RAREFACTION CURVES
vided by Heck et al. [7], who also considered
BACKGROUND
When one samples information about a
population of people, machines, butterflies,
stamps, trilobites, and nearly anything else,
it is a natural tendency that as more objects
are obtained, the number of distinct kinds
of objects increases. Rarefaction is a sam-
pling technique used to compensate for the
effect of sample size on the number of groups
observed in a sample and can be important in
comparisons of the diversity of populations.
Starting from a sample of units classified into
groups, the rarefaction technique provides
the expected number of groups still present
when a specified proportion of the units are
randomly discarded. In this way a large sam-
ple can be rarefied, or made smaller, to
facilitate comparison with a smaller sample.
For example, suppose that I spend two
months collecting specimens and find 102
distinct species represented among the 748
individuals collected. If you then spend a
week collecting specimens at another loca-
tion and find only 49 species among 113
specimens, can we conclude by comparing
your 49 to my 102 species that your popula-
tion was less diverse than mine? Of course
not. We need to correct for sample size to do
a proper comparison because if you had col-
lected for a longer time, you would probably
have obtained a larger number of samples
and of species. Applying the techniques of
rarefaction to the detailed data (the method
needs to know how many individuals are in
each species), the rarefied number of species
in my sample might turn out to be 50.14
species for subsamples of size 113. When com-
pared to your count of 49 species, we could
then conclude that the population diversities
are not very different.
A brief history of rarefaction begins with
work by Sanders [12], who developed a
technique to compare deep-sea diversity to
shallow-water habitats. Problems of overesti-
mation were noted by Hurlbert [8], Fager [3],
and Simberloff [14], with an improved formu-
lation given by Hurlbert and by Simberloff.
A formula for the variance of the number of
Encyclopedia of Statistical Sciences, Copyright 2006 John Wiley & Sons, Inc.
groups present in a rarefied sample was pro-
the determination of sufficient sample size
for data collection .
Sampling properties of the rarefaction
measure were explored by Smith and Grassle
[15]. Rarefaction has been applied exten-
sively in the study of diversity throughout
the fossil record by Raup [10,11] and by oth-
ers. Some criticisms and suggestions relating
to the application of rarefaction methods
were made by Tipper [18]. Upper and lower
bounds on rarefaction curves were devel-
oped by Siegel and German [13]. Because
it is based on sampling from the data, rar-
efaction is related to the bootstrap method
of Efron [1]; this connection is explored by
Smith and van Belle [16].
Rarefaction is related to the ideas of diver-
sity and evenness in populations. A recent
overview of diversity measurement may be
found in Patil and Taillie [9] with discussion
by Good [4] and Sugihara [17]. Rarefaction
may be considered as an interpolation pro-
cess, compared to the more difficult problem
of extrapolation as considered by Good and
Toulmin [5] and by Efron and Thisted [2], in
which the goal is to estimate the number of
additional groups that would be observed if a
larger sample could be obtained.
DEFINITION AND PROPERTIES OF
RAREFACTION
Suppose that we have a situation in which
N items are classified into K groups in such
a way that each item is in exactly one group
and each group contains at least one item.
For example, the items might be individual
specimens that have been collected and the
groups might represent the various species
present; for analysis at a higher taxonomic
level, the items might be species grouped
according to genus.
To describe the situation completely, let
the number of items in group i be denoted Ni .
1
2 RAREFACTION CURVES
The data may be described as follows:
N = total number of items
K = total number of groups
Ni = number of items in group i
(i = 1, . . . , K).
To facilitate computation, we will define Mj
to be the number of groups containing exactly
j units (j
1):
Mj = number of Ni equal to j.
From these definitions, it follows that


