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Home , Community (ecology), Ecosystem ecology

and and ecology provide two different perspectives on ecological , their structure, their functioning, their dynamics and their . While population and community ecology takes as its starting point the population and its interactions with other , is mainly concerned with the flows of matter and in the overall composed of biological and their abiotic environment. Linking Community and Ecosystem Ecology (LINKECOL) However, populations and do An ESF scientific programme not exist in isolation; they are parts of , and, as such, they are subjected to constraints arising from ecosystem functioning, in particular energy dissipation and cycling. At the same time, ecosystems do not exist without their biologi- cal components; the latter impose their own constraints on ecosystem processes, as the disruptions generated by some biological invasions attest. And in the face of the growing threat of a massive loss of biologi- cal diversity, interest is increasing concern- ing the role of in ecosystem processes. This has created an urgent need to integrate the two subdisciplines and it is the principal aim of the LINKECOL programme to unify these different perspectives. Such integration is essential not only to advance our fundamental understanding of natural and managed ecosystems but also to provide answers to more applied questions such as the impacts of biodiversity loss or The European invasions on ecosystem sustainability. Foundation acts as a catalyst for the development of science by bringing together leading scientists and funding agencies to debate, plan and implement pan-European initiatives. 1 This separation of the two subdisci- plines is understandable as they partly address issues at different hierarchical levels and different spatial and temporal scales. But it is harmful insofar as it is an obstacle to their unity and mutual enrichment. Populations and communities do not exist in isolation; they are parts of ecosystems, and, as such, they are subjected to constraints arising from ecosystem functioning, in particular energy dissipation and nutrient cycling. These constraints can deeply alter the nature of species interac- tions and community properties such Scientific background as food-web stability. On the other hand, ecosystems do not exist with- The need for integration of popula- out their biological components; the tion/community and ecosystem latter impose their own constraints ecology on ecosystem processes, as the The vigorous growth in ecology from disruptions generated by some its origins in the early years of the biological invasions attest. In the face 20th century has been accompanied of the growing threat of a massive by the creation of numerous subdis- loss of biological diversity, interest is ciplines. Although specialisation may increasing concerning the role of be inevitable, it also creates problems biodiversity in ecosystem processes. because conceptual frameworks in There is today an urgent need for different subdisciplines often diverge integration of the two subdisciplines. over time. This is nowhere more Such an integration is already emerg- apparent than between two of the ing on a world-wide scale as well as in major subdisciplines of ecology: Europe, as a result of both basic and population and community ecology applied scientific questions: How will on the one hand and ecosystem changes in species and genetic ecology on the other. These two diversity affect ecosystem processes subdisciplines have grown largely and the related services they provide independently, each having its own to humankind? How will these concepts, theories and methodolo- changes affect the stability of ecosys- gies. Population and community tems, and their ability to withstand ecology is mainly concerned with the natural and anthropogenic perturba- dynamics, evolution, diversity and tions? How do biological organisms complexity of the biological compo- and abiotic factors interact to regu- nents of ecosystems; its starting late the flow of energy, the structure point is the population and its of the and the cycling of interactions with other populations. chemical elements in aquatic and Ecosystem ecology is mainly con- terrestrial ecosystems? On a large cerned with the functioning of the time scale, are , and overall system composed of biologi- micro-organisms involved in mutu- cal organisms and their abiotic alistic relationships mediated by environment; its starting point is the material cycling despite their appar- flow of matter or energy among ent immediate antagonism? Can functional compartments.

2 indirect interactions evolve by and significantly influence species traits? These are some examples of the questions that have recently received growing attention. Both population, community and ecosystem ecology have a long history in Europe, but, as mentioned above, these subdisciplines have had largely separate developments, with different strengths in different countries and laboratories. Attempts towards the goal of integration of the subdisciplines are developing rapidly through new experimental knowledge and expertise, and attract The effects of species and theoretical approaches, but they new groups and researchers to the diversity on ecosystem processes are still dispersed, both scientifically field. It will also provide a European are investigated using field experiments in European and geographically. European ecology contribution to the Diversitas and grasslands. would greatly benefit from a co- IGBP-GCTE international © N. Buchmann ordinated effort to stimulate ex- programmes. The programme will be changes of ideas, of new theoretical centred on the three main scientific insights, of new experimental themes: systems, and of researchers among 1. and ecosystem European countries. processes; 2. Ecology and evolution of indirect effects in ecosystems; The ESF programme 3. Biological invasions and ecosystem processes.

