Et Al 2009 Climate Change and The Future Odiversity in Europe

Climate Change and the Future of Freshwater Biodiversity in Europe: A
Primer for Policy-Makers

Author(s): Brian Moss, Daniel Hering, Andy J. Green, Ahmed Aidoud, Eloy Becares, Meryem
Beklioglu, Helen Bennion, Dani Boix, Sandra Brucet, Laurence Carvalho, Bernard Clement,
Tom Davidson, Steven Declerck, Michael Dobson, Ellen van Donk, Bernard Dudley, Heidrun
Feuchtmayr, Nikolai Friberg, Gael Grenouillet, Helmut Hillebrand, Anders Hobaek, Kenneth
Irvine, Erik Jeppesen, Richard Johnson, Iwan Jones, Martin Kernan, Torben L. Lauridsen,
Marina Manca, Mariana Meerhoff, Jon Olafsson, Steve Ormerod, Eva Papastergiadou, W. Ellis
Penning, Robert Ptacnik, Xavier Quintana, Leonard Sandin, Miltiadis Seferlis, Gavin Simpson,
Cristina Triga, Piet Verdonschot, Antonie M. Verschoor and Gesa A. Weyhenmeyer.
Source: Freshwater Reviews, 2(2):103-130. 2009.
Published By: Freshwater Biological Association
URL: http://www.bioone.org/doi/full/10.1608/FRJ-2.2.1
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103
Article
Climate change and the future of freshwater biodiversity

in Europe: a primer for policy-makers
*Brian Moss1,31, Daniel Hering2, Andy J. Green3, Ahmed Aidoud4, Eloy Becares5, Meryem Beklioglu6,
Helen Bennion7, Dani Boix8, Sandra Brucet9, Laurence Carvalho10, Bernard Clement4, Tom Davidson7,
Steven Declerck11, Michael Dobson12, Ellen van Donk13, Bernard Dudley10, Heidrun Feuchtmayr1,14,
Nikolai Friberg9,15, Gael Grenouillet16, Helmut Hillebrand17, Anders Hobaek18, Kenneth Irvine19, Erik
Jeppesen9, Richard Johnson20, Iwan Jones21, Martin Kernan7, Torben L. Lauridsen9, Marina Manca22,
ȱ ě9,23, Jon Olafsson24, Steve Ormerod25, Eva Papastergiadou26, W. Ellis Penning27,
Robert Ptacnik28,32, Xavier Quintana8, Leonard Sandin20, Miltiadis Seferlis29, Gavin Simpson7, Cristina
Trigal5, Piet Verdonschot30, Antonie M. Verschoor13 and Gesa A. Weyhenmeyer20.
1
School of Biological Sciences, University of Liverpool, Liverpool L69 3GS, UK; 2 University of Duisberg-Essen, Dept of Applied
Zoology/Hydrobiology, D-45141, Essen, Germany; 3 Dept of Wetland Ecology, Estación Biológica de Doñana-CSIC, Americo Vespucio,
41092 Sevilla, Spain; 4 UMR CNRS, EcoBio, University of Rennes 1, 35042, Rennes, France; 5 Area de Ecologia, Facultad de Biologia,
Universidad de Leon, Spain; 6 Middle East Technical University, Biological Department, TR06531, Ankara, Turkey; 7 Environmental
Change Research Centre, Dept of Geography, University College, London, UK; 8 Institute of Aquatic Ecology, Campus Montilivi,
Faculty of Sciences, 17071, University of Girona, Girona, Spain; 9 Dept of Freshwater Ecology, National Environment Research Institute,
Aarhus University, Silkeborg, Denmark; 10 NERC Centre for Ecology and Hydrology, Edinburgh, UK; 11 Laboratory of Aquatic Ecology
12
and Evolution, Katholieke Universiteit Leuven, Leuven 3000, Belgium; Freshwater Biological Association, Ferry Landing, Far
Sawrey, Ambleside, Cumbria, UK; 13 Dept of Aquatic Foodwebs, Centre for Limnology (NIOO-KNAW), 3631, AC Nieuwersluis, The
Netherlands; 14 Centre for Ecology and Hydrology, NERC, Lancaster, UK; 15 Catchment Management Group, Macaulay Land Research
16
Institute, Craigiebuckler, Aberdeen, UK; Laboratoire Evolution et Diversité Biologique, UMR 5245, ENSAT, Castanet-Tolosan,
17
France; Institute for Chemistry and Biology of the Marine Environment, Schleusenstrasse 1, 26382, Wilhelmshaven, Germany;
18
ȱȱȱȱǰȱȱĜȱǰȱȱ¡ȱŘŖŘŜǰȱǰȱśŞŗŝǰȱǰȱ ¢ȱȱȱȱ
¢ǰȱ¢ȱȱǰȱǯǯȱ¡ȱŝŞŖřǰȱśŖŘŖȱǰȱ ¢ǲȱ19 School of Natural Sciences, Zoology, Trinity College, Dublin
2, Ireland; 20 Dept of Aquatic Sciences and Assessment, Swedish University of Agricultural Sciences, Uppsala, Sweden; 21 Centre for
Ecology and Hydrology, NERC, Dorset, UK; 22ȱȱȱȱȱȱȱǰȱȱ£ǰȱ¢ǲȱ 23 Facultad de
Ciencias Universidad de la Republica, Montevideo, Uruguay; 24 Institute of Freshwater Fisheries, Vagnhofdi 7, 105 Reykjavik, Iceland;
25
ȱȱȱ ǰȱěȱȱȱǰȱěȱ¢ǰȱěǰȱ ǲȱ26ȱȱȱ¢ǰȱ¢ȱȱǰȱ
ŘŜśŖŖǰȱǰȱ ǲȱ27ȱǰȱĞǰȱȱǲȱ28 Norwegian Institute for Water Research, Gaustadalléen 21, N-0349,
29 30
Oslo, Norway; Institute of Botany, Aristotle University of Thessalonika, Thessalonika, Greece; Alterra, Dept of Ecology and
Environment, NL-6700, AA Wageningen, The Netherlands; 31 Institute for Sustainable Water, Integrated Management and Ecosystem
Research (SWIMMER), Nicholson Building, University of Liverpool, Liverpool, UK; 32 ǰȱǯȱȱǰȱȱŗǰȱ
ȬŘŜřŞŘȱǰȱ ¢ǯ
ȘȱȱȱȱǻȓǯǯǼǯ
Received 30 January 2009; accepted 26 May 2009; published 14 September 2009
DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009
104 Moss, B. et al
Abstract
Earth’s climate is changing, and by the end of the 21st century in Europe, average temperatures
ȱ¢ȱȱȱȱ¢ȱȱȱŘȱǚǰȱȱȱ¢ȱŚȱǚǰȱ ȱȱěȱȱĴȱȱ
ȱȱȱ¢ȱȱ¡ȱ ȱǯȱȱĴȱȱ¢Ȭȱȱȱ
ȱ ȱȱȱȱǰȱ ȱȱȱȱěǰȱ ȱȱȱȱȱȱ
on which societies depend and how to adapt human societies to the changes. Natural systems are still
seen, through a long tradition of conservation management that is largely species-based, as amenable
to adaptive management, and biodiversity, mostly perceived as the richness of plant and vertebrate
ǰȱ Ğȱ ȱ ȱ ȱ ȱ ǯȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ
changes will be possible only in a minority of cases but that prediction of trends in general structure
ȱȱȱȱȱ ȱ¢ȱǻȱǰȱȱĚȱǰȱ
shallow lakes and deep lakes) in three broad zones of Europe (Mediterranean, Central and Arctic-
Boreal) is practicable. Maintenance and rehabilitation of ecological structures and operations will
inevitably and incidentally embrace restoration of appropriate levels of species biodiversity. Using
expert judgement, based on an extensive literature, we have outlined, primarily for lay policy makers,
the pristine features of these systems, their states under current human impacts, how these states are
likely to alter with a warming of 2 °C to 4 °C and what might be done to mitigate this. We have avoided
technical terms in the interests of communication, and although we have included full referencing as
in academic papers, we have eliminated degrees of detail that could confuse broad policy-making.
¢ ǱȱǲȱǲȱĚǲȱǲȱǲȱ¢¢ǲȱ¢ǲȱȱǯ
Introduction 2007) whilst exotic species are appearing where they did
not occur before (Lodge, 1993; Dukes & Mooney, 1999)
If the biota, in the course of aeons, has built something we like and relationships between them are changing, with
but do not understand, then who but a fool would discard consequences throughout the ecosystem (Winder &
seemingly useless parts? To keep every cog and wheel is the Schindler, 2004).
ęȱȱȱȱǯȱ The stage of the environmental theatre (Hutchinson,
Aldo Leopold (1938) 1965) is undergoing a substantial change of scene. The
plot of the evolutionary play, a perpetual action of
The climate of Earth is changing rapidly (IPCC, 2007) with natural selection, competition, predation, immigration
human activities mostly responsible for this and many and emigration to cope with environmental change, will
other global changes (Vitousek et al., 1997; Sala et al., stay the same, but the future identity of its players, the
2000). Glaciers are melting, ice covers on lakes form later biodiversity, is very uncertain. Biodiversity has become a
ȱ ȱ ǰȱ ȱ Ĵȱ ȱ ȱ ȱ ȱ widely used word in political agenda. Ecologists used the
Ě ȱȱǯȱȱ
ȱĚǰȱȱ ȱȱȱ simpler term ‘diversity’ until the concept was politicised
are becoming more frequent. Rouse et al. (1997) and ȱ ȱ ȱ ȱ ǯȱ ȱ ȱ ȱ Ğȱ ǰȱ
Schindler (2001) have reviewed the changes in physico- outside professional ecology, as the sole key measure of
chemical characteristics of fresh waters, but there are also the state of nature. Particular species are seen as icons,
ȱ ȱ ȱ ȱ ¢ȱ ǻħȱ ȱ and degradations or improvements are perceived as
al., 2005; Jeppesen et al., 2009) with likely feedbacks to the their exit from, or entry onto, the stage. The perception of
physical processes. Many species are declining (Pimm professional ecologists, however, is more profound. There
et al., 1995; Brown et al., 2007; Durance & Ormerod, are deep questions about what biodiversity means, what its
© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

Climate change and biodiversity in Europe 105
ęȱȱȱȱȱȱȱǰȱȱ ȱ ǰȱǰȱȱȱȱȱȱȱęǰȱ
to measure it. Ultimately, biodiversity is a component of the but not necessarily most important. An ecologist will point
functioning of ecosystems, but it is the arrangement of the out that there is also a plethora of bacterial and microbial
ȱȱĴȱȱȱȱȱȱȱǯ diversity (Torsvik et al., 2002; Weisse, 2006), which is at
Whilst climate modellers predict the physical states of least as crucial, and probably more so, to the functioning of
the water cycle from place to place as temperatures increase, the planet’s chemical cycles; that a species comprises much
albeit with many uncertainties of detail, ecologists are being additional necessary genetic diversity within its apparent
asked to predict what will happen to biodiversity in the uniformity; that there is an important element of relative
face of these changes. But there are few simple and general ȱ ȱ ȱ ěȱ ȱ ȱ ȱ ȱ
models for biodiversity (Balian et al., 2008) comparable communities; and that there are concepts of local diversity
with those for climate and atmospheric chemistry. Living (for example, in a particular river), habitat diversity (for
systems embody the complexities of physical systems example, among rivers) and regional diversity (collective
whilst adding further, much more variable, layers of over large areas or distinctive tracts of terrain). These are
their own. In this paper we ask how whole freshwater respectively also called alpha, beta and gamma diversity
ecosystems might change, in Europe, in response to changes ǻĴǰȱŗşŝŘǼǯȱ
in the greenhouse gas composition of the atmosphere. Our ȱ ȱ ę¢ȱ ȱ ȱ ȱ ȱ ȱ ȱ
approach has been to examine the issues as a professional of predicting future regional biodiversity than local
group, using expert judgment based on existing experience biodiversity. The undisturbed communities of individual
and information. The discussions were conducted in a lakes or rivers certainly have some order in their local
three-day workshop in Bristol, UK in July 2007, under the biodiversity, but they have a great deal of unexplained
auspices of the EU EUROLIMPACS programme. Our variation too. The order comes from the limited number
target readership is not our fellow scientists, although ȱ ȱ ȱ ȱ Ěȱ ¢Ǳȱ ȱ
we have referenced the paper thoroughly in conformity ability, competition, biotic interactions such as grazing or
with our conventions; it is the non-scientists involved predation, and physical disturbance. The unpredictability
in government and policy-making. Increasingly they comes from random population oscillations that may make
come from an education and experience that has had less a species locally extinct purely by chance, and from the
immersion in natural systems (Louv, 2008) than is desirable ¢ȱ ȱ ȱ ¢ǰȱ ȱ Ğȱ ¢ǰȱ
when the world’s major problems are those of its natural Ěȱ ȱ ȱ ȱ ȱ ȱ ȱ ¢ȱ ¢ǯȱ
environment. To such problems, the more commonly If we take a series of communities in a similar
emphasised economic and social problems are subsidiary category, the invertebrates of small upland streams,
and derivative. An equable state of the biosphere is ȱ ȱ ȱ Ĵȱ ȱ ȱ ǰȱ ȱ ¡ǰȱ
the sine qua non, without exception, of everything else. and use sophisticated statistical methods that correlate
environmental conditions with the species present,
The meaning of biodiversity relationships will emerge, but typically lower than 20 % of
ȱěȱ ȱȱȱǰȱȱ ȱȱȱ
For a policy maker, a sophisticated understanding of number of environmental conditions are measured. Any
biodiversity is desirable and thus we discuss some of the single factor, nutrients for instance, will account for only a
important issues about the meaning and measurement of few per cent of the variation, and temperature even less.
¢ȱȱȱȱ¢ȱěȱȱȱ ȱ ¡ȱ ȱ ¢ȱ ȱ Ĵȱ ȱ
change on it. To the lay person, biodiversity is an easy unmeasured physico-chemical features and to biological
concept. It is the number of species of animals, usually interactions, which are generally not incorporated into the
meaning charismatic vertebrates, and higher plants. To be analysis largely because they are too numerous and too

