ecological indicators 9 (2009) 651658

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A geometrical index for measuring species diversity

Diogenes Campos *, Jose Fernando Isaza
Faculty of Natural Sciences, University Jorge Tadeo Lozano, Bogota, Colombia

article info

abstract

Article history:

During the past decades several biodiversity indices have been proposed and employed in

Received 16 March 2008

ecological literature. Although each one has been partially justified on practical or quasi-

Received in revised form

theoretical grounds, recommendations of ecological theorists differ from describing which

4 July 2008

index to use. The goal of this article is to introduce a new index for measuring biological

Accepted 4 July 2008

diversity that is sensitive to the number of different species (species richness, S), and the
relative abundance of them. We take advantage from the mathematical relation between
Simpson index and the geometrical concept of a S-dimensional sphere of radius r, where r is

Keywords:

the square root of the Simpson index. Full applications of the method are developed, first

Biological diversity

with hypothetical communities and then with real data for 1761 specimens of 82 weevil

Biodiversity index

species collected in several forest types [Ohsawa, M., 2005. Species richness and composi-

Simpson index

tion of Curculionidae (Coleopters) in a conifer plantation, secondary forest, and old-growth

forest in the central mountainous region of Japan. Ecological Research. 20, 632].
# 2008 Elsevier Ltd. All rights reserved.

Classification codes:
400.000
400.010
400.050

1.

Introduction

Biodiversity or biological diversity is a concept that covers

genes, species, ecosystems and ecosystem functions. This
paper will focus on species diversity that indicates the status
of the ecosystem and the quality of the living environment
(Izsak, 2007).
During the past decades several biodiversity indices
have been proposed and employed in ecological and
biological literature. Although each one has been partially
justified on quasi-theoretical grounds (Kempton, 1979)
there are limitations and difficulties with the use of
species diversity measures, and recommendations of
ecological theorists differ at describing which index to
use, see e.g. (Rennolls and Laumonier, 2000; Magurran,
1974). The paper entitled Through the jungle of biological
diversity suggests that no proper, generally agreed-on
definition and biodiversity measurement techniques have

been formulated and accepted amongst ecologists (Ricotta,

2005).
In this paper a new index for measuring biological diversity
is proposed that is sensitive to the number of different species
and the relative abundance of them. Consider a sample region
A with fixed boundaries, and a S-species community labeled
as f1; 2; . . . ; Sg. Denote by fN1 ; N2 ; . . . ; NS g the number of
P
organisms of each species, N : Sn1 Nn the total number of
organisms and p f p1 ; p2 ; . . . ; pS g the probability distribution
representing the relative abundance, with pn : Nn =N and
PS
n1 pn 1. That is, pn denotes the (theoretical) probability of
an individual belonging to the n th species.
A measure of homogenity or concentration of any finite
probability distribution, p, is the Simpson index D (Izsak and
Papp, 2000),
r2 :

S
X
pn 2 ;

D : r2 ;

n1

* Corresponding author. Tel.: +57 1 2834610.

E-mail addresses: josefernando.isaza@utadeo.edu.co, dcamposr@utadeo.edu.co (D. Campos).
1470-160X/$ see front matter # 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.ecolind.2008.07.007

(1)

652

ecological indicators 9 (2009) 651658

that ranges in interval 0  D  1, while the Gini-Simpson index

P
DGS 1  D Sn1 pn 1  pn is a measure of heterogeneity or
diversity, in that, values of D near zero representing high diversity and values near one corresponding to low diversity. While
Simpson index D describes the probability that two organisms
drawn randomly and independently from a population belonging to the same species, DGS is the probability that two randomly
sampled individuals are of two different classes.
In this paper, we propose a new set of biodiversiy indices
Bk S; r by using the fact that (1) describes a sphere of radius
p
r D in an euclidian space of S dimensions, with coordinate
P
axes p1 , p2 ,. . ., pS . The conditions 0  pn  1 and nS
n1 pn 1
on the probability distribution p f p1 ; p2 ; . . . ; pS g also imply
the existence of an hyperplane in S dimensions. This hyperplane intersects each coordinate axis at the point 1 and the
point P p1 ; p2 ; . . . ; pS , representing p, always lies in the first
quadrant of S-dimensional space on a sphere of radius r  1
centered at origin (see Fig. 1).
Note that almost six decades after Edward H. Simpson
(1949) proposed D as a measure of species concentration, as far
as we know, no person has taken advantage of the fact that
Eq. (1) represents a S-dimensional sphere for which the
formula for finding its volume is known (Kubo, 1974). Therefore, the major contribution of this paper consists in using this
property for introducing a new biodiversity index that is based
on a mathematical comparison of the volumes of two spheres,
both having the same Simpson index D but differing in the
number of species, S and S k, respectively (k a non-negative
integer).
A major practical advantage of index Bk S; r ak S bk r is
that it can be partitioned as the product of two components, a
function ak S dependent of species richness and a function
p
bk r D dependent of the Simpson index. It follows the
relation ln Bk S; r ln ak S ln bk r that allows to interpret
changes in biodiversity as the combination of two contributions, species richness and abundance distribution.

