56 Introduction

CHAPTER 3.2 RANK-ABUNDANCE DISTRIBUTIONS

The distribution of the relative abundance of species in local assemblages follows regularities that can be described by mathematical distribution models: If sorted by rank, species frequencies tend to follow a hollow curve (Robinson 1998, May 1975) with a few very abundant, some medium-abundant, and many rare species. This pattern is particularly obvious in species-rich samples (e.g. insects from the tropics), but is almost ubiquitous in its general trend (Hayek & Buzas 1997). Fisher et al. (1943) have successfully proposed the distribution model of the logarithmic series (empirically based on Lepidoptera catches in Southeast-Asia), of which the diversity index Fishers is derived (Kempton & Taylor 1974), while Preston (1948) alternatively proposed the lognormal distribution as a suitable description of the rankabundance relationships. Frequency distributions are not only a pivotal point in the measurement of species diversity (evenness is basically a measure of this distribution, Weiher & Keddy 1999, see also Southwood & Henderson 2000), but may also give clues about the underlying mechanisms of community assemblage. Based on a model of niche partitioning, McArthur (1965) proposed the broken stick distribution, which has received little empirical support (Hubbell 2001), while the empirically derived logseries and lognormal distributions (May 1975, Tokeshi 1993) fit most samples of organisms that were ever taken. Many more phenomenologically or mechanistically derived distributions have been proposed since then to provide the best fit to species abundance distributions (see e.g. Tokeshi 1993, Bell 2000, Plotkin et al. 2000, McGill 2003, Sol et al. 2004, Nummelin & Kaitala 2004). The logseries was often viewed as the better-fitting distribution (e.g. Kempton & Taylor 1974), but recently available, very large samples are better described by the lognormal (see Hubbell 2001 for data and further references). Despite an apparent competition between these two models, several attempts were made to show that both distributions might be special cases of a general distribution model (e.g. Diserud & Engen 2000, Hubbell 2001). Attempts to derive the empirical distributions, including the skewed lognormal form in many very large data sets (Nee et al. 1991, Hubbell 2001), from biologically meaningful first principles (see also Basset et al. 1998) are increasingly successful for neutral models (e.g. Bell 2000). Hubbell (2001) proposed the zero-sum multinomial distribution (zsm) based on his neutral model of population growth, which derives lognormal-like patterns for local assemblages, while infinitely large samples of the metacommunity (Hubbell 2001) yield a logseries-like distribution pattern (but see Alonso & McKane 2004, also McGill & Collins 2003, Bell 2000, Mouillot et al. 2000, Engen & Lande 1996a, b, Hengeveld & Stam 1978, Kempton & Taylor 1974, May 1975, Preston 1962a, b for alternative approaches). Despite the compilation of many supporting examples by Hubbell (2001), his model has so far received little independent empirical support (e.g. McGill 2003, Ulrich & Ollik 2004, Leigh et al. 2004). In an empirical approach, comparisons of rank-abundance data with the standard models of species-abundance relations can give important clues of the structuring of the assemblage under investigation (Southwood & Henderson 2000). Deviations from the models can give hints about the unusual ecology of species that cause the deviation. For example, plots of the rank abundance relationship of Geometrid moth samples from a traditionally cultivated region in Borneo identified several species which were clearly commoner than predicted by the