106 Moss, B. et al
subtle for ready measurement. A typical natural water, of these small patches will change greatly as climate
for example, will have thousands, possibly millions, changes. But because of the numbers of species, the mutual
ȱ ěȱ ȱ ȱ ȱ ǰȱ ¢ȱ ȱ ȱ Ěȱ¢ȱȱȱȱȱǰȱ
organic compounds, whilst routine analyses cover but and the inevitable patchiness of their distributions,
a handful of mainly inorganic substances. Biological future local changes are impossible to predict accurately.
ȱ ȱ ȱ ȱ ¢ȱ ęǯȱ ȱ ¡ȱ ȱ ěȱ ȱ Ĵȱ ǯȱ ȱ ȱ ȱ ȱ ȱ
variation may also come from sampling error. It is millions of square kilometres, the size of several European
rarely possible to sample a community very thoroughly countries combined. Future climate models already make
because the intricacies of individual species distributions reasonable predictions for this sort of area, and past climatic
make this practicably and economically impossible. An indicators, for example pollen analysis of sediments and
ȱ ȱ ȱ ȱ ȱ ȱ Ğȱ ȱ ȱ ȱ ǰȱ ȱ ȱ ȱ Ĵȱ ȱ ȱ ¢ȱ
¢ȱȱ¢ȱȱȱěȱȱȱȱȱ (Drescher-Schneider et al., 2007; Miroslaw-Grabowska
large regions may be obscured by this great complexity. & Niska, 2007). A region is big enough to smooth out a
The richness of undisturbed biological communities great deal of local variation. We decided it was practicable
will come as a surprise to many policy makers, especially to consider Europe in terms of the Mediterranean
if they are more familiar with agriculture and forestry. region, the central land mass, and the Boreal-Arctic zone.
In a small European mountain brook in central Europe, The next issue was which general habitats to consider
the Breitenbach stream, which has been continuously in such regions. The Water Framework Directive (WFD)
investigated for more than 40 years (Illies, 1975, 1979; (European Council, 2000) requires a typology, or pigeon-
Zwick, 1992; Obach et al., 2001), more than 1000 invertebrate holing, of habitats to be drawn up within categories of lakes,
animal species have been recorded, but still many more rivers, estuaries (transitional areas) and coastal waters, and
might occur. A typical community might overall have then the ecological quality of each type determined on a
some tens to hundreds of thousands of species, ranging scale whose anchor is a reference state that is essentially
from microorganisms to vertebrates. Where even just pristine (‘ȱ ȱ ȱ Ěȱ ȱ ȱ ¢’). The
potential paired biological interactions among these are ȱ¢ȱȱȱęȱȱę¡ȱȱȱ
considered, the possible number is enormous. There is also ǰȱȱȱȱ¢ȱȱ¢ȱĚȱȱ
a natural turnover in the list of species, brought about by nature, but which are independent of ecological quality.
immigration and local extinction. The species list is never The WFD suggests a few such parameters, but even a
ǰȱȱȱȱȱȱĚȱ¢ȱ¢ȱǯȱȱ few produces hundreds to thousands of combinations.
ȱȱ¢ȱȱȱȱȱȱȱǯȱȱȱĚ¢ǰȱ Three sizes of catchment area, three general sorts of
but the majority not very far. Crawling invertebrates, such geology, three altitude bands and 24 biogeographic
ȱĚ ǰȱȱȱȱȱȱȱęȱȱȱ regions in Europe produce 648 sorts of rivers, and for
range following a glaciation that ended 10 000 years ago lakes, where area and depth are also potentially involved,
(Reynoldson, 1983). A gap of only a few hundred metres another order of magnitude ensues. Such a system may
ȱěȱǰȱȱȱȱȱęǰȱ¢ȱȱȱȱ ȱ ȱ ęȱ ȱ ȱ ȱ ǯȱ ȱ ȱ ȱ
of riparian forest bats or birds, despite their aerial dexterity. ȱ ěȱ ȱ ȱ ǰȱ ȱ ȱ ¢ȱ ǯȱ
Naturalists delight in the nuances of local biodiversity, We decided that four general categories were sensible:
knowing that a particular species may be found here but erosive rivers, depositional floodplain rivers, their associated
not there, and recognising the subtleties of patches of shallow lakes and wetlands, and deep lakes. We recognise,
environment that are associated with particular animals or however, that discrete types of any system do not, in reality,
plants. Most of these relationships are sensed but not fully ǯȱ ȱ ȱ ȱ ęȱ ǰȱ ȱ ǰȱ ȱ
¡ǰȱȱȱȱĞȱ¢ǯȱȱȱĴȱ confound any typology. Considering only very broad

categories, as we have, is a realistic option, for it enfolds the Moreover, there is much uncertainty about what
continua. It also allows inclusion of small bodies of water, constitutes a species, especially in microscopic organisms.
such as ponds, that can be excluded from the Directive, ȱ ȱ ȱ ȱ ȱ ęȱ ȱ ȱ ȱ
despite their overwhelming dominance of the world’s what appear to be generally similar cells. Molecular
freshwater area (Oertli et al., 2002; Downing et al., 2006). studies increasingly underline the long-realised
Large bodies of water are not more important than small, understanding that almost all individuals within a
where either collective area or species richness is concerned. ȱ ȱ ěǰȱ ȱ ȱ ¢ȱ ȱ ȱ
existence of several species where only one was previously
Measurement of biodiversity recognised (Petrusek et al., 2008). They have demonstrated
an enormous degree of variation within apparently the
The issue then arose of how to measure biodiversity. same species in adjacent localities (Weider & Hobaek,
Typically, biodiversity studies concentrate on one group of 2003). Such understanding means that biodiversity will
organisms: in fresh waters, usually plants, phytoplankton, probably always be underestimated where such detailed
£ǰȱ ǰȱ ęȱ ȱ ǯȱ ȱ ¢ȱ work is not possible, during routine monitoring surveys,
rarely consider the less well-known groups, among them for example. Furthermore, many of the apparent species
the bacteria, actinomycetes, microfungi, nematodes, of small but very abundant invertebrates, particularly
harpacticoids, ciliates, benthic algae, and the parasites nematodes and insects, may not have been described or
that are probably much more diverse than the free-living may be distinguishable only by increasingly fewer experts.
organisms (Torsvik et al., 2002; Balian et al., 2008). It is Counting species is thus not as simple as it sounds.
Ğȱȱȱ¢ȱȱȱȱȱȱȱ One possible resolution of these issues, therefore, is not
groups, that a habitat rich in birds will be rich in ciliates to consider species or even genera or families of organisms
and vice-versa. Where several groups have been studied at all but to think in terms of functional diversity (Wilson,
together, sometimes this is the case (for example, in small ŗşşşǲȱǰȱŘŖŖŗǲȱ¢ȱǭȱ ǰȱŘŖŖŘǰȱŘŖŖŜǲȱěǰȱ
farmland ponds, Declerck et al., unpublished manuscript), 2003; Hooper et al., 2005). Functional diversity is a measure
ȱĞȱȱȱȱǻ ȱȱǯǰȱŘŖŖŘǲȱȱȱǯǰȱŘŖŖśǼȱ of the traits that organisms have, or what they do in the
and where it is, the reasons for a particular diversity may ecosystem. Several organisms may share the same group
ȱȱȱȱȱěȱȱǻ
ȱǭȱǰȱ of traits and perhaps functional diversity overcomes the
2003; Hawkins et al., 2003; Rodrigues & Brooks, 2007). need to recognise species and implies that some species are
A given habitat will also show seasonal changes and may dispensable if others with similar traits persist. Alas, where
also vary naturally from year to year, sometimes very ȱȱȱȱȱȱȱĴȱ
strikingly in the Mediterranean region (Bonis et al., 1995; to demonstrate this, it would appear that the ultimate
Gafny & Gasith, 1999). Shallow lakes, for example, may functional units are still the species (Petchey et al., 2007).
switch from a diverse plant-dominated community to a Groups of traits are unique to species; they are only partly
less diverse algal dominated system, with less structure, as shared. It would seem that there is meaning to the degree
a result of human pressures, but they may also do this from of diversity that the very powerful process of natural
¢ȱȱ¢ȱȱȱȱȱȱĚȱȱ ȱ selection has produced. A system whose parts are tested to
(Uhlmann, 1980) or water level (van Geest et al., 2005; ȱȱ¢ȱȱȱȱĚǯȱȱ
Beklioglu et al., 2006; Valdovinos et al., 2007) and many Every cog and wheel has meaning. Yet we can only
organisms may not be common to the two states. Is the broadly estimate the list of cogs and wheels in any system.
biodiversity of such lakes that at any one time or the sum of If functional diversity measures are ultimately still a
the several years and states? measure of species diversity, perhaps operational diversity
is a possible means to assess the intactness of an ecosystem

108 Moss, B. et al
and a surrogate for species diversity, whilst also telling and mineralisers, and predators and parasites to police the
us more about the ecological quality than a species list activities of other guilds. It will require nutrients, but will be
alone, however complete, can do. Operational diversity ȱĜȱȱȱȱȱȱ¢ǰȱ¢ȱ¢ȱ
is about processes in a system. The cogs are all there, but with great parsimony rather than requiring continued
in the right relationships and numbers to keep the wheels outside subsidy. A fully operational system will be big
turning smoothly. If operational diversity is intact, as in and interconnected with other similar systems through
a pristine system, we argue that an appropriate species corridors or tracts of other natural systems to ensure the
diversity must inevitably be present to support it. Over opportunity for immigration of new species that gives
long periods, natural selection has produced species that resilience, and an area large enough to maintain populations,
ȱȱ¢ȱȱȱȱȱȱęȱȱȱ even of its largest organisms, that do not become inbred
species that comprise them. A selection process at a higher and genetically limited. There are many examples that
level among colonisers of a habitat operates similarly to ȱ ȱ ¢ȱ ęȱ ȱ ȱ ěȱ
ȱ ȱ ȱ ȱ ȱ ȱ ęȱ ȱ ȱ ¢ȱ ȱȱȱ¢ǰȱȱ¢ȱ¢ȱĚ¢ȱ
(Janzen, 1985; Kawecki & Ebert, 2004; Wilkinson, 2004). ȱȱ¢ȱȱęǰȱȱȱȱǻȱ
ȱ ȱ ȱ ȱ ¢ȱ ȱ ¢ȱ ęȱ ȱ ȱ & Nakano, 1998; Amezaga et al., 2002; Naiman et al., 2002;
environmental circumstances, maintaining itself and Knight et al., 2005; Fukui et al., 2006; Klaassen & Nolet, 2007).
ȱ ȱ Ěȱ ȱ ȱ ȱ ȱ Operational diversity would be best expressed in
resistance to change, and of resilience when change terms of processes (photosynthesis, respiration, carbon
ȱȱǯȱȱȱĴȱ¢ȱ¢ȱȱȱȱ storage, nutrient turnover, turnover rates of species,
ability to substitute previously minority species (those ȱ ȱǼȱȱ ȱȱȱĞȱȱ
waiting in the wings in the theatrical analogy) in response comprehensive and comparable data on these to know
to change (Carpenter et al., 1995). A fully operational ȱ ȱ ȱ ȱ ȱ ȱ ȱ ęȱ ǯȱȱ
system is self-maintaining. Systems that require external Paradoxically we therefore have to fall back on a surrogate
management for their continuation are damaged systems of broadly operational but taxonomic groups (Murphy et
and the greater the management needed, the greater has al., 1994), coupled with our experience of working with
been the damage. To be sure, some traditionally managed a variety of systems operating with high, if not pristine,
systems, such as open fens used for grazing or the ȱ ¢ȱ ȱ Ĝ¢ǯȱ ȱ ȱ ȱ ȱ ȱ
production of hay, maintain high diversity by preserving information on many groups, we decided in this exercise to
ȱ ¢ȱ ȱ ěȱ ȱ ȱ ȱ ȱ ǰȱ ȱ ȱ think in terms of plankton, aquatic plants, benthic macroin-
management has been no detriment locally. The sense here ǰȱ ęȱ ȱ ȱ ȱ ȱ ȱ ǯȱ
is of big, regional systems that have been over-exploited or ȱȱȱ ȱȱȱěȱȱȱ
seriously disturbed and in which species have been lost or ȱȱȱ¡ȱȱȱ¢ȱȱęȱ¢ȱ
invasive species gained such that management is needed to operational groups in three regions. Climate change is not
prevent the loss of others. Such management may be very ȱȱȬěȱ ǯȱȱȱȱȱȱǰȱ ȱȱ
expensive, whereas self-managing systems cost nothing. band of possible temperatures predicted for the future
Operational diversity is then a measure of the century, from perhaps at least a 2 °C increase to maybe
complexity of the system needed for its independent 6 °C or more, especially in the Arctic. We decided that we
ǯȱȱȱ ȱȱĚȱȱȱȱȱȱȱ ȱ ȱ ȱ ȱ ȱ ěȱ ȱ ěȱ ȱ
organisms but not necessarily, beyond a need for trees in a along this gradient but could indicate general trends.
forest, in a list of particular species. It may be manifested
by a need for functional groups (guilds) of organisms:
¢ȱǰȱ£ǰȱěȱȱȱȱ

Determinants of natural ȱ ¢ȱ ȱ ¢ǰȱ Ğȱ ȱ Ȭ¢ȱ ȱ ȱ
biodiversity: existing patterns ȱȱ¢ȱȱǻĴȱǭȱǰȱŘŖŖŖǼǯȱ
Actual evapotranspiration rates summarise the
There remained two further issues before we could come combined temperature and moisture gradients. The
to a view about how operational freshwater diversity will general trend of biodiversity with evapotranspira-
change in Europe as climate changes. The first was to decide tion in Europe will be a bell-shaped curve, with low
what determines the baseline diversity in systems not values where evapotranspiration is low because of low
ȱ¢ȱȱ ȱĚȱȱȱ¢ȱ temperatures, despite abundant water in the north,
based human society, because change is only absolutely and low because of high temperature but extreme
measurable against a fundamental reference. The second shortage of water and higher salinity in the south (Fig. 1).
ȱ ȱȱȬěȱȱȱȱȱȱ¢ȱ Then there is a well-established link between nutrient
ȱ ȱ ȱ ȱ ȱ ǯȱ ȱ ȱ ęȱ ȱ ȱ ¢ȱȱ¢ǰȱĞȱ ȱȱȱȱ
was the easier to contemplate but was still fraught with communities (Stevens et al., 2004; Crawley et al., 2005) but
Ĝ¢ǯȱ ȱ ¢ȱ ¢ȱ ȱ Ĵȱ ȱ ȱ also a feature of fresh waters (Dodson et al., 2000; James
ȱȱȱȱǯȱȱȱȱĞȱȱȱ et al., 2005). Habitats with modest nutrient supplies,
Ĵȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ above some minimal threshold below which the habitat
ȱǻ
ȱȱǯǰȱŘŖŖřǼǯȱȱȱȱȱĴȱȱ is so nutrient poor that it is extreme, are associated with
ȱěȱȱȱȱȱȱȱǰȱě¢ȱ high diversity. Conversely, nutrient-enriched conditions
having destroyed habitats so that the polar and boreal favour vigorous, competitive species that exclude smaller,
zones are in an early recolonisation phase in which species more specialist and less aggressive ones and result in
are still returning. Additionally, the longer growing seasons ȱ ¢ȱ ǻȱ ȱ ǯǰȱ ŗşşşǲȱ Ĵȱ
and higher temperatures, allowing more generations and
productivity towards the Equator, may
have promoted speciation and allowed
ěȱ ȱ ȱ ǯȱ ȱ ȱ
climatic conditions in some tropical
regions have remained relatively steady
for some million years, large parts of the
boreal zone have been ice-free for only a
few thousand years.
There is also a moisture gradient,
with endorheic, closed habitats where
water leaves only by evaporation,
having more extreme salinity and
temperature conditions than open,
exorheic systems, where the basins
ȱȱĚ ȱȱĚ ȱȱȱ
water. Most closed systems in Europe
are in the Mediterranean region but
cold closed systems occur in the dry
Austro-Hungarian plain. Closed Fig. 1. Relationship between biodiversity (in a wide sense) and evapotranspiration as a
systems have specialist species, resilient general climate indicator.

