Journal of Vegetation Science && (2011)
Too good to be true: pitfalls of using mean Ellenberg
indicator values in vegetation analyses
David Zeleny ´ & Andre ´ P. Schaffers
Keywords
Bio-indication; Circularity of reasoning;
Compositional similarity; Environmental
calibration; Null models; Ordination analysis;
Randomization; Species richness; Weighted
averaging
Abbreviations
ANOVA = analysis of variance; CA =
correspondence analysis; CCA = canonical
correspondence analysis; DCA = detrended
correspondence analysis; EIVs = Ellenberg
indicator values; TWINSPAN = two-way
indicator species analysis
Received 2 September 2010
Accepted 14 October 2011
Co-ordinating Editor: Michael Palmer
Zeleny ´ , D. (corresponding author,
zeleny@sci.muni.cz): Department of Botany
and Zoology, Masaryk University, Kotla ´r ˇska ´ 2,
CZ-611 37, Brno, Czech Republic
Schaffers, A.P. (Andre.Schaffers@wur.nl):
Department of Environmental Sciences, Nature
Conservation and Plant Ecology Group,
Wageningen University, PO BOX 47, 6700 AA,
Wageningen, The Netherlands
Abstract
Question: Mean Ellenberg indicator values (EIVs) inherit information about
compositional similarity, because during their calculation species abundances
(or presence–absences) are used as weights. Can this similarity issue actually be
demonstrated, does it bias results of vegetation analyses correlating mean EIVs
with other aspects of species composition and how often are biased studies
published?
Methods: In order to separate information on compositional similarity possibly
present in mean EIVs, a new variable was introduced, calculated as a weighted
average of randomized species EIVs. The performance of these mean randomized
EIVs was compared with that of the mean real EIVs on the one hand and random
values (randomized mean EIVs) on the other. To demonstrate the similarity issue,
differences between samples were correlated with dissimilarity matrices based
on various indices. Next, the three mean EIV variables were tested in canonical
correspondence analysis (CCA), detrended correspondence analysis (DCA),
analysis of variance (ANOVA) between vegetation clusters, and in regression on
species richness. Subsequently, a modified permutation test of significance was
proposed, taking the similarity issue into account. In addition, an inventory was
made of studies published in the Journal of Vegetation Science and Applied Vegeta-
tion Science between 2000 and 2010 likely reporting biased results due to the simi-
larity issue.
Results: Using mean randomized EIVs, it is shown that compositional similarity is
inherited into mean EIVs and most resembles the inter-sample distances in cor-
respondence analysis, which itself is based on iterative weighted averaging. The
use of mean EIVs produced biased results in all four analysis types examined:
unrealistic (too high) explained variances in CCA, too many significant correla-
tions with ordination axes in DCA, too many significant differences between
cluster analysis groups and too high coefficients of determination in regressions
on species richness. Modified permutation tests provided ecologically better
interpretable results. From 95 studies using Ellenberg indicator values, 36
reported potentially biased results.
Conclusions: No statistical inferences should be made in analyses relating mean
EIVs with other variables derived from the species composition as this can pro-
duce highly biased results, leading to misinterpretation. Alternatively, a modi-
fied permutation test using mean randomized EIVs can sometimes be used.
Introduction
Ellenberg indicator values (EIVs; Ellenberg et al. 1992) and
their geographic alternatives (e.g. Landolt 1977; Borhidi
1995; Hill et al. 1999; Pignatti 2005; Lawesson et al. 2009)
are frequently used by European vegetation scientists as
surrogates for measured environmental variables. Several
studies have demonstrated that the calculated mean of
EIVs for species present in the vegetation sample are often
a good estimate of real environmental conditions, even if
this relationship may be limited to certain parts of a given
gradient or to a particular vegetation type (for a detailed
Journal of Vegetation Science
Doi: 10.1111/j.1654-1103.2011.01366.x © 2011 International Association for Vegetation Science 1