View source for A review of biodiversity-ecosystem function research


{{Review
|name= Norbert Dankers
|AuthorID=10939
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Human needs and actions have, and will continue to, extensively alter ecosystems and [[biodiversity]] on a global scale <ref name = "Vitousek"> Vitousek, P.M., Mooney, H.A., Lubchenco, J. and Melillo, J.M. (1997). Human domination of earth’s ecosystems. ''Science'' 277: 494-499</ref>. Predictions of changes in biodiversity, not only in marine, but also terrestrial and freshwater ecosystems <ref name = "Sala"> Sala, O.E., Chapin, F.S., Armesto, J.J., Berlow, E., Bloomfield, J., Dirzo, R., Huber-Sanwald, E., Huenneke, L.F., Jackson, R.B., Kinzig, A., Leemans, R., Lodge, D.M., Mooney, H.A., Oesterheld, M., Poff, N.L., Sykes, M.T., Walker, B.H., Walker, M. and Wall, D.H. (2000). Biodiversity - Global biodiversity scenarios for the year 2100. ''Science'' 287: 1770-1774. </ref>, have raised substantial concern over the consequences of biodiversity loss on ecosystem processes and [[ecosystem function]], which subsequently affect the provision of ecosystem goods and services, and ultimately affect human well-being <ref name = "Diaz"> Diaz, D., Fargione, J., Chapin, F.S. III and Tilman, D. (2006). Biodiversity loss threatens human well-being. ''PLOS Biology'' 4: 1300-1305.</ref>.
 
Since the early 1990’s a portfolio of evidence obtained from the development of theory, laboratory experiments, field experiments and observational studies has shown that, irrespective of the system under study, increasing biodiversity tends to have positive effects on ecosystem properties, although the pattern of response may vary depending on the [[ecosystem]] and species investigated.


[[Image:Flow diagramm.jpg|thumb|right|450px|Figure 1: Summary of research approaches adopted to address the relationship between biodiversity and ecosystem function in the peer-reviewed scientific literature. Yellow boxes represent theoretical studies, green boxes represent experimental manipulations of species diversity, red boxes represent the observational studies and the purple boxes represent those studies in which BEF concepts are linked to, or applied, in the real world. Modified from Godbold, J.A. (2008). Marine benthic biodiversity-ecosystem function relations in complex systems. Ph.D. Thesis, University of Aberdeen.]]

== Emergence of a new paradigm ==
 
The central thesis that guided early community ecology saw patterns in the distribution and abundance of species merely as an expression of the [[abiotic]] (chemical and physical) and [[biotic]] (species interactions) components of the environment, giving a predictive understanding of species distribution and abundance within ecosystems <ref name="Pearson"> Pearson, T.H. and Rosenberg, R. (1978). Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. ''Oceanography and Marine Biology: An Annual Review'' 16: 229-311</ref>. In the early 1990’s, however, an increasing number of ecologists began to challenge this view and, instead, started to examine, how ecosystem properties are mediated by the biota <ref name="Schulze"> Schulze, E.D. and Mooney, J.A. (1993). Biodiversity and ecosystem function. Springer-Verlag, Berlin</ref>. A wide range of hypotheses were developed describing the form of the biodiversity-ecosystem function relationship and which collectively formed a framework within which this relationship could be tested experimentally <ref name="Schmid 2002"> Schmid, B., Hector, A. and Huston, M.A. (2002). The design and anlalysis of biodiversity experiments. In Loreau, M., Naeem, S. and Inchausti, P. (eds) Biodiversity and ecosystem functioning: Synthesis and perspectives. Oxford University Press. pp. 61-75.</ref>. 