Fig. 1 Geometrical interpretation of Eq. (1) for the case of

three species, S 3. The plane passing through the points
1; 0; 0, 0; 1; 0 and 0; 0; 1 is given by p1 p2 p3 1.

We are also able to get a visual and geometrical

representation of biodiversity by using a cartesian bidimenp
sional plane, hereafter called biodiversity plane. It has r D as
x-axis, biodiversity index Bk S; r as y-axis. Biodiversity
conditions of the region under study can be always summarized in a standard diagram, as those shown in examples below
(see Figs. 3 and 4). In similarity to thermodynamical isothermal
curves (Greek isos, equal; thermos, heat), the r- Bk S; r
biodiversity plane is arranged in layers of isonumber of
species curves (S-curves) resulting in an onion-like structure
(structure composed of a set of S-curves). This feature allows
for comparison of communities: e.g. if they have the same
species richness, all Bk S; r values belong to same S-curve.
Other means of measuring biological diversity is the
Shannon index, also called Shannon entropy or Shannon
information index (Izsak and Papp, 2000),

H p H p1 ; p2 ; . . . ; pS : 

S
X

pn ln pn ;

(2)

n1

where, if any pn is assumed to be zero then 0ln 0 is to be

interpreted as zero. The quantity H p is a function of distribution p and the unit depends on the base b of logarithm
used (ln ln b ): b 2, bit; b e, nat; b 3, trit; b 10, hartley.
H p is the information gained on the biological diversity
in the region, assuming that the total number of species S
and the probability distribution p of them are known. As
0  H p  ln S, there are two extreme situations: (i) minimum
biological diversity occurs when only one species exists (all pn
are zero but one takes value 1, H p 0). (ii) Maximum biological diversity occurs when the probability distribution p is
homogeneous (all pn take the same value pn 1=S, and therefore Hmax p ln S).
Further motivation for introducing a new biological
diversity index is concerned with a remark by Kolasa and
Biesladka (1984). Computing the Shannon index or Simpson
index constitutes an error because they combine levels
(number of species and distribution of their abundances) into
one value of unknown ecological meaning, and in this
procedure information is lost. By taking advantage of the
descomposition ln Bk S; r ln ak S ln bk r, we conclude
that this is not the case with the index introduced in present
paper. In fact, for a given numerical couple S; r, the Bk S; rvalue (say g) is located on a well-defined unique S-curve (e.g.
see Figs. 3 and 4). The reverse process, that in the bidodiversity
plane starts from a fixed S-curve and the value g, also leads to
a unique value r. Another way of preserving the full
information is to record the couple ak S; bk r.
Index Bk S; r is also an ordering system. Index value
associated with S; p be greater than the value corresponding
to S 0 ; p0 if, either D D0 (same value of r) and S > S 0 (S-curve
above S 0 curve), or S S 0 (just the same S-curve) and D < D0
(that is, r < r0 ).
In Section 2, the promised diversity index Bk S; r is
introduced and the method is illustrated by considering
examples with hypothetical communities. For a practical
application of the method a full example is developed (Section
3) by using data from the weevil (Coleoptera: Curculionidae)
species richness, published by Ohsawa (2005) and discussed
further by Ito (2007). In Section 4, we conclude with some

ecological indicators 9 (2009) 651658

remarks and discussion, including the relation between the

new index and the Renyi information of order 2.

where subindex 0 is used for recalling that D0 and r0 are the

smaller values of D and r, respectively.

2.2.

2.

Biodiversity index of order k

2.1.

