J. theor. Biol.

(1970)129, 151-154

A Test for Comparing Diversities Based on the

Shannon Formula

Several indices of dispersion have been suggested by ecologists, the most

commonly used being the measure corresponding to the entropy concept
defined by
s

H = - C pi ln pi
i=l

where pi 2 , Cpi = 1. Thus the statistic is

h = -i$ h/n) ln Cniln)

and this can be regarded as an estimate of H being given a random sample

of n observations from data falling into the s categories c,(l, 2,. . . , s) with
probabilities ~~(1, 2,. . . , s). It is well known that h and H lie between zero
and Ins, the latter being achieved when the observations (or category
probabilities) are uniformly distributed (since x In x -+ 0 as x + 0 it is
customary to take ni In ni as zero when 12i = 0).
The distributional properties of h must be known before one can compare
the diversities of populations. Much has been done recently toward establish-
ing the distribution of h as well as other measures of diversity by Bowman,
Hutcheson, Odum & Shenton (1969).
Beforelooking at the distribution of h, we record the exact values of the first
two moments of h. From Hutcheson (1969) and Bowman et al. (1969),

Eh = In n --(n - l)P,(ln 2 - In 1) -

(::_:)P(lnn
- (“i’) ln(n-l)+...+(-l)“-‘In1 >
(1)

where

P, = i p:.
j=l
151
152 K. HUTCHESON

For example,
11= 2 Eh = (l-P,)In2
11= 3 Eh = -2(P,-P,)-ln2+(1-P,)In3.

where B = integer part of (n-b-2)/2.

These expressions can be used as long as n and s are not excessively large
(say, S, n, up to about 200). Even with large computers the time to compute
Var h can be significant.
In order for one to make use of the moments of h in a practical situation
one must find good estimates of Eh and Var h. From Bowman et al. (1969)
the series expansions are,
Eh=-Cpilnpi-(s-1)/2n+(l-Cp,~‘)/
12n2+C(p;‘-p,y2)/12n3+. .. (3)
Var h = [C pi ln2 pi - (C pj In pJ’]/n + (S- l)/2n2 +
+(-1+~p~~‘-~p,~11np~+~p,~‘~p~Inpi)/6n3+.... (4)
It is shown, by Bowman et al. (1969), that the distribution of h is asymp-
totically normal. If the categories are equiprobable the distribution is x2.
It is well known that the sampling distribution of the statistic
z = G1-~2)-(P1 -P2>

(of/n, + a$/n,)“’
is normal with mean 0 and variance 1. This is approximately true for
sufficiently large values of nl and n2 even if the populations are not normal.
If the values of c1 and g2 are not known and if it is known that each
population is normally (or nearly normal) distributed, then the statistic
obtained by substituting the estimates Var h, and Var h, for CJ~and e2
yields a statistic which is approximately t-distribution with degrees of
freedom
(Var h,)’ (Var h,)’
d.f. = [Var h1 +Var h,]’ ____- +
nl n2
As noted in section 1 exact calculations are very unwieldy for n, s, 2200.
However, the equation (4) above gives a good approximation to the sample
 LETTERS TO THE EDITOR 153

variance and asymptotically approaches the sample variance. Thus,

h--h,-(C11P2)
t = (Var h, + VBr h2) “’
is asymptotically t with d.f. given above.
Using data from Stoddard & Norris (1967) we will compare the diversity
of the bird casualties of January 1964 to that of January 1965. The number
of individuals killed in January 1964 was 57 with 16 species represented.
The number of individuals killed in .January 1965 was 23 with 11 species
represented. For 1964,
II, = 2.44155 Vdr 11, = 0.0148724
for 1965,
II, = 1.80785 Var 11~= 0.0620399

with 35 degrees of freedom.

We would reject the hypothesis of equal diversities at the 5 “J level.
The true variance for 1965 was found to be
Var 11~= 0.0547087, thus the above test is conservative.
The data from Table 1 of Pielou (I 966) gives for plot 6 earlier,
species = 6, individuals == 496. I1 = 0.97931
plot 6 later,
species = 6, individuals := 443, /I = 1.02472.
To test for a difference in diversity we have
1.02472 - 0.9793 1
= 0.8973
’ = (0~0012556+0~0013055)“2
with degrees of freedom large. Thus, we cannot reject the hypothesis of equal
diversity.

I would like to acknowledge the help of Professors L. R. Shenton, J. B. Douglass

and M. C. Carter.

Department of Statistics and Institrrte of Ecology, KERMIT HUTCHESON

University of Georgia,
Athens, Georgia, U.S.A.

(Received 27 February 1970)

154 K. HUTCHESON

REFERENCES
BOWMAN, K. O., HUTCHESON, K., ODUM, E. P. & SHENTON, L. R. (1969). Znrernational
Symposium on Stutisticd Ecology. Vol. 3. University Park: Pennsylvania State University
Press.
HUTCHESON, K. (1969). Ph.D. Dissertation.
PIELOU, E. C. (1966). J. theor. Biol. 10, 370.
STODDARD, H. L. & NORRIS, R. A. (1967). Bull. Tall Timbers Res. Stn 8, 1.