Measurement of Faunal Similarity in Paleontology

Author(s): David M. Raup and Rex E. Crick

Source: Journal of Paleontology, Vol. 53, No. 5 (Sep., 1979), pp. 1213-1227
Published by: SEPM Society for Sedimentary Geology
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 JOURNALOF PALEONTOLOGY,V. 53, NO. 5, P. 1213-1227, 9 TEXT-FIGS., SEPTEMBER 1979

MEASUREMENT OF FAUNAL SIMILARITY IN

PALEONTOLOGY

DAVID M. RAUP AND REX E. CRICK1

Department of Geology, Field Museum of Natural History, Chicago, Illinois 60605 and
Museum of Paleontology, University of Michigan, Ann Arbor, Michigan 48109

ABSTRACT-A probabilistic index of faunal similarity is proposed which compares the number of taxa
common to two faunas with the number that would be expected to be in common if the taxa were
distributed randomly. Departures of observed from expected numbers in common express the level
of similarity or dissimilarity. The frequency of taxa in the whole data set is used to adjust for the
differing probability of occurrence of taxa (cosmopolitan versus endemic). The new index can be used
to determine whether similarities or dissimilarities between faunas are statistically significant.
The index is tested with 1) modern biogeography of echinoids, 2) environmental distribution of
modern foraminifera in Santa Monica Bay, and 3) Ordovician biogeography of nautiloids. In each
case, the proposed index is more effective than traditional indexes of faunal similarity (Simpson,
Jaccard, and Dice coefficients) in addition to the advantage of making possible rigorous assessment
of statistical confidence. The index should also be useful in a biostratigraphic context. The computer
program used for calculating the index is available from the authors.

INTRODUCTION applied in evaluating the sizes of the assem-

A COMMON PROBLEM throughout paleoecolo-
gy, biostratigraphy, and paleobiogeography is
that of comparing faunal lists to evaluate their
similarities and differences. If two lists have
no taxa in common, it can be assumed that
something was different. The possible causes
vary from ecological differences (marine vs.
fresh water; shallow vs. deep water, etc.) to
temporal differences (complete evolutionary
turnover) to biogeographic differences (provin-
ciality, separation by geographic barriers,
etc.). If the two lists are identical, on the other
hand, ecological, temporal, and biogeographic
unity is assumed (at some scale, at least). The
problems arise when some but not all taxa are
shared. Ordinarily, two rather different ap-
proaches are taken in this intermediate situa-
tion: 1) assessment of similarity on the basis of
well informed intuition, and 2) computation of
a numerical index of similarity.
The intuitive approach is not without value.
The experienced practitioner recognizes that
some taxa are more common than others and
gives less weight to their joint occurrence in
two or more assemblages. Many paleobiogeog-
raphers, for example, discount cosmopolitan
forms. In biostratigraphy, the joint occurrence
of long-ranging taxa is given much less weight
than that of short-ranging taxa. Judgment is
I Present address: Dept. Geology, Univ. Texas,
Arlington 76019.
Copyright ? 1979, The Society of Economic
Paleontologists and Mineralogists
blages being compared. For example, if two
assemblages have 15 taxa each, the presence
of 10 taxa in common clearly indicates a great-
er fundamental similarity than if the two as-
semblages had had 100 taxa each. Questions
and ambiguities occur, however, and these
have given rise to attempts to quantify the pro-
cess of comparison.
Many quantitative measures of faunal sim-
ilarity have been proposed and several are in
common use by paleontologists. Perhaps the
one most widely applied is the Simpson Coef-
ficient (Simpson, 1943, 1947) which may be
defined as follows:
S = 100k/B,
where: k = the number of taxa common to
two assemblages A and B,
B = the total taxa found in the
smaller assemblage (B < A).
The Simpson Coefficient varies from zero to
100 and thus implies percentage similarity.
Although it is an easily managed index, au-
thors have pointed out (most recently, Hen-
derson and Heron, 1977), that the Simpson
and comparable indexes have important short-
comings. Text-figure 1 illustrates one of the
prime difficulties. A series of Venn diagrams
shows several cases which yield identical val-
ues of 'S' yet which are intuitively very differ-
ent. In each case, the area of the largest circle
1213 0022-3360/79/0053-12
13$03.00

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1214 DAVID M. RAUP AND REX E. CRICK

20 20
.2011 .02
.20 .04

1. 2.

;= 20 20
= .11 .11
)= .20 .20

3. 4.

SIMPSON = ro100 JACCARD= -k DICE = 2k

B A+B- k A+B
TEXT-FIG. 1-Venn diagrams showing hypothetical cases wherein two faunal assemblages (A and B) are
drawn from a pool of taxa (N). The number of taxa (k) common to A and B is indicated by the overlap
of the two smaller circles. For each case, the Simpson, Jaccard, and Dice similarity measures have been
calculated.

indicates the total pool (N) of taxa which could
occur in an assemblage and the two smaller
circles (A and B) are assemblages drawn from
this pool. The overlap zone (k) between the
smaller circles represents the number of taxa
found in both assemblages. In all cases, the
Simpson Coefficient is 20 yet one's intuition
suggests that the four cases do not indicate the
same similarity in the sense of process (mean-
ing ecological, temporal or geographic similar-
ity).
In recognition of these and other difficulties,
many authors have proposed alternate
schemes. Two of the coefficients most com-
monly applied in paleontology are the Jaccard
and Dice. Cheetham and Hazel (1969) have
provided an excellent comparative review of
these and about 20 other similarity coeffi-
cients. Other critical reviews of selected coef-
ficients exist in the literature (see, for example,
papers by Henderson and Heron, 1977, and
Simberloff, 1978). Most of these authors have
emphasized that it is important to have valid
measures of faunal similarity because of the
increasing use of multivariate statistical anal-
ysis of large masses of distributional (presence/
absence) data. The multivariate analysis can
be only as good as the matrix of similarity val-
ues that forms its input!
At the risk of oversimplification, one can

