Professional Documents
Culture Documents
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.
lakes?
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
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 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.
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
Ice
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
Warm
and nitrogen. As a consequence of the low productivity of
monomictic
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).
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.
Riparian
Pelagic Littoral
Piscivorous fish
Macrophytes Planktivorous
fish
Macroinvertebrates
Zooplankton Periphyton
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.
Piscivorous fish
Piscivorous fish
Other fish
Shrimp (omnivorous)
Other fish
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). 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),
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
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-
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
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).
Chambers PA and Kal J (1985) Depth distribution and biomass Lucassen EC, Smolders AJ and Roelofs JG (2012) Liming induces
of submersed aquatic macrophyte communities in relation to changes in the macrophyte vegetation of Norwegian softwater
Secchi depth. Canadian Journal of Fisheries and Aquatic Sci- lakes by mitigating carbon limitation: results from a eld
ences 42: 701709. experiment. Applied Vegetation Science 15: 166174.
Cousyn C, De Meester L, Colbourne JK et al. (2001) Rapid, local Meerho M, Clemente JM, Teixeira- de Mello F et al. (2007a) Can
adaptation of zooplankton behavior to changes in predation warm climate-related structure of littoral predator assemblies
pressure in the absence of neutral genetic changes. Proceedings weaken the clear water state in shallow lakes? Global Change
of the National Academy of Sciences of the United States of Biology 13: 18881897.
America 98: 62566260. Meerho M, Fosalba C, Bruzzone C et al. (2006) An experimental
De Meester L, Gomez A, Okamura B et al. (2002) The Mono- study of habitat choice by Daphnia: plants signal danger more
polization Hypothesis and the dispersalgene ow paradox in than refuge in subtropical lakes. Freshwater Biology 51: 1320
aquatic organisms. Acta Oecologica 23: 121135. 1330.
Ellis BK, Stanford JA, Goodman D et al. (2011) Long-term eects Meerho M, Iglesias C, Teixeira-de Mello F et al. (2007b) Eects
of a trophic cascade in a large lake ecosystem. Proceedings of the of habitat complexity on community structure and predator
National Academy of Sciences of the United States of America avoidance behaviour of littoral zooplankton in temperate ver-
108: 10701075. sus subtropical shallow lakes. Freshwater Biology 52: 1009
Fryer G (1991) Comparative aspects of adaptive radiation and 1021.
speciation in Lake Baikal and the great rift lakes of Africa. Meerho M, Teixeira-de Mello F, Kruk C et al. (2012) Environ-
Hydrobiologia 211: 137146. mental warming in shallow lakes: a review of potential changes
Goldschmidt T, Witte F and Wanink J (1993) Cascading eects of in community structure as evidenced from space-for-time sub-
the introduced Nile perch on the detritivorous/phytoplankti- stitution approaches. Advances in Ecological Research 46: 259
vorous species in the sublittoral areas of Lake Victoria. Con- 349.
servation Biology 7: 686700. Moss B, Kosten S, Meerho M et al. (2011) Allied attack: climate
Gonzalez-Bergonzoni I, Meerho M, Davidson TA et al. (2012) change and nutrient pollution. Inland Waters 18: 101105.
Meta-analysis shows a consistent and strong latitudinal pattern Moss B, Stanseld J and Irvine K (1990) Problems in the
in sh omnivory across ecosystems. Ecosystems 15: 492503. restoration of a hypertrophic lake by diversion of a nutrient-
Hansen K (1962) The dystrophic lake type. Hydrobiologia 19: rich inow. Verhandlungen der Internationale Vereinigung fur
183190. Theoretische und Angewandte Limnologie 24: 568572.
Hansson L-A (1992) The role of food chain composition and Naumann E (1929) The scope and chief problems of regional
nutrient availability in shaping algal biomass development. limnology. Internationale Revue der gesamten Hydrobiologie
Ecology 73: 241247. und Hydrographie 22: 423444.
Horppila J and Kairesalo T (1992) Impacts of bleak (Alburnus Nystrom PER and Strand J (1996) Grazing by a native and an
alburnus) and roach (Rutilus rutilus) on water quality, sedi- exotic craysh on aquatic macrophytes. Freshwater Biology 36:
mentation and internal nutrient loading. Hydrobiologia 243: 673682.
323331. Pace ML, Cole JJ, Carpenter SR et al. (2004) Whole-lake carbon-
Jeppesen E, Lauridsen TL, Kairesalo T et al. (1998) Impact of 13 additions reveal terrestrial support of aquatic food webs.
submerged macrophytes on sh-zooplankton interactions in Nature 427: 240243.
lakes. In: Jeppesen E, Sndergaard M, Sndergaard M and Pauwels K, De Meester L, Michels H et al. (2014) An evolutionary
Christoersen K (eds) The Structuring Role of Submerged Mac- perspective on the resistance of Daphnia to the epizoic rotifer
rophytes in Lakes, pp. 91114. New York, NY: Springer-Verlag. Brachionus rubens. Freshwater Biology 59: 12471256.