In a series of phases, biodiversity-ecosystem function research has steadily improved to make experimental designs and model predictions more realistic (Figure 1). The timing of publications from each phase shows that, although different approaches have been used within the biodiversity-ecosystem function framework almost since the first influential paper was published in 1994 <ref name="Naeem 1994"> Naeem, S., Thompson, L.J., Lawler, S.P., Lawton, J.H. and Woodfin, R.M. (1994). Declining biodiversity can alter the performance of ecosystems.  ''Nature'' 368: 734-736.</ref>, there has been a general trend for experimental and theoretical studies to incorporate more natural environmental complexities (Reality filter, Figure 1).

== Phase 1 - Laboratory experiments ==
Initially, biodiversity-ecosystem function relationships were investigated by manipulating biodiversity under controlled conditions in the laboratory (Figure 2a, b) <ref name = "Covich"> Covich, A.P., Austen, M.C., Barlöcher, F., Chauvet, E., Cardinale, B.J., Biles, C.L., Inchausti, P., Dangles, O., Solan, M., Gessner, M.O., Statzner, B. and Moss, B. (2004). The role of Biodiversity in the functioning of freshwater and marine benthic ecosystems. ''BioScience'' 54: 767-775.</ref> <ref name = "Hooper"> Hooper, D.U., Chapin, F.S. III, Ewel, J.J., Hector, A., Inchausti, P., Lavorel, S., Lawton, J.H., Lodge, D.M., Loreau, M., Naeem, S., Schmid, B., Setala, H., Symstad, A.J., Vandermeer, J., Wardle, D.A. (2005). Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. ''Ecological Monographs'' 75: 3 - 35.</ref>. In such studies, simple, single species communities are randomly assembled and their effects on ecosystem function determined. Then diversity is increased by constructing multi-species assemblages comprising the single species that have already been characterised and the effects of these multi-species assemblages on ecosystem function is determined (Figure 2c). If the observed response of the multi-specific assemblages differs from the response predicted by summation of the single species responses, then it is concluded that diversity has had an effect.

 [[Image:Figure 2a.jpg|thumb|left|350px|Figure 2a: Mesocosms used to investigate the effect of invertebrate biodiversity loss on sediment nutrient release, e.g. in Emmerson ''et al.'' (2001). Consistent patterns and the idiosyncratic effects of biodiversity in marine ecosystems. ''Nature'' 411: 73-77]]
 [[Image:Figure 2b.jpg|thumb|350px|none|Figure 2b: Mesocosms used to investigate the effect of biodiversity loss on bioturbation and nutrient release, e.g. in Ieno ''et al''. (2006). How biodiversity affects ecosystem functioning: roles of infaunal species richness, identity and density in the marine benthos. ''Marine Ecology Progress Series'' 311: 263-271.]]  


Although these experiments successfully articulated biodiversity-ecosystem function hypotheses and allowed for the unambiguous interpretation of cause-effect relationships, critics were quick to assert that such studies lack realism because they tend to only include a few species (representing only a subset of the total community), often only from one trophic level, and they invariably assume that species loss is random. In addition, these types of experiments attracted further criticism because ecosystem function is measured infrequently and in the absence of the appropriate environmental context, thus making applicability to the real world questionable <ref name = "Srivastava"> Srivastava, D.S. and Vellend, M. (2005). Biodiervsity-ecosystem function research: is it relevant to conservation? ''Annual review of Ecology, Evolution and Systematics'' 36: 267-294.</ref>.

[[Image:Figure 2c.jpg|thumb|right|450px|Figure 2c: Experimental design commonly adopted in biodiversity-ecosystem function experiments. Red boxes indicate treatments with one species (Single species), two species and three species (Species mixtures). Total biomass between treatments is kept constant.]]