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 FAUNAL SIMILARITY
argue that most existing similarity coefficients
suffer from two main problems. First, they
have not been derived in a mathematically rig-
orous way: that is, they have been 'thought
up' rather than built on sound mathematical
principles. Their validity has all too often been
tested by whether they seem to work in prac-
tice. Second, they have not been tied to clearly
defined null hypotheses; as a result, statisti-
cally meaningful comparisons between values
of a coefficient are impossible. It has been im-
possible to say whether two assemblages are
similar (or dissimilar) at the 95% level of con-
fidence, for example. The discussion by Sim-
berloff (1978) includes a particularly good
treatment of this point.
Henderson and Heron (1977) recognized
and discussed many of the problems just de-
scribed and made an attempt to produce a rig-
orous and statistically valid similarity mea-
sure. The present effort takes a slightly
different tack in the hope of developing a yet
more robust approach to the similarity ques-
tion. Our approach is similar to that of Sim-
berloff (1978), but our objectives and the re-
sulting technique are substantially different.
THE APPROPRIATE NULL HYPOTHESIS
Suppose that taxa are sprinkled randomly
in space and time and that species lists are
made up from the taxa that happen, by
chance, to fall in certain areas and in certain
stratigraphic intervals. Most of the species lists
will differ from one another just because of
the vagaries of sampling but they will have an
average similarity which is predictable from
the numbers of taxa, areas, and stratigraphic
intervals involved. As will be shown below,
it is possible under the random sprinkling hy-
pothesis to predict how many species should
be expected to be shared ('k') and the expected
variation in this number. The expected 'k' and
its probable variation constitute the appropri-
ate null hypothesis for assessing faunal simi-
larity.
In the real world, the distribution of taxa in
space and time is generally non-random. Tri-
lobites are confined to a small portion of geo-
logic history (the Paleozoic), echinoderms are
confined to marine environments, and so on.
When we use the temporal confinement of tri-
lobites or the ecological confinement of echi-
noderms to make other interpretations, we are 1 and assume, as before, that the areas of the
tacitly rejecting the null hypothesis of random circles correspond to the numbers of species in
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1215
sprinkling. In the trilobite and echinoderm ex-
amples the departure from chance expecta-
tions is so obvious that sophisticated statistical
testing is unnecessary. But most cases of in-
terest are more subtle and rigorous treatment
is obligatory-which is to say that one cannot
rely on intuition alone.
In actual cases, it makes no difference
whether the null hypothesis of randomness is
rejectable 10% or 90% of the time. We wish
to use it only as a standard of comparison and
as a means of assessing the probability that
two assemblages had different ecologic, tem-
poral, or geographic settings. When dealing
with assemblages from radically different fa-
cies or from different continents, one would
expect to be able to reject the null hypothesis
most of the time. On the other hand, when
dealing with assemblages from the same for-
mation in a local area, one would expect not
to reject the null hypothesis and to conclude
that the compositional differences between as-
semblages are just the result of chance differ-
ences in sampling.
We propose to use a comparison between
the observed number of taxa common to two
assemblages (or faunas) and the probability
distribution of the expected number of com-
mon taxa as a measure of the similarity of the
two assemblages. Assemblages which are
more similar than predicted by the null hy-
pothesis will be interpreted as indicating a pos-
itive bias in the make-up of the assemblages.
That is, ecologic, temporal, or geographic fac-
tors must have limited the taxa available for
those assemblages. Conversely, assemblages
less similar than predicted will be interpreted
as indicating a negative bias.
Simberloff (1978) had a quite different ob-
jective. Working with modern species distri-
butions in the Galapagos Islands, he was ask-
ing whether the total distribution represents
a significant departure from the null hypoth-
esis of random sprinkling. That is, he was ask-
ing whether the array of species lists is consis-
tent with the proposition that differences in
composition result solely from sampling error
(in dispersal of species) and not from real bio-
geographic effects.
METHODS
Consider the Venn diagrams in Text-figure
1216 DAVID M. RAUP AND REX E. CRICK

.8
>..
I. 2.
_ .6 .6'
_J
-4

<:.4 .4
(a
0
a. .2 .2

0 0
0 1 2 3 4 5 0 3 4 5
kexp
.4 .4
3. 4.
.3 .3

.2 .2

.1 .1. k bs
i I
0 f I
f
mt
_ - - i.