The present programme aims at 1. Species diversity and ecosystem unifying the perspectives of commu- processes nity and ecosystems ecology. This is Until recently little attention has an important scientific challenge, been given to the ecological conse- both to improve our fundamental quences of the losses of biodiversity. understanding of natural and man- Most concerns over biodiversity aged ecosystems and to provide losses have focused on the important appropriate answers to more applied ethical and aesthetic dimensions of questions such as the impacts of biotic impoverishment or on the biodiversity loss or species invasions losses of potentially economically on ecosystem sustainability. The goal valuable crops or drugs. There is now of the programme is to promote the growing recognition, however, that development and integration of ecosystems operate in a manner that research linking community and provides free ecological services to ecosystem ecology across Europe. . These services are derived The programme will strengthen from the normal functioning of interactions among groups from ecosystems, raising the important different countries and different question whether depauperate scientific traditions, stimulate the ecosystems perform differently or development of ideas, support the less efficiently than the more species- initiation of collaborative research rich systems from which they are projects, facilitate the exchange of derived.

3 cycling, thus affecting ecosystem functioning. This part of LINKECOL has the following aims: . To synthesise current experimental and theoretical advances in the area of the effect of biodiversity on the magnitude and stability of ecosystem processes. . To develop innovative approaches in this area. . To contribute to an understanding of the mechanistic basis of the response – or lack of response – of ecosystem processes to species diversity. The loss of plant Several hypotheses have been pro- To investigate the functional species diversity posed on this topic, ranging from one . properties, at the aggregated ecosys- reduces primary extreme stating that all species are production in tem level, of mechanistic theories of unique in their contribution to European interspecific and coex- ecosystem performance to the other grasslands istence based on consump- (Hector et al., extreme stating that most species in a tion, space occupancy or response to Science functional group are redundant in 286:1123- disturbances. their ecosystem impact. Another 1127, 1999). possibility is the “insurance hypoth- . To investigate, both theoretically esis” : High may not and experimentally, the interplay always play a significant role in between ecosystem processes and maintaining ecosystem processes community assembly. under constant or benign environ- To examine how environmental mental conditions, but it may . changes interact with changes in nevertheless be important for main- biological diversity in determining taining their stability under chang- ecosystem processes. ing environmental conditions. Global or more local environmental 2. Ecology and evolution of indirect changes such as increased atmo- effects in ecosystems Interactions between organisms and spheric CO2 concentration or N deposition are likely to interact with between organisms and their abiotic changes in biodiversity and ecosys- environment are not only direct, but tem processes. These environmental they also have a great number of changes may have profound effects indirect effects on other organisms on ecosystem processes such as and processes. Some of these indirect , acidification and effects have a considerable impact on nutrient mineralisation, which affect the functioning of ecosystems. Well- the competitive balance between known examples are keystone preda- plant species and result in changes in tion, indirect mutualisms between species diversity and composition. plants and predators, or indirect These changes in species diversity facilitation by ecosystem engineers. and composition in turn may have important impacts on soil organic matter dynamics and nutrient

4 Reciprocally, some ecosystem pro- . To understand the relationship cesses mediate significant indirect between the spatial scale considered, effects among organisms and thereby the spatial heterogeneity of the modify the nature of species interac- environment and species interac- tions. In particular, nutrient cycling tions, and the extent and magnitude plays a crucial role as a circular causal of indirect effects. pathway which transmits indirect . To develop new theories and effects to all ecosystem components. experiments on the evolutionary A clear if controversial example of significance of indirect effects in this is to be found in plant—herbi- ecosystems. vore interactions. While the direct trophic interaction between plants and is negative for the plants, the positive indirect effect of herbivory due to increased nutrient cycling by the herbivores may result in a net gain in productivity for the plants, thus generating an indirect between plants and herbivores. Another example con- cerns indirect effects generated by differential recycling of mineral elements due to differences in elemental constraints (C/N/P ratios) 3. Biological invasions and among trophic levels. ecosystem processes Many indirect effects in ecosystems There has recently been increased are still poorly understood, both attention on the magnitude and theoretically and experimentally. Yet impact of biological invasions. Alien they may be critical for our under- represent a serious standing of the long-term impact of problem internationally affecting species losses. Their evolutionary health, agricultural potential, implications may also be consider- biodiversity, and the structure and able, but the evolution of indirect functioning of natural ecosystems. interactions is an area of evolution- This problem will probably become ary that poses complex more acute with increasing global problems and is only starting to trade, global environmental change receive some attention. Research into and changing land use patterns. this area may well drastically change The ecological and economic conse- our understanding of the evolution quences of invasives are considerable. of species under natural conditions. Biological invasions by exotic species Some aims of this sub-programme clearly alter the composition and are: community structure of invaded areas. They often lead to irreversible