New biodiversity index

In statistical mechanics it is well known that volume of

sphere (1) of radius r in a S-dimensional space is equal to
(Kubo, 1974)
VS r

pS=2
rS ;
GS=2 1

(3)

where Gx is the Gamma function, with following properties:

p
p
Gu 1 uGu, G1 1, G1=2 p, G3=2 1=2 p,
p
G5=2 3=4 p, and so on.
Following Hill (1973), for the purposes of community
description we should express measures of diversity on a
uniform scale. That is to say, we should use the reciprocal of
Simpson index DRS 1=D but not the Gini-Simpson index
DGS 1  D. He also remarks that there is good reason for
favoring diversity numbers over entropies, e.g. favoring DRS
over H p. To be consistent with these criteria, let us define
biodiversity index of order k by the formula
Bk S; r :

VS r
a S
k k ak S bk r;
VSk r
r

r 6 0;

(4)

653

Examples with hypothetical communities

For the purpose of illustration and for comparing the present

indices Bk S; r with the Shannon index we create a set of
hypothetical communities L 1; 2; 3; . . . (Fig. 2). In the x axis,
consider S species labeled as f1; 2; . . . ; Sg, and in the y axis,
assume that the maximum number of organisms of each
species is Nmax . Each community (L) is labeled by the number
written at the point of intersection of the two orthogonal
dashed lines shown in the Fig. 2. The community is arranged
by assuming that each rectangle represents one organism, if
the rectangle is to the left or below the dashed lines (black
rectangles), otherwise assign the value zero (white rectangles).
Let us start with the community L 1 (low diversity) and
increase successively L. For each new value of L, the
corresponding L-community has a more equitable distribution of species (more black rectangles). Finally, the community
L S has the maximum diversity (SNmax black rectangles).
Note that there is a clear gradient of diversity of the
communities, from L 1 (low diversity) to L S (maximum
diversity), For the L-community the total number of organisms is NL LNmax LS  L and for the s th species:
Ns L Nmax , if 1  s  L; Ns L L, if L 1  s  S; and
Ns L 0, if S  s. Thus, ps L : Ns L=NL denotes the

where k is a non-negative integer (mathematically S and k are

not limited to integral values) and bk r : 1=rk 1=Dk=2 . For a
fixed value of k, the coefficient

ak S :

GS k 2=2
pk=2 GS 2=2

(5)

increases as function of species richness, S. From definition

(4), it is clear that Bk S; r compares a system with S species
with a system with S k species, using for comparison welldefined geometrical entities, namely: volumes of spheres of
radius r existing in S- and S k-dimensional euclidian
spaces, respectively.
As particular cases let us consider k 1 and k 2, for getting


GS 3=2 1 a1 S

;
B1 S; r p
r
pGS 2=2 r
(6)


S
1
a2 S
1
B2 S; r
2 :
2
pr2
r
Due to Eq. (1), indices Bk S; r are also functions of the
Simpson index, D r2 .
Note that the remarkably simple result a2 S S=2 1=p
leads to B2 S; r a2 SDRS , where DRS is the reciprocal of
Simpson index.
Because 0  pn  1, then 0  D  1. Value D r2 1 is
obtained if one of the species has unit probability. If all species
have the same probability pn 1=S (with n 1; 2; . . . ; S), from
(1) we get
D0 r20

1
;
S

1
r0 p ;
S

Fig. 2 Scheme for creation of hypothetical communities

L f1; 2; . . . ; Sg with S species labeled as f1; 2; . . . ; Sg, the
maximum number of organisms of s th species is Nmax .
For the L-community each rectangle to the left or below
the two orthogonal dashed lines represents one organism
(in this figure L 1, black rectangles) and otherwise zero
organisms (white rectangles). By increasing L we
progressively change from a community (L 1) with low
diversity to a community (L S) with maximum diversity.