110 Moss, B. et al
et al., 2001). This, also, theoretically gives a bell-shaped ȱ ȱ ȱ ȱ ¢ǰȱ ȱ ȱ ¢ȱ ěȱ
distribution (Fig. 2) of biodiversity with nutrient availability. of high evapotranspiration may give low local diversity.
¢ǰȱ ȱ Ěȱ ¢ȱ ǻǰȱ Seeing these links is easiest in continents that are
1978). Considerable disturbance, such as glaciation, leads large and comparatively simple in outline shape so that
to extinction. A modest amount of local disturbance climatic gradients are steady and latitudinal barriers to
leads to a mosaic of sub-habitats that preserve patches movement of organisms, following a disturbance, are
ȱ ěȱ ȱ ȱ ȱ ȱ ¢ȱ absent. Examples are East Asia, the Americas and Africa
high local diversity (Bonis et al., 1995; Frisch et al., 2006a). but there are irregularities even there. Europe is a small
No disturbance may lead to the relative uniformity of and very irregular continent. It has mountain barriers like
climax communities, but there is never a complete lack the Alps, Carpathians and Pyrenees, and marine barriers
of some sort of disturbance. However, at a regional in the Baltic, North Sea and North Atlantic that cut across
scale, lack of severe disturbance leads to speciation over climate gradients; in addition, it is a continent riddled with
long periods, and increasing diversity. In a predictable, peninsulas and islands that develop their own idiosyncrasies.
Ĵȱ ȱ ǰȱ ȱ ȱ ¢ȱ ȱ Simple climatic gradients therefore do not entirely
Ĝ¢ȱȱ¢ȱȱȱȱȱȱȱ ȱ ȱȱȱ¢ȱĴȱȱǰȱ
develop mechanisms to avoid competition with each and the antecedents for species’ origin and colonisation
other. With time since a major disturbance, such as a ȱȱ¢ȱěȱȱȱȱǯȱȱȱ
glaciation, biodiversity thus increases steadily, tending of northern Europe was covered by ice sheets up to
to, but never quite reaching, a plateau where natural 10 000 years ago; almost no distinctive new species have
extinction rates are only a fraction smaller than speciation developed there as a result of recent evolutionary processes
rates (Fig. 3). In the Mediterranean region this process has (although there is much evidence of speciation in progress
(Ferguson & Taggart, 1991; Sandlund et al., 1992; Weider &
Hobaek, 2000)), and almost all species
occurring have immigrated in the last
10 000 years. Usually these have been
Ě¡ǰȱ ¢ȱ ȱ ǰȱ
capable of coping with a variety of
habitat conditions, with high dispersal
rates. During the ice ages, central
Europe was located between the main
ice masses covering northern Europe
and the Alps. Similar to northern
Europe, only very few species have
their origin in central Europe, but
immigration from refuges in the
Mediterranean zone was simpler, since
distances were relatively short and a
variety of habitats developed shortly
Ğȱȱȱǯȱȱȱȱȱ
zone many species survived the ice
Fig. 2. Relationship between biodiversity (in a wide sense) and key nutrient availability ȱ ȱ ǰȱ Ğȱ ȱ ȱ
(largely N and P). Ombrotrophic refers to supply by direct rainfall; minerotrophic refers to
¢ȱĞȱȱȱȱ ȱȱȱǯ further species and subspecies as a

is a gradient from ombrotrophic

(rain-fed nutrient) conditions to
minerotrophic (ground and soil-water
ȱ Ǽȱ ǰȱ Ěȱ ȱ
only local geology but its interaction
with climate. Poorly weathered
igneous rocks have waters dominated
by rainwater chemistry; weathering
of more soluble sedimentary rocks
increases the ionic concentrations
substantially, but can be overridden in
very wet areas. The surface vegetation
even on limestone becomes rainwater-
dominated if precipitation is high
enough, as in western Ireland. There
is a general relationship between lower
nutrient availability in ombrotrophic
conditions and higher in minerotrophic
Fig. 3. General relationship between biodiversity (in a wide sense) and time since a major
disruption or disturbance as a result of a combination of processes including invasion and conditions.
speciation. The third axis is time elapsed since
result of isolation (Cosswig, 1955; Pauls et al., 2006). Many a major disturbance, major being one
of these species have a restricted range and have not (yet) ȱ ȱ ěȱ ȱ ȱ ǰȱ ȱ ȱ ȱ
managed to colonise other parts of Europe. Most groups ice age or extensive volcanic activity. Combinations of
of plants and animals are thus characterised by a high the relationships shown in Figs 1, 2 & 3 gave the three
ȱ ȱ Ğȱ ȱ ȱ ȱ ȱ ȱ ȱ dimensional picture of Fig. 4 for Analoguesia and allowed
Mediterranean, and a low number of more generalist us to envisage how climate change might impact on this
species in northern Europe, with central Europe being picture. For example, trends can be traced as evapotran-
ǯȱȱȱȱĴȱȱ¢ȱȱȱȱ spiration changes, as habitats become more minerotrophic
of ice disturbance, rather than present climatic conditions, when evaporation rates increase and ombrotrophism
although the two phenomena are obviously linked. becomes less prominent, and as regional droughts and, in
ȱ ǰȱ ȱ Ěȱ ȱ ȱ ǯ
Analoguesia
Mitigation
Since existing climatic gradients in Europe seem to be less
ȱȱ¡ȱȱ¢ȱĴǰȱȱ The second issue of possible mitigation by human
as they are by major past events, we needed a conceptual societies, as temperatures increase in future, is much more
model to predict future changes and so invented a regular speculative. Humans have already greatly altered natural
ȱ ȱȱǯȱȱȱȱ¡ȱęȱ Ĵȱȱ¢ȱǻȱȱǯǰȱŗşşŝǲȱǰȱ
ȱǯȱȱȱęȱȱȱȱǰȱȱ ŘŖŖŝǼǯȱ ȱ ȱ ȱ ȱ ¢ȱ Ěȱ
ȱ ȱ ǰȱ ȱ ě¢ȱ ȱ ¢ȱ¢ȱȱȱȱ ȱȱȱęȱ
ȱ Ěȱ ȱ ȱ ȱ ȱ ¢ȱ ȱ could determine the local vegetation, the nature of hunting
incorporates rising salinity in endorheic areas. The second habitats and vulnerability of prey. Technological societies,

112 Moss, B. et al
generating energy such as

tidal generation could have
ȱ ěȱ ȱ ȱ
ȱ ȱ ȱ Ěȱ
sections of rivers. Expansion
in hydroelectric power
ȱ ȱ ě¢ȱ
block the runs of migratory
ęȱ ȱ ȱ ȱ ǰȱ
despite the incorporation of
ęȱȱȱǰȱ ȱ
ȱ Ğȱ ȱ ¢ȱ Ĝǯȱȱ
Reservoirs also cause release of
methane, a potent greenhouse
gas, as the former terrestrial
soils are waterlogged.
Generation of electricity by
biomass burning could turn
Fig. 4. ȱȱěȱȱǰȱȱ¢ȱȱȱȱȱ¢ȱ huge swathes of land into
(in a wide sense) in a hypothetical regular continent, Analoguesia, in which there are no latitudinal
monocultures of willow. These
barriers to movement of organisms. The greater the depth of shading, the greater the biodiversity.
might be less heavily fertilised
however, now dominate biodiversity and there can be than food crops and hence reduce the nutrient pollution of
ȱ ȱ ȱ ȱ ȱ ȱ Ěȱ ȱ ȱ ǯȱȱ fresh waters, but the reverse could be the case if herbaceous
In Europe, the concept of even a nearly pristine habitat, crops, like sugar cane or rapeseed, were used. Changes in
the gold standard of high ecological quality for the Water agriculture will be inevitable as already hot environments
Framework Directive, is that of the Holy Grail. Beyond become deserted, environmental refugees move polewards,
a local increase in the diversity of patches of land owing import of food becomes restricted and more home-
to light, traditional agricultural management, where this grown food is required. Intensive agriculture and dense
ȱ ǰȱ ȱ Ěȱ ȱ ȱ ȱ ¢ȱ Ĵȱ ȱ ȱ ȱ ȱ ¢ȱ ȱ ¢ǯȱ
ȱ ¢ȱ ¢ȱ ǯȱ ȱ ęǰȱ ǰȱ Introduced species might become an even more severe
toxic pollution, urban development, river engineering ǯȱ ȱ ȱ ȱ Ğȱ ȱ ȱ ȱ
and drainage all impoverish habitats, and deliberate or introductions but if commercially or recreationally
accidental introductions of species usually have the same important species disappear there may be a tendency
ěȱ ȱ ȱ ȱ ȱ ȱ to introduce replacements irrespective of their legality.
native species (Lodge, 1993; Lodge et al., 1998; Dukes & Introduction of one species generally brings in many
Mooney, 1999; Sala et al., 2000). ǰȱ ȱ ¢ȱ ȱ ǰȱ ȱ Ĵȱ ȱ ȱ
A great deal of uncertainty as to the impacts of intended species. Future agriculture and aquaculture,
climate change on biodiversity hinges on how human demanding higher production to compensate for loss
ȱȱȱȱȱȱĴȱȱȱȱ of farmland in lower latitudes, might also make greater
ěǯȱȱȱȱȱǰȱǰȱ ȱ ȱ ¢ȱ ęȱ ǰȱ ȱ ¢ȱ
ȱȱȱǰȱȱȱȱĞȱ ȱ similar risks to those that come from introduced species.
determinants of current biodiversity. New methods of

Therefore our, and indeed anyone’s, prediction of the The essence of a pristine system is that it manages itself and
future biodiversity can only be very general. Elaboration ¢ȱȱȱȱȱ¢ȱȱȱęǯ
of apparently objective ‘horizon scanning’ techniques,
formalised into ostensible credibility by elaborate computer Change in erosive river systems
ǰȱȱĴȱȱȱȱȱȱȱ¢ȱ
££¢ǯȱȱęȱȱȱȱȱ¢ȱ A river forms a continuum, along which many things
on a very broad scale is as honest an approach as is possible. systematically change: discharge, current, sediment
load, temperature and food sources as well as species
Scenarios composition. ‘Erosive river systems’ include small
and medium-sized rivers, largely located in hilly and
Bearing all these things in mind, we have condensed ȱǰȱ ȱȱȱěȱȱ ȱĚ ȱȱ
ȱ ȱ ȱ ǯȱ ȱ ȱ ȱ ęȱ Ě¢ȱ to abrade material from the bed and move it downstream.
reconstructed a scenario for pristine (high quality) There is a smooth transition to depositional rivers, with
conditions for each of our four habitat types. Our the transition zone being particularly rich in habitats and
impression of much current policy-making is that there species.
ȱ¢ȱĴȱȱȱ ȱȱȱ ȱǯȱȱ Pristine erosive river systems have wild waters
With every successive generation of policy-makers, their cascading in a single channel through narrow valleys,
experience, based on growing up in a world being damaged especially in spring when snow melts from the uplands.
at unprecedented rates, becomes more and more separated Though tundra-bounded at their highest altitudes and
from ecological reality. Secondly, we have indicated how latitudes, they become dominated by forested riparian
this pristine scenario has changed with current human (bankside) zones southwards and towards sea level.
impacts. We have then projected onto this a temperature The forest and the stream are one system, for the main
change of about 2 °C to 4 °C, with associated hydrological energy source for the aquatic community is the terrestrial
changes as predicted by the IPCC (2007). It seems unlikely Ĵȱ ȱ ȱ ȱ ȱ ȱ ǰȱ ȱ ȱ ȱ ¢ȱ
now that we will avoid a 2 °C rise and with current political rocks and woody forest debris that also naturally falls
and social resistance to making severe enough changes in in. Invertebrate groups, the leaf shredders and wood
ways of life everywhere, a much greater increase is likely ǰȱȱęȬȱȱȱȱǰȱȱ
in the foreseeable future. Our use of three broad zones of ȱ Ĵǰȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ
Europe – the boreal and polar, central, and Mediterranean ȱ ȱ ȱ Ĵȱ ȱ ȱ ǯȱ ȱ ȱ ȱ ȱ
Ȯȱȱǰȱȱ ȱȱȱǰȱȱ£ǰȱęȱ widens, rocks in its unshaded centre become colonised
biological components and three scenarios, we have still by algae and mosses that are fed upon by invertebrates
ȱȱȱȱȱȱŗŞŖȱěȱȱȱȱȱȱ scraping the surfaces. In turn the invertebrate processors
ȱȱȱȱĜ¢ȱȱĴȱȱ ȱ ȱ ¢ȱ ǰȱ ęȱ ȱ ȱ ǯȱ
in any greater detail. Lastly, we have indicated what might Nutrients are scarce, for pristine forest systems have
be done to mitigate the likely changes to our already powerful conservation mechanisms that limit losses from
ęȱ¢ǰȱȱȱȱȱȱȱȱ their soils, but supplemental sources come from the ocean
operation by retaining a characteristic biodiversity. For ȱ ¢ȱ ȱ ęǰȱ ȱ ȱ Ğȱ
those few systems, in the remote areas of the continent, spawning, and the excreta of bears that feed upon them
where relatively undamaged systems still remain, the only ǻȱȱǯǰȱŘŖŖŘǼǯȱȱȱĴȱ¢ȱȱȱȱ
prescription can be to do nothing other than, if possible, of the immediate riparian zone. The debris of these trees,
extend their areas, particularly in a north-south direction. falling into the water, retains the carcasses, and the nutrients
that eventually decompose from them, within the river

114 Moss, B. et al
stretch. Aerial insects trapped at the water surface provide (Molles & Gosz, 1980) and towns has increased nutrient
ȱȱęȱ ȱȱȱǰȱǰȱȱȱ Ě ǰȱ ȱ ȱ ȱ ȱ ȱ ęȱ ȱ ȱ
birds may depend on the emergence of stream insects (Kato rocks and a change in the macroinvertebrate community
et al., 2003; Fukui et al., 2006). In the more open stretches, from detritivores towards herbivores. Dams have also
thickets of willow may form, the food of moose and other changed downstream features (Palmer et al., 2008), with a
deer, whose numbers may be controlled by wolves that ȱȱȱȱȱǰȱȱ ȱęȱȱ
are essential to maintenance of the plant and bird diversity Ğȱ ȱ ȱ ǰȱ ȱ ȱ ȱ ȱ
in the river corridor (Ripple & Beschta, 2004a,b; Beschta ȱȱȱ ǯȱȱȱȱĚ ȱȱȱȱ
ǭȱǰȱŘŖŖŞǼǯȱȱ
ȱĚ ȱȱ¢ȱȱǰȱ much more build up of sediments in semi-permanent
and gravel and sand bars separate the braids of what beds and a simpler, less dynamic geomorphological
becomes a structurally complex system, with successions structure. Woody debris is scarce, for modern forestry
to woodlands of poplar, willow and alder on the islands turns everything into a commercial resource and is even
in the stream. In Mediterranean regions, similar systems depriving some systems of essential calcium (Jeziorski et
show their particular characteristics in generally lower, but al., 2008). But the nutrient parsimony of the former system
ĚȱĚ ǰȱȱȱ ȱȱȱȱȱ is generally replaced by a nutrient surplus that leads to
stream channel in the hot dry summers (Acuña et al., 2005). loss of biodiversity and sometimes dense beds of plants
Like all other aquatic ecosystems in Europe, erosive ȱ ȱ ȱ ¢ȱ ȱ ȱ ȱ ȱ Ěȱ
river systems have already been changed by many impacts. of a riparian zone that is now agricultural or built upon.
There have been large amounts of acidity delivered in ¢ȱȱȱȱȱȱȱȱĞȱȱȱ
rain and snow, but many small streams have not been former characteristics of the river continuum. Removal of
so severely polluted in other ways, so a comparatively riparian vegetation increases water temperatures to values
high share of invertebrate species has been conserved. typical for downstream regions; organic pollution and
ǰȱ ¢ȱ ȱ ȱ ȱ ȱ ěȱ eutrophication reduce the oxygen content of small streams
almost all European streams and rivers, and disrupt the to that typically found in medium-sized rivers. Processes
river continuum and the links between the stream and and biota change accordingly. In terms of temperature,
its terrestrial surroundings. Most of the rivers, even in oxygen and food resources, the majority of erosive
Scandinavia, have been straightened and the channels river sections today resemble the original state of their
cleared of woody debris and large rocks to ease the transport downstream reaches, but at the same time lack characteristic
of timber from upstream logging to the coastal sawmills features of larger rivers, such as the high discharge.
(Törnlund & Östlund, 2002), or they have been dammed for Climate change will worsen this situation by further
hydroelectric generation. Intact medium-sized rivers can increasing water temperatures and associated features. It
only now be seen in northern Russia and in a few National will contribute to a general upstream movement of river
ȱȱȱǰȱȱȱĴȱȱ¢ȱȱ zones, particularly obliterating species bound to small
of their large mammals. The typical central European streams and springs, which cannot move further upstream.
ȱȱȱ¢ȱ¢ȱȱȱȱĚ ǰȱ ȱ ęȱ ȱ ȱ ǰȱ ¢ȱ ȱ ǰȱ ȱ
ȱȱȱȱȱ ȱǰȱĚȱȱ cool- or cold-adapted, and we may expect their loss with
or hydroelectric generation. Its riparian zone may still be ȱ¢ȱ ȱ ȱ¢ȱǻȱęǼȱ ȱ
ǰȱȱȱȱȱĞȱȱȱǰȱȱȱ the system is large enough and still not dammed from
of exotic, fast-growing planted conifers. The interactions its lowland reaches to prevent movement. Changes in
with large mammals have been lost, as have been the hydrology that lead to reduced summer rainfall may
former prodigious movements of the Atlantic salmon. promote further damming for water supply and storage
Development of the catchments as pastureland, ski slopes and further disruption. Extreme rainfall events, which are

