Ecological Archives E086-055-A5

Colleen K. Kelly and Michael G. Bowler. 2005. A new application of storage dynamics: differential sensitivity, diffuse competition, and temporal niches. Ecology 86:1012–1022.

Appendix E. More on measuring deviation. A pdf file of this appendix is also available.

How might fluctuations best be quantified, assuming that we have determined comparable intervals of time and a curve against which to compare the species involved in the lottery we have described?  Let  be the number expected in bin i according to, for example, an exponential distribution and  the number actually observed.  One measure of deviation from the expected smooth curve is


This quantity is well defined but will not have a  distribution unless  is normally distributed about .  In Kelly et al. (2001), the measure

was employed, which is an analog of  for Poisson distributed  (Baker and Cousins 1984).

A measure of the fractional deviation from an expected (or hypothesised) smooth curve is


Both (E.1) and (E.2) can be divided into interesting and uninteresting pieces, as follows:


where  is an interesting deviation (in this paper, a fluctuation in recruitment caused by an environmental effect) and  is a statistical fluctuation (which is not interesting).  Then to the extent that  and  are uncorrelated


If  is Poisson distributed about ,  and  fluctuates.

Thus                               and                  .

In (E.3) the second (statistical) term is independent of the sample size or shape of the expected distribution (but depends on the number of bins) but the first (interesting) term depends on the shape and the sample size.  In (E.4) the situation is reversed: the first (interesting) term is independent of the shape or sample size, whereas the second (statistical) term depends on both – but can be calculated for a given expected distribution.

A measure of the  type can be used to establish deviation from some hypothesized distribution beyond the merely statistical, but care may be necessary in interpreting differences in the estimator of which is


The measure we are after for comparison with the lottery model is the measure of fractional deviation for which we took the estimator


There are two caveats that should be borne in mind.  The first is that the statistical terms have a distribution and the corrections in (E.5) and (E.6) are the mean values of the statistical terms.  The second caveat is that the populations of individual bins may not be Poisson distributed.

            The variance on (E.5) due to statistical fluctuations about  which are Poisson distributed is

and the variance on (E.6) due to such statistical fluctuations is

            We do not think it possible (or necessary) to extract a better estimate of the variance V on  than


without a rather complete understanding of the details of each two component system and the larger forest environment within which they operate.



Baker, S., and R. D. Cousins. 1984. Clarification of the use of chi-square and likelihood functions in fits to histograms. Nuclear Instruments and Methods in Physics Research 221:437–442.

Kelly, C. K., H. Banyard Smith, Y. M. Buckley, R. Carter, M. Franco, W. Johnson, T. Jones, B. May, R. Perez Ishiwara, A. Perez-Jimenez, A. Solis Magallanes, H. Steers, and C. Waterman. 2001. Investigations in commonness and rarity: a comparative analysis of co-occuring, congeneric Mexican trees. Ecology Letters 4:618–627.

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