0 5 10 15 20 0 5 10 15 20
TEXT-FIG. 2-Curves showing solutions to equation (4) for all possible values of 'k' in the four cases
illustratedin Text-figure1. The point markedkobsis the numberof taxa observedin commonin Text-
figure1. Note that the relationshipsare not continuousfunctions:only integervaluesof 'k' are possible.

the total pool and two assemblages drawn
from that pool. Assume further that all species
in the pool have an equal chance of being cho-
sen for each of the assemblages. This is a sim-
plistic assumption because it is well known
that species vary in their abundance so that
some have a much higher probability of oc-
curring in any given assemblage than others.
But this is a convenient scenario with which
to introduce a methodology and is the one used
by Henderson and Heron (1977) and by Sim-
berloff (1978).
The situation just presented can also be
thought of in the classic context of an 'urn
problem.' Assume that a large urn contains
many balls, each numbered differently, and
that this collection of balls constitutes the pool
of species (N). Now draw 'A' balls at random
from the urn to define assemblage A. Then
replace them and draw 'B' balls to form as-
semblage B. The question is: how many of the
same balls will be found in both A and B?
This problem was solved by Henderson and
Heron (1977) by a logical series of steps cul-
minating in their equation (4) and in a slightly
different form by Simberloff (1978), (Null Hy-
pothesis I). The solution presented here is
more straightforward and more flexible than
either of the previous efforts.
The total number of different 'A' assem-
blages that can be drawn from the pool (N) is
the number of combinations of N things taken
A at a time, or
NcA N= - (1)
(N A)! A!(

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 FAUNAL SIMILARITY 1217

TABLE 1-Probabilities calculated from equation (4) for the four cases shown in Text-figure 2: 'kobs' is the number of
species observed to be in common and 'kexp'is the number expected to be in common on the assumption of random
sprinkling of species.

Cases:
1 2 3 4
Probability that kexpis less than kobs: .77 .07 .39 .005
Probability that kexpequals kobs: .21 .26 .24 .012
Probability that kexpexceeds kobs: .02 .67 .37 .983
1.00 1.00 1.00 1.000

Similarly, the number of possible 'B' as- Prob [k species in common] =

semblages is NCB. Therefore, the total num- A! B! (N - A)! (N - B)!
ber of different pairs of assemblages is the N! k! (A - k)! [(N - B) - (N - k)]! (B -
k)!'
product:
(4)
NCA-NB. For any set of A, B, and N values, this equa-
The probability that some particular number tion can be solved for the series 'k' values rang-
of species ('k') will occur in both assemblages ing from 0 to B so that a precise probability
may be expressed as follows: distribution can be developed for variation in
the expected number of species in common.
Prob [k species in common] = This is illustrated in Text-figure 2 for the four
total ways of obtaining k species in common cases shown in Text-figure 1. The equation
total number of different assemblage pairs has also been applied to the cases treated by
Henderson and Heron (1977, fig. 3) and the
(2)
results are comparable though not identical to
The denominator in this expression is the theirs.
product developed above. The numerator can In Text-figure 2, the values of 'k' observed
be constructed by inspection of the Venn in Text-figure 1 are indicated by 'kobs'and the
diagrams in Text-figure 1, as follows: all several theoretically possible values of 'k' by
species in the 'k' area must also belong to The ordinate is the probability of each
'B' which, in turn, must belong to 'N.' There- 'kexp.'
particular kexpoccurring by chance. Note that
fore, all possible compositions of the 'k' area the distributions are of markedly different
can be expressed by: shapes depending on the size of the pool and
NCB B k. the sizes of the two assemblages. The distri-
butions are often highly skewed, in contrast to
There remain only the species that belong to the examples shown by Henderson and Heron.
'A' but not to 'k.' All possible compositions of In fact,
skewing is probably typical of real
this group can be expressed by: world data because N is generally much larger
than A or B.
(N - B)(A - k).
The numerical data from Text-figure 2 are
The numerator of the probability expression is summarized in Table 1. For Case 1, there is
thus the product: only a .02 probability of finding more than the
- B)C(A - k). observed number of species in common follow-
NC'Bck(N
ing the hypothesis of random sprinkling. For
The terms for NCB in numerator and denomi- Case 4, on the other hand, the observed value
nator cancel and we are left with: will be exceeded by chance more than 98% of
Prob [k species in common] = the time. These relations can be used to define
a similarity measure, as follows:
BCk(N - B)C(A -k) = 1 minus the prob-
INDEXOFSIMILARITY
NCA
ability that kexp
When this is evaluated using factorials as in will be greater
equation (1), the result is: than kobs.