ȱȱȱȱ¢ǰȱ ȱȱĞȱȬȱ Depositional rivers

the structure of the system. Summer droughts may reduce
Ě ȱ ¢ȱ ¢ȱ ȱ ȱ ȱ ȱ ¢ȱ ȱ ȱ ȱ ȱȱȱȱȱǰȱȱ ȱĚȱ
Mediterranean zone, where many rivers will be completely courses of rivers have already been seriously damaged
dry for large parts of the summer. Rivers process organic by human activities (Postel et al., 1996) so that almost no
Ĵȱ ȱ ¢ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ¢ȱȱȱĚȱȱȱ¡ȱ
to cope with raw sewage. In general the extremes of in Europe. They develop naturally where the catchment
that problem, and the consequent huge reductions in has reached a size that the amount of water entering at
¢ȱ ȱ ęȱ ȱ ǰȱ ȱ ¢ȱ ȱ Ě ȱ ȱ ȱ ȱ ¢ȱ ȱ ȱ ȱ
ǰȱȱȱȱȱȱȱĴȱȱȱǯȱȱ ȱȱȱȱ ȱȱĚ ǯȱȱȱȱ
ȱ ȱ ȱ Ě ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱȱȱȱȱ¢ȱȱȱĚ ȱȱ
will cause increased deoxygenation, threatening animal occur in winter and spring will spill onto a wider channel,
communities and the ability of the river to deal with urban ȱĚǯȱȱȱĚȱȱȱ ȱȱ
and agricultural wastes. Arctic streams will be particularly on the size of the river and its catchment, and therefore
vulnerable (Prowse et al., 2006) for temperatures are its outer reaches may be dry in some years, giving the
predicted to rise much more than in central Europe. Arctic ȱȱȱȱȱ¢ȱǰȱȱĚǰȱ
ęȱȱȱȱȱȱȱȱȱȱȱȱ rather than the reality of a sometimes-dry river bed. At
ȱȱȱ¢ȱȱȱȱȱȱęȱ¢ǯȱȱ ȱȱĚȱȱ ȱȱ¡ȱ¢ȱ¢ȱȱȱ
Declines in Arctic charr have already been noted and ȱĚȱ¢ǯȱȱȱ ȱĞȱȱȱ
ȱ ȱ ȱ ȱ ȱ ǻęȱ ȱ ǯǰȱ ŘŖŖŞǼǯȱ of silt and other sediment from upstream and will create
ȱ ȱ ȱ ěȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ĚǱȱ ǰȱ
be combined with measures to improve ecological backwater lagoons, ox-bow lakes where meanders are cut
quality under the Water Framework Directive. Hitherto, ěǰȱȱȱȱȱȱ£ȱȱ ȱ
improvements in water quality in English streams have from a permanent swamp close to the summer channel
sometimes masked climate-induced changes (Durance & to progressively less wet woodlands and grasslands at the
ǰȱŘŖŖşǼȱȱȱ ȱȱę¢ȱȱȱǯȱȱ Ěȱȱǻ¢ȱȱǯǰȱŗşşŜǲȱȱȱǯǰȱŘŖŖśǼǯȱȱ
The lessons to be learned from intact systems are that Large European rivers, like the Rhine, Vistula or Danube,
connectivity, both along the course and with the terrestrial ¢ȱȱĚȱȱȱ ǰȱ ȱ
surroundings, reduced nutrient inputs and increased formed a mosaic of main and abandoned channels, side
ěȱ ȱ ȱ ȱ ǻ ȱ ȱ ȱ ȱ ǰȱ ȱ ǰȱ ȱ Ěȱ ȱ ȱ ȱ ¢ȱ
stream as well as provide its energy source) are the three successional stages.
¢ȱȱȱȱȱěȱȱǰȱ In its pristine state, this complex structure harboured
ȱ ǰȱ ęǰȱ ȱ ȱ ȱ animals that fed on the emergent swamp vegetation,
climate change. Removal of dams may be impracticable, woody debris from the swamp forest, and organic
ȱ ȱ ȱ ȱ ȱ Ě ȱ ȱ ȱ deposits brought from upstream or derived from the
¢ȱ ǰȱ ȱ Ĵȱ ȱ ȱ ęȱ ȱ ȱ high productivity of the swamps. The summer channel
ȱ ęȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ Ȭȱ ǰȱ ȱ ęȱ
of the dam, will help. Above all, existing measures to a zooplankton community in the turbid water, and
ȱ ¡ȱ ǰȱ ȱ ȱ ęǰȱ ȱ sometimes a phytoplankton community where water
be continued, even increased. Climate change does not was retained long enough for the community not to be
replace existing problems; generally it will worsen them. ȱ ȱǻ¢ǰȱŘŖŖŖǲȱȱǭȱĴǰȱ
2002). In the swamp lagoons and ox bow lakes, the water

116 Moss, B. et al
cleared and a lagoon community developed so that the ȱ ȱ ȱ ęȱ ȱ ȱ Ȭȱ
Ěȱ ȱ ȱ ȱ ȱ ȱ ȱ ¢ǯȱȱ ȱ ȱĞȱȱǰȱȱȱȱȱ
At low water there was much interaction with the forests do not at all resemble the original state (Gherardi, 2007).
ȱȱĚȱȱȱ ȱȱȱȱ We do not expect a great deal of further decline in
moved in to graze on the lush grasslands, followed by their biodiversity that will be consequent on warming of
predators. There was a rich community of water birds. Ěȱ ȱ ȱ ȱ ȱ ȱ ȱ
Fish migrated both up and down river and outwards from zones. They have already been comprehensively wrecked.
the summer channel into the swamps to feed and spawn as ȱ ȱ ȱ ȱ Ěȱ ǰȱ ȱ
water levels increased in spring. Many traditional human might bring changes to the community of migratory birds
cultures have used the rhythm of water level change to (Sutherland, 1998) and to the dominant swamp plants.
ȱȱȱȱȱęȱȱȱ Ĵȱȱ Egrets, formerly only occasional in the UK, are now becoming
and temporary cultivation or grazing in the drier period. common (Combridge & Parr, 1992) as temperatures rise,
Much of this structure and function has been destroyed ȱ ęȱ ȱ ȱ ȱ ȱ ȱ ŗşşśǯȱ ȱ ȱ Ȭ ȱ
ȱȱĚȱȱȱȱȱ stilts arrive in the UK from the Mediterranean in warm
ȱȱĚȱȱȱȱȱȱȱ years (Figuerola, 2007). Warming will reduce oxygen
ęȱȱȱǰȱȱȱȱȱȱȱȱȱ concentrations in the backwaters and swamps, but swamp
farmed. Floodplain soils are very fertile when drained until ȱȱ¢ȱęȱȱ¡¢ȱȱȱ
their peats oxidise to an infertile acidity. The river system able to cope with this have developed, usually by adopting
ȱȱĞȱȱȱȱȱȱǰȱ some form of air rather than gill breathing. The expected
ȱ ȱ Ğȱ ȱ ȱ ȱ ȱ ȱ rise in temperature will only modestly change this situation.
of water downstream. Water is pumped into it from the What may be a far more prominent consequence,
ȱĚȱȱȱȱȱȱǯȱȱȱ however, is the expected change in hydrology and the
ȱȱȱȱ ȱĞȱȱȱȱ ȱȱȱȱ frequency of extreme events (Michener et al., 1997). Storms
river as the former organic soils have oxidised and even have already led to reclamation of part of the Mississippi
the inorganic soils have shrunk on drying. The pumped Ěȱȱ ȱȱȱȱŘŖŖŝȱȱ¢ȱ
ȱȱĞȱȱȱ¢ǰȱ ȱȱȱȱȱ ¡ȱĚȱȱȱȱ ǰȱȱȱǯȱȱȱ
traces of biocides used in the agriculture that develops. response to this has been to contemplate even larger
ǰȱȱȱȱ ȱȱȱȱĚȱ ȱ ǯȱ ȱ ȱ Ěȱ ȱ ȱ ¢ȱ ȱ
have led to greater rates of passage of water downstream decrease biodiversity greatly for there is comparatively
ȱ ȱ ȱ ȱ ȱ ȱ ȱ Ěȱ ȱ ĴȱĞȱȱ¢ȱȱ¢ǯȱȱȱȱȱ
in the reaches closer to the sea. Once there has been a long-term approach, however, would be to reconstitute
ȱ ȱ Ěȱ ȱ ȱ ȱ ȱ ȱ Ěȱ ȱ ȱ ȱ ¡ǰȱ ¢ȱ
ȱȱĚȱȱȱȱĞȱȱȱȱȱȱǰȱ from the upstream reaches to the lower; this would be
ȱȱȱ¢ȱĴȱȱȱȱȱȱ accompanied by greatly enhanced biodiversity and
Ěȱ¢ǰȱĚȬȱȱ ȱęȱ Ĵȱ ȱĚȱȱȱȱȱ ǯȱ
ǯȱ ȱ ȱ ¢ȱ ǰȱ ȱ ¢ȱ Ěȱ In the Arctic there will be serious damage. Here,
mosaic of terrestrial and aquatic habitats, has been lost. especially in the big river deltas of northern Scandinavia
The channel itself will continue to act as a habitat for ȱ ǰȱ ȱ ȱ ȱ ¡ȱ ȱ ȱ Ěǯȱȱ
aquatic organisms. Many large rivers were, however, Floods are to some extent dependent on ice dams that form
so severely polluted in the 20th century that there was as the rivers melt northwards in spring, and the permafrost
almost the complete disappearance of organisms other system provides freeze-thaw mechanisms that give rise
ȱ ǯȱ ȱ Ğȱ ȱ ȱ ǰȱ ȱ ȱ to millions of small lakes and lagoons. As temperatures

increase, permafrost melts and these features will be et al., 2001). Permanent Mediterranean lakes will be very
progressively lost in a more amorphous swampy landscape, ȱ ȱ ȱ ȱ ęȱ ȱ ȱ ȱ ȱ ¢Ȭ
possibly lacking the isolated small bodies of water that intolerant Cladocera, leading to reduced zooplankton
¢ȱȱĞȱ¢ȱěȱȱ ȱ grazing and higher phytoplankton densities. Natural
short distances (Hobaek & Weider, 1999). Roads and changes in water level lead to the presence of extensive
¢ȱ ȱ ȱ ǰȱ ¢ȱ ȱ Ě¢ȱ plant beds in Lake Kinneret in Israel in some (low water)
and mosquitoes will increase and existing economic years and complete absence when levels are higher (Gafny
activities in these regions may have to be abandoned. & Gasith, 1999).
Human activities have frequently switched these
Shallow lakes systems to a turbid state lacking plants and with reduced
¢ȱǻȱȱǯǰȱŗşŞşǲȱěȱȱǯǰȱŗşşřǲȱȱ
Shallow lakes, in the sense used here, are those where the et al., 2005), although the process can also occur naturally
predominant primary production comes from algal and ȱěȱ ȱ ȱȱǻǰȱ
ȱȱȱȱ ȱȱĴǰȱ 1980) or weather-induced changes in water level (Blindow
rather than from the phytoplankton (Moss et al., 1996; et al., 1993). A combination of increased nutrient loading
Lachavanne & Juge, 1997). In practice this means lakes ȱęȱȱȱ¢ȱȱȱǻȱĴǰȱ
with a mean depth less than about 3 m. In their pristine herbicides, introduction of exotic grazing birds such as
state, such lakes are rich in biodiversity, except sometimes ȱǰȱęȱȱȱȱǰȱȱȱ
in highly peaty regions where the water is stained deep ȱ ȱ ȱ ȱ ȱ ¢ęȱ ǻ£ȱ ȱ ǯǰȱ
brown and light penetration is impeded. Otherwise, a 2003)) or those that disrupt the mechanisms that maintain
community of perhaps 10–30 submerged macrophyte the water clarity (for example, toxicity to zooplankton
species (James et al., 2005) supports a diverse periphyton through pesticide residues, increased salinity or heavy
ȱ Ȭȱ ȱ ǻ ħ ȱ ȱ ǯǰȱ metals) is responsible. There has been considerable
ŗşşŖǼǯȱȱȱȱ¢ȱęȱȱ ȱǯȱȱȱȱ ěȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ
community is also present, but dominance by individual systems because of their conservation and biodiversity
phytoplankton species is avoided through grazing by the importance (Moss et al., 1996; Jeppesen et al., 1999, 2005).
£ȱȱęȱȱȱȱęȱȱ ȱ¢ȱȱȱěǯȱȱȱ¢ȱ
within the plant beds. The water remains clear as a result, enhance the symptoms of eutrophication by increasing the
ȱȱę¢ȱȱȱǰȱȱ rate of release of phosphorus from sediments (McKee et al.,
in the pristine state, with a catchment covered by natural 2003), increasing phytoplankton growth rates (van Donk
vegetation, nutrient inputs will be low. In ponds, which also & Kilham, 1990; Reynolds, 1997; Howard & Easthope,
ȱ ȱȱ¢ǰȱȱ ȱĚȱ ȱ 2002) and encouraging the growth of exotic plant species
ęȱȱȱǻęȱȱǼǰȱȱȱ¢ȱȱȱ that displace native species (McKee et al., 2002), not least
some small ponds will also support a diverse community Ěȱ ȱ ǻ¢ȱ ȱ ǯǰȱ ŘŖŖşǼǰȱ ȱ ȱ ȱ
of unusual invertebrates capable of aestivating under such such as Salvinia and spp. causing major problems.
conditions (Jakob et al., 2003). Mediterranean shallow lakes In Sweden, a prolonged growing season appears to
that dry out in summer may be individually less diverse be favouring the invasion of a mucilaginous alga,
than those in central Europe for they are strongly limited Gonyostomum semen, which dominates the phytoplankton
by changes in salinity (Green et al., 2002a, 2005; Frisch et biomass with reductions in overall diversity. Warming
al., 2006a,b; Beklioglu & Tan, 2008) but support a specialist will decrease oxygen concentrations as a result of purely
fauna derived from a high regional diversity well adapted physical relationships at least, but disproportionately
to that (Giudicelli & Thierry, 1998; Bazzanti et al., 1996; Boix from increased biological activity, leading to changed