K




Ni = N, Mj = K,
i=1 j=1
Now consider a rarefied sample, con-
structed by choosing a random subsample of
n from N items without replacement. Some
of the groups may be absent from this sub-
sample. Let Xn denote the (random) number
of groups that still contain at least one item
from the rarefied sample:
Xn = number of groups still present
in a subsample of n items.
It must be true that Xn  K with strict
inequality whenever at least one group is
missing from the rarefied sample.
The rarefaction curve, f(n), is defined as
the expected number of groups in a rarefied
sample of size n, and can be computed in
several ways:
 1

K  
  N N Ni
f (n) = E Xn =K
n n
i=1

  1
Nj N
=K Mj . 2. Efron, B. and Thisted, R. (1976). Biometrika,
n n
j=1
It is always true that 0  f (n)  K, f (0) =
0, f (1) = 1, and f (N) = K. Moreover, f is mo-
notone increasing and concave downward.
Because these binomial coefficients can
become large and overflow when computers
are used, it is preferable to compute directly
with the ratio of the two coefficients, which is
always between 0 and 1. This ratio may easily
be updated using a multiply and a divide to
obtain successive terms:
   
N (j + 1) N 1
n n
   
Nnj N j N 1
= .
Nj n n
The rarefaction values f (n) are often dis-
played as a continuous curve even though
they are actually discrete values. Consider,
for example, the rarefaction curve for N =
748 units (species) within K = 102 groups
(families) of bivalves from Siegel and Ger-
man [13], Fig. 1. (Data were collected by
Gould and are described in ref. 6.)
jMj = N.
j=1
SAMPLING PROPERTIES
In many situations it is more realistic to sup-
pose that the observed values of items and
groups are not fixed but instead represent a
sample from a multinomial population. The
expected number of groups represented in a
sample of n items from this population can be
used as a measure of the population diversity.
Based on the observed data, the rarefaction
curve value f (n) can be used as an estimate
of this population diversity measure. Within
this context, Smith and Grassle [15] have
proven that the rarefaction value is a mini-
mum variance unbiased estimate (MVUE).
They also provide an unbiased estimate of
the variance of the estimate which takes into
account the sampling variability of the pro-
cess that generated the data.
REFERENCES
1. Efron, B. (1982). The jackknife, the Bootstrap,
and Other Resampling Plans. SIAM, Philadel-
phia.
63, 435448.
3. Fager, E. W. (1972). Amer. Naturalist, 106,
293310.
4. Good, I. J. (1982). J. Amer. Statist. Ass., 77,
561563.
5. Good, I. J. and Toulmin, G. H. (1956).
Biometrika, 43, 4563.
6. Gould, S. J., Raup, D. M., Sepkoski, J. J.,
Schopf, T. J. M., and Simberloff, D. S. (1977).
Paleobiology, 3, 2340.
 RAREFACTION CURVES 3

Figure 1.

7. Heck, K. L., Jr., van Belle, G., and Sim-

berloff, D. S. (1975). Ecology, 56, 14591461.
8. Hurlbert, S. H. (1971). Ecology, 52, 577586.
9. Patil, G. P. and Taillie, C. (1982). J. Amer.
Statist. Ass., 77, 548561, 565567.
10. Raup, D. M. (1975). Paleobiology, 1, 333342.
11. Raup, D. M. (1979). Science, 206, 217218.
12. Sanders, H. L. (1968). Amer. Naturalist, 102,
243282.
13. Siegel, A. F. and German, R. Z. (1982). Bio-
metrics, 38, 235241.
14. Simberloff, D. S. (1972). Amer. Naturalist,
106, 414418.
15. Smith, W. and Grassle, J. F. (1977). Biomet-
rics, 33, 283292.
16. Smith, E. P. and van Belle, G. (1984). Biomet-
rics, 40, 119129.
17. Sugihara, G. (1982). J. Amer. Statist. Ass., 77,
564565.
18. Tipper, J. C. (1979). Paleobiology, 5, 423434.

See also BOOTSTRAP; DIVERSITY INDICES; ECOLOGICAL STATISTICS;

and LOGARITHMIC SERIES DISTRIBUTION.

ANDREW F. SIEGEL