654

ecological indicators 9 (2009) 651658

Table 1 Number of organisms in seven hypothetical communities L 1; 2; . . . ; 7 with seven species (S 7), and a
common coefficient a1 S 128=35p  1:1641
Community L

s th species

1
2
3
4
5
6
7

17
17
17
17
17
17
17

1
17
17
17
17
17
17

1
2
17
17
17
17
17

1
2
3
17
17
17
17

1
2
3
4
17
17
17

1
2
3
4
5
17
17

1
2
3
4
5
6
17

Entropy H p
(bits)
1.502
2.074
2.366
2.548
2.668
2.751
2.807

Index B1 S; r

1.559
2.095
2.441
2.684
2.860
2.988
3.080

The communities are ordered from low diversity to high diversity, property that is confirmed by the values of entropy H p and the index
B1 S; r.

probability of an individual belonging to the s th species of

community L.
Tables 1 and 2 show examples with S 7 and S 10
species, assuming Nmax 17. We get two sets of seven and ten
communities, respectively. On Fig. 3, we plot the values of
B1 S; r-index and observe that they are arranged along two
layers of S-curves. In the biodiversity plane these layers arise
because, according to the first Eq. (6), we get B1 S; r by
multiplying the square root of DRS by a1 S, and this is a
function of the species richness. At this point it is worth noting
that the biodiversity plane can be used as a standard plotting
method for the comparison of different biodiversity data sets.
As remarked by e.g. (Magurran, 1974), the value of Shannon
index obtained from empirical data usually falls between 1.5
and 3.5 and rarely surpasses 4. The fact that the Shannon
index is so narrowly constrained in most circunstances can
make interpretation difficult. The index Bk S; r ak S bk r is
a better approach because the factor ak S explicitly depends
on species richness, S.

3.
data

Application of the method by using real

To provide an example, we use M. Ohsawas data corresponding to a 4-year study with weevils conducted in forests in the

central mountainous region of Japan (Ohsawa, 2005). In the

selected region 44 stands were chosen comprising of 24 larch
plantations. A Malaise trap was set in each of the stands to
capture insects; weevils were separated from among the
trapped material. In total 1761 specimens of 82 species were
captured.
Because of some problems in Ohsawas study, Ito (2007)
recalculated the biodiversity indices using combined species
diversity values of five forest types: (a) larch middle-aged
(stands 110), (b) larch thinned (stands 1218), (c) larch long
rotation (stands 2024), (d) secondary forest (stands 2529 and
3134) and (e) old-growth forest (stands 3740 and 4244). The
other stands (4, 11, 19, 30, 35, 36) were excluded from Ito
calculations because they were located in places distant from
other stands of each forest type.
Now, biodiversity indices B1 S; r and B2 S; r are calculated
for the five forest types considered by Ito. Let us consider a
given forest type with S species and M stands. We organize
data from Table 12 in the Appendix A of Ohsawas paper in an
array gnm , with n 1; 2; . . . ; S, and m 1; 2; . . . ; M, where entry
gnm is the number of specimens of n th species in m th stand.
Notice that two different kinds of probabilities enter into the
analysis of data:
 For fixed m, define the set f p1m ; p2m ; . . . ; pSm g, where
P
pnm : gnm = Sn1 gnm is the probability of finding in m th
stand a specimen of n th species.

Table 2 Number of organisms in ten hypothetical communities L 1; 2; . . . ; 10 with ten species (S 10), and a common
coefficient a1 S 693=512  1:3535
Community L

1
2
3
4
5
6
7
8
9
10

s th species
1

10

17
17
17
17
17
17
17
17
17
17

1
17
17
17
17
17
17
17
17
17

1
2
17
17
17
17
17
17
17
17

1
2
3
17
17
17
17
17
17
17

1
2
3
4
17
17
17
17
17
17

1
2
3
4
5
17
17
17
17
17

1
2
3
4
5
6
17
17
17
17

1
2
3
4
5
6
7
17
17
17

1
2
3
4
5
6
7
8
17
17

1
2
3
4
5
6
7
8
9
17

Entropy H p
(bits)
2.028
2.544
2.812
2.981
3.095
3.176
3.234
3.275
3.303
3.322

Index B1 S; r

2.039
2.740
3.196
3.519
3.758
3.935
4.068
4.165
4.234
4.280

The communities are ordered from low diversity to high diversity, property that is confirmed by the values of entropy H p and the index
B1 S; r.

