Diversity

In the previous chapter we covered ways of describing samples of benthos, but specifically did not include diversity. We can talk of primary community variables, such as abundance (A), species richness (S) and biomass (B), and derived variables from these such as true diversity indices, evenness indices, and ratios indicating the relationship between species richness and abundance (A/S, the abundance ratio or the average abundance per species) and between biomass and abundance (B/A, the biomass ratio or the mean biomass per individual). Diversity is not just simply about the number of species found in a sample or area, but also uses data on the abundances of individuals among the species and the way those abundances are distributed among the species within the assemblage. There are many ways of describing diversity. Here we give a summary of the most important ones and reference sources of recent literature on the subject (see also the data analysis summary in Chapter 11). In the following section we consider simple indices (univariate) as measures of diversity; multivariate methods of analysing patterns will be covered in Chapter 7 on the effects of disturbance. The simplest way to measure diversity is the number of species found in a sample, called the species richness (S or SR). Yet diversity is not just about numbers of species; it is also concerned with the distribution of numbers of individuals per species. For example, if one assemblage has 50 individuals of each of 2 species A and B whereas another assemblage has 99 individuals of species A and 1 individual of species B, then both have the same species richness but the first assemblage is the more diverse. Thus a measure of diversity (an index) must take into account not only the number of species, but also the number of individuals per species. To distinguish this from species richness, the combination of individuals per species and number of species is called heterogeneity diversity. In fact there are a large number of diversity indices, and we do not propose to consider them all here (Magurran 2004 gives an excellent and detailed account and others are mentioned in the summary in Chapter 11).