118 Moss, B. et al
ęȱ ȱ ȱ ȱ ęȱ ȱ ȱ ¢ȱ ǯȱȱ that inevitably have been brought in from the warmer
The piscivorous Northern pike (¡ȱ ) is expected regions and which will therefore thrive, and misguided
to show some of the most severe decreases (Reist et al., ȱ ¢ȱ ȱ Ĵȱ ȱ ȱ ȱ ęȱ
ŘŖŖŜǼǯȱ ȱ ȱ ęȱ ȱ ¢ȱ ȱ ȱ ¢ȱ ȱ ȱ stocks demanded by the more competitive of anglers.
in a shallow lake, but the introduced, rather damaging Lakes have the problem (and sometimes advantage)
common carp (Cyprinus carpio) will, and indeed may ȱȱȱȱȱ¢ȱȱȱĚ¢ȱ
ȱȱě¢ȱȱȱȱȱȱȱȱȱ insects will readily colonise a new lake, migration of
does at present. The native tench (Tinca tinca) and crucian species from the south to replace those that do not survive
carp (Carassius carassius) may also thrive. In addition, warming will be impeded by isolation (Reist et al., 2006).
there may be life history changes, leading to dominance Individual populations may become extinct in any case if
of small, fast reproducing species and individuals, with migration is too limited and the residual populations lack
major consequences for abundance and diversity of lower genes that would allow them to adjust to climate change.
ȱȱǻěȱȱǯǰȱŘŖŖŝǲȱȱȱǯǰȱŘŖŖşǼǯ Nonetheless, birds are major vectors of both invertebrates
ȱȱȱęȱ¢ȱȱȱȱȱ and plants (Green et al., 2002b; Green & Figuerola, 2005;
ȱęȱȱ ¢ȱȱȱ¢ǰȱȱȱȱ Frisch et al., 2007) and migratory birds may become very
other species with the intentions of sport rather than balanced important in restocking biota depleted by warming.
¢ǯȱ ȱ ȱ ȱ ¢ȱ ěȱ ȱ ȱ ȱ ȱ ȱ ęȱ ȱ ȱ ȱ ǰȱ ǰȱ
drying out in warmer summers and in the Mediterranean where major barriers of mountain or sea intervene.
zone many will disappear (Blondel & Aronson, 1999). The prognosis for the distal islands of Europe such as
Those shallow Mediterranean lakes that persist, already Iceland, the Faeroes, Ireland and the United Kingdom, is
stressed by variable and high salinity, will become more especially bad. Pressures to introduce species deliberately
saline, more extreme environments, with reduced diversity will be greater than on the mainland and since there
ǻĴȱǭȱǰȱŘŖŖŖǲȱȱȱǯǰȱŘŖŖřǲȱȱȱ is a high chance of other species (associated algae and
al., 2005; Green et al., 2005, Beklioglu et al., 2007; Boix et al., Ǽȱȱȱ ȱȱȱęȱȱ
2008). With reduced water levels, the former open water plant introductions, there is a risk that although biodiversity
of Lake Chameiditida in Greece has become dominated might be maintained in terms of species richness, there will
¢ȱȱĚȱĞȱȱȱȱǻ¢ȱTypha) since be major problems in the structure of the communities
the 1970s. Many amphibians will become locally, perhaps ȱǯȱȱħȱȱǯȱǻŘŖŖśǼȱȱȱ ȱ ȱ
regionally extinct. Small positives might include reduced exacerbate the loss of submerged plant communities.
nutrient loading from decreased precipitation, but greater Mescosm experiments (McKee et al., 2002; Feuchtmayr et
release of nutrients from the sediments and increased al., 2009) do not support this but suggest a major increase
frequency of complete drying out will cancel any such ȱ Ěȱ ȱ ȱ ȱ ȱ ęȱ ǯȱȱ
advantage (Beklioglu & Tan, 2008; Jeppesen et al., 2009). Comparative analyses of lakes in Europe, Florida and
ȱ ȱ ȱ ěȱ ¢ȱ ȱ ȱ Uruguay, however, indicate an increase in many symptoms
ȱȱȱȱȱȱȱěȱ of eutrophication, even if plants are present, and loss of
to reduce phosphorus loads (Cooke et al., 2005), remove ȱ¢ȱǻȱȱǯǰȱŘŖŖŝǰȱěȱȱǯǰȱŘŖŖŝǼǯȱȱȱ
ȱęȱȱǻȱȱȱǯǰȱŗşşŖǲȱ
ȱ secondary spread of common carp could prove devastating
et al., 1998; Mehner et al., 2002) and limit ingress of toxic ȱȱǰȱȱħȱȱǯȱǻŘŖŖśǼȱ¢ȱǯ
pollutants. Mitigation against warming can only mean a
ȱěȱȱȱȱȱ ȱȱȱ
¢ȱȱ¢ȱȱęȱȱȱȱ
local food or biomass production, a spread of exotic species

Deep lakes membranes and interference with reproduction, and to

bring about less diverse plant communities as Sphagnum
Deep lakes are not sharply distinct from shallow lakes, comes to predominate. Precipitation of phosphorus by
but in general their productivity is dominated by aluminium mobilised from the catchment and removal of
ȱ ȱ ¢ȱ ȱ ȱ ¢ȱ ĴȬ bicarbonate by reduced pH may result in a greater diversity
living communities of submerged plants and associated of certain algal groups, such as the Chrysophyceae and
ǯȱȱ¢ȱĞȱȱȱ¢ȱȱĚȱ ȱ ǰȱ ȱ ȱ ȱ Ěȱ ȱ ȱ ¢ȱ ȱ
water (epilimnia) and colder denser water (hypolimnia) ¢ȱǯȱȱĚȱȱȱȱ£ȱȱ ȱ
during the summer. In Europe, deep lakes are widespread ȱȱȱȱ£ȱȱĴȱȱęȱ
in upland regions or regions of poorly weathered rock. In are not unusual. In the Mediterranean zone there are few
their pristine states they have very low nutrient loadings deep lakes that are not man-made by damming rivers.
and clear waters, are dominated by a sparse plankton, and ȱǰȱ ȱȱěȱȱȱ ȱ
Ğȱ¢ȱęȱȱȱȱȱȱǯȱȱ by sheer volume of water than shallow lakes, are already
ȱ ȱ ȱ ȱ ȱ Ĵȱ £ǰȱ Ğȱ ȱ ȱ ȱ ȱ ěȱ ȱ ȱ ȱ ȱ ȱ ȱ ¡ȱ
ȱȱ ȱǰȱȱ ȱȱȱȱĴȱ ȱ ȱ ęǯȱ ȱ ȱ ȱ ȱ ȱ ȱ ȱ
is much limited by the depth of the basin. Large size may melts earlier in spring. They have responded to warmer
give an increased overall richness not least because the winter and early spring temperatures by shallower mixing
existence of sheltered bays and exposed shores in the same ȱ ȱȱȱȱȱęǰȱȱ
water body leads to a variety of inshore communities. increased density and biomass of plankton. Zooplankton
Dependent on absolute depth and continentality, which growth, however, also occurs earlier and there may be earlier
increases the stability of the layering because of higher suppression of phytoplankton by zooplankton grazing
surface temperatures, hypolimnia may be naturally poor (Straile, 2000; Anneville et al., 2005). Earlier warming and
in oxygen or even anaerobic, but typically are big enough ȱȱ ȱȱĞȱȱȱȱȱ
to retain a substantial oxygen store, for example in the large ȬȱęǯȱȱȱȱȱȱȬȱ
sub-alpine lakes. The deeper waters may provide summer dark hypolimnion, in which zooplankton prey may be
ȱȱȬȱęȱȱȱȱȱȱ ȱȱęȱȱȱȱȱȱ¢ȱęȱǻȱ
salmonids, and the open waters provide hunting grounds ȱǯǰȱŘŖŖŝǼǰȱȱ ȱȱȱȱȱęȱȱ
ȱȱȱęȱǰȱ¢ǰȱȱȱȱ that require cool, well-oxygenated water in summer. This
that shun smaller, shallower lakes and are part of the ¢ȱȱȱȱȱȱȱęȱȱ
pelagic food web. are cold-water adapted and more common in the Arctic
ȱ ȱ Ĵȱ ȱ ȱ ȱ ǰȱȱȱȱȱȱęȱȱȬȱ
ȱ Ğȱ ȱ ȱ ȱ ȱ ȱ ȱ ǰȱ ǲȱ¢ȱ¢ȱȱęȱȱȱǯȱȱ¢ȱ
in contrast to the situation for many shallow lakes in the invertebrate species in this region are probably highly
lowlands. Eutrophication leads to greater deoxygenation cold-adapted and will become extinct but may be replaced
of hypolimnia, to blooms (Ibelings et al., 2003) (in the strict through rapid evolutionary mechanisms by similar forms.
sense of cyanobacterial populations that migrate within Birds may bring in new plants and invertebrates, for many
the water column sometimes to form a surface scum – the migrants move through the entire latitude of Europe.
Ǽǰȱ ȱ ¢ȱ ȱ ȱ ȱ Ȭ ȱ ęȱ ȱȱȱĚȱȱȱȱȱȱȱ
species that require high oxygen concentrations and which lakes perhaps have a less potent, though still measurable,
previously found cool oxygenated refuges in summer ěȱȱȱ ȱȱȱ ȱǯȱȱȱȱȱȱ
ȱ ȱ ¢ǯȱ ȱ ęȱ ȱ ȱ ȱ ȱęȱȱȱȱȱȱȱȱ ȱǰȱ
many animal species through damage to gill and other when oxygen concentrations may fall greatly, the deeper

120 Moss, B. et al
lakes may prove more resilient. Where lakes contain a ȱ ȱȱȱȱȱĴǰȱȱȱ

¡ȱȱ Ȭȱ¢ȱęȱȱȬȱ ¡ȱȱȱ ȱȱȱĝȱǯȱȱ
esocids, salmonids and coregonids, we expect a spread of Conversely, increased costs of oil (for reasons other than
cyprinids throughout the lake and this may have major ȱ Ǽȱ ȱ Ĵȱ ȱ ȱ ȱ ȱ
ȱȱȱȱȱęȱȱ (because of carbon dioxide release) may lead to lowered
usually favour the former groups. Mediterranean deep NOx emissions and hence some reduction in nitrogen
ȱ ȱěȱ ȱ ȱǰȱȱȱǻ ȱ loading on upland lakes. This could well be completely
ȱǯǰȱŗşşŜǼȱȱȱȱȱȱȱĴȱ£ȱ ěǰȱ ǰȱ ¢ȱ ȱ ȱ ȱ ȱ ȱ Ĵǯ
with a major reduction in biodiversity. Their hypolimnia
will deoxygenate but only to a slightly greater degree, for Conclusions
they already become seriously deoxygenated in summer.
ȱ¢ȱěȱȱȱȱ¢ȱȱȱ We approach future climate change from a baseline of
because upland regions that presently do not support crop freshwater habitats that are already seriously damaged,
agriculture may be increasingly brought into cultivation, with the damage being greatest in the central and
ȱ ȱ ȱ Ȭěǯȱ ȱ ǰȱ ȱ ȱ ȱ£ǰȱȱȱęȱȱȱ¡ȱ
north. Only for very large,
deep lakes do we have
more than a handful of sites
in central and southern
Europe that can remotely
approximate to pristine
conditions. Six thousand
years of progressive
ȱ ȱ Ĵȱ
have rendered the concept
of ‘High ecological status’
ęȱ ¢ȱ ȱ ȱ
Framework Directive a
largely hypothetical one,
ȱ ȱ ęȱ ȱ
ȱȱ¢ȱĜȱ
bodies (Moss, 2008). In
ȱ ȱ ěȱ ȱ
warming and associated
changed hydrology, we
thus start from a basis
where many interacting
factors have already
Fig. 5.ȱ ȱ ȱ Ěȱ ȱ ȱ ȱ ȱ ȱ ¢ȱ ǻȱ ȱ ȱ Ǽȱ ȱ ȱ ȱ
temperature, European latitude (taking into account the possibilities of invasion from the south) and reduced biodiversity and
residence time. The higher the temperature increase, the greater likelihood of decline due to lack of time continue to threaten it.
for adjustment to occur. Invasion of the Mediterranean region from the south is hindered by marine
We approach changes
barriers and although invasion of the Boreal/Arctic is possible, it will be counteracted by extinction
of cold-adapted species. Long residence time gives a degree of resistance because it implies a larger in biodiversity in the
system.

that the major changes will

be in the extreme north and
in the south, that they will
be greater the more marked
the climate change, and
that they will impact river
systems most and deep
lakes least. Finally, Tables
1, 2 & 3 summarise current
major human impacts,
additional impacts of
warming and appropriate
mitigation measures for
maintenance of biodiversity
in freshwater ecosystems.
There is increasing
experimental evidence
that the degree of pristine
biodiversity has meaning
and that we must view
Fig. 6.ȱȱȱȱěȱȱȱȱ ȱȱȱȱȱ ȱ¢ȱȱ
geographical location in Europe on degree of decline of biodiversity (in a wide sense). Position within
any substantial loss of it as
the three-dimensional box indicates the degree of change in biodiversity from highest to lowest with serious. The operation of
denser shading indicating greatest change. the engines of ecosystems
knowledge that with tens of thousands of species occurring is a function of availability of an appropriate range of
in potentially trillions of combinations, with elements components (Tilman et al., 1996; Hooper et al., 2005),
of randomness as well as determinism both by physico- with much built-in substitution (spare parts) to cope
chemical and biological mechanisms, and with continuous ȱ ¢ȱ Ěȱ ǯȱ ȱ ȱ ȱ ȱ
evolutionary change possible in every species, we have understanding of earth-systems-science states that
no hope of making sensible predictions beyond those our existence on this planet depends on the continued
that are very general. When human intervention is also functioning of natural systems that regulate atmospheric
programmed into this, even the generalities become very and oceanic composition, we must be very concerned that
uncertain. It is like juggling with a trillion balls. The task these systems are being damaged so extensively. There
is impossible. All that we can be certain of is that almost are political hopes that climate change can be mitigated
all of our habitats are severely damaged already and that ȱȱǰȱȱȱęȱ ȱȱȱǯȱȱ
warming is likely to worsen their quality in many ways. Our fears as professional ecologists, however, are that
Fig. 5 summarises the likely degrees of change in biodiversity political institutions have isolated themselves so much
to be expected for the three broad European zones we from knowledge of the fundamental driving engines
have considered, in terms of increased temperature (and of the biosphere that they leave us with an increasingly
associated alterations in hydrology) and retention time of defunct vehicle. Yet there remain skilled mechanics in
the water mass, as we move from upland erosive streams ȱȱȱ¢ȱȱȱȱĴȱ
ȱĚȱǰȱ ȱȱȱȱǯȱȱFig. 6 advice than those to whom policy-makers currently turn.
combines these three variables into a picture that suggests

122
Table 1. Summary of current major human impacts, additional impacts of warming and appropriate mitigation measures, where possible, for maintenance of biodiversity in Boreal-Arctic
freshwater ecosystems. The lists are not comprehensive.
Present major impacts ěȱȱ Mitigation

Boreal-Arctic erosive rivers Relatively small in the Arctic; ȱĚ ȱ ȱȱǲȱȱȱ Maintenance of intact forested corridors
most Boreal rivers are dammed Ȭ ȱęȱǻ¢ȱȱȱ in the Boreal and continuation of
and catchments disrupted by coregonids) and other animals; increased ȱȱȱęǯȱȱ
forestry with toxic pollution likelihood of damming for power generation. Problems need largely to be tackled
(paper industry), eutrophication at source (i.e. severe restriction of
ȱęǯȱȱȱ greenhouse gas emissions).

mercury in peaty areas.
Boreal-Arctic depositional As for erosive rivers in this zone. ȱĴȱȱĚ ȱ ȱȱ Maintenance of intact forested corridors
Ěȱ melt and faster spring melt. Disruption of in the Boreal, greater nutrient restriction
¢ȱęȱǯȱȱȱȱȱȱ and continued measures to combat
greater coalescence of associated wetlands. ęǯȱȱȱȱȱȱ
ȱȱȬ ȱȱȱęǯ tackled at source (i.e. severe restriction of
greenhouse gas emissions).
Boreal-Arctic shallow lakes As for erosive rivers in this zone. Increased growth season for plants. Reduced Greater nutrient restriction and
season of ice cover and reduced winter continued measures to combat
deoxygenation. Greater nutrient input. ęǯȱȱȱ¢ȱȱȱ
ȱȱ¢ȱęȱȱ¢ȱ extensive for much individual
ęȱǯȱȱȱȱȬ ȱęǯȱȱ mitigation. Problems need to be tackled
by exotic species. Increased release of carbon at source (i.e. severe restriction of
dioxide and methane from sediments and greenhouse gas emissions).
peats.
Boreal-Arctic deep lakes As for erosive rivers in this zone. Increased growth season. Greater hypolimnial Greater nutrient restriction and
deoxygenation. Greater nutrient input. continued measures to combat
Mismatches between algal, zooplankton and ęǯȱȱȱȱȱ
ęȱ ȱȱȱęǯȱȱȱȱ undesirable exotics may be possible.
Ȭ ȱęǯȱȱȱ¢ȱ¡ȱǯ Problems need to be tackled at source
(i.e. severe restriction of greenhouse gas
emissions).
DOI: 10.1608/FRJ-2.2.1
Moss, B. et al
Table 2. Summary of current major human impacts, additional impacts of warming and appropriate mitigation measures, where possible, for maintenance of biodiversity in Central-
European freshwater ecosystems. The lists are not comprehensive.

