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CHAPTER 3.

2 RANK-ABUNDANCE DISTRIBUTIONS

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region in correct proportions. Rather, logistics are probably the most important factor on sample site choice. In Borneo, data from all major habitat types are available (although some did not find their way into analysis due to low specimen counts, e.g. mangrove or heath forest), whereas samples from the rest of Southeast-Asia might be non-representative chance samples. As a consequence, average frequencies derived from this data might be poor in representing mean population density across the continent even without the potential biases discussed in the paragraph above. Figure 3.13 exemplifies the combined effects of these potential flaws in data quality: In a perfect world, where all samples are from the same habitat type, and there are no differences in local productivity or trapping efficiency, absolute sample size of a species, the number of samples where a species is present, and a species frequency were perfectly correlated. This is obviously not the case, although correlations are significant (Spearman rank correlations, R>0,352, p<0,001). Species frequencies vary considerably over their range, probably at least partly as a result of varying local habitat conditions. No obvious geographic patterns (e.g. a decline in local frequency near the limit of a species range) could be observed in the most common species, for which such mapping was possible (data not shown). The biases could be expected to blur whatever pattern could be found in the distribution of mean population densities. On the other hand, good models of species-abundance relations on a geographically large scale have to consider, or be at least robust to, habitat heterogeneity. It will still take a long time until flawless data for tropical invertebrates become available on a sub-continental geographical scale, so for the moment whatever data are there should be used to explore patterns and hypotheses albeit with caution and an open eye for potential biases (Lawton 2000).

Patterns of species frequencies distributions and the neutral model Despite the limitations due to data quality which are discussed above, patterns of mean frequencies in regional and local assemblages strongly resemble those predicted by Hubbells (2001) theory of non-specific population growth of species (i.e. ecological neutrality; for discussion of the theory, see also Ricklefs 2003, Hubbell 2003). Furthermore, deviations from predictions (e.g. the tailing-off of rare species even among the metacommunity samples) or differences between subsets of data (e.g. Southeast-Asia vs. Borneo) can be readily explained and interpreted by the proposed effect of dispersal limited populations. Thus, on the not-so-rigorous level of graphic resemblance (e.g. McGill 2003a, b) Sphingid data support Hubbells (2001) suggestions on a far larger geographical scale than the examples presented in his book. However, correct prediction of observed patterns does not necessarily prove the correctness of underlying assumptions, i.e. the ecological neutrality of species in Hubbells (2001) theory (e.g. Purves & Pacala, in press). Sphingid assemblages are clearly influenced by habitat conditions (disturbance, elevation; see chapter 3.1), which indicates that species have differential success in different habitat types (i.e., niches). On the other hand, predicted patterns persisted even though very heterogeneous data from various habitat types were used. Hubbell (2001) concludes that possibly the specialization of species does not have much impact on many community-level patterns of the system an interpretation that is supported

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