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1218 DAVID M. RAUP AND REX E. CRICK
This is equivalent to:
INDEX OF SIMILARITY= the probability
that kexp will
be less than or
equal to kobs.
The four values of the INDEX are thus .98,
.33, .63, and .02, respectively.
At this point, we can ask if any of the above
figures are statistically significant in the sense
that the null hypothesis of random sprinkling
can be rejected. Questions concerning statis-
tical significance must be framed carefully. It
is tempting to say that the INDEX OF SIMI-
LARITY in Case 1 allows for the rejection of
the null hypothesis with 98% confidence. But
note that the observed number in common in
Case 1 is expected to occur 21% of the time
under the null hypothesis so that the existence
of this number in common is not startling. In
Case 4, on the other hand, a 'k' greater than
that observed is expected more than 98% of
the time and we can conclude that the null
hypothesis can be rejected. We can go further
and say that because kobs is significantly low,
the two assemblages are significantly dissimi-
lar-which is to say that something influenced
the selection of species from the pool such that
fewer than expected occur in common. None
of the other cases show significant departure
from chance expectations at the 95% level of
confidence. A significant similarity would be
represented by a number greater than or equal
to 0.95 in the first row of Table 1. Thus, the
INDEXOF SIMILARITY as defined here cannot
be used as a direct test of statistical signifi-
cance but the data contributing to it can be so
used.
The foregoing scheme is unfortunately not
appropriate for general application because it
uses the simplifying assumption that all
species are present in equal numbers in the
pool and thus have an equal chance of occur-
ring in any assemblage. When equation (4) is
used with actual assemblage pairs, the ob-
served 'k' values are usually much higher than
would be expected. Most assemblages appear
to be significantly similar to each other. This
is because most related faunas contain a few
common, nearly ubiquitous species which
have the effect of elevating the observed 'k'
values. This problem can be avoided by ex-
plicitly accounting for differences in the rela-
tive probability of each species occurring in an
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assemblage. In other words, the species pool
can be constructed with species of differing
frequencies. This is analogous to an urn which
has more of some kinds of balls than of others.
Simberloff (1978) solved this problem in the
formulation of his Null Hypothesis II. He used
the actual frequencies of species in a region as
an explicit measure of the probability of these
species appearing in any one assemblage
drawn from that region. We will follow the
same approach.
The formation of the pool can be illustrated
by using one of the examples of actual data
that we will employ later in this paper as a
test of the methodology. We will use data on
the present biogeography of the 222 genera of
living echinoid echinoderms. Their distribu-
tion is expressed in terms of their presence or
absence in 40 sampling areas in the present-
day oceanic world. Some of the areas are ar-
bitrarily defined and some are based on tra-
ditional biogeographic divisions. The basic
data set thus consists of lists of genera for each
of the 40 sampling areas. Some genera are en-
demic to a single area while others are found
in many areas. In this data set, 60 of the 222
genera are endemics and the most 'cosmopol-
itan' genus is found in 20 of the 40 areas. The
number of genera occurring in a given area
ranges from 1 to 120.
We will assume, a la Simberloff, that the
probability of occurrence of a genus in a sam-
pling area is directly proportional to the num-
ber of areas in which that genus occurs. There-
fore, a genus which occurs in only one area is
seen as having a probability of 1/40 of occur-
ring in any given area. A genus which is found
in 10 areas has a probability of 10/40 of oc-
curring, and so on. There is a definite hint of
circularity in this reasoning but Simberloff
(1978) has presented convincing arguments for
the lack of significant circularity. The main
point is that the reasoning does not make any
demands on the spatial distribution of occur-
rences within the whole region under study: it
does not preclude a concentration of occur-
rences in one part of the region. Therefore, the
null hypothesis of random sprinkling is a valid
null hypothesis and can be falsified.
The pool of taxa from which local faunas
are formed has as many occurrences of each
genus as there are occurrences of that genus
in the total data set. When assemblages are
selected at random from such a pool, the cos-
 FAUNAL SIMILARITY
mopolitan genera are more likely to occur and
are thus more likely to be genera common to
both members of a pair of assemblages. Local
endemics (those with probabilities of 1/40, in
the echinoid case) can occur in two assem-
blages but the probability of this event is low.
One would naturally like to be able to derive
an equation equivalent to equation (4) which
would predict values of kexpunder the condi-
tions described above. We have been unable
to derive this equation. Therefore, we have
had to rely on monte carlo simulations-just
as Simberloff did for his purposes. Our meth-
od is as follows:
1) For each pair of assemblage sizes in the
real world data set construct an imaginary pair
of assemblages (A and B) by drawing species
from the pool. This is accomplished by com-
puter with a random number generator. Hav-
ing made the two assemblages, the lists of gen-
era are compared and the number of genera
in common is recorded. This number is one
outcome of sampling under the random sprin-
kling hypothesis: that is, one point in a kexp
probability distribution.
2) The same procedure is repeated many
times with the number of taxa shared by each
pair of assemblages being recorded.
3) A frequency distribution of the results is
an estimate of the probability distribution of
kexpunder the specified conditions of A and B.
4) The number of taxa actually shared (kobs)
by assemblages of these sizes in the real world
is compared with the monte carlo generated
distribution and the INDEXOF SIMILARITY is
computed as in the simplified case described
earlier.
An actual example of this procedure is il-
lustrated in Text-figure 3 for a pair of sam-
pling areas in the echinoid data set: these areas
had 19 and 36 genera, respectively. A fre-
quency distribution of 50 simulated assem-
blage pairs is shown in Text-figure 3 along
with the actual number observed in common
(5). In this case, kobsfalls nearly at the center
of the simulated distribution and the null hy-
pothesis cannot be rejected. But the percent-
age of simulations having 'k' less than or equal
to kobs may be used as the INDEX OF SIMILAR-
ITY.
When this procedure was followed with the
entire echinoid data set, most cases fell be-
tween the 5% tails of their distributions (as in
Text-fig. 3).
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1219
co 30
20- \ A= 36
L.
0
B 19
0 5 10 15 19
kexp
TEXT-FIG. 3-Example of treatmentof a compar-
ison between two echinoid faunas. kohs is the
numberof generaactuallyobservedto be in com-
mon. The curveshowsthe percentof simulations
yielding each value of kexp.The ruled portion
indicates the numberof simulationshaving 'k'
values less than or equal to the observedvalue.
A special problem arises where the smaller
assemblage (B) is very small. In one echinoid
pair, for example, assemblage B contained
only two genera and thus the only possible
values of 'k' are 0, 1, and 2. Fifty simulations
produced the following k's:
k=0 40 (80%),
k= 1 9 (18%),
k= 2 1 (2%).
50
There were no genera actually common to the
two areas (kobs= 0). Thus, using the proce-
dure described earlier, computing the INDEX
would yield a value of 0.80. But this implies
a higher similarity than may exist. In other
words, we do not know where the value of
kobs falls within the 80%. Therefore, an arbi-
trary convention was adopted: the INDEX is
computed on the basis of the midpoint of the
string of simulated 'k' values which are equal
to the observed 'k.' The INDEXin this case is
recorded as 0.40. The same convention was
followed throughout. In the case illustrated in
Text-figure 3, the INDEXwas recorded as 0.39.
This was arrived at by summing the percent-
ages of the simulations that gave 'k' values less
than kobs (2 + 6 + 20 = 28) and adding one-
half the percentage of simulations where 'k'
equaled kobs(?2 x 22 = 11).
1220 DAVID M. RAUP AND REX E. CRICK

