• diagnostic species;
  • environmental indication;
  • indicator species;
  • indicator value;
  • vegetation classification;
  • vegetation types


  1. Indicator species are often determined using an analysis of the relationship between the species occurrence or abundance values from a set of sites and the classification of the same sites into site groups (habitat types, community types, disturbance states, etc.). It may happen, however, that a particular site group has no indicator species even if its sites have a community composition that is clearly distinct from the sites of other site groups. This motivates an exploration of the indicator value of not only individual species but also species combinations.
  2. Here, we present a novel statistical approach to determine indicators of site groups using species data. Unlike traditional indicator value analysis, we allow indicators to be species combinations in addition to single species. We require that all the species forming the combination must occur in the site to use the combination as an indicator. We present a simple algorithm that identifies the set of indicators (each one being either a single species or a species combination) that show high positive predictive value for the target site group. Moreover, we demonstrate the use of the percentage of sites of the site group where at least one of its valid indicators occurs to determine whether the group can be reliably predicted throughout its range.
  3. Using a simulation study, we show that if two species are not strongly correlated and their frequency in the data set is larger than the frequency of sites belonging to the site group, the joint occurrence of the two species has higher positive predictive value for the site group than the two species taken independently.
  4. We illustrate the proposed method by determining which combinations of vascular plants can be used as indicators for 29 shrubland and forest vegetation types of New Zealand.
  5. The proposed methodology extends traditional indicator value analyses and will be useful to develop multispecies ecological or environmental indicators. Further, it will allow newly surveyed sites to be reliably assigned to previously defined vegetation types.