Despite debates over experimental designs and applicability, syntheses of the available evidence suggest that, irrespective of the system studied, increased biodiversity tends to have a positive effect on ecosystem properties, such as [[primary production]], [[nutrient]] flux and [[decomposition]], but the pattern of response varies depending on the ecosystem and species investigated <ref name = "Balvanera"> Balvanera, P., Pfisterer, A.B., Buchmann, N., He, J.S., Nakashizuka, T., Raffaelli, D. and Schmid, B. (2006). Quantifying the evidence for biodiversity effects on ecosystem functioning and services. ''Ecology Letters'' 9: 1146-1156.</ref> <ref name = "Cardinale"> Cardinale, B.J., Srivastava, D.S., Duffy, J.E., Wright, J.P., Downing, A.L., Sankaran, M., and Jouseau, C. (2006). Effects of biodiversity on the functioning of trophic groups and ecosystems. ''Nature'' 443: 989-992.</ref>. This variability between results suggests that ecosystems respond differently to biodiversity loss, as natural ecosystems are complex, open systems that are composed of interconnected gradients, patches and networks between, and within which, organisms move and interact <ref name = "Levinton"> Levinton, J. and Kelaher, B. (2004). Opposing forces of deposit-feeding marine communities. ''Journal of Experimental Marine Biology and Ecology'' 300: 65-82.</ref>. The high degree of control required in experiments and the short time periods under which they are conducted means, however, that the environmental variation and biological interactions that occur in natural systems are largely controlled for. The inherent complexity of environmental systems resulted in increasing calls to incorporate more environmental realism into experimental designs <ref name ="Hooper"/> <ref name = "Naeem 2008"> Naeem, S. (2008). Advancing realism in biodiversity research. ''Trends in Ecology and Evolution''. 23: 414-416.</ref> as it is still unclear whether the currently observed patterns in the biodiversity-ecosystem function relationship will hold, for example, for realistic extinction scenarios, in multi-trophic communities, and over larger spatial and temporal scales <ref name ="Srivastava"/> <ref name = "Raffaelli"> Raffaelli, D.G. (2006). Biodiversity and ecosystem functioning: issues of scale and trophic complexity. ''Marine Ecology Progress Series 311:285-294''.</ref>.

== Phase 2 – Inclusion of a subset of environmental variation ==
 [[Image: BEF_Seagrass.jpg|thumb|left|250px| Figure 3: Outdoor mesocosms used in experiments investigating the effects of gazer diversity on ecosystem functioning in seagrass beds, e.g. in Duffy ''et al''.(2003). Grazer diversity effects on ecosystem functioning in seagrass beds. ''Ecology Letters'' 6: 637-645.]] 

Diversity manipulations in outdoor mesocosms Figure 3 <ref name = "Duffy"> Duffy, J.E., Richardson, J.P. and Canuel, E.A. (2003). Grazer diversity effects on ecosystem functioning in seagrass beds. ''Ecology Letters'' 6: 637-645.</ref> are subjected to natural fluctuations in light, temperature and rainfall and thus represent better analogues of natural systems than more controlled laboratory conditions. 

In-situ experiments or species addition/removal field experiments have been particularly valuable for determining biodiversity effects under naturally fluctuating environmental conditions (Figure 1)<ref name = "O'Connor"> O’Connor, N.E. and Crowe, T.P. (2005). Biodiversity loss and ecosystem functioning: Distinguishing between number and identity of species. ''Ecology'' 86: 1783-1796</ref> <ref name = "Moore"> Moore, T.N. and Fairweather, P.G. (2006). Decay of multiple species of seagrass detritus is dominated by species identity, with an important influence of mixing litters. ''Oikos'' 114: 329-337.</ref> <ref name = "Godbold"> Godbold, J.A., Solan, M. and Killham, K. (2009)Consumer and resource diversity effect on marine macroalgal decomposition. "Oikos" 118: 77-86.</ref>. These experiments largely involve synthetically assembled communities in plots, mesocosms (Figures 4a, 4b) or mesh-bags, in which biodiversity is controlled, but natural fluctuations in, for example, tidal cycles, rainfall or temperature, occur (Figure 4c). Although such studies incorporate more environmental variation, the spatial and temporal scales of the experimental designs largely do not relate to the dynamics of the study organisms, particularly in terms of their size or generation time. One way to incorporate such population dynamics at small and large scales is to adapt elements of meta-population theory <ref name = "France"> France, K.E. and Duffy, J.E. (2006). Diversity and dispersal interactively affects predictability of ecosystem function. ''Nature'' 441: 1139-1143.</ref>. In such studies, the importance of dispersal, the mechanism that regulates and maintains species coexistence (especially in fragmented habitats), in mediating the biodiversity-ecosystem function relationship, is recognised. At the same time, theoretical models have been extended to include environmental fluctuations <ref name = "Ives"> Ives, A.R. and Cardinale, B.J. (2004). Food-web interactions govern the resistance of communities after non-random extinctions. ''Nature'' 429: 174-177.</ref>.