The use of monte carlo methods calls for distributional data all come from Mortensen's
considerable computation time-much more Monograph of the Echinoidea (1928-51)
than would be required if an analytical expres- which provides a consistent and authoritative
sion comparable to equation (4) were avail- taxonomic base. Of the forty geographic sam-
able. But the results are just as accurate, given pling areas used for the present study, most
enough simulations. In the echinoid case we are relatively shallow water coastal or insular
used 50 simulations for each pair of assem- areas where distributions of taxa tend to re-
blage sizes. 100 or 1,000 simulations per pair flect regional climate. The others have uni-
would have produced more precise distribu- formly cold water faunas: the non-insular
tions but 50 was chosen as the best compro- ocean areas of the North Pacific, South Pacif-
mise with the limitations of computer budgets. ic, North Atlantic, Central Atlantic, and
The important point is that the simulation South Atlantic and the Arctic and Antarctic
technique does not sacrifice rigor unless the Oceans.
number of simulations is too low. As indicated earlier, the data set consists of
The computer program used for the echi- 222 genera which range from local endemics
noid and other analyses is available from the to those found in as many as 20 of the 40 sam-
authors. It is a relatively expensive program pling areas. In keeping with the philosophy of
to run. The cost depends on the number of the method, no data were discarded because
assemblages and the variation in their sizes. of endemism or cosmopolitanism and no areas
The echinoid data set described here is unusu- were excluded because of small sample size.
ally large and requires about 26 cpu minutes The basic computer program was run to as-
on an IBM 360/65 to produce the similarity sess similarity between the members of all pos-
matrix plus a complete record of the 19,750 sible pairs of the 40 generic lists. Fifty simu-
simulations required for the job. A variety of lations were used for each pair of assemblage
techniques could be used to reduce the cost sizes. The output consisted of 1) the tabulated
but they would sacrifice accuracy. results of all simulations (number of genera in
common) and 2) a matrix of values of the com-
APPLICATIONS
puted INDEX OF SIMILARITY. Various analy-
The method described in this paper can be ses were performed on the output, some of
applied to any data set consisting of presence which will be described below.
and absence of taxa. In other words, any sit- Text-figure 4 shows how one of the sam-
uation which yields floral or faunal lists is ap- pling areas compares with the other thirty-
propriate. Each list may represent a single col- nine. The reference area (marked by an 'X')
lecting locality or a composite of information on the west coast of Central America was cho-
from a group of geographically, ecologically or sen arbitrarily and other choices produce com-
stratigraphically related localities. parable results. Values of the INDEX OF SIM-
In order to test the methodology, we will ILARITYare contoured and show decrease in
present three quite different examples: 1) glob- similarity with distance from the reference
al biogeography of living echinoid echino- area. Contouring was straightforward; that is,
derms, 2) distribution of benthic foraminifera extreme contortion of contour lines was not
in Santa Monica Bay, California, and 3) global necessary. Furthermore, the resulting pattern
biogeography of Ordovician nautiloid cepha- is plausible and interpretable in biogeographic
lopods. It should be emphasized that the pro- terms. The map shows clearly that the echi-
posed INDEX OF SIMILARITY, like all other noids of the Eastern Pacific are much more
similarity measures, is a purely descriptive similar to those of the Western Atlantic than
tool. Its purpose is to measure similarities and to those of the Western Pacific and Indian
differences between taxonomic lists and to as- Ocean regions. In fact, the presence of the
sess the statistical significance of these simi- Central American barrier is not evident in the
larities and differences. It does not interpret pattern. (This would not be the case at the
the results in the sense of telling us the biolog- species level where virtually no echinoid
ical or geological factors responsible for the species are common to both sides of the Isth-
similarities or differences. mus of Panama.) While some details of the
Echinoid biogeography.-The data set used pattern may reflect sampling error, there is no
for this test was presented briefly above. The reason to believe that Text-figure 4 is not a

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 FAUNAL SIMILARITY 1221

TEXT-FIG.4-Analysis of echinoidbiogeographicdata. Dotted outlinesseparatethe samplingareas:the

numbersindicatethe INDEXOF SIMILARITYof each area with respectto an arbitrarilychosenreference
area(markedwith X: Pacificcoastof CentralAmerica).Numbersenclosedin circlesindicatesignificant
similarity to the referencearea; numbersin boxes indicatesignificantdissimilarityto the reference
area. Intermediatevalues of the INDEXare contoured.

