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Coastal biodiversity is complex due to the large numbers of species interacting in space and time at many different scales. Because little is known of the invertebrates which make up most of this biodiversity, conserving biodiversity is a difficult task. Robust analytical techniques must be developed that can measure natural changes in biodiversity before one will be able to assess whether biodiversity is changing in response to anthropogenic disturbances. Many attempts have been made to reduce the complexity of a species x sample data-matrix, which is the basic measure of biodiversity, into simpler measures, which are more amenable to analysis. These are not generally successful because they capture only part of the variation in assemblages of organisms. This paper introduces the basic components into which measures of biodiversity can be “decomposed” to allow existing methods of analyses and ecological knowledge to understand measures of and changes to biodiversity.

Agresti, A. 1990 Categorical Data Analysis. Wiley, New York.
Andersen, A. N. 1995. Measuring more of biodiversity: genus richness as a surrogate for species richness in Australian ant fauna. Biological Conservation 73: 39-43.
Anderson, M. J. 2001. A new method for non-parametric multivariate analysis of variance. Austral Ecology 26: 32-46.
Andrewartha, H. G. and Birch, L. C. 1984 The Ecological Web. University of Chicago Press, Chicago.
Bayne, B. E., Clarke, K. R. and Gray, J. S. (eds.) 1988. Biological effects of pollutants. Results of a practical workshop. Marine Ecology Progress Series Special Issue 46: 1-278
Caro, T. M. and O'Doherty, G. 1999. On the use of surrogate species in conservation biology. Conservation Biology 13: 805-14.
Chapman, M. G. 1998. Relationships between spatial patterns of benthic assemblages in a mangrove forest using different levels of taxonomic resolution. Marine Ecology Progress Series 162: 71-8.
Connell, J. H. 1972. Community interactions on marine rocky intertidal shores. Annual Review of Ecology and Systematics 3: 169-92.
Coull, B. C., Hicks, G. R. F. and Wells, J. B. J. 1981. Nematode/copepod ratios for monitoring pollution: a rebuttal. Marine Pollution Bulletin 12: 378-81.
Dayton, P. K. 1984. Processes structuring some marine communities: Are they general? Pp 181-97 in Ecological Communities: Conceptual Issues and the Evidence, edited by D. R. Strong, D. Simberloff, L. G. Abele, and A. B. Thistle. Princeton University Press, Princeton, New Jersey.
Darwin, C. 1859 On the Origin of Species. Murray, London.
Fairweather, P. G. 1991. Statistical power and design requirements for environmental monitoring. Australian Journal of Marine and Freshwater Research 42: 555-68.
Gray, J. S. 2000. The measurement of marine species diversity, with an application to the benthic fauna of the Norwegian continental shelf. Journal of Experimental Marine Biology and Ecology 250: 23-49.
Gray, J. S., Clarke, K. R., Warwick, R. M. and Hobbs, G. 1990. Detection of the initial effects of pollution on marine benthos: an example from the Ekofisk and Eldfisk oilfields, North Sea. Marine Ecology Progress Series 66: 285-99.
Green, R. H. 1979 Sampling Design and Statistical Methods for Environmental Biologists. Wiley, Chichester.
Herman, P. M. J. and Heip, C. 1988. On the use of meiofauna in ecological monitoring: who needs taxonomy? Marine Pollution Bulletin 19: 665-8.
Johnson, C. R. and Field, C. A. 1993. Using fixed-effects model multivariate analysis of variance in marine biology and ecology. Annual Review of Oceanography and Marine Biology 31: 177-222.
Jones, G. P. and Kaly, U. L. 1996. Criteria for selecting marine organisms in biomonitoring studies. Pp 29-48 in Detecting Ecological Impacts: Concepts and Applications in Coastal Habitats, edited by R. J. Schmitt, and C. W. Osenberg. Academic Press, San Diego.
Kastendiek, J. 1982. Competitor-mediated coexistence: interactions among three species of benthic macroalgae. Journal of Experimental Marine Biology and Ecology 62: 201-10.
Lawton, J. H., Naeem, S., Thompson, L. J., Hector, A. and Crawley, M. J. 1998. Biodiversity and ecosystem functioning: getting the Ecotron experiment in its correct context. Functional Ecology 12: 848-52.
Lewis, J. R. 1964 The Ecology of Rocky Shores. The English Universities Press, London.
Morrisey, D. J., Howitt, L., Underwood, A. J. and Stark, J. S. 1992a. Spatial variation in soft-sediment benthos. Marine Ecology Progress Series 81: 197-204.
