. Wagner and Adrian, 2011).
The abiotic local environment is also strongly inuenced
by the nutrient loading to the lake. In the early twentieth
century, the European limnologists Einar Naumann and
August Thienemann (Thienemann, 1921; Naumann, 1929)
introduced the concept of trophic states, mainly based on
the abundance of phytoplankton communities and nutri-
ent concentrations, but also on sediment-living midge
larvae (chironomids). Oligotrophic lakes are typically
clear-water lakes with low productivity of phytoplankton
due to low concentrations of nutrients, mostly phosphorus
and nitrogen. As a consequence of the low productivity of
organisms at the base of the food web (e.g. phytoplankton
or periphyton) the productivity of organisms higher up in
the food web, such as herbivorous and predatory macro-
invertebrates and sh, is also low in oligotrophic lakes.
With increasing nutrient loading, lakes become more
eutrophic and consequently have higher primary and
secondary production. Many eutrophic lakes may have
widespread and productive stands of macrophytes, emer-
gent and submerged, as well as oating-leafed plants. Free-
oating plants may occur when ambient temperatures are
high enough and the lake or lake arms are wind-sheltered.
At the highest nutrient levels, typically in lakes that have
been polluted by humans, the submerged macrophytes
often disappear. The lake is then characterised by a high
phytoplankton biomass, resulting in highly turbid water,
lower biodiversity and even blooms of potentially toxic
cyanobacteria (Moss et al., 1990; Scheer et al., 1993).
3
 Lake Communities

Chances on cyanobacterial dominance are especially high
in warm lakes (Kosten et al., 2012). The more oligotrophic
situation is generally referred to as the clear water or
submerged macrophytes dominated state, whereas the
eutrophic situation is referred to as a turbid or phyto-
plankton dominated state. However, various feedback
mechanisms within the lake may cause a hysteresis eect
leading to a considerable range of nutrient loadings in
which a lake can be in either alternative state. Once, for
instance, submerged macrophytes have disappeared,
enhanced resuspension of sediment may cause the under-
water light conditions deteriorate. Nutrient loading then
has to be reduced beyond the loading at which the mac-
rophytes disappeared to diminish phytoplankton growth
and in this way improve light conditions for the return of
the macrophytes.
Some lakes get a high input of organic carbon from the
surrounding terrestrial environment. Such lakes generally
have low productivity, are brown coloured and are often
acidic due to high amounts of humic substances entering
from the catchment area. These lakes are called dys-
trophic lakes (Hansen, 1962).
The abiotic environment in the individual lake thus
ultimately determines which species from the regional
species pool can potentially succeed in colonizing the lake
and which are ltered out (Figure 1). Most species have a
rather narrow range of environmental conditions within
which they thrive, that is, their ecological niche. If, for
example, the winter or summer temperature is very low or
high, respectively, or if pH levels decrease below a thresh-
old value, many organisms will be unable to grow, repro-
duce and, eventually, survive in that habitat. For example,
nutrient status (productivity) and pH are particularly
important in determining dominance within the phyto-
plankton community, and the major phytoplankton
groups may be associated to points along gradients of pH
and productivity (Bronmark and Hansson, 2005).
Within-lake Communities
A lake can be divided into three major zones or habitats:
the pelagic, the littoral and the benthic or profundal
(Figure 3). Each zone typically has its own community of
organisms that are adapted to the special conditions that
exist there. Even though globally most lakes are shallow
and small, resulting in strong interactions among these
habitats, traditional limnology has mostly focused on
processes occurring in the pelagic.

Riparian

Pelagic Littoral
Piscivorous fish

Macrophytes Planktivorous
fish

Macroinvertebrates

Zooplankton Periphyton

Euphotic zone Phytoplankton

Profundal or benthic
Benthivorous fish

Figure 3 The different habitats of a lake, including the near-shore littoral zone, the open-water pelagic zone and the benthic or profundal zone where low-
light levels inhibit the growth of primary producers (aphotic zone). Representative organisms from the main communities of the classical food web and
their feeding interactions are shown. Arrows indicate the major fluxes of nutrients among lakes zones and with the surrounding terrestrial environment
through the riparian zone. Subsidies from the lake to the land may be locally important. Drawing by M. Meerhoff & T. Christensen.