valid description of generic distribution in this faunas in a rigorous probabilistic way. In

group of echinoderms.
Similar plots have been made using the
Simpson, Dice, and Jaccard similarity index-
es. The results are approximately the same
although contouring was more difficult. (In
particular, the Simpson Coefficient data con-
tained several unexplained anomalies.) The
good results produced by the conventional in-
dexes are not surprising in view of the ex-
tremely high quality of Mortensen's distribu-
tional data and the basic simplicity of echinoid
biogeography at the generic level.
If the production of a map such as Text-
figure 4 were the only purpose, the other in-
dexes would probably be adequate and the
saving in computation time would be substan-
tial. But the similarity measure proposed here
allows one to evaluate differences between
Text-figure 4, seven of the similarities are sig-
nificantly high (at the 95% level of confidence):
these are the areas whose INDEX values are
circled. In these cases, the number of genera
observed to be in common is greater than in
at least 48 of the 50 simulations. The values
of the INDEXcontained in squares are signif-
icantly low: that is, the number of taxa ob-
served in common is less than in at least 48 of
the 50 simulations. The distribution of circled
and boxed similarity values is the expected
one. The intermediate cases indicate a prob-
ability of faunal similarity but do not lead to
the rejection of the null hypothesis. We can
say, for example, that the Alaskan echinoids
appear to be different from those of Central
America but the difference is not statistically
significant and thus could be caused by chance

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1222 DAVID M. RAUP AND REX E. CRICK

_ _____ _____

A:
COASTAL
)
J
TROPICAL/SUBTROPICAL 0 /
INDO-PACIFIC /
NORTHATLANTIC
I.
U 0

SEA OF JAPAN

SEA OF JAPAN
0

SOUTH PACIFIC 0 SEA OF OKHOTSK

0

0 ANTARCTIC

PC I
TEXT-FIG.5-Multivariate analysisof echinoidbiogeographicdata. The firsttwo principalcomponents
(PCI and PCII)are plottedfor the 40 samplingareas. PCI separatescoastalareas of the Indo-Pacific
from otherareasand fromthe cold water, open ocean areas. PCII reflectswatertemperature.

sampling error. The orderliness of the contour
lines strongly suggests, of course, that the
Alaskan and Central American echinoids are
in fact different in the sense that they do not
represent random sprinkling from the same
pool.
Text-figures 5 and 6 show 2-dimensional
ordination plots of the first three principal
components axes representing 98.5 percent of
the variation in the data set. The principal
components, PCI, PCII, PCIII, account for
51, 28, and 19.5 percent of the variation, re-
spectively. The sampling areas which form
tight, natural groups are shown as solid dots
and the groups are labeled. Others are shown
as open circles and identified individually.
Text-figure 5 is an ordination of PCI and
PCII. PCI clearly separates the coastal areas
of the Indo-Pacific region from those of the
Eastern Pacific and the Atlantic. This sepa-
ration is the result of the East Pacific Barrier
(Ekman, 1953), an 1,810 km expanse of open
ocean separating the islands of Outer Polyne-
sia and the tropical/subtropical coast of Amer-
ica. Under ordinary oceanic conditions, echi-
noid larvae are not capable of crossing this
barrier and faunas on either side of the barrier
are significantly different below the family
level. Separation of the South Australian and
New Zealand regions from the tropical/sub-
tropical regions of the Indo-Pacific illustrates
cold-temperate character of the South Austra-
lian and New Zealand faunas. Although geo-
graphically proximal, the echinoid faunas of
the Sea of Japan and the Sea of Okhotsk are
remarkably different. The echinoid fauna of
the Sea of Japan consists of shallow water,
warm-temperate genera derived from the sub-
tropical Indo-Pacific via the warm Kuroshio
Current while the echinoid fauna of the Sea of

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 FAUNAL SIMILARITY 1223

Il -

ATLANTIC

INDIAN OCEANO

IU NORTHATLANTIC ARCT \

0
CENTRALATLANTIC0O SOUTH ATLANTIC
SEA OF OKHOTSK
Q
^ \~~~ SOUTH PACIFIC
S
?* OSEA OF JAPAN
/COASTAL
TROPICAL/SUBTROPICAL COASTAL
INDO-PACIFIC * \ C APACIFIC
EASTERN

( C NORTHPACIFI ORH PACIFIC 0

NORTH PATAGONIA

COASTAL ANTARCTIC
WESTERNATLANTIC

PC I
TEXT-FIG. 6-Multivariate analysisof echinoidbiogeographicdata. The first and third principalcom-
ponents(PCI and PCIII)are plotted for the 40 samplingareas. PCIII serves to separatethe coastal
areasof the EasternPacificand Atlanticinto distinctregions.

Okhotsk consists of cold-temperate genera de-
rived from the north via the cold Oyashio Cur-
rent. Any chance mixing of the faunas is re-
duced by a shallow submarine sill separating
the two bodies of water. Deep water and high
latitude faunas tend to cluster in the lower
right corner of Text-figure 5. Text-figure 6
shows that the coastal areas of the Eastern
Atlantic, Western Atlantic, and Eastern Pa-
cific are separated along PCIII. Naturally, sta-
tistical significance cannot be assessed in the
results of the multivariate analysis but the or-
dination plots yield considerable information
of biogeographic interest.
Foraminifera in Santa Monica Bay.-In a
superbly detailed study, Zalesny (1959) record-
ed and interpreted the distribution of the fo-
raminifera in the bottom sediments of Santa
Monica Bay, California. Seventy bottom sam-
pies were analyzed to produce distributional
data on 96 foraminiferal species and subspe-
cies. The samples covered an area of approx-
imately 100 square kilometers in water depths
ranging from 10 to 828 meters. All occurrence
data (in terms of percentage abundance) were
tabulated in the Zalesny paper. For the pres-
ent study, these were converted to simple pres-
ence and absence of taxa and the INDEX OF
SIMILARITY was computed for all pairs of the
70 sampling localities.
Text-figure 7 shows a contour map of raw
similarity data comparable to that for echi-
noids (Text-fig. 4). The reference fauna (Za-
lesny's sample #3110) is in the left central part
of the map and is indicated by an 'X.' Con-
tours reflect decreasing similarity of the other
69 localities with respect to the reference lo-
cality. Numerical values of the INDEX OF SIM-