[[Image: Figure 4a.jpg|thumb|left|150px|Figure 4a: Mesocosm design to hold invertebrate communities and algal resources. Blue grids represent fine mesh, allowing tidally associated environmental fluctutations within the mesocosms. Used in Godbold ''et al.''(2009) Consumer and resource diversity effect on marine macroalgal decomposition. ''Oikos'' 118: 77-86.]]

[[Image: Figure 4c.jpg|thumb|right|350px|Figure 4b: Mesocosms containing invertebrate communities and algal resources pushed 10 cm into the mud, Godbold ''et al.'' (2009).]] 

Most biodiversity-ecosystem function studies, use only a subset of the natural community to construct gradients of diversity, often selectively choosing those species that are either highly abundant or that may give the greatest response <ref name ="Raffaelli"/>. A move away from random extinction scenarios has relied primarily on modelling approaches to predict the consequences of diversity loss <ref name = "Solan"> Solan, M., Cardinale, B.J., Downing, A.L., Engelhardt, K.A.M., Ruesink, J.L. and Srivastava, D.S. (2004). Extinction and ecosystem function in the marine benthos. ''Science'' 306: 1177-1180.</ref> <ref name = "McIntyre"> McIntyre, P.B., Jones, L.E., Flecker, A.S. and Vanni, M.J (2007). Fish extinctions alter nutrient recycling in tropical freshwaters. ''Proceedings of the National Academy of Sciences of the USA'' 104: 4461-4466.</ref>. The derived scenarios use extinction drivers that are explicitly associated with trait-based extinction probabilities and thereby provide a more direct way of assessing the possibilities of ecosystem responses to biodiversity loss. Such modelling approaches have the potential for exploring the consequences of biodiversity loss at greater spatial and temporal scales by making use of large data sets which are already available for many marine and terrestrial areas <ref name ="Raffaelli"/>.

== Phase 3 – Incorporating environmental variation ==
Despite the fact that there has been a steady increase in complexity of experimental and theoretical investigations of the biodiversity-ecosystem function relationship, it is still not clear whether the same patterns observed in experimental systems are just as strong and clear in natural systems. Studies based on field observations (Figure 4), have given some insight into the biodiversity-ecosystem function relationship in natural systems <ref name = "Troumbis"> Troumbis, A.Y. and Memtsas, D. (2000). Observational evidence that diversity may increase productivity in Mediterranean shrublands. ''Oecologia'' 125: 101-108.</ref> <ref name = "Cardinale"> Cardinale, B.J., Palmer, M.A., Ives, A.R. and Brooks, S.S. (2005). Diversity-productivity relationships in streams vary as a function of the natural disturbance regime. ''Ecology'' 86: 716-726.</ref> <ref name = "Worm"> Worm, B., Barbier, E.B., Beaumont, N., Duffy, J.E., Folke, C., Halpern, B.S., Jackson, J.B.C., Lotze, H.K., Micheli, F., Palumbi, S.R., Sala, E., Selkoe, K.A., Stachowicz, J.J. and Watson, R. (2006). Impacts of biodiversity loss on ocean ecosystem services. ''Science'' 314: 787-790.</ref>, however such studies are generally correlative in nature, lack direct experimental control and do not allow for replication. In addition, the inability to control confounding factors restricts the determination of cause-effect relationships between biodiversity loss and ecosystem function <ref name = "Troumbis"> Troumbis, A.Y. and Memtsas, D. (2000). Observational evidence that diversity may increase productivity in Mediterranean shrublands. ''Oecologia'' 125: 101-108.</ref> <ref name = "Cardinale"> Cardinale, B.J., Palmer, M.A., Ives, A.R. and Brooks, S.S. (2005). Diversity-productivity relationships in streams vary as a function of the natural disturbance regime. ''Ecology'' 86: 716-726.</ref> <ref name = "Worm"> Worm, B., Barbier, E.B., Beaumont, N., Duffy, J.E., Folke, C., Halpern, B.S., Jackson, J.B.C., Lotze, H.K., Micheli, F., Palumbi, S.R., Sala, E., Selkoe, K.A., Stachowicz, J.J. and Watson, R. (2006). Impacts of biodiversity loss on ocean ecosystem services. ''Science'' 314: 787-790.</ref>. Despite efforts to overcome such problems (e.g. selecting sites of similar abiotic conditions<ref name = "Troumbis"/>; or collecting data on additional environmental variables<ref name = "Cardinale"/>), results are still open to alternative interpretations and have subsequently not shown any consistent results. 