4 eLS & 2014, John Wiley & Sons, Ltd. www.els.net


Pelagic
The pelagic zone is dominated by planktonic organisms, that
is, microscopic primary producers (phytoplankton) and
animals (zooplankton) that live suspended in the water col-
umn and have no or restricted capabilities of movement in a
world perceived as highly viscous due to their small body size.
Phytoplankton comprises of microalgae and cyanobacteria
of diverse phylogenetic origins and a diverse range in size and
shape, from single-celled species a few micrometres long to
colonial or lamentous forms that can easily be seen by the
naked eye. Phytoplankton morphologies range from simple,
circular shapes, such as in many centric diatoms, to highly
specialized spined or agellated species (e.g. Reynolds, 2006).
The variability and diversity make phytoplankton a com-
mon and successful life form in all lakes, since there are
always some species that t in a specic constellation of
environmental features. The most common groups are green
algae, diatoms, dinoagellates and cyanobacteria.
Phytoplankton are typically consumed by zooplankton
the major herbivores in the pelagic community. Two
important groups, based on their grazing capacity, are the
Cladocera, of which the waterea (Daphnia) is a common
example in many lakes, and Copepoda. Zooplankton species
have a wide variety of food sources; and although most graze
on phytoplankton, many are omnivorous. Some species
include bacteria in their diet whereas others are predators.
Fishless lakes and ponds are often dominated by large
zooplankton species, whereas in eutrophic temperate lakes,
and in many warm climate lakes, the zooplankton assem-
blage is dominated by small-bodied groups with low grazing
capacity, such as rotifers, due to the high, size-selective
predation pressure from sh. Through their consumption of
bacteria, zooplankton, especially the ciliates and hetero-
trophic nanoagellates, form the link between the classical
food web (e.g. Figure 3) and the microbial loop. Bacteria use
dead organic matter suspended or dissolved in the water
column as a carbon source. The organic matter consumed by
bacteria and secondary producers was traditionally
thought to be produced within the lake (autochtonous car-
bon); recent studies, however, show that a substantial part of
the organic matter actually originates from the surrounding
area (allochtonous carbon) (Pace et al., 2004).
Benthic or profundal
The lake bottom under the pelagic zone is called the benthic
or profundal zone. The zoobenthic community, that is,
organisms that live on solid substrates or in the sediments,
typically consists of chironomid larvae, dierent oligochaete
worms and occasional mussels. The profundal is typically
rich in detritus food sources but provides an environment
that is rather inhospitable to most organisms due to the
often low levels of oxygen. The benthic zone also stores
nutrients that are released back to the pelagic zone (i.e.
internal nutrient load) when sediment is resuspended by
wind or by organisms that dwell on or inside the sediment.
Additionally, anoxic conditions may facilitate the release of
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Lake Communities
phosphorus (e.g. Horppila and Kairesalo, 1992; Snder-
gaard et al., 2003; Figure 3). Depending on lake depth and
water characteristics that enhance the attenuation of
incoming light (such as water colour and amount of sus-
pended matter), the profundal may be aphotic, that is, light
is no longer sucient to allow photosynthesis by benthic
primary producers (in general referred to as periphyton).
Photosynthesis by periphyton (as by the other primary
producers), is thus restricted to the euphotic zone (typically
estimated as the depth with 41% of the incoming light).
Littoral
The littoral zone is the near-shore area where there is
typically enough light available for rooted submerged
macrophytes to grow. The littoral connects the riparian
(landwater interface) with the inner lake communities and
processes. The extent of the littoral zone depends on the
transparency of the water and the slope of the bottom. In
shores with a gently sloping bottom the littoral zone
extends further out into the lake. On wind-exposed shores,
a zone devoid of macrophytes and with few macro-
invertebrate species may exist due to constant sediment
resuspension or erosion (exposing a rocky shore) caused by
wave action (Chambers, 1987; Chambers and Kal, 1985).
Macrophytes are often covered with attached microalgae
(periphyton) that provide a rich food source for many
herbivorous macroinvertebrates, such as snails, insect
larvae (mayies and caddis ies) and crustaceans (amphi-
pods and isopods). In shallow lakes, submerged plants
often represent a diurnal refuge for predation-sensitive
organisms, such as cladoceran zooplankton, against visual
sh predators. This may facilitate the survival of grazing
zooplankton (Timms and Moss, 1984; Jeppesen et al.,
1998). The refuge capacity of plants, however, varies with
climate, likely as a consequence of the assemblage structure
and size distribution of predators (Meerho et al., 2007a,b;
Kosten et al., 2009b). Predatory macroinvertebrates, such
as dragony and damsely larvae and leeches, and many
sh species (particularly of small body size) also use the
structurally complex habitat of the littoral zone to repro-
duce, feed or hide from their own predators.
Habitat coupling
Lake food webs are often compartmentalised based on the
described within-lake habitats. Organisms in the pelagic
and benthic habitat are then assumed to belong to discrete
food webs with negligible transfer of energy between them.
This assumption holds, for instance, for some sh species
that have a specialized diet and are associated with one lake
zone only example, planktivorous sh in the pelagic zone.
Recent studies, however, have shown that the benthic and
pelagic food chains are often strongly linked. Analyses of
food sources of sh species (and of swimming macro-
invertebrates such as shrimps) caught in the pelagic clearly
illustrates this benthicpelagic coupling; a major propor-
tion of their diet often consists of benthic (and littoral)
5
 Lake Communities