. To identify indirect effects gener- species extinctions. As a matter of ated by species interactions that have fact, invasive species are considered a consistently significant impact on to be the second greatest threat to Mesocosm experiments ecosystem processes. show the importance biodiversity globally ( destruc- of indirect effects in . To develop new theories and tion being the first). They can result the response of experiments on indirect effects in enormous short-term economic lake ecosystems to mediated by material cycling or other losses within managed ecosystems pertubations such as (crops, in particular). Their control nutrient enrichment in ecosystem processes. France. © Lacroix

5 using chemicals or physical measures quences are likely to be. In addition, often has environmentally damaging invasive species provide a good model consequences, with longer-term for integrating ecophysiology, economic impacts. , community ecology and ecosystem ecology. They There is increasing evidence that can reveal the ecosystem impact of invading species can alter properties individual species, as well as their of whole ecosystems, including indirect effects on other species and productivity, nutrient cycling, and community processes. soil fertility. For example, a preda- ceous flatworm from New Zealand is Primary aims of this part of currently spreading in Scotland and LINKECOL are: Ireland. It preys preferentially on . To identify the ecological at- earthworms, thus indirectly threat- tributes of invasive species that ening soil structure and fertility, contribute to the success or failure of processes, and bird their invasion in an ecosystem. communities. Exotic grasses that have invaded semi-arid shrublands . To investigate, theoretically and and woodlands have increased the experimentally, the properties of probability and severity of fires in communities and ecosystems that many areas. make them more resistant or resilient to invasions. We still have limited understanding of what makes species successful . To develop comparative investiga- potential invaders, what increases the tions of the of of communities or ecosys- invasive organisms in their native tems to invasions, and what impacts and exotic areas, and identify re- biological invasions have on commu- search protocols for such investiga- nities and ecosystems. Such an tions. understanding would allow us to . To develop the ability to predict the predict when and where biological community and ecosystem impacts invasions are more likely to occur, of biological invasions. and what their ecological conse-

Activities

Workshops The programme will fund workshops dealing with aspects of the scientific themes. Some workshop themes will be suggested by the steering commit- tee, but there will also be funding available for other workshops on themes suggested by individuals in the participating countries. Calls for proposals for workshops will be made annually (October 1999, March 2000). In 2000 two workshops are scheduled:

6 . Elemental constraints on energy transfer in food webs . Biodiversity and ecosystem func- tioning Workshops will be attended by about 40—50 scientists with a significant proportion of young researchers. They will be open to researchers from all European countries. The Scien- tific Steering Committee will also promote smaller workshops on newly emerging research topics.

Exchange visits The exchange visits scheme will support working visits of up to 6 complementary skills from different The functional significance months by young or leading countries. of forest species diversity is researchers visiting laboratories explored with a canopy Applications will be granted on the crane in Switzerland. outside their home country to work basis of letters of reference, past © C. Körner on collaborative research on one of performance, quality of the project the research themes of the proposal and compatibility with the programme. Exchange visits are scientific themes of the programme. important to increase mobility Details are available on the between European centres, promote programme’s web site the coherence of research in Europe, (http://www.esf.org/linkecol). and bring together experts with

