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Lake Communities Introductory article

Sarian Kosten, Department of Aquatic Ecology and Environmental Biology, Institute for Article Contents
. Introduction
Water and Wetland Research, Radboud University Nijmegen, Nijmegen, The Netherlands
. Within-lake Communities
Mariana Meerhoff, Centro Universitario Regional Este, Universidad de la Republica, . Local Biotic Interactions
Maldonado, Uruguay . Current Threats to Lake Communities

Based in part on the previous version of this eLS article Lake Commu- Online posting date: 17th November 2014
nities (2007) by Christer Bronmark and Lars-Anders Hansson.

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

eLS & 2014, John Wiley & Sons, Ltd. 1

Lake Communities

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

regional climate

Regional species pool b c e f g h

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

lake characteristics

c e f g

Biotic interactions:

competition, predation, tro

parasitism, mutualism

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

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Lake Communities

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

eLS & 2014, John Wiley & Sons, Ltd. 3

Lake Communities

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


Pelagic Littoral
Piscivorous fish

Macrophytes Planktivorous


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.

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Pelagic phosphorus (e.g. Horppila and Kairesalo, 1992; Snder-

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

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Piscivorous fish

Piscivorous fish

Other fish
Shrimp (omnivorous)
Other fish

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

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

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

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

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their growth at the water surface may lead to outshade both and conrmed in a large number of experimental manip-
submerged plants and phytoplankton. In sh, intraspecic ulations at dierent scales in dierent parts of the world,
competition has strong negative eects on growth and and are most pronounced in cold and temperate lakes
fecundity. In lakes with many sh species, there may be (Jeppesen et al., 2012). Fish evidently play a strong struc-
competition for zooplankton and macroinvertebrates as a turing role in lakes. Their size-selective predation aects
food source between planktivorous and benthivorous sh pelagic, benthic as well as littoral organisms and inuences
and juveniles of sh species that are specialized piscivores species composition as well as the size-structure of lower
as adults. This competition may be suciently strong to trophic levels. See also: Experimental Systems in Aquatic
severely restrict the recruitment of the predator species to Ecology; Trophic Cascades
the piscivorous stage. Strong competition for the better- In most studies showing strong cascading eects in
quality animal resources is one mechanism that might freshwater food chains, the eects have been caused by
explain the dominance of the omnivory strategy among direct (lethal) predatorprey interactions. However, pre-
tropical and subtropical shes. dators may also inuence the behaviour of their prey, for
Although competition may be a strong structuring force example, by reducing their feeding activity or by changing
in many lake communities, predation and herbivory are their habitat/refuge use. Predator-mediated changes in the
major causes of mortality for many freshwater organisms habitat use of an intermediate consumer may aect its diet
and may thus reduce population densities to levels where and thus change the strength and direction of interactions
competition for resources is no longer limiting. For in food chains (Romare and Hansson, 2003; Meerho
example, when piscivorous sh are introduced, intra- and et al., 2006).
interspecic competition among planktivorous sh may no Predators and parasites not only inuence current prey
longer be important. Besides, predators are often size- and host populations but may trigger changes in future
selective preferring to feed on larger prey because of the generations as well. Natural selection by sh on zoo-
higher energy return, and this may have strong con- plankton, for instance, can cause genetic changes inuen-
sequences for the species composition and size-structure of cing the phototactic behaviour of zooplankton. This rapid
the prey populations. A classical whole-lake experiment in evolution improves the predator-avoidance of the zoo-
the 1960s clearly illustrated this. During the experiment, plankton (Cousyn et al., 2001). Similar rapid evolutionary
sh were introduced into a shless lake causing a dramatic adaptations have been found in reaction to exposure to
shift in the zooplankton community, from large-bodied to parasites (Pauwels et al., 2014). See also: Ecological
smaller species (Brooks and Dodson, 1965). The mechan- Genetics
ism explaining this pattern was coined the sizeeciency
hypothesis. At low sh densities, large zooplankton
dominate because of their ecient grazing and their ability Current Threats to Lake Communities
to feed on particles with a wider size range. Large clado-
ceran zooplankton such as Daphnia, however, are more So far we have described the dierent interacting factors
vulnerable to sh predation, which results in a dominance that shape natural lake communities. Unfortunately, we
of small-bodied, less vulnerable zooplankton at high sh can no longer analyse and understand ecological processes
densities. The sizeeciency hypothesis further predicts in isolation of human activities. Anthropogenic eects may
that at high planktivorous sh densities, the phyto- originate from activities within the lakes catchment, but
plankton biomass will be high, whereas at low planktivore some disturbances have a very distant source, both in space
densities, large zooplankton will eciently reduce phyto- and time. The anthropogenic eects aect lake commu-
plankton biomass. Brooks and Dodson thus not only nities at dierent scales, by altering dierent species sort-
postulated a mechanism for how sh predation aects ing lters (Figure 1). Humans may aect both the lakes
zooplankton population density, species composition and abiotic environment and the biotic environment, often by
size-structure, but also a mechanism for how predation directly altering species composition. Here, we will only
eects may propagate to lower trophic levels, that is, an briey mention some of the threats that have had and will
explanation of cascading trophic interactions. continue to have strong adverse eects on lake commu-
A trophic cascade is an indirect interaction character- nities. Although we present them independently, it is
istic of linear food chains where a predator species has an important to bear in mind that these processes may occur
indirect positive eect on a primary producer species by simultaneously and interact with one another in nonlinear
reducing the abundance of the herbivore species feeding on ways, leading to often unpredictable emergent properties.
the primary producer. In lakes, changes in sh populations
typically the top trophic level are expected to cascade Eutrophication
down to lower trophic levels and nally aect the primary
producers typically phytoplankton via zooplankton The eutrophication process of lakes is characterised by
(Carpenter et al., 1987). Besides, sh eects can also cas- increased nutrient (mostly nitrogen and phosphorus)
cade down on periphyton through predation on macro- concentrations due to input of, for example, human sewage
invertebrate grazers (Jones and Sayer, 2003). The or fertilizer run-o from agricultural areas. This addition
predictions of the trophic cascade concept have been tested triggers a chain of events starting with a massive increase in