Piscivorous fish

Piscivorous fish

Other fish
Shrimp (omnivorous)
Other fish

Lit. invertebrates (c.) Pel. invertebrates (c.)

Lit. invertebrates (h.) Lit. invertebrates (c.)

Cladocerans
Lit. invertebrates (h.) Pel. invertebrates (c.)
Phyto
Periphyton Cladocerans

Phyto
Periphyton
Figure 4 Simplified scheme of trophic interactions among main trophic groups in temperate and warm shallow lakes with comparable phytoplankton
(phyto) biomass. The densities in the subtropics are expressed relative to those in the temperate lakes (considered as the unit, for being the most known
web). Shrimp relative density is dotted due to typical shrimp absence in temperate lakes. The trophic groups were classified as primary producers
(periphyton and phytoplankton), intermediate herbivores (h.) such as cladocerans, and other invertebrates of littoral (Lit.) and pelagic (Pel.) zones,
intermediate carnivores (c.), intermediate omnivores, and top carnivores (piscivorous fish). Except fish, the same taxa shared the same trophic classification
in both climate zones. Modified with permission from Meerhoff et al. (2007a). Copyright & 2007, John Wiley and Sons.

animals (Vadeboncoeur et al., 2002; Solomon et al., 2011).
Many species swim across habitat boundaries daily or
switch habitats during their development (ontogeny).
Many piscivorous shes feed on zooplankton in the littoral
zone and on macroinvertebrates in the benthic zone as
juveniles and switch to a (partially pelagic) sh diet when
they are large enough to attack sh prey.
Energetic links from the littoral and benthic food webs to
the pelagic food web are thus important for the whole-lake
energy dynamics. However, the energy pathway may also go
the other direction, that is, from the pelagic food web to
organisms in the benthic habitat. For example, much of the
production of benthic organisms, such as chironomid larvae
and oligochaetes, in the profundal soft sediments depend on
the downfall of detritus from the pelagic. The importance of
each habitat for the whole-lake primary and secondary pro-
duction varies with lake area, depth and trophic state
(Vadeboncoeur et al., 2002; Liboriussen and Jeppesen, 2003).
Habitat use and food sources of lake organisms vary
substantially among climatic regions. In subtropical and
tropical lakes omnivorous sh are more common than in
similar temperate lakes (Teixeira-de Mello et al., 2009;
Gonzalez-Bergonzoni et al., 2012). Besides, sh more fre-
quently use the littoral and benthic habitats in warm lakes.
These dierences in sh community structure result in
substantial dissimilarities in habitat coupling and lake
communities in dierent climatic regions (Figure 4;
Meerho et al., 2007a).
Local Biotic Interactions
Although not all species that are able to colonise a lake and
tolerate the environmental constraints persist and form
viable populations, founder events combined with rapid
local adaptation are important processes determining
which species (or genotypes) become dominant in a lake.
Rapidly adapting and fast growing species can fully exploit
the resources of new areas and high population densities
may prevent the successful invasion of other species (De
Meester et al., 2002). Species composition and diversity is
further inuenced by biotic interactions such as competi-
tion, predation, parasitism and mutualism. The outcome
of the competition for light and nutrients among dierent
primary producers, for instance, largely determines which
kind of primary producer (submerged macrophytes,
benthic microalgae, free-oating macrophytes or phyto-
plankton) will dominate the lake. This has direct con-
sequences for the relative importance of the dierent
habitats within the lake (see Within-lake communities). In
lakes where competition for nutrients is important due to
nutrient limitation in the water, submerged macrophytes
are at an advantage as most can take up nutrients from
both the water and the sediments and can thus outcompete
non-rooted primary producers. However, when nutrients
in the water column are plentiful and phytoplankton
growth is unlimited by nutrient availability, phyto-
plankton biomass can reach levels where the water gets so
turbid that light does not penetrate to the lake bottom.
Light then becomes a limiting resource for submerged
macrophytes (and benthic algae), limiting their depth dis-
tribution and eventually making them disappear com-
pletely. At a smaller scale, microalgae that live attached to
the surface of macrophytes may reduce the growth of their
macrophyte host also by competition for light and nutri-
ents (Phillips et al., 1978). Alternatively, when conditions
are favourable for free-oating plants (i.e. high nutrient
levels, nonfreezing temperatures and wind protection),