Morrisey, D. J., Underwood, A. J., Howitt, L. and Stark, J. S. 1992b. Temporal variation in soft-sediment benthos. Journal of Experimental Marine Biology and Ecology 164: 233-45.
Naeem, S., Thompson, L. J., Lawler, S. P., Lawton, J. H. and Woodfin, R. M. 1995. Empirical evidence that declining species diversity may alter the performance of terrestrial ecosystems. Philosophical Transactions of the Royal Society Series B 347: 249-62.
Paine, R. T. 1974. Intertidal community structure: experimental studies on the relationship between a dominant competitor and its principal predator. Oecologia 15: 93-120.
Paine, R. T. 1994 Marine Rocky Shores and Community Ecology: An Experimentalist's Perspective. Ecology Institute, Oldendork/Luhe, Germany.
Phillips, D. J. H. 1978. The use of biological indicator organisms to quantitate organo-chlorine pollutants in aquatic environments: a review. Environmental Pollution, 16: 167-229.
Pielou, E. C. 1974 Population and Community Ecology: Principles and Methods. Gordon and Breach, New York. Pimm, S. L. 1982 Food Webs. Chapman and Hall, London.
Potter, I. C., Bird, D. J., Claridge, P. N., Clarke, K. R., Hyndes, G. A. and Newton, L. C. 2001. Fish fauna of the Severn Estuary. Are there long-term changes in abundance and species composition and are the recruitment patterns of the main marine species correlated? Journal of Experimental Marine Biology and Ecology 258: 15-37.
Quinn, G. P. and Keough, M. J. 2002 Experimental Design and Data Analysis for Biologists. Cambridge University Press, Cambridge.
Raimondi, P. T. and Reed, D. C. 1996. Determining the spatial extent of ecological impacts caused by local anthropogenic disturbances in coastal marine habitats. Pp 179-98 in Detecting Ecological Impacts: Concepts and Applications in Coastal Habitats, edited by R. J. Schmitt, and C. W. Osenberg. Academic Press, San Diego.
Steneck, R. S. 1982. A limpet-coralline alga association: adaptations and defenses between a selective herbivore and its prey. Ecology 63: 507-22.
Stewart-Oaten, A., Murdoch, W. M. and Parker, K. R. 1986. Environmental impact assessment: ‘pseudoreplication’ in time? Ecology 67: 929-40.
Tilman, D. 1999. The ecological consequences of changes in biodiversity: A search for general principles. Ecology 80: 1455-74.
Underwood, A. J. 1979. The ecology of intertidal gastropods. Advances in Marine Biology 16: 111-210.
Underwood, A. J. 1985. Physical factors and biological interactions: the necessity and nature of ecological experiments. Pp. 371-90 in The Ecology of Rocky Coasts, edited by P. G. Moore, and R. Seed. Hodder & Stoughton, London.
Underwood, A. J. 1986. What is a community? Pp. 351-67 in Patterns and Processes in the History of Life. Dahlem Konferenzen, edited by D. M. Raup, and D. Jablonski. Springer-Verlag, Berlin.
Underwood, A. J. 1991. Beyond BACI: experimental designs for detecting human environmental impacts on temporal variations in natural populations. Australian Journal of Marine and Freshwater Research 42: 569-87.
Underwood, A. J. 1992. Beyond BACI: the detection of environmental impact on populations in the real, but variable, world. Journal of Experimental Marine Biology and Ecology 161: 145-78.
Underwood, A. J. 1994. Spatial and temporal problems with monitoring. Pp. 101-23 in Rivers Handbook, Volume 2, edited by P. Calow and G. E. Petts. Blackwell Scientific Publications, London.
Underwood, A. J. 1997 Ecological Experiments: Their Logical Design and Interpretation Using Analysis of Variance. Cambridge University Press, Cambridge.
Underwood, A. J. 1999. Physical disturbances and their direct effect on an indirect effect: responses of an intertidal assemblage to a severe storm. Journal of Experimental Marine Biology and Ecology 232: 125-40.
Underwood, A. J. 2000. Experimental ecology of rocky intertidal habitats: what are we learning? Journal of Experimental Marine Biology and Ecology 250: 51-76.
Underwood, A. J. and Keough, M. J. 2001. Supply-side ecology: the nature and consequences of variations in recruitment of intertidal organisms. Pp 183-200 in Marine Community Ecology, edited by M. D. Bertness, S. D. Gaines, and M. E. Hay. Sinauer Asssociates Inc., Sunderland, Maine.