Central erosive rivers Already very severe ȱȱȱǰȱĚ ȱȱ¢ȱ ȱ ěȱȱ ȱȱȱ ȱ
ęǯȱȱęǰȱ exacerbate most existing problems. Farming at higher native trees. Nutrient management on
DOI: 10.1608/FRJ-2.2.1
damming, eutrophication elevations will exacerbate eutrophication. Replacement farms. Restrictions on fuel consumption
from atmospheric nitrogen of cool water species by warmer water species. More and land fertilisation may alleviate nitrogen
sources and upland farming, damming for power generation, and for water storage for ęǯ
and deforestation of catchment. irrigation and domestic supply. Some invasion of exotic Reservoir management must incorporate
ěȱȱȱ ȱ species. ȱȱĚ ȱȱ ȱȱȱȱ
Climate change and biodiversity in Europe
¡ȱĞ ȱȱȱ protect downstream systems.

sources. Channel engineering.
Central depositional Already devastated by More summer deoxygenation in engineered channels and ȱȱĚȱ¢ȱ
Ěȱ engineering works involving ęȱǯȱȱ ȱ¢ȱȱǯȱȱȱȱ from upstream to downstream. Nutrient
drainage and channelisation. ¢ȱęȱǯȱȱȱȱ¡ȱ¢ȱȱȱǯ management. Full enactment of Water
ȱĚȱȱǯ Framework Directive to restore systems to
good ecological quality.
Central shallow lakes Severe eutrophication. Increased symptoms of eutrophication. Increased Extensive nutrient management. Fisheries
Introduction of damaging deoxygenation, greater but less diverse plant growth, risk management to exclude undesirable
ęȱǻǯǯȱȱǼǯȱȱ ȱȱęȱǰȱ¢ȱȱǯȱȱȱ invaders and maintain balanced community
Recreational damage by boats. release of carbon dioxide and methane from sediments. with piscivores. Irrigation restrictions to
Ȭěȱȱȱǯȱ
ȱȱȱȱ¢ȱęǯȱȱȱ¢ȱ maintain water supply. Intensive nutrient
undesirable exotics. Increased nutrient loading from management for agriculture and urban
more intensive farming for food and biofuels. Water sources. Restoration of construction of
level reduction from increased irrigation in general area. surrounding wetlands. Probably too
Flooding by sea in coastal areas as sea levels rise and expensive to protect coastal lagoonal systems
complete loss of some lagoonal systems. from sea-level rise.
Central deep lakes ęȱȱǰȱ As for Boreal-Arctic deep lakes but with markedly Intensive nutrient management. Fisheries
eutrophication almost increased symptoms of eutrophication; greater management to exclude invaders and
everywhere. Engineering to hypolimnial deoxygenation and almost complete loss of maintain balanced community with
raise and control levels for ȱȱȱęǯȱȱ ȱȱȱȱ piscivores. Restriction of vehicle usage
ȱ¢ȱȱȱȱĴȱ ¢ȱęǯȱȱȱȱȱȱ ȱǯȱ ȱȱȱęȱȱ
Freshwater Reviews (2009) 2, pp. 103-130

123
zones eutrophication problems. Full enactment of

provisions of Water Framework Directive.
124
Table 3. Summary of current major human impacts, additional impacts of warming and appropriate mitigation measures, where possible, for maintenance of biodiversity in Mediterranean
freshwater ecosystems. The lists are not comprehensive.

Mediterranean erosive Eutrophication, exacerbated by upland ȱěȱ ȱȱȱȱ ȱȱĚ ȱȱ ȱ
rivers ȱęǯȱȱȱȱȱȱ Ě ǯȱȱ¢ȱ¢ȱȱ¢ȱ storage reservoirs. Re-establishment
corridors to farming. Damming dry with loss of biodiversity. Severe of forest corridors and rigorous control
for water storage and inadequate deoxygenation and increased salinity in ȱȬȱęǯȱȱȱȱ

ȱĚ ǯȱȱȱ¡ȱ ȱȱȱ ȱȱȱȱęȱ agriculture in arid areas. Problems need to
species causing severe problems, and invertebrates. More invasion of exotic be tackled at source (i.e. severe restriction
¢ȱęȱȱ¢ęǯ species. of greenhouse gas emissions).
Mediterranean depositional Already devastated by engineering More summer deoxygenation in Restriction of irrigated agriculture in arid
Ěȱ works involving drainage and ȱȱȱęȱǯȱȱ ȱ areas. Removal of dams. Restoration of
ǯȱȱȱĚȱȱ symptoms of eutrophication. Greater Ěȱ¢ȱȱȱȱ
intact. ęǯȱȱȱȱ¢ȱęȱǯȱȱ downstream. Nutrient management. Full
Invasion of exotics likely to be severe. enaction of Water Framework Directive to
restore systems to good ecological quality.
Mediterranean shallow Increased salinity owing to loss of More will completely dry out in summer, Nutrient management and greater
lakes natural water supply to irrigation. becoming intensely saline as they do so restriction on use of water for irrigation
Complete drying out in many cases. with restriction, then loss of the biota. will help, as will full enaction of Water
Eutrophication from farming. ȱȱȱȱęȱȱȱ Framework Directive to restore systems to
species. Eutrophication symptoms will good ecological quality. Problems need to
increase (particularly cyanobacterial be tackled at source (i.e. severe restriction
blooms) in those that persist. of greenhouse gas emissions).
Mediterranean deep lakes There are few of these and most are ęȱȱ¡ȱǰȱ Ĵȱȱǰȱȱȱ
ęȱǰȱěȱȱ ¢ȱȱȱĴȱ£ȱȱ restrictions on irrigation and urban
¡ȱĚȱȱ ȱȱȱ ęȱ ȱȱ ȱȱ¡ȱ development. Problems need to be
severe eutrophication, except in the drawdown. More intense and frequent tackled at source (i.e. severe restriction of
uplands. algal blooms as temperatures rise. greenhouse gas emissions).
DOI: 10.1608/FRJ-2.2.1
Moss, B. et al
References composition of Espolla pond (NE Iberian peninsula): the

neglected biodiversity of temporary waters. Wetlands 21, 577-
Acuña, V., Muñoz, I., Giorgi, A., Omella, M., Sabater, F. & Sabater, 592.
S. (2005). Drought and postdrought recovery cycles in an Boix, D., Gascón, S., Sala, J., Badosa, A., Brucet, S., López-Flores,
Ĵȱ ȱ Ǳȱ ȱ ȱ ȱ ǯǰȱ¢ǰȱǯǰȱ ǰȱǯȱǭȱǰȱǯǯȱǻŘŖŖŞǼǯȱȱĴȱȱ
aspects. Journal of the North American Benthological Society 24, composition and species richness of crustaceans and aquatic
919-933. insects along environmental gradients in Mediterranean water
Amezaga, J.M., Santamaria, L. & Green, A.J. (2002). Biotic wetland bodies. Hydrobiologia 597, 53-69.
connectivity – supporting a new approach for wetland policy. Bonis, A., Lepart, J. & Grillas, P. (1995). Seed bank dynamics and
Acta Oecologica – International Journal of Ecology 23, 213-222. coexistence of annual macrophytes in a temporary and variable
ǰȱǯȱǭȱĴǰȱ ǯȱǻŘŖŖŘǼǯȱȱ¢ȱȱ¡¢ȱ habitat. Oikos 74, 81-92.
ȱ ȱ ȱ ȱ Ěǯȱ ȱ Freshwater Biology 47, Brown, L.E., Hannah, D.M. & Milner, A.M. (2007). Vulnerability of
761-776. alpine stream biodiversity to shrinking glaciers and snowpacks.
Anneville, O., Gammeter, S. & Straile, D. (2005). Phosphorus Global Change Biology 13, 958-966.
decrease and climate variability: mediators of synchrony in Brucet, S., Boix, D., López-Flores, R., Badosa, A., Moreno-Amich,
phytoplankton changes among European peri-alpine lakes. R. & Quintana, X. (2005). Zooplankton structure and dynamics
Freshwater Biology 50, 1731-1746. in permanent and temporary Mediterranean salt marshes:
Balian, E.V., Segers, H., Lévèque, C. & Martens, K. (2008). The taxon-based and size-based approaches. Archiv für Hydrobiologie
freshwater animal diversity assessment: an overview of the 162, 535-555.
results. Hydrobiologia 595, 627-637. Carpenter, S.R., Frost, T.M., Persson, L., Power, M. & Soto, D.
Bazzanti, M., Baldoni, S. & Seminara, M. (1996). Invertebrate (1995). Freshwater ecosystems: linkages of complexity and
macrofauna of a temporary pond in Central Italy: composition, processes. In: ȱȱȱ¢Ǳȱȱ ȱ
community parameters and temporal succession. Archiv für (eds H. Mooney et al.), pp. 96-105. John Wiley & Sons, N.Y.
Hydrobiologie 137, 77-94. Ĵǰȱ ǯȱ ǭȱ ǰȱ ǯȱ ǻŘŖŖŖǼǯȱ ȱ ¢ȱ ȱ ȱ
Beklioglu, M. & Tan, C.O. (2008). Restoration of a shallow mechanisms in brackish water fauna. ȱȱȱ
Mediterranean lake by biomanipulation complicated by 40, 7-14.
drought. Fundamental and Applied Limnology 171, 105-118. ǰȱǯȱǭȱǰȱǯȱǻŗşşŘǼǯȱȱĚ¡ȱȱĴȱȱȱȱȱ
Beklioglu, M., Altinayar, G. & Tan, C.O. (2006). Water level control Ireland in 1989. British Birds 85,16-21.
ȱȱ¢ȱȱȱęȱ ȱȱ Connell, J.H. (1978). Diversity in tropical rain forest and coral
of Mediterranean Turkey. Archiv für Hydrobiologie 166, 535-556. reefs. Science 199, 1302-1310.
Beklioglu, M., Romo, S., Kagalou, I., Quintana, X. & Bécares, Cooke, G.D., Welch, E.B., Peterson, S.A. & Nichols, S.A. (2005).
E. (2007). State of the art in the functioning of shallow Restoration and Management of Lakes and Reservoirs, 3rd edition.
Mediterranean lakes: workshop conclusions. Hydrobiologia 584, Taylor & Francis, London & New York. 548 pp.
317-326. Cosswig, K. (1955). Zoogeography of the near East. Systematic
Beschta, R.L. & Ripple, W.J. (2008). Wolves, trophic cascades, and Zoology 40, 49-96.
rivers in the Olympic National Park, USA. Ecohydrology 1, 118-130. Crawley, M.J., Johnston, A.E., Silvertown, J., Dodd, M., de
Blindow, I., Andersson, G., Hargeby, A. & Johansson, S. (1993). Mazancourt, C., Heard, M.S., Henman, D.F. & Edwards, G.R.
ȬȱĴȱȱȱȱȱȱ ȱ ȱ (2005). Determinants of species richness in the Park Grass
eutrophic lakes. Freshwater Biology 30, 159-167. experiment. The American Naturalist 165, 179-192.
Blondel, J. & Aronson, J. (1999). Biology and Wildlife of the Cumming, G.S. (2007). Global biodiversity scenarios and
ȱǯ Oxford University Press, Oxford. landscape ecology. Landscape Ecology 22, 671-685.
Boix, D., Sala, J. & Moreno-Amich, R. (2001). The faunal Davidson, T.A., Sayer, C.D., Bennion, H., David, C., Rose, N.