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their growth at the water surface may lead to outshade both
submerged plants and phytoplankton. In sh, intraspecic
competition has strong negative eects on growth and
fecundity. In lakes with many sh species, there may be
competition for zooplankton and macroinvertebrates as a
food source between planktivorous and benthivorous sh
and juveniles of sh species that are specialized piscivores
as adults. This competition may be suciently strong to
severely restrict the recruitment of the predator species to
the piscivorous stage. Strong competition for the better-
quality animal resources is one mechanism that might
explain the dominance of the omnivory strategy among
tropical and subtropical shes.
Although competition may be a strong structuring force
in many lake communities, predation and herbivory are
major causes of mortality for many freshwater organisms
and may thus reduce population densities to levels where
competition for resources is no longer limiting. For
example, when piscivorous sh are introduced, intra- and
interspecic competition among planktivorous sh may no
longer be important. Besides, predators are often size-
selective preferring to feed on larger prey because of the
higher energy return, and this may have strong con-
sequences for the species composition and size-structure of
the prey populations. A classical whole-lake experiment in
the 1960s clearly illustrated this. During the experiment,
sh were introduced into a shless lake causing a dramatic
shift in the zooplankton community, from large-bodied to
smaller species (Brooks and Dodson, 1965). The mechan-
ism explaining this pattern was coined the sizeeciency
hypothesis. At low sh densities, large zooplankton
dominate because of their ecient grazing and their ability
to feed on particles with a wider size range. Large clado-
ceran zooplankton such as Daphnia, however, are more
vulnerable to sh predation, which results in a dominance
of small-bodied, less vulnerable zooplankton at high sh
densities. The sizeeciency hypothesis further predicts
that at high planktivorous sh densities, the phyto-
plankton biomass will be high, whereas at low planktivore
densities, large zooplankton will eciently reduce phyto-
plankton biomass. Brooks and Dodson thus not only
postulated a mechanism for how sh predation aects
zooplankton population density, species composition and
size-structure, but also a mechanism for how predation
eects may propagate to lower trophic levels, that is, an
explanation of cascading trophic interactions.
A trophic cascade is an indirect interaction character-
istic of linear food chains where a predator species has an
indirect positive eect on a primary producer species by
reducing the abundance of the herbivore species feeding on
the primary producer. In lakes, changes in sh populations
typically the top trophic level are expected to cascade
down to lower trophic levels and nally aect the primary
producers typically phytoplankton via zooplankton
(Carpenter et al., 1987). Besides, sh eects can also cas-
cade down on periphyton through predation on macro-
invertebrate grazers (Jones and Sayer, 2003). The
predictions of the trophic cascade concept have been tested
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Lake Communities
and conrmed in a large number of experimental manip-
ulations at dierent scales in dierent parts of the world,
and are most pronounced in cold and temperate lakes
(Jeppesen et al., 2012). Fish evidently play a strong struc-
turing role in lakes. Their size-selective predation aects
pelagic, benthic as well as littoral organisms and inuences
species composition as well as the size-structure of lower
trophic levels. See also: Experimental Systems in Aquatic
Ecology; Trophic Cascades