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1224 DAVID M. RAUP AND REX E. CRICK

50 fm

80-
100,
0

* 0
0
x
#3110
0

I
I

\"' /40 ,/
II
II
I

TEXT-FIG. 7-Analysis of foraminiferal assemblages from Santa Monica Bay. Solid contours indicate
variation of the INDEX OF SIMILARITYwith respect to an arbitrary reference fauna (#3110). Triangles
represent assemblages which are significantly similar to the reference fauna: open circles are assemblages
significantly dissimilar to the reference fauna; solid circles are assemblages not significantly similar or
dissimilar to the reference fauna.

ILARITY are not shown in this case but the
location of each site is shown by a small sym-
bol. Those indicated by triangles are the ones
that are significantly similar to sample #3110,
those indicated by open circles are significant-
ly dissimilar, and the solid dots represent lo-
calities which are not significant in either di-
rection. Also included are the bathymetric
contours for 10, 50, and 100 fathoms. The
shelf edge is well defined in Santa Monica Bay
and the continental slope is steep. The shelf is
indented by two major submarine canyons:
Santa Monica Canyon (near the reference lo-
cality) and the Redondo Canyon (southeast
corner of the mapped area).
The contours of faunal similarity follow the
bathymetry with remarkable faithfulness. The
shelf edge is clearly defined and both canyons
are evident. The one major anomaly is the
small 'bump' on the inner shelf produced by
sample #3348. Similarity between this sample
and the deep water reference fauna is substan-
tially higher than is found between the other
shelf faunas and the reference fauna. It is not,
however, a statistically significant anomaly. In
fact, when #3348 is compared with the three
closest localities, statistically significant simi-
larity is found! #3348 may therefore by a sim-
ple chance departure from the overall pattern
of the contoured similarity surface or it may

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 FAUNAL SIMILARITY 1225

TEXT-FIG. 8-Analysis of Arenigianbiogeographyof nautiloids.Symbolsare as in Text-figure7 and

show the locationsof 52 samplingareas, one of which (Siberia)was used as the referencefauna. The
paleogeographicreconstructionis an interpolationbetweenpublishedmaps(Scotese,et al., 1979)pro-
vided by C. R. Scoteseand A. M. Ziegler.

result from a minor habitat difference between
#3348 and other shelf sites. The latter sug-
gestion is likely in view of the fact that Zales-
ny's sediment maps show that #3348 comes
from a small patch of silt on the shelf surface
otherwise covered with sand, gravel, or rock.
All the deep water sampling sites are in silty
sediments. It is not surprising therefore, that
the silt patch on the shelf should yield an as-
semblage with relatively high similarity to the
deep-water reference fauna.
It should be emphasized that Text-figure 7
does not in itself require a bathymetric or sed-
iment interpretation. As Zalesny (1959) points
out, many other ecological parameters such as
temperature and salinity parallel changes in
depth and sediment type. The contoured IN-
DEX OF SIMILARITY only provides a statistical
framework for interpretation.
Text-figure 7 can be used also to investigate
another aspect of faunal similarity. The null
hypothesis of random sprinkling predicts that
about 5% of the sites should appear to be sig-
nificantly similar to the reference fauna and
that about 5% should be significantly dissim-
ilar and that these cases of apparent statistical
significance should be randomly distributed
over the area. In this instance, therefore, 10%
or about seven of the 69 assemblages should
show statistical significance in one direction or
the other. In fact, 21 or 30% show statistical
significance and their distribution is obviously
non-random over the geographic area. This
means that the null hypothesis of random
sprinkling can be rejected when considering
foraminiferal assemblages of the whole bay.
This is not surprising in the Santa Monica Bay
case and is certainly substantiated by Zales-
ny's detailed analysis of the distributions of
individual taxa. It illustrates how the method
being presented here can be used to explore
the question of whether distribution of taxa is
purely stochastic or whether it is biased by
deterministic biological and/or physical fac-
tors. Even though the stochastic model can be
rejected easily in this case, the INDEX based
on the null hypothesis of random distribution
is still a valuable aid to ecological interpreta-
tion.
Multivariate analysis was also carried out
on the foraminiferal data. Bivariate ordination
plots are eminently contourable and follow
bathymetry.
Ordovician nautiloid biogeography.-This
data set consists of 182 genera of Arenigian
nautiloid cephalopods which range from en-
demics to those found in as many as 20 of 52
sampling areas. The data are taken from a
broader study of Ordovician biogeography
(Crick, 1978). The location of sampling areas

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1226 DAVID M. RAUP AND REX E. CRICK

TEXT-FIG.
9-Multivariate analysisof Arenigianbiogeographicdata. A plot of the first two principal
components (PCI and PCII) separates the major geographic elements of the early Ordovician.