Warwick, R. M. and Clarke, K. R. 1995. New ‘biodiversity’ measures reveal a decrease in taxonomic distinctness with increasing stress. Marine Ecology Progress Series, 129: 301-05.
Wootton, J. T. 1977. Estimates and tests of per capita interaction strength: diet, abundance, and impact of intertidally foraging birds. Ecological Monographs 67: 45-64.
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References

Agresti, A. 1990 Categorical Data Analysis. Wiley, New York.
Andersen, A. N. 1995. Measuring more of biodiversity: genus richness as a surrogate for species richness in Australian ant fauna. Biological Conservation 73: 39-43.
Anderson, M. J. 2001. A new method for non-parametric multivariate analysis of variance. Austral Ecology 26: 32-46.
Andrewartha, H. G. and Birch, L. C. 1984 The Ecological Web. University of Chicago Press, Chicago.
Bayne, B. E., Clarke, K. R. and Gray, J. S. (eds.) 1988. Biological effects of pollutants. Results of a practical workshop. Marine Ecology Progress Series Special Issue 46: 1-278
Caro, T. M. and O'Doherty, G. 1999. On the use of surrogate species in conservation biology. Conservation Biology 13: 805-14.
Chapman, M. G. 1998. Relationships between spatial patterns of benthic assemblages in a mangrove forest using different levels of taxonomic resolution. Marine Ecology Progress Series 162: 71-8.
Connell, J. H. 1972. Community interactions on marine rocky intertidal shores. Annual Review of Ecology and Systematics 3: 169-92.
Coull, B. C., Hicks, G. R. F. and Wells, J. B. J. 1981. Nematode/copepod ratios for monitoring pollution: a rebuttal. Marine Pollution Bulletin 12: 378-81.
Dayton, P. K. 1984. Processes structuring some marine communities: Are they general? Pp 181-97 in Ecological Communities: Conceptual Issues and the Evidence, edited by D. R. Strong, D. Simberloff, L. G. Abele, and A. B. Thistle. Princeton University Press, Princeton, New Jersey.
Darwin, C. 1859 On the Origin of Species. Murray, London.
Fairweather, P. G. 1991. Statistical power and design requirements for environmental monitoring. Australian Journal of Marine and Freshwater Research 42: 555-68.
Gray, J. S. 2000. The measurement of marine species diversity, with an application to the benthic fauna of the Norwegian continental shelf. Journal of Experimental Marine Biology and Ecology 250: 23-49.
Gray, J. S., Clarke, K. R., Warwick, R. M. and Hobbs, G. 1990. Detection of the initial effects of pollution on marine benthos: an example from the Ekofisk and Eldfisk oilfields, North Sea. Marine Ecology Progress Series 66: 285-99.
Green, R. H. 1979 Sampling Design and Statistical Methods for Environmental Biologists. Wiley, Chichester.
Herman, P. M. J. and Heip, C. 1988. On the use of meiofauna in ecological monitoring: who needs taxonomy? Marine Pollution Bulletin 19: 665-8.
Johnson, C. R. and Field, C. A. 1993. Using fixed-effects model multivariate analysis of variance in marine biology and ecology. Annual Review of Oceanography and Marine Biology 31: 177-222.
Jones, G. P. and Kaly, U. L. 1996. Criteria for selecting marine organisms in biomonitoring studies. Pp 29-48 in Detecting Ecological Impacts: Concepts and Applications in Coastal Habitats, edited by R. J. Schmitt, and C. W. Osenberg. Academic Press, San Diego.
Kastendiek, J. 1982. Competitor-mediated coexistence: interactions among three species of benthic macroalgae. Journal of Experimental Marine Biology and Ecology 62: 201-10.
Lawton, J. H., Naeem, S., Thompson, L. J., Hector, A. and Crawley, M. J. 1998. Biodiversity and ecosystem functioning: getting the Ecotron experiment in its correct context. Functional Ecology 12: 848-52.
Lewis, J. R. 1964 The Ecology of Rocky Shores. The English Universities Press, London.
Morrisey, D. J., Howitt, L., Underwood, A. J. and Stark, J. S. 1992a. Spatial variation in soft-sediment benthos. Marine Ecology Progress Series 81: 197-204.
Morrisey, D. J., Underwood, A. J., Howitt, L. and Stark, J. S. 1992b. Temporal variation in soft-sediment benthos. Journal of Experimental Marine Biology and Ecology 164: 233-45.