In most studies showing strong cascading eects in
freshwater food chains, the eects have been caused by
direct (lethal) predatorprey interactions. However, pre-
dators may also inuence the behaviour of their prey, for
example, by reducing their feeding activity or by changing
their habitat/refuge use. Predator-mediated changes in the
habitat use of an intermediate consumer may aect its diet
and thus change the strength and direction of interactions
in food chains (Romare and Hansson, 2003; Meerho
et al., 2006).
Predators and parasites not only inuence current prey
and host populations but may trigger changes in future
generations as well. Natural selection by sh on zoo-
plankton, for instance, can cause genetic changes inuen-
cing the phototactic behaviour of zooplankton. This rapid
evolution improves the predator-avoidance of the zoo-
plankton (Cousyn et al., 2001). Similar rapid evolutionary
adaptations have been found in reaction to exposure to
parasites (Pauwels et al., 2014). See also: Ecological
Genetics
Current Threats to Lake Communities
So far we have described the dierent interacting factors
that shape natural lake communities. Unfortunately, we
can no longer analyse and understand ecological processes
in isolation of human activities. Anthropogenic eects may
originate from activities within the lakes catchment, but
some disturbances have a very distant source, both in space
and time. The anthropogenic eects aect lake commu-
nities at dierent scales, by altering dierent species sort-
ing lters (Figure 1). Humans may aect both the lakes
abiotic environment and the biotic environment, often by
directly altering species composition. Here, we will only
briey mention some of the threats that have had and will
continue to have strong adverse eects on lake commu-
nities. Although we present them independently, it is
important to bear in mind that these processes may occur
simultaneously and interact with one another in nonlinear
ways, leading to often unpredictable emergent properties.
Eutrophication
The eutrophication process of lakes is characterised by
increased nutrient (mostly nitrogen and phosphorus)
concentrations due to input of, for example, human sewage
or fertilizer run-o from agricultural areas. This addition
triggers a chain of events starting with a massive increase in
7
 Lake Communities
Density
Collapse of both density and biomass
Biomass
Body size
Lake productivity
Figure 5 Conceptual scheme of changes in the fish community structure
(mean body size, density and biomass) along a productivity gradient
(eutrophication process). Often, a decrease in oxygen concentration as a
consequence of very high nutrient levels may promote sudden massive fish
kills. Drawing by M. Meerhoff and T. Christensen.
the growth of primary producers since these are generally
growth-limited by nitrogen and mostly by phosphorus in
freshwater ecosystems. Periphyton and submerged mac-
rophytes, and in warm locations free-oating macro-
phytes, often also show an increase in biomass. However,
phytoplankton, especially cyanobacteria and green algae,
typically take over as the dominant primary producers by
reducing the amount of light penetrating down to periph-
yton and submerged macrophytes. The high production
results in an increasing amount of dead organisms accu-
mulating as sediment. Bacteria that break down this
organic material need large amounts of oxygen, leading to
reduced oxygen concentrations and sometimes anoxia.
This is especially pronounced during stratication, when
oxygen-rich surface water does not mix with the water in
the deeper layer. After an initial increase in sh biomass,
average body size and density at the onset of eutrophica-
tion, the composition of the sh assemblage changes and
the amount of small-bodied cyprinid shes generally