[[Image:SPI images.jpg|thumb|centre|500px|Figure 4c: Sediment profile images used to investigate the effects of biodiversity loss on sediment mixing in natural systems. From Godbold, J.A. (2008). Marine benthic biodiversity - ecosystem function relations in complex systems. Ph.D. Thesis, University of Aberdeen.]]

This third phase (Figure 1) is currently in its infancy, however it heralds a point in biodiversity-ecosystem function research history where the discipline has matured and a full suite of evidence (i.e. theory, methodology, laboratory and field experiments, or field observations) exists, providing insight into the likely ecosystem consequences of biodiversity loss. The biodiversity-ecosystem function relationship has been found to vary depending on the relative contribution of dominant and minor species <ref name = "Emmerson"> Emmerson, M.C., Solan, M., Emes, C., Paterson, D.M. and Raffaelli, D. (2001). Consistent patterns and the idiosyncratic effects of biodiversity in marine ecosystems. ''Nature'' 411: 73-77.</ref>, environmental context <ref name = "Biles"> Biles, C.L., Solan, M., Isaksson, I., Paterson, D.M., Emes, C. and Raffaelli, D.G. (2003). Flow modifies the effect of biodiversity on ecosystem functioning: an in situ study of estuarine sediments. ''Journal of Experimental Marine Biology and Ecology'' 285: 165-177. </ref> <ref name = "Lecerf"> Lecerf, A. Risnoveanu, G., Popescu, C., Gessner, M.O. and Chauvet, E. (2007). Decomposition of diverse litter mixtures in streams. ''Ecology'' 88: 219-227.</ref>, density dependence and species interactions <ref name = "O'Connor"/> <ref name = "Godbold"/> <ref name = "Dyson"> Dyson, K.E., Bulling, M.T., Solan, M., Hernandez-Milian, G., Raffaelli, D.G., White, P.C.L., Paterson, D.M. (2007). Influence of macrofaunal assemblages and environmental heterogeneity on microphytobenthic production in experimental systems. ''Proceedings of the Royal Society B-Biological Sciences'' 274: 2547-2554.</ref>, but few studies have explicitly incorporated those structuring abiotic (environmental heterogeneity) and biotic (movement, dispersal) features that are key to species co-existence and vital for the maintenance of species diversity <ref name = "Loreau 2003"> Loreau, M., Mouquet, N. and Gonzalez, A. (2003). Biodiversity as spatial insurance in heterogeneous landscapes. ''Proceedings of the National Academy of Sciences of the USA'' 100: 12765-12770.</ref>. In addition, biodiversity effects on ecosystem properties are significantly weaker under less-well controlled conditions <ref name = "Balvanera"/>, suggesting that the effect of biodiversity on ecosystem properties may be masked by abiotic factors in natural systems <ref name = "Huston"> Huston, M.A. and McBride, A.C. (2002). Evaluating the relative strengths of biotic versus abiotic controls on ecosystem process. In: Loreau, M., Naeem, S. and Inchausti, P. (eds) Biodiversity and Ecosystem Functioning: Synthesis and Perspectives. Oxford University Press.</ref>. At present there is insufficient empirical evidence to determine the modifying effects of environmental factors, such as nutrient concentration, heterogeneity or elevated CO<sub>2</sub> on biodiversity and community dynamics and, subsequently, ecosystem properties <ref name = "Balvanera"/>. Even fewer attempts have been made, however, to establish and distinguish the relative importance of biodiversity and environmental factors in modifying ecosystem properties. Thus, a main challenge for the biodiversity-ecosystem function community is to demonstrate whether the observed importance of biodiversity in controlled experimental systems also persists in natural systems.