Jeppesen E, Sndergaard M, Lauridsen TL et al. (2012) Bioma- Phillips GL, Eminson D and Moss B (1978) A mechanism to
nipulation as a restoration tool to combat eutrophication: account for macrophyte decline in progressively eutrophicated
recent advances and future challenges. Advances in Ecological fresh waters. Aquatic Botany 4: 103126.
Research 47: 411488. Rahel FJ and Olden JD (2008) Assessing the eects of climate
Jones JI and Sayer CD (2003) Does the sh-invertebrate-periph- change on aquatic invasive species. Conservation Biology 22:
yton cascade precipitate plant loss in shallow lakes? Ecology 84: 521533.
21552167. Reynolds C (2006) The Ecology of Phytoplankton. Cambridge:
Kosten S, Huszar VLM, Becares E et al. (2012) Warmer climate Cambridge University Press.
boosts cyanobacterial dominance in lakes. Global Change Romare P and Hansson L-A (2003) A behavioral cascade: top-
Biology 18: 118126. predator induced behavioral shifts in planktivorous sh and
Kosten S, Huszar VLM, Mazzeo N et al. (2009a) Lake and zooplankton. Limnology and Oceanography 48: 19561964.
watershed characteristics rather than climate inuence nutrient Scheer M, Hosper SH, Meijer ML et al. (1993) Alternative
limitation in shallow lakes. Ecological Applications 19: 1791 equilibria in shallow lakes. Trends in Ecology and Evolution 8:
1804. 275279.
Kosten S, Lacerot G, Jeppesen E et al. (2009b) Eects of sub- Solomon CT, Carpenter SR, Clayton MK et al. (2011) Terrestrial,
merged vegetation on water clarity across climates. Ecosystems benthic, and pelagic resource use in lakes: results from a three-
12: 11171129. isotope Bayesian mixing model. Ecology 92: 11151125.
Liboriussen L and Jeppesen E (2003) Temporal dynamics in epi- Sndergaard M, Jensen J and Jeppesen E (2003) Role of sediment
pelic, pelagic and epiphytic algal production in a clear and a and internal loading of phosphorus in shallow lakes. Hydro-
turbid shallow lake. Freshwater Biology 48: 418431. biologia 506509: 135145.
Teixeira-de Mello F, Meerho M, Pekcan-Hekim Z et al. (2009) Carpenter SR and Kitchell JF (1993) The Tropic Cascade in Lakes.
Substantial dierences in littoral sh community structure and Cambridge: Cambridge University Press.
dynamics in subtropical and temperate shallow lakes. Fresh- Cooke GD, Welch EB, Peterson S and Nichols SA (2005)
water Biology 54: 12021215. Restoration and Management of Lakes and Reservoirs. Boca
Thienemann A (1921) Seetypen. Naturwissenschaften 9: 343346. Raton, FL: CRC Press.
Timms R and Moss B (1984) Prevention of growth of potentially Lampert W and Sommer U (1997) Limnoecology: The Ecology of
dense phytoplankton populations by zooplankton grazing, in Lakes and Streams. New York, NY: Oxford University Press.
the presence of zooplanktivorous sh, in a shallow wetland Likens GE (2010) Lake Ecosystem Ecology: A Global Perspective.
ecosystem. Limnology and Oceanography 29: 472486. Waltham, MA: Academic Press.
Vadeboncoeur Y, Vander Zanden MJ and Lodge DM (2002) Moss BR (2009) Ecology of Fresh Waters: Man and Medium, Past
Putting the lake back together: reintegrating Benthic pathways to Future. West Sussex, England: John Wiley & Sons.
into Lake Food Web Models Lake ecologists tend to focus their Moss B, Madgwick J and Phillips G (1996) A Guide to the
research on pelagic energy pathways, but, from algae to sh, Restoration of Nutrient-Enriched Shallow Lakes (Wetlands
benthic organisms form an integral part of lake food webs. International Publication), 180 pp. Norwich, UK: Broads
BioScience 52: 4454. Authority.
Wagner C and Adrian R (2011) Consequences of changes in Scheer M (1998) Ecology of Freshwater Lakes. London: Chap-
thermal regime for plankton diversity and trait composition in a man and Hall.
polymictic lake: a matter of temporal scale. Freshwater Biology Wetzel RG (2001) Limnology: Lake and River Ecosystems. New
56: 19491961. York, NY: Academic Press.
Further Reading
Bronmark C and Hansson L-A (2006) The Biology of Ponds and
Lakes, 2nd edn. Oxford: Oxford University Press.