increases considerably during the eutrophication process
(Figure 5). Additionally, in hypereutrophic systems massive
sh kills are frequent due to the oxygen decit. Since
cyprinids are ecient predators on zooplankton and
macroinvertebrates, the abundance and size of these preys
decreases. Hence, the eutrophication process leads to
considerable changes in the structure of the lake ecosystem,
which in turn aect the previously associated ecosystem
services (e.g. recreation, shing and use as a source of
drinking water). See also: Eutrophication of Lakes and
Rivers
Acidification
Acid rain, mainly as a consequence of industrial activities,
has strong eects on lake communities by reducing the pH
8 eLS & 2014, John Wiley & Sons, Ltd. www.els.net
of the water. Especially soft water lakes, that is, lakes with a
low buering capacity, are at risk. Already at pH levels of
56, the phytoplankton species diversity decreases con-
siderably, mainly because cyanobacteria and diatoms dis-
appear and dinoagellates and chrysophytes become
dominant. The decrease in the number of phytoplankton
species, as well as their biomass, leads to higher water
transparency, a characteristic feature of acidied lakes.
Acid-tolerant groups, such as Sphagnum mosses, may
ourish. The fauna also becomes less diverse as acidica-
tion proceeds. In northern temperate lakes, when the pH
drops to between 5 and 6, amphipods, craysh and shes
such as roach (Rutilus rutilus) are lost from the system,
whereas pike (Esox lucius) and perch (Perca uviatilis), as
well as mayies, remain even at pH levels below 5. Further,
one of the most signicant features of acidied lakes is the
disturbed reproduction of sh, resulting in sh populations
being dominated by large, old individuals. As these indi-
viduals eventually die, the lake becomes shless (Bronmark
and Hansson, 2005). Although dierent (industrial and
agricultural) management measures have considerably
reduced the acid rain in large parts of Europe and the
United States, many lakes in these regions still suer the
consequences of past acidication (e.g. Lucassen et al.,
2012; Angeler and Johnson, 2012). Moreover, acid rain is a
major current threat in regions with upcoming economies
and increasing industries.
Invasive species
The intentional or accidental introduction of species to
lakes where they do not naturally occur (often called
exotic or invasive species indistinctively, although con-
cepts dier slightly) can have strong eects on lake com-
munities. Eects of the many exotic species on native
communities are copious and may work through dierent
pathways such as habitat modication, predation, her-
bivory, competition, hybridisation and the transmission of
pathogens. Often a combination of dierent pathways
plays a role (e.g. Ellis et al., 2011). When water plant-eating
cray-sh are introduced, for instance, their herbivory also
aects the habitat of many littoral species (Nystrom and
Strand, 1996). Although there are many dierent types of
organisms that have been introduced to lakes by humans
(e.g. plants, mussels, crabs and shrimps), sh are the most
notorious group. Worldwide, exotic shes are introduced
in lakes to subsequently be shed upon. This can have
severe ecological (and often economic and social) con-
sequences. A well-known example is the introduction of
Nile perch in Lake Victoria, where it caused a severe
reduction in the native cichlid biodiversity (Goldschmidt
et al., 1993). The eects of invasive species are often highly
unpredictable, and as removal of invasive species from the
lake is hard and often even impossible, prevention of the
introduction of invasive species and decreasing the inva-
sibility of the system by removing other external pressures
are important management measures.
 Lake Communities