Naeem, S., Thompson, L. J., Lawler, S. P., Lawton, J. H. and Woodfin, R. M. 1995. Empirical evidence that declining species diversity may alter the performance of terrestrial ecosystems. Philosophical Transactions of the Royal Society Series B 347: 249-62.
Paine, R. T. 1974. Intertidal community structure: experimental studies on the relationship between a dominant competitor and its principal predator. Oecologia 15: 93-120.
Paine, R. T. 1994 Marine Rocky Shores and Community Ecology: An Experimentalist's Perspective. Ecology Institute, Oldendork/Luhe, Germany.
Phillips, D. J. H. 1978. The use of biological indicator organisms to quantitate organo-chlorine pollutants in aquatic environments: a review. Environmental Pollution, 16: 167-229.
Pielou, E. C. 1974 Population and Community Ecology: Principles and Methods. Gordon and Breach, New York. Pimm, S. L. 1982 Food Webs. Chapman and Hall, London.
Potter, I. C., Bird, D. J., Claridge, P. N., Clarke, K. R., Hyndes, G. A. and Newton, L. C. 2001. Fish fauna of the Severn Estuary. Are there long-term changes in abundance and species composition and are the recruitment patterns of the main marine species correlated? Journal of Experimental Marine Biology and Ecology 258: 15-37.
Quinn, G. P. and Keough, M. J. 2002 Experimental Design and Data Analysis for Biologists. Cambridge University Press, Cambridge.
Raimondi, P. T. and Reed, D. C. 1996. Determining the spatial extent of ecological impacts caused by local anthropogenic disturbances in coastal marine habitats. Pp 179-98 in Detecting Ecological Impacts: Concepts and Applications in Coastal Habitats, edited by R. J. Schmitt, and C. W. Osenberg. Academic Press, San Diego.
Steneck, R. S. 1982. A limpet-coralline alga association: adaptations and defenses between a selective herbivore and its prey. Ecology 63: 507-22.
Stewart-Oaten, A., Murdoch, W. M. and Parker, K. R. 1986. Environmental impact assessment: ‘pseudoreplication’ in time? Ecology 67: 929-40.
Tilman, D. 1999. The ecological consequences of changes in biodiversity: A search for general principles. Ecology 80: 1455-74.
Underwood, A. J. 1979. The ecology of intertidal gastropods. Advances in Marine Biology 16: 111-210.
Underwood, A. J. 1985. Physical factors and biological interactions: the necessity and nature of ecological experiments. Pp. 371-90 in The Ecology of Rocky Coasts, edited by P. G. Moore, and R. Seed. Hodder & Stoughton, London.
Underwood, A. J. 1986. What is a community? Pp. 351-67 in Patterns and Processes in the History of Life. Dahlem Konferenzen, edited by D. M. Raup, and D. Jablonski. Springer-Verlag, Berlin.
Underwood, A. J. 1991. Beyond BACI: experimental designs for detecting human environmental impacts on temporal variations in natural populations. Australian Journal of Marine and Freshwater Research 42: 569-87.
Underwood, A. J. 1992. Beyond BACI: the detection of environmental impact on populations in the real, but variable, world. Journal of Experimental Marine Biology and Ecology 161: 145-78.
Underwood, A. J. 1994. Spatial and temporal problems with monitoring. Pp. 101-23 in Rivers Handbook, Volume 2, edited by P. Calow and G. E. Petts. Blackwell Scientific Publications, London.
Underwood, A. J. 1997 Ecological Experiments: Their Logical Design and Interpretation Using Analysis of Variance. Cambridge University Press, Cambridge.
Underwood, A. J. 1999. Physical disturbances and their direct effect on an indirect effect: responses of an intertidal assemblage to a severe storm. Journal of Experimental Marine Biology and Ecology 232: 125-40.
Underwood, A. J. 2000. Experimental ecology of rocky intertidal habitats: what are we learning? Journal of Experimental Marine Biology and Ecology 250: 51-76.
Underwood, A. J. and Keough, M. J. 2001. Supply-side ecology: the nature and consequences of variations in recruitment of intertidal organisms. Pp 183-200 in Marine Community Ecology, edited by M. D. Bertness, S. D. Gaines, and M. E. Hay. Sinauer Asssociates Inc., Sunderland, Maine.
Warwick, R. M. and Clarke, K. R. 1995. New ‘biodiversity’ measures reveal a decrease in taxonomic distinctness with increasing stress. Marine Ecology Progress Series, 129: 301-05.
Wootton, J. T. 1977. Estimates and tests of per capita interaction strength: diet, abundance, and impact of intertidally foraging birds. Ecological Monographs 67: 45-64.
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