Funding

ESF scientific programmes are The Netherlands; Norges principally financed by the Forskningsråd, Norway; Polska Foundation’s Member Organisations Akademia Nauk, Poland; Instituto de on an à la carte basis. LINKECOL is Cooperação Ciêntifica e Tecnológica supported by: Internacional, Portugal; Comisión Fonds National de la Recherche Interministerial de Ciencia y Scientifique, Fonds voor Tecnología – Oficina de Ciencia y Wetenschappelikjk Onderzoek - Tecnología, Consejo Superior de Vlaandern, Belgium; Akademie ved Investigaciones Científicas, Spain; Cesk˘ é republiky, Grantová agentura Naturvetenskapliga Forskningsrådet, Cesk˘ é republiky, Czech Republic; Sweden; Schweizerischer Statens Naturvidens-kabelige Nationalfonds zur Förderung der Forskningsråd, Denmark; Suomen wissenschaftlichen Forschung/Fonds Akatemia/Finlands Akademi, Fin- National Suisse de la Recherche land; Centre National de la Recher- Scientifique, Switzerland; Natural che Scientifique, France; Hermann Environment Research Council, von Helmholtz-Gemeinschaft United Kingdom. Deutscher Forschungszentren, Germany; Nederlandse organisatie voor Wetenschappelijk onderzoek,

7 LINKECOL Steering Committee

Professor Michel Loreau (Chairman) Professor Dag O. Hessen Alternate members: École Normale Supérieure University of Oslo Laboratoire d’Écologie Department of Biology Professor Ernst-Detlef Schulze UMR 7625 Div. Limnology Dr. Nina Buchmann 46 rue d’Ulm Box 1027 Blindern Max-Planck-Institute for 75230 Paris Cedex 05 0316 Oslo France Norway Postfach 10 01 64 Tel: +47 22 85 45 53 07701 Jena Tel: +33 1 44 32 37 09 Germany Fax: +33 1 44 32 38 85 Fax: +47 22 85 44 38 E-mail: [email protected] E-mail: [email protected] Tel: +49 3641 643721 Fax: +49 3641 643710 E-mail: [email protected] Professor Jan Bengtsson Professor Guy Josens Dept. Ecology and Crop Université Libre de Bruxelles Production Science Laboratoire d’Écologie Animale Professor January Weiner CP 160/13 Dept. of Ecosystem Studies Swedish University of Institute of Environmental Agricultural Sciences Avenue F.D. Roosevelt 50 1050 Bruxelles Jagellonian University Box 7043 Ul. Ingardena 6 750 07 Uppsala Belgium Tel: +32 2 650 2259 30 060 Krakow Sweden Poland Tel: +46 18 67 15 16 Fax: +32 2 650 2231 Tel: +48 12 633 6377 ext. 2447/2618 Fax: +46 18 673430 E-mail: [email protected] Fax: +48 12 6341 978 E-mail: [email protected] Professor Seppo Kellomäki E-mail: [email protected] Professor Frank Berendse University of Joensuu Wageningen Agricultural University Faculty of Dr. Annette Moth-Wiklund Department of Environmental Sciences PO Box 111 ESF Senior Scientific Secretary Nature Conservation and 80101 Joensuu Finland Mrs. Catherine Lobstein Group ESF Administrative Assistant Bornsesteeg 69 Tel: +358 13 2513630 6708 PD Wageningen Fax: +358 13 2514444 European Science Foundation The Netherlands E-mail: [email protected] 1 quai Lezay-Marnésia 67080 Strasbourg Cedex Tel: +31 317 484973 Fax: +31 317 484845 Professor Christian Körner France E-mail: [email protected] University of Basel WWW Home Page: Institute of http://www.esf.org Professor Michael J. Crawley Schonbeinstrasse 6 Tel: +33 (0)3 88 76 71 30 Department of Biology 4056 Basel Fax: +33 (0)3 88 37 05 32 Imperial College at Silwood Park Switzerland E-mail: [email protected] Ascot SL5 7PY Tel: +41 61 267 35 10 United Kingdom Fax: +41 61 267 35 04 Tel: +44 1344 623 911 E-mail: [email protected] For the latest information on Fax: +44 1344 294 339 this programme consult the E-mail: [email protected] Dr. Jens Mogens Olesen LINKECOL home page: Department of Ecology and http://www.esf.org/linkecol Professor Carlos M. Herrera Institute of Biological Sciences Estación Biológica de Doñana Aarhus University CSIC Building 540 Cover picture: © C. Körner Avenida Maria Luisa s/n Ny Munkegade Pabellón del Perú 8000 Aarhus C 41013 Sevilla Denmark Spain Tel: +45 89 42 32 67 Tel: +34 95 423 23 40 Fax: +45 86 12 71 91 Fax: +34 95 462 11 25 E-mail: [email protected] E-mail: [email protected]

Professor Joao S. Pereira ordre 000052

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