is shown in Text-figure 8 on a reconstruction
of Ordovician paleogeography developed by
Scotese et al. (1979).
The faunal relationships were measured
with the computer program used in the pre-
ceding examples. Contouring of the similarity
values with respect to Siberia as an arbitrary
reference area (Text-fig. 8) shows the expected
decrease in similarity away from the reference
area. Contouring the same data on a map of
modern geography (not shown) reveals sub-
stantial anomalies which reflect the differences
between modern and Ordovician geography.
For example, the Bear Island fauna is signif-
icantly similar (at the 95% level) to faunas
from Arctic Canada and Scotland but it is not
significantly similar to Norway, Sweden, or
Estonia. The latter three areas are closest to
Bear Island at the present time but were sep-
arated (as part of Baltica) from Bear Island in
the Ordovician.
Patterns in 2-dimensional ordinations of the
principal component axes are not quite as eas-
ily interpreted as were comparable plots of
Recent echinoid and foraminiferal data. This
reflects loss of information about physical en-
vironments and a certain amount of geograph-
ic uncertainty. However, information on as-
sociated faunas and sediments, along with
knowledge of tectonic setting, does make the
multivariate plots understandable. Text-figure
9 shows a plot of PCI and PCII for the 52
sampling areas. Clusters showing the principal
geographic regions (plates) are indicated. Plots
including PCIII (not shown) show separation
of two important Ordovician facies; the plat-

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 FAUNAL SIMILARITY 1227

form facies characterized by shelly faunas and or biogeographic proximity rather than tem-
the slope deposits (graptolitic facies). More de- poral identity. But this is an ever-present
tail on this aspect is given elsewhere (Crick, problem in biostratigraphy which must be
1978). dealt with regardless of the method used to
assess similarity. In the biostratigraphic con-
DISCUSSION text, tests of statistical significance could be
The similarity measure presented here is performed in the manner of the echinoid and
somewhat cumbersome and expensive because foraminiferal examples.
of the simulation technique. The rewards may
be worth the extra effort, however. These may ACKNOWLEDGMENTS
be summarized as follows: This work was supported in part by the
1) Distributional data are weighted on the Earth Sciences Section, National Science
basis of frequency so that widespread taxa do Foundation, NSF Grant DES75-03870. We
not have a disproportionate influence on mea- would also like to thank Richard K. Bambach
surement of similarity. and Alan H. Cheetham for helpful reviews of
2) There is no need to discard taxa on the the manuscript.
a priori grounds that they are too widespread
or too localized. REFERENCES
3) The similarity or dissimilarity of any two Cheetham,A. H. and J. E. Hazel. 1969. Binary
faunas can be tested for statistical significance. (presence-absence) similaritycoefficients.J. Pa-
Such tests are robust assuming that enough leontol. 43:1130-1136.
simulations have been run. Crick, R. E. 1978. Ordoviciannautiloidbiogeog-
raphy:a probabilisticand multivariateanalysis.
4) Because the evaluation of similarity does Ph.D. Dissertation,Univ. Rochester,166 p.
not presume any particular shape for the prob- Ekman, S. 1953. Zoogeographyof the Sea. Sedg-
ability distribution of expected numbers of wick & JacksonLtd., London,417 p.
taxa in common, the results may be considered Henderson, R. A. and M. L. Heron. 1977. A prob-
precise and not dependent upon generaliza- abilistic method of paleobiogeographic analysis.
tions drawn from computed variances of the Lethaia 10:1-15.
Mortensen, T. 1928-1951. A Monograph of the
probability distribution. Echinoidea. C. A. Reitzel, Copenhagen. 5 vols.,
5) An entire faunal realm or data set can be 4469 p.
investigated for significance of the observed Rohlf, F. J., J. Kishpaugh and D. Kirk. 1971. NT-
departures from a random sprinkling (stochas- SYS. Numerical Taxonomy System of Multi-
tic) model of taxon distribution. variate Statistical Programs. Tech. Rep. State
The three examples that have been de- Univ. New York at Stony Brook, New York.
Scotese, C. R., R. K. Bambach, C. Barton, R. Van
scribed do not include one in a biostratigraphic Der Voo and A. M. Ziegler. 1979. Paleozoic
context but biostratigraphicapplications should base maps. J. Geol. 87:217-277.
be straightforward and follow logically from Simberloff, D. S. 1978. Using island biogeographic
the biogeographic/ecological cases used here. distributions to determine if colonization is sto-
For example, the probable stratigraphic posi- chastic. Am. Naturalist 112:713-726.
tion of a new fossil assemblage could be as- Simpson, G. G. 1943. Mammals and the nature
sessed by comparing it with a large number of of continents. Am. J. Sci. 241:1-31.
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assemblages in a standard (possibly composite) tinental relationships during the Cenozoic. Geol.
sequence. This could be done in the fashion of Soc. Am. Bull. 58:613-688.
the contour maps of Text-figures 4, 7, and 8 Zalesny, E. R. 1959. Foraminiferal ecology of
except that it would be a one-dimensional in- Santa Monica Bay, California. Micropaleontol.
stead of a two-dimensional problem. The 5:101-126.
highest INDEX OF SIMILARITYwould be cen-
tered on the assemblages in the standard se- MANUSCRIPT RECEIVED FEBRUARY 17, 1979
REVISED MANUSCRIPT RECEIVED APRIL 12, 1979
quence most similar to the new assemblage.
This would not demand that a temporal cor- The Field Museum of Natural History contributed
relation be made at that point, of course, be- $500 in support of this article.
cause the similarity might be due to ecological

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