Warming Changes in precipitation

g
in e
ad tim
M lo
or in ce
e es en

Due igher

Com yan
m g id
in an res

ion
er Ch nd

of c
h
to g temp
al

pet obac
tat
iza a

ntr nd
tio
ap

itive
ol
na

row
ad n
ati ctio

th o ratures
co

adv ria
al

Inc
gic

red du
on rat rea

anta
te
ptim
es

e
More and sed g
olo

s p pro

gre ro
ysi

ate wth

ge
nutrients

um
r
Ph

re

r su
Mo

rviv

at
fish
l es

more al
food, ling More
More recyc
nu trient floating vegetation
More nced
Enha r
small fish fo
on ht
i lig
More cyanobacteria tit d
pe an
Hi

m ts
g

higher algal biomass co

he

en
tri
rp

Com
nu
re

g petit
grazin
da

nutr ion
Less ient for
tio

s an
n

Less d lig
ht Less
large zooplankton submerged
vegetation
More
mineralization

Figure 6 Some relationships now established that link climate change and an enhancement of eutrophication symptoms (Moss et al., 2011, reproduced by
permission of the Freshwater Biological Association. Drawing by A.R. Joyner).

Climate change processes, enhancing the release of phosphorus from the

sediment due to favourable redox conditions and by
Climate change represents one of the most recent threats to accelerating the decomposition of organic matter (e.g.
freshwater ecosystems, both directly by aecting, for Moss et al., 2011). In particular, it is expected that climate
instance, the water balance (precipitation and evaporation warming will promote the occurrence of bloom forming
regimes), heat retention, stratication patterns, and the cyanobacteria, increase water turbidity and threaten the
thermal biology of organisms, and indirectly by aecting tness particularly of large-bodied predatory shes
the need for irrigation and water level control in many (Figure 6).
regions worldwide. Climate change eects will interact with other external
The sensitivity of particular lakes and their biota to local stressors currently aecting the state and function of lakes,
changes in climate depends again on the lake location and with consequences on community structure (Meerho
associated climate regime, the characteristics of the et al., 2012) and overall lake functioning not yet
catchment, morphometric characteristics of the lake (such predictable.
as area, depth and fetch), lake history and nutrient status.
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Further Reading
Bronmark C and Hansson L-A (2006) The Biology of Ponds and
Lakes, 2nd edn. Oxford: Oxford University Press.
eLS & 2014, John Wiley & Sons, Ltd. www.els.net
Lake Communities
Carpenter SR and Kitchell JF (1993) The Tropic Cascade in Lakes.
Cambridge: Cambridge University Press.
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Restoration and Management of Lakes and Reservoirs. Boca
Raton, FL: CRC Press.
Lampert W and Sommer U (1997) Limnoecology: The Ecology of
Lakes and Streams. New York, NY: Oxford University Press.
Likens GE (2010) Lake Ecosystem Ecology: A Global Perspective.
Waltham, MA: Academic Press.
Moss BR (2009) Ecology of Fresh Waters: Man and Medium, Past
to Future. West Sussex, England: John Wiley & Sons.
Moss B, Madgwick J and Phillips G (1996) A Guide to the
Restoration of Nutrient-Enriched Shallow Lakes (Wetlands
International Publication), 180 pp. Norwich, UK: Broads
Authority.
Scheer M (1998) Ecology of Freshwater Lakes. London: Chap-
man and Hall.
Wetzel RG (2001) Limnology: Lake and River Ecosystems. New
York, NY: Academic Press.
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