== See also ==
Further detailed reviews on the relationship between biodiversity and ecosystem function include:

Balvanera, P., Pfisterer, A.B., Buchmann, N., He, J.S., Nakashizuka, T., Raffaelli, D. & Schmid, B. (2006). Quantifying the evidence for biodiversity effects on ecosystem functioning and services. ''Ecology Letters'' 9: 1146-1156.

Cardinale, B.J., Srivastava, D.S., Duffy, J.E., Wright, J.P., Downing, A.L., Sankaran, M., & Jouseau, C. (2006). Effects of biodiversity on the functioning of trophic groups and ecosystems. ''Nature'' 443: 989-992.

Covich, A.P., Austen, M.C., Barlöcher, F., Chauvet, E., Cardinale, B.J., Biles, C.L., Inchausti, P., Dangles, O., Solan, M., Gessner, M.O., Statzner, B. & Moss, B. (2004). The role of Biodiversity in the functioning of freshwater and marine benthic ecosystems. ''BioScience'' 54: 767-775. 

Hooper, D.U., Chapin, F.S. III, Ewel, J.J., Hector, A., Inchausti, P., Lavorel, S., Lawton, J.H., Lodge, D.M., Loreau, M., Naeem, S., Schmid, B., Setala, H., Symstad, A.J., Vandermeer, J. & Wardle, D.A. (2005). Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. ''Ecological Monographs'' 75: 3 - 35.

Hughes, J.B. & Petchey, O.L. (2001). Merging perspectives on biodiversity and ecosystem functioning. ''Trends in Ecology and Evolution'' 16: 222-223.

Loreau, M, Naeem, S., Inchausti, P., Bengtsson, J, Grime, J.P., Hector, A., Hooper, D.U., Huston, M.A., Raffaelli, D., Schmid, B., Tilman, D. & Wardle, D.A. (2001). Biodiversity and ecosystem functioning: Current knowledge and future challenges. ''Science'' 294: 804-808.

Solan, M., Godbold, J.A., Symstad, A., Flynn, D.F.B. & Bunker, D. (2009). Biodiversity-ecosystem function research and biodiversity futures: early bird catches the worm or a day late and a dollar short? In: Biodiversity and human impacts. Ecological and societal implications. Naeem, S., Bunker, D.E., Hector, A., Loreau, M. & Perrings, C. (Eds.). Oxford University Press.

Stachowicz, J.J., Bruno, J.F. & Duffy, J.E. (2007). Understanding the effects of marine biodiversity on communities and ecosystems. ''Annual Review of Ecology, Evolution and Systematics'' 38: 739-766.

The influence of the lugworm (''Arenicola marina'') on biodiversity and ecosystem functioning in an intertidal mudflat [http://www.marbef.org/outreach/newsletter.php]<p>

==References==
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{{author
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|AuthorFullName=Godbold, Jasmin
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[[Category:Coastal and marine ecosystems]]   
[[Category:Marine Biodiversity‏‎]]