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of the water. Especially soft water lakes, that is, lakes with a
Density low buering capacity, are at risk. Already at pH levels of
56, the phytoplankton species diversity decreases con-

Collapse of both density and biomass

Biomass 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
Body size 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,
Lake productivity one of the most signicant features of acidied lakes is the
disturbed reproduction of sh, resulting in sh populations
Figure 5 Conceptual scheme of changes in the fish community structure
(mean body size, density and biomass) along a productivity gradient
being dominated by large, old individuals. As these indi-
(eutrophication process). Often, a decrease in oxygen concentration as a viduals eventually die, the lake becomes shless (Bronmark
consequence of very high nutrient levels may promote sudden massive fish and Hansson, 2005). Although dierent (industrial and
kills. Drawing by M. Meerhoff and T. Christensen. 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
the growth of primary producers since these are generally consequences of past acidication (e.g. Lucassen et al.,
growth-limited by nitrogen and mostly by phosphorus in 2012; Angeler and Johnson, 2012). Moreover, acid rain is a
freshwater ecosystems. Periphyton and submerged mac- major current threat in regions with upcoming economies
rophytes, and in warm locations free-oating macro- and increasing industries.
phytes, often also show an increase in biomass. However,
phytoplankton, especially cyanobacteria and green algae,
typically take over as the dominant primary producers by Invasive species
reducing the amount of light penetrating down to periph-
yton and submerged macrophytes. The high production The intentional or accidental introduction of species to
results in an increasing amount of dead organisms accu- lakes where they do not naturally occur (often called
mulating as sediment. Bacteria that break down this exotic or invasive species indistinctively, although con-
organic material need large amounts of oxygen, leading to cepts dier slightly) can have strong eects on lake com-
reduced oxygen concentrations and sometimes anoxia. munities. Eects of the many exotic species on native
This is especially pronounced during stratication, when communities are copious and may work through dierent
oxygen-rich surface water does not mix with the water in pathways such as habitat modication, predation, her-
the deeper layer. After an initial increase in sh biomass, bivory, competition, hybridisation and the transmission of
average body size and density at the onset of eutrophica- pathogens. Often a combination of dierent pathways
tion, the composition of the sh assemblage changes and plays a role (e.g. Ellis et al., 2011). When water plant-eating
the amount of small-bodied cyprinid shes generally cray-sh are introduced, for instance, their herbivory also
increases considerably during the eutrophication process aects the habitat of many littoral species (Nystrom and
(Figure 5). Additionally, in hypereutrophic systems massive Strand, 1996). Although there are many dierent types of
sh kills are frequent due to the oxygen decit. Since organisms that have been introduced to lakes by humans
cyprinids are ecient predators on zooplankton and (e.g. plants, mussels, crabs and shrimps), sh are the most
macroinvertebrates, the abundance and size of these preys notorious group. Worldwide, exotic shes are introduced
decreases. Hence, the eutrophication process leads to in lakes to subsequently be shed upon. This can have
considerable changes in the structure of the lake ecosystem, severe ecological (and often economic and social) con-
which in turn aect the previously associated ecosystem sequences. A well-known example is the introduction of
services (e.g. recreation, shing and use as a source of Nile perch in Lake Victoria, where it caused a severe
drinking water). See also: Eutrophication of Lakes and reduction in the native cichlid biodiversity (Goldschmidt
Rivers et al., 1993). The eects of invasive species are often highly
unpredictable, and as removal of invasive species from the
Acidification lake is hard and often even impossible, prevention of the
introduction of invasive species and decreasing the inva-
Acid rain, mainly as a consequence of industrial activities, sibility of the system by removing other external pressures
has strong eects on lake communities by reducing the pH are important management measures.

8 eLS & 2014, John Wiley & Sons, Ltd.

Lake Communities

Warming Changes in precipitation

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

Due igher

Com yan
m g id
in an res

er Ch nd

of c
to g temp

pet obac
iza a

ntr nd


ad n
ati ctio

th o ratures

adv ria


red du
on rat rea


More and sed g

s p pro

gre ro

ate wth




r su


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

m ts

higher algal biomass co




g petit

nutr ion
Less ient for

s an

Less d lig
ht Less
large zooplankton submerged

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.
Some lakes may be formed or increase their volume due to
ice melting, whereas others may become temporary or even
disappear. The structure of the local trophic webs seems References
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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. 11