126 Moss, B. et al
& Wade, M.P. (2005). A 250 year comparison of historical, Moss, B., Harvey, I. & Atkinson, D. (2009). Global warming
macrofossil and pollen records of aquatic plants in a shallow ȱǱȱěȱȱ ȱ¢ȱȱȱ
lake. Freshwater Biology 50, 1671-1686. communities in experimental, hypertrophic, shallow lake
Declerck, S., Vandekerhove, J., Johansson, L., Muylaert, K., Conde- mesocosms. Journal of Applied Ecology 46, 713-723.
Porcuna, J.M., van der Gucht, K., Pérez-Martinez, C., Lauridsen, Figuerola, J. (2007). Climate and dispersal: black-winged
T., Schwenk, K., Zwart, G., Rommens, W., López-Ramos, A., stilts disperse further in dry springs. ȱ 2(6), e539.
Jeppesen, E., Vyverman, W., Brendonck, L. & De Meester, doi:10.1371/journal.pone.0000539.
L. (2005). Multi-group biodiversity in shallow lakes along Frisch, D., Moreno-Ostos, E. & Green, A. (2006a). Species richness
gradients of phosphorus and water plant cover. Ecology 86, and distribution of copepods and cladocerans and their relation
1905-1915. to hydroperiod and other environmental variables in Doñana,
ǰȱ ǯǯǰȱ Ĵǰȱ ǯǯȱ ǭȱ Ĵǰȱ ǯǯȱ ǻŘŖŖŖǼǯȱ ȱ ȱ south-west Spain. Hydrobiologia 556, 327-340.
relationship in lake communities between primary productivity Frisch, D., Rodríguez-Pérez, H. & Green, A.J. (2006b). Invasion of
and species richness. Ecology 81, 2262-2679. ęȱȱȱÛȱȱǰȱ ȱǰȱ¢ȱ
Downing, J.A., Prairie, Y.T., Cole, J.J., Duarte, C.M., Tranvik, L.J., an exotic estuarine copepod. Aquatic Conservation: Marine and
Striegl, R.G., McDowell, W.H., Kortelainen, P., Caraco, N.F., Freshwater Ecosystems 16, 483-492.
Melack, J.M. & Middelburg, J.J. (2006). The global abundance Frisch, D., Green, A.J. & Figuerola, J. (2007). High dispersal capacity
and size distribution of lakes, ponds, and impoundments. of a broad spectrum of aquatic invertebrates via waterbirds.
Limnology and Oceanography 51, 2388-2397. Aquatic Sciences 69, 568-574.
Drescher-Schneider, R., de Beaulieu, J.-L., Magny, M., Walter- ǰȱǯǰȱǰȱǯǰȱǰȱǯȱǭȱǰȱǯȱǻŘŖŖŜǼǯȱȱěȱȱ
Simonnet, A.-V., Bossuet, G., Millet, L., Brugiapaglia, E. & emergent aquatic insects on bat foraging in a riparian forest.
Drescher, A. (2007). Vegetation history, climate and human Journal of Animal Ecology 75, 1252-1258.
impact over the last 15,000 years at Lago dell’Accesa (Tuscany, Gafny, S. & Gasith, A. (1999). Spatially and temporally sporadic
Central Italy). Vegetation History and Archaeobotany 16, 279-299. ȱȱ¢ȱȱȱĴȱ£ȱȱȱ ǰȱ
Dukes, J.S. & Mooney, H.A. (1999). Does global change increase Israel: taking advantage of a window of opportunity. Aquatic
the success of biological invaders? Trends in Ecology and Botany 62, 249-267.
Evolution 14, 135-139. Gherardi, F. (2007). ȱǱȱęǰȱȱȱ
ȱ ǯȱ ǭȱ ǰȱ ǯǯȱ ǻŘŖŖŝǼǯȱ ȱ ȱ ȱ ěȱ ȱ ǯȱ Springer, Berlin. 733 pp.
upland stream macroinvertebrates over a 25-year period. Global Giudicelli, J. & Thierry, A. (1998). La faune des mares temporaires,
Change Biology 13, 942-957. son originalité et son intérêt pour la biodiversité des eaux
Durance, I. & Ormerod, S.J. (2009). Trends in water quality continentales méditerranéennes. Ecologia Mediterranea 24, 135-
ȱ ȱ ȱ Ȭȱ ȱ ěȱ ȱ ȱ 143.
macroinvertebrates. Freshwater Biology 54, 388-405. Green, A.J. & Figuerola, J. (2005). Recent advances in the study
European Council (2000). Directive 2000/60/EC of the European of long-distance dispersal of aquatic invertebrates via birds.
Parliament and of the Council of 23 October 2000 establishing Diversity and Distributions 11, 149-156.
ȱ ȱȱ¢ȱȱȱȱęȱȱ ȱ¢ǯȱȱ Green, A.J., Fox, A.D., Hilton, G., Hughes, B., Yarar, M. & Salathé,
Ĝȱȱȱȱȱ L 327ǰȱŗȬŝřǯȱȱĜȱȱ T. (1996). Threats to Burdur Lake ecosystem, Turkey and
Ĝȱȱȱȱȱǰȱǯ its waterbirds, particularly the white-headed duck ¡¢ȱ
ǰȱǯȱǭȱǰȱǯǯȱǻŗşşŗǼǯȱȱ ȱěȱȱ leucocephala. Biological Conservation 76, 241-252.
the sympatric brown trout (ȱĴ) populations of Lough Green, A.J., El Hamzaoui, M., El Agbani, M.A. & Franchimont, J.
Melvin, Ireland. Biological Journal of the Linnean Society 43, 221- (2002a). The conservation status of Moroccan wetlands with
237. particular reference to waterbirds and to changes since 1978.
¢ǰȱ
ǯǰȱ ǰȱ ǯǰȱ
Ĵǰȱ ǯǰȱ ǰȱ ǯǰȱ
¢ǰȱ ǯǰȱ Biological Conservation 104, 71-82.

Green, A.J., Figuerola, J. & Sánchez, M.I. (2002b). Implications of Illies, J. (1979). Annual and seasonal variation of individual
waterbird ecology for the dispersal of aquatic organisms. Acta weights of adult water insects. Aquatic Insects 1, 153-163.
Oecologica – International Journal of Ecology 23, 177-189. IPCC (2007). Summary for Policymakers. In: Climate Change 2007:
Green, A.J., Fuentes, C., Moreno-Ostos, E. & da Silva, S.L.R. (2005). ȱ¢ȱȱǯȱȱȱȱȱ ȱȱȱȱ
ȱĚȱȱȱȱȱȱ ȱȱȱȱȱȱȱȱȱ
in brackish lakes in Eastern Spain. Annales De Limnologie Change (eds S.Q.D. Solomon, M. Manning, Z. Chen, M. Marquis,
– International Journal of Limnology 41, 73-81. K.B. Averyt, M. Tignor & H.L. Miller), pp. 1-22. Cambridge
ǰȱ ǯȬǯǰȱ Ĵǰȱ

ǯǰȱ ǰȱ ǯǰȱ
ǰȱ ǯǯǰȱ University Press, Cambridge, UK.
Jeppesen, E., Kairesalo, T., Luokkanen, E., Nilsson, P.-Å., Irvine, K., Moss, B. & Balls, H. (1989). The loss of submerged
Søndergaard, M. & Strand, J. (1998). Biomanipulation as an ȱ ȱȱǯȱȱȱ ȱęȱȱ
application of food-chain theory: constraints, synthesis and zooplankton in a set of experimental ponds, and conclusions.
recommendations for temperate lakes. Ecosystems 1, 558-574. Freshwater Biology 22, 89-107.
Hawkins, B.A. & Porter, E.E. (2003). Does herbivore diversity Jakob, C., Poizat, G., Veith, M., Seitz, A. & Crivelli, A.J. (2003).
ȱȱȱ¢ǵȱȱȱȱȱȱĴĚǯȱȱ Breeding phenology and larval distribution of amphibians in
American Naturalist 161, 40-49. a Mediterranean pond network with unpredictable hydrology.
Hawkins, B.A., Field, R., Cornell, H.V., Currie, D.J., Guégan, Hydrobiologia 499, 51-61.
ǯǯǰȱ ǰȱ ǯǯǰȱ ǰȱ ǯǯǰȱ Ĵǰȱ ǯ ǯǰȱ ěǰȱ James, C., Fisher, J., Russell, V., Collings, S. & Moss, B. (2005).
T., O’Brien, E.M., Porter, E.E. & Turner, J.R.G. (2003). Energy, Nitrate availability and hydrophyte species richness in shallow
ǰȱȱȬȱȱĴȱȱȱǯȱȱ lakes. Freshwater Biology 50, 1049-1063.
Ecology 84, 3105-3117. £ǰȱǯ
ǯȱǻŗşŞśǼǯȱȱȱȱęĴǯȱȱOikos 45, 308-310.
Hobaek, A. & Weider, L.J. (1999). A circumpolar study of Arctic Jeppesen, E., Søndergaard, M., Kronvang, B., Jensen, J.P., Svendsen,
¢Ǳȱ ¢ȱ Ĵȱ ȱ ȱ ȱ ¡ȱ L.M. & Lauridsen, T.L. (1999). Lake and catchment management
complex. Ambio 28, 245-250. in Denmark. Hydrobiologia 395/396, 419-432.
Hooper, D.U., Chapin, F.S., Ewel, J.J., Hector, A., Inchausti, P., Jeppesen, E., Søndergaard, M., Jensen, J.P., Havens, K.E., Anneville,
Lavorel, S., Lawton, J.H., Lodge, D.M., Loreau, M., Naeem, S., O., Carvalho, L., Coveney, M.F., Deneke, R., Dokulil, M.T., Foy,
Schmid, B., Setälä, H., Symstad, A.J., Vandermeer, J. & Wardle, B., Gerdeaux, D., Hampton, S.E., Hilt, S., Kangur, K., Köhler, J.,
ǯǯȱǻŘŖŖśǼǯȱȱěȱȱ¢ȱȱ¢ȱǱȱ Lammens, E.H.H.R., Lauridsen,T.L., Manca, M., Miracle, M.R.,
a consensus of current knowledge. Ecological Monographs 75, Moss,B., Nõges, P., Persson, G., Phillips, G., Portielje, R., Romo,
3-35. S., Schelske, C.L., Straile, D., Tatrai,I., Willen, É. & Winder,
Howard, A. & Easthope, M.P. (2002). Application of a model to M. (2005). Lake responses to reduced nutrient loading – an
ȱ¢ȱ ȱĴȱȱȱȱȱ analysis of contemporary long-term data from 35 case studies.
change at Farmoor Reservoir, Oxfordshire, UK. Science of the Freshwater Biology 50, 1747-1771.
Total Environment 282, 459-469. ǰȱ ǯǰȱ èǰȱ ǯǰȱ ěǰȱ ǯǰȱ ǰȱ ǯǯȱ
Hutchinson, G.E. (1965). The Ecological Theater and the Evolutionary & Jensen, J.P. (2007). Shallow lake restoration by nutrient
¢ǯ Yale University Press, New Haven. 139 pp. ȱȱȮȱȱȱęȱȱȱǯȱȱ
Ibelings, B.W., Vonk, M., Los, H.F.J., van der Molen, D.T. & Hydrobiologia 584, 239-252.
ħǰȱǯǯȱǻŘŖŖřǼǯȱȱ££¢ȱȱȱ¢ȱȱ Jeppesen, E., Kronvang, B., Meerhof, M., Søndergaard, M.,
waterblooms: validation with NOAA-AVHRR satellite images. Hansen, K.M., Andersen, H.E., Lauridsen, T.L., Liboriussen, L.,
Ecological Applications 13, 1456-1472. Beklioglu, M., Ozen, A. & Olesen, J.E. (2009). Climate change
ǰȱǯȱǻŗşŝśǼǯȱȱȱ ȱĴȱȱȱȱȱȱ ěȱȱěǰȱȱȱǰȱȱȱ
waters. Verhandlungen der Internationalen Vereinigung für state and potential adaptations. Journal of Environmental Quality
theoretische und angewandte Limnologie 19, 1703-1711. (in press).

128 Moss, B. et al
Jeziorski, A., Yan, N.D., Paterson, A.M., DeSellas, A.M., Turner, Response of freshwater microcosm communities to nutrients,
ǯǯǰȱ ěǰȱ ǯǯǰȱ ǰȱ ǯǰȱ ǰȱ ǯǯǰȱ ǰȱ ǯ ǯǰȱ ęǰȱ ȱ ȱ ȱ ȱ ȱ ȱ ǯȱȱ
Palmer, M.E., McIver, K., Arseneau, K., Ginn, B.K., Cumming, Limnology and Oceanography 48, 707-722.
B.F. & Smol, J.P. (2008). The widespread threat of calcium ěǰȱ ǯǰȱ ǰȱ ǯǯǰȱ ¡ȱ ȱ ǰȱ ǯǰȱ ǰȱ ǯǰȱ
decline in fresh waters. Science 322, 1374-1377. Pedersen, A.R. & Jeppesen, E. (2007). Can warm climate-related
Kato, C., Iwata, T., Nakano, S. & Kishi, D. (2003). Dynamics of ȱȱĴȱȱȱ ȱȱȱ ȱ
ȱȱĚ¡ȱěȱȱȱȱ Ȭȱ state in shallow lakes? Global Change Biology 13, 1888-1897.
spiders. Oikos 103, 113-120. Mehner, T., Benndorf, J., Kasprzak, P. & Koschel, R. (2002).
Kawecki, T.J. & Ebert, D. (2004). Conceptual issues in local Biomanipulation of lake ecosystems: successful applications
adaptation. ¢ȱĴ 7, 1225-1241. and expanding complexity in the underlying science. Freshwater
Klaassen, M. & Nolet, B.A. (2007). The role of herbivorous water Biology 47, 2453-2465.
birds in aquatic systems through interactions with aquatic Michener, W.K., Blood, E.R., Bildstein, K.L., Brinson, M.M. &
macrophytes, with special reference to the Bewick’s Swan Gardner, L.R. (1997). Climate change, hurricanes and tropical
– Fennel Pondweed system. Hydrobiologia 584, 205-213. storms, and rising sea level in coastal wetlands. Ecological
Knight, T.M., McCoy, M.W., Chase, J.M., McCoy, K.A. & Holt, R.D. Applications 7, 770-801.
(2005). Trophic cascades across ecosystems. Nature 437, 880- Miroslaw-Grabowska, J. & Niska, M. (2007). Reconstruction of
884. environmental conditions of Eemian palaeolake at Studzieniec
ħà ǰȱǯǰȱ ǰȱǯǯȱǭȱȱǰȱǯȱǻŗşşŖǼǯȱȱ
¢¢Ȭ (Central Poland) on the basis of stable isotope and Cladocera
macroinvertebrate interactions in Zwemlust, a lake undergoing analyses. Quaternary International 162, 195-204.
biomanipulation. Hydrobiologia 200, 467-474. Ĵǰȱ ǯ ǯǰȱǰȱǯǯǰȱǰȱǯǯǰȱ ǰȱ ǯǯǰȱ¢ǰȱ
Lachavanne, J.B. & Juge, R. (1997). Biodiversity in Land-inland H.L., Waide, R.B., Willig, M.R., Dodson, S.I. & Gough, L. (2001).
Water Ecotones (eds J.-B. Lachavanne & R. Juge). Man and the What is the observed relationship between species richness and
Biosphere series. UNESCO, The Parthenon Publishing Group, productivity? Ecology 82, 2381-2396.
Paris. 308 pp. ǰȱǯǯȱǭȱ £ǰȱǯǯȱǻŗşŞŖǼǯȱȱěȱȱȱȱȱȱȱ ȱ
Leopold, A. (c.1938). Conservation. From ‘A Survey of quality and invertebrates of a mountain stream. Water, Air, &
Conservation’. Manuscript. Revised and combined with other ȱȱ14, 187-205.
materials in A Sand County Almanac (1966). Oxford University ħǰȱǯǯǰȱ
ûǰȱǯǰȱȱȱǰȱǯǯǰȱǰȱǯǯǰȱ
Press. 240 pp. Bodelier, P.L.E., Boers, P.C.M., Pires, L.M.D., Gons, H.J., Ibelings,
Lodge, D.M. (1993). Biological invasions: lessons for ecology. B.W., Noordhuis, R., Portielje, R., Wolfstein, K. & Lammens,
Trends in Ecology & Evolution 8, 133-137. E.H.R.R. (2005). The impact of climate change on lakes in the
Lodge, D.M., Stein, R.A., Brown, K.M., Covich, A.P., Brönmark, Netherlands: a review. Aquatic Ecology 39, 381-400.
C., Garvey, J.E. & Klosiewskt, S.P. (1998). Predicting impact of Moss, B. (2008). The Water Framework Directive: total environment
freshwater exotic species on native biodiversity: challenges in or political compromise. Science of the Total Environment 400, 32-
spatial scaling. Austral Ecology 23, 53-67. 41.
Louv, R. (2008). Last Child in the Woods. Algonquin Books of Moss, B., Madgwick, J. & Phillips, G. (1996). A Guide to the
Chapel Hill, Chapel Hill, N.C., USA. 390 pp. ȱ ȱ Ȭȱ ȱ ǯȱ Environment
Maltby, E., Hogan, D.V. & McInnes, R.J. (1996). Functional Analysis Agency, Broads Authority & European Union Life Programme,
ȱ ȱ ȱ ¢ǰȱ ȱ ŗǻǼǯȱ ȱ ¢ȱ Norwich. 179 pp.
ȱǯ European Commission, Luxemburg. 448 pp. Murphy, K.J., Castella, E., Clement, B., Hills, J., Obrdlik, P., Pulford,
ǰȱ ǯǯǰȱ ǰȱ ǯȱ ǭȱ ǰȱ ǯǯȱ ǻŘŖŖŝǼǯȱ ȱ Ğȱ ȱ I.D., Schneider, E. & Speight, M.C.D. (1994). Biotic indicators
phenology of Bythrotrephes longimanus and its modern success of riverine wetland ecosystem functioning. In: Global Wetlands:
in Lake Maggiore as a result of changes in climate and trophy. Old World and New (ed. W.J. Mitsch), pp. 659-682. Elsevier,
ȱȱȱȱ29, 515-525. Amsterdam.
ǰȱ ǯǰȱ
Ĵǰȱ ǯǰȱ ǰȱ ǯǯǰȱ ǰȱ ǯǰȱ ǰȱ ǰȱ ǯǯǰȱ ¢ǰȱ ǯǯǰȱ ǰȱ ǯǯȱ ǭȱ
ęǰȱ ǯǯȱ ǻŘŖŖŘǼǯȱȱ
ǯǰȱ
¢ǰȱ ǯǰȱ ǭȱ ǰȱ ǯȱ ǻŘŖŖŘǼǯȱ ȱ ěȱ ȱ ȱ ęȱǰȱȱȱȱ¢ȱȱ ȱȱ
climate warming on macrophytes in freshwater microcosm riparian ecosystems. Ecosystems 5, 399-417.
communities. Aquatic Botany 74, 71-83. Nielsen, D.L., Brock, M.A., Rees, G.N. & Baldwin, D.S. (2003).
McKee, D., Atkinson, D., Collings, S.E., Eaton, J.W., Gill, A.B., ěȱ ȱ ȱ ¢ȱ ȱ ȱ ¢ȱ ȱ

¢ǰȱǯǰȱ
Ĵǰȱ ǯǰȱ
¢ǰȱǯǰȱǰȱǯȱǭȱǰȱǯȱǻŘŖŖřǼǯȱȱ Australia. Australian Journal of Botany 51, 655-665.

Obach, M., Wagner, R., Werner, H. & Schmidt, H.H. (2001). Montana, USA. Forest Ecology and Management 200, 161-181.
ȱȱ¢ȱȱȱȱ ȱęȱ Ripple, W.J. & Beschta, R.I. (2004b). Wolves and the ecology of fear:
neural networks. Ecological Modelling 146, 207-217. can predation risk structure ecosystems? BioScience 54, 755-766.
Oertli, B., Auderset, J.D., Castella, E., Juge, R., Cambin, D. & Rodrigues, A.S.L. & Brooks, T.M. (2007). Shortcuts for biodiversity
ǰȱǯȬǯȱǻŘŖŖŘǼǯȱȱȱ£ȱĴǵȱȱȱȱ ȱǱȱȱěȱȱǯȱȱAnnual
between pond area and biodiversity. Biological Conservation 104, Review of Ecology Evolution and Systematics 38, 713-737.
59-70. Ç£ǰȱ ǯǰȱ ·ǰȱ ǯȱ ǭȱ ¤£Ȭ¤£ǰȱ ǯȱ ǻŘŖŖřǼǯȱ ȱ Ğȱ
Palmer, M.A., Reidy Liermann, C.A., Nilsson, C., Florke, M., from clear to turbid phase in Lake Chozas (NW Spain) due to
Alcamo, J., Lake, P.S. & Bond, N. (2008). Climate change and ȱȱȱȱȱ ȱ¢ęȱǻȱ
the world’s river basins: anticipating management options. clarkii). Hydrobiologia 506, 421-426.
Frontiers in Ecology and the Environment 6, 117-128. Rouse, W.R., Douglas, M.S.V., Hecky, R.E., Hershey, A.E., Kling,
Pauls, S.U., Lumsch, H.T. & Haase, P. (2006). Phylogeography of G.W., Lesack, L., Marsh, P., McDonald, M., Nicholson, B.J.,
ȱ ȱ Ě¢ȱ Drusius discolor: evidence for multiple ǰȱǯǯȱǭȱǰȱǯǯȱǻŗşşŝǼǯȱȱěȱȱȱȱȱȱ
refugia and periglacial survival. Molecular Ecology 15, 2153- freshwaters of Arctic and subarctic North America. Hydrological
2169. 11, 873-902.
Petchey, O.L. & Gaston, K.J. (2002). Functional diversity (FD), ǰȱ ǯǯǰȱ ȱ ǰȱ ǯǯǰȱǰȱ ǯǯǰȱ ǰȱ ǯǰȱ ęǰȱ
species richness and community composition. ¢ȱĴȱ J., Dirzo,R., Huber-Sanwald, E., Huenneke, L.F., Jackson,
5, 402-411. R.B., Kinzig, A., Leemans, R., Lodge, D.M., Mooney, H.A.,
Petchey, O.L. & Gaston, K.J. (2006). Functional diversity: back to ǰȱǯǰȱěǰȱǯǯǰȱ¢ǰȱǯǯǰȱǰȱǯ
ǯǰȱǰȱǯȱ
basics and looking forward. ¢ȱĴ 9, 741-758. & Wall, D.H. (2000). Global Biodiversity Scenarios for the Year
Petchey, O.L., Evans, K.L., Fishburn, I.S. & Gaston, K.J. (2007). 2100. Science 287, 1770-1774.
Low functional diversity and no redundancy in British avian Sandlund, O.T., Gunnarsson, K., Jónasson, P.M., Jonsson, B.,
assemblages. Journal of Animal Ecology 76, 977-985. Lindem, T., Magnússon, K.P., Malmquist, H.J., Sigurjónsdóttir,
ǰȱǯǰȱ
ǰȱǯǰȱǰȱǯǯǰȱǰȱǯǰȱ+Ćǰȱǯǰȱǰȱ H., Skúlason, S. & Snorrason, S.S. (1992). The Arctic charr,
N. & Schwenk, K. (2008). A taxonomic reappraisal of the Salvelinus alpinus in Thingvallavatn. Oikos 64, 305-351.
European Daphnia longispina complex (Crustacea, Cladocera, ěǰȱǯǰȱ
ǰȱǯ
ǯǰȱħǰȱǯȬǯǰȱǰȱǯȱǭȱǰȱǯȱ
Anomopoda). Zoologica Scripta 37, 507-519. (1993). Alternative equilibria in shallow lakes. Trends in Ecology
ǰȱǯǯǰȱǰȱ ǯǯǰȱ ĴǰȱǯǯȱǭȱǰȱǯǯȱǻŗşşśǼǯȱȱȱ and Evolution 8, 275-279.
future of biodiversity. Science 269, 347-350. ǰȱǯǯȱǻŘŖŖŗǼǯȱȱȱȱěȱȱȱ ȱ
Postel, S.L., Daily, G.C. & Ehrlich, P.R. (1996). Human appropriation and other human stresses on Canadian freshwaters in the new
of renewable fresh water. Science 271, 785-788. millennium. Canadian Journal of Fisheries and Aquatic Sciences 58,
Prowse, T.D., Wrona, F.J., Reist, J.D., Hobbie, J.E., Lévesque, L.M.J. 18-29.
& Vincent, W.F. (2006). General features of the Arctic relevant to Steiger, J.,Tabacchi, E., Dufour, S., Corenblit, D. & Peiry, J.L. (2005).
climate change in freshwater ecosystems. Ambio 35, 330-338.
¢ȱ ȱ ěȱ ȱ ȱ ȱ
Reist, J.D., Wrona, F.J., Prowse, T.D., Power, M., Dempson, J.B., ȱ ȬĚȱ ȱ ¢Ǳȱ ȱ ȱ ȱ ȱ
ǰȱ ǯǯȱ ǭȱ ǰȱ ǯǯȱ ǻŘŖŖŜǼǯȱ ȱ ȱ ȱ ȱ ěȱ ȱ temperate zone. River Research and Management 21, 689-691.
climate change on selected Arctic freshwater and anadromous Stevens, C.J., Dise, N.B., Mountford, J.O. & Gowing, D.J. (2004).
ęǯȱȱAmbio 35, 381-387. Impact of nitrogen deposition on the species richness of
Reynolds, C.S. (1997). ȱȱȱȱǱȱȱȱȱ grasslands. Science 303, 1876-1879.
¢ȱ¢ǯȱ Excellence in Ecology, Book 9, ed. O. Kinne. Straile, D. (2000). Meteorological forcing of plankton dynamics in
International Ecology Institute, Oldendorf/Luhe, Germany. 371 a large and deep continental European lake. Oecologia 122, 44-
pp. 50.
Reynolds, C.S. (2000). Hydroecology of river plankton: the role of ǰȱǯǯȱǻŗşşŞǼǯȱȱȱȱĚ¡¢ȱȱȱȱ
¢ȱȱȱĚ ǯȱȱ
¢ȱȱ14, 3119-3132. migration systems. Journal of Avian Biology 29, 441-446.
Reynoldson, T.B. (1983). The population biology of Turbellaria Tilman, D. (2001). Functional diversity. In: Encyclopedia of
with special reference to the freshwater triclads of the British Biodiversity (ed. S.A. Levin), pp. 109-120. Academic Press, San
Isles. Advances in Ecological Research 13, 235-326. Diego, CA.
Ripple, W.J. & Beschta, R.I. (2004a). Wolves, elk, willows, and Tilman, D., Wedin, D. & Knops, J. (1996). Productivity and
trophic cascades in the upper Gallantin Range of Southwestern ¢ȱ Ěȱ ¢ȱ ¢ȱ ȱ ȱ

130 Moss, B. et al
ecosystems. Nature 379, 718-720. understanding of phytoplankton ecology? Freshwater Biology

Törnlund, E. & Östlund, L. (2002). Floating timber in northern 48, 1669-1675.
ǯȱȱȱȱȱĚ ¢ȱȱȱȱ Ĵǰȱ ǯ
ǯȱ ǻŗşŝŘǼǯȱ ȱ ȱ ȱ ȱ ȱ ȱ
rivers. Environment and History 88, 85-106. diversity. ¡ 21, 213-251.
Torsvik, V., Øvreås, L. & Thingstad, T.F. (2002). Prokaryotic Wilkinson, D.M. (2004). The parable of Green Mountain,
diversity – magnitude, dynamics, and controlling factors, ȱǰȱ¢ȱȱȱȱęĴǯȱȱ
Science 296, 1064-1066. Journal of Biogeography 31, 1-4.
Uhlmann, D. (1980). Stability and multiple steady states of Wilson, J.B. (1999). Guilds, functional types and ecological groups.
hypereutrophic ecosystems. In: Hypertrophic Ecosystems, Oikos 86, 507-522.
ȱȱ
¢¢ȱǯȱŘ (eds J.Barica & L. Mur), pp. Winder, M. & Schindler, D.E. (2004). Climate change uncouples
235-248. Dr W. Junk. trophic interactions in an aquatic ecosystem. Ecology 85, 2100-
Urabe, H. & Nakano, S. (1998). Contribution of woody debris 2106.
ȱ ȱ ȱ ęȱ ȱ ȱ ȱ ȱ ¢ȱ ęǰȱǯǯǰȱǰȱǯǯȱǭȱǰȱǯǯȱǻŘŖŖŞǼǯȱȱȱȱȱ
deciduous forest, northern Japan. Ecological Research 13, 335- (Salvelinus alpinus) populations of Windermere, UK: population
345. trends associated with eutrophication, climate change and
Valdovinos, C., Moya, C., Olmos, V., Parra, O., Karrasch, B. & increased abundance of roach (Rutilus rutilus). Environmental
ĴǰȱǯȱǻŘŖŖŝǼǯȱȱȱȱȱ ȬȱĚȱȱ Biology of Fishes 83, 25-35.
the conservation of shallow water benthic macroinvertebrates: Zwick, P. (1992). Stream habitat fragmentation – a threat to
an example in the Andean zone of Chile. Biodiversity & biodiversity. Biodiversity and Conservation 1, 80-97.
Conservation 16, 3095-3109.
ȱǰȱǯȱǭȱ ǰȱǯǯȱǻŗşşŖǼǯȱȱȱěȱȱȬ
limited and phosphorus-limited growth and competitive
interactions among three diatoms. ȱȱ¢¢ 26, 40-
50.
van Donk, E., Grimm, M.P., Gulati, R.D. & Klein Breteler, J.P.G.
(1990). Whole-lake food-web manipulation as a means to study
community interactions in a small ecosystem. Hydrobiologia
200/201, 275-289. ȱę
van Geest, G.J., Wolters, H., Roozen, F.C.J.M., Coops, H.,
Brian Moss: Fresh waters have always been a major
ħǰȱǯǯǯǰȱǰȱǯǯȱǭȱěǰȱǯȱǻŘŖŖśǼǯȱȱȬ
part of Brian’s life, since learning about the dramatic
ȱ Ěȱ ěȱ ¢ȱ ȱ ȱ Ěȱ
lakes. Hydrobiologia 539, 239-248. story of how lakes stratify in his undergraduate
Vitousek, P.M., Mooney, H.A., Lubchenco, J. & Melillo, J.M. (1997). career nearly half a century ago, and the stimulation
Human domination of Earth’s ecosystems. Science 277, 494- of visiting, as a postgraduate, the enormously
499. ¢ȱȱ¢ȱȱǯȱĞȱ
ǰȱǯǯǰȱǰȱǯǯǰȱǰȱǯǯǰȱĴǰȱ ǯǰȱ ǰȱǯǯǰȱ
he has carried out freshwater research in Africa,
Dodson, S.I., Juday, G.P. & Parmenter, R. (1999). The relationship
the United States and Europe and has taught
between productivity and species richness. Annual Review of
Ecology and Systematics 30, 257-300. freshwater science on almost all the continents. The
Weider, L.J. & Hobaek, A. (2000). Phylogeography and arctic ȱ ȱ ȱ ¢ȱ ǰȱ ȱ ęȱ
biodiversity: a review. Annales Zoologici Fennici 37, 217-231. observations to experiments in the lab, in mesocosms,
Weider, L.J. & Hobaek, A. (2003). Glacial refugia, haplotype experimental tanks and whole lakes and has centred
distributions, and clonal richness of the ȱ¡ complex
ȱǰȱȱȱȱȱěȱȱ
in arctic Canada. Molecular Ecology 12, 463-473.
climate change. He is presently Holbrook Gaskell
Weisse, T. (2006). Biodiversity of freshwater microorganisms
– achievements, problems, and perspectives. ȱȱȱ Professor of Botany at the University of Liverpool,
Ecology 54, 633-652. UK and President of the International Society for
ěǰȱ ǯȱ ǻŘŖŖřǼǯȱ ȱ ȱ ȱ ȱ ȁȱ ȱ ¢Ȃȱ ¢ǯȱȱȱȱȱ¢ȱĴȱ ȱȱ
and ‘functional diversity’ in lake phytoplankton – a new muddy, he plays the double bass and writes poems.

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Climate Change and the Future of Freshwater Biodiversity in Europe: A

Primer for Policy-Makers

Climate change and the future of freshwater biodiversity

Received 30 January 2009; accepted 26 May 2009; published 14 September 2009

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

is a gradient from ombrotrophic

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

generating energy such as

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

ȱȱȱȱ¢ǰȱ ȱȱĞȱȬȱ Depositional rivers

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

Deep lakes membranes and interference with reproduction, and to

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

lakes may prove more resilient. Where lakes contain a ȱ ȱȱȱȱȱĴǰȱȱȱ

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

that the major changes will

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

Present major impacts ěȱȱ  Mitigation

© Freshwater Biological Association 2009

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¡ȱĞ ȱȱȱ protect downstream systems.

Freshwater Reviews (2009) 2, pp. 103-130

zones eutrophication problems. Full enactment of

Present major impacts ěȱȱ  Mitigation

© Freshwater Biological Association 2009

References composition of Espolla pond (NE Iberian peninsula): the

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

ǰȱ ǯȬǯǰȱ Ĵǰȱ

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

DOI: 10.1608/FRJ-2.2.1 Freshwater Reviews (2009) 2, pp. 103-130

ecosystems. Nature 379, 718-720. understanding of phytoplankton ecology? Freshwater Biology

© Freshwater Biological Association 2009 DOI: 10.1608/FRJ-2.2.1

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Present major impacts ěȱȱ Mitigation

Present major impacts ěȱȱ Mitigation

¡ȱĞ ȱȱȱ protect downstream systems.

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