655

ecological indicators 9 (2009) 651658

hri

M
X

Wm rm ;

m1

v
uM
uX
Wm rm  hri2 ;
Dr t
m1

(8)
hBk i

M
X

Wm Bk S m ; rm ;

m1

v
uM
uX
Wm Bk S m ; rm  hBk i2 ;
DBk t
m1

Fig. 3 The r- B1 S; r biodiversity plane is arranged in

layers of isonumber of species curves (S- curves) that
create on an onion like structure. Here, graphical
comparison of two sets of hypothetical communities with
S 7 and S 10 species, respectively. Each community is
characterized by the square root of Simpson index (radius
r of a S-dimensional sphere) and by the value of index
B1 S; r. This value is represented by the black point
located on the continuous curve B1 S; r a1 S=r. Vertical
lines are used as a visual help for the correspondence
between the radius r and the value of B1 S; r.

where h. . .i and D symbols are average value and uncertainty

value of the associated variable. Note that the indices Bk S; r
and above statistical quantities are a composite of the number
of species in the sample (S), the number of specimens of each
species in the sample (represented in pnm ) and the speciess
statistical weight (Wm ).

3.1.

In order to illustrate the method with an example consider the

data for the Larch-middle-aged forest that includes stands
1  10, S 29 species and N 607 specimens (Ohsawa, 2005;
Ito, 2007). Biodiversity indices B1 S m ; rm and B2 S m ; rm , radius
rm for m th stand and statistical calculations over ensemble of
10 stands are calculated by using formulae (6), (1) and (8).
Table 3 shows the obtained results including values for
Shannon entropy, Simpson index and also the data for a virtual
stand (row 11$ ) arranged with the accumulated data of all
other stands.
Fig. 4 shows the characterization of larch middle-aged
forest by using biodiversity indices B1 S m ; rm and B2 S m ; rm as
p
function of r (recall that r D is a measure of species relative
abundance). Each stand (say m th) is associated with a unique
curve characterized by species richness, S m . That is, the
positive quadrant plane is shelled by curves that differ from
each other by S m value.
The chosen forest has associated an uncertainty window
hri  1=2Dr; hBk i  1=2DBk that is represented in Fig. 4 by a
rectangle with a central point hri; hBk i (black big point), base
Dr, height DBk , and area Dr DBk (k 1 or k 2).

 For the ensemble of M stands as a whole, assign the set

fW1 ; W2 ; . . . ; WM g of non-negative numbers Wm , normalized
P
to one, M
m1 Wm 1, assuming that Wm is statistical weight
of m th stand. Generally, value of Wm depends on experimental factors and a standard assumption is that all stands
have equal probability, Wm 1=M (m 1; 2; . . . ; M). Since
Ohsawa reports the total number of species captured in each
stand it seems the appropriate choice
S m =S
Sm
PM
;
Wm PM
m1 S m =S
m1 S m

Calculations for Larch-middle-aged forest

(7)

where S m =S appears to be the probability of collecting in m th

stand S m species, accepting that in the forest there are S
different species. Since the same species can be collected in
P
two or more different stands, M
m1 S m 6 S.
Statistical calculations over the ensemble of M stands
proceed with the help of the following standard relations:

Table 3 Larch middle-aged forest

Stand m th
1
2
3
4
5
6
7
8
9
10
11

H (bits)

Sm

rm

B1 S m ; rm

1.911
1.252
1.591
1.536
3.160
1.172
1.401
0.720
1.335
1.586
2.040

0.311
0.500
0.393
0.432
0.134
0.668
0.517
0.827
0.487
0.412
0.434

6
3
5
6
11
8
5
10
5
7
29

0.558
0.707
0.627
0.657
0.366
0.817
0.719
0.910
0.698
0.642
0.659

1.962
1.200
1.626
1.664
3.86
1.505
1.417
1.488
1.460
1.813
3.34

Stand 110, S 29, N 607. Row 11$ represents a virtual stand arranged with the accumulated data of all other stands.

B2 S m ; rm
4.10
1.59
2.84
2.95
15.5
2.38
2.15
2.31
2.29
3.47
11.4

656

ecological indicators 9 (2009) 651658

Fig. 4 The r- Bk S; r biodiversity plane is arranged in

layers of isonumber of species curves (S- curves) that
create on the plane an onion-like structure. In these
figures, data is used for the characterization of larch
middle-aged forest by using biodiversity indices B1 S m ; rm
(above) and B2 S m ; rm (below). From left to right,
m; S 5; 11, (1, 6), (3, 5), (10, 7), (4, 6), 11$ ; 29, (9, 5), (2,
3), (7, 5), (6, 8), (8, 10), where couple m; S refers to m thstand and number of species captured in it and 11$
corresponds to virtual stand. Forests biodiversity index is
represented by the big black point located in the center of
uncertainty rectangular window.

Fig. 5 Comparison of biodiversity indices and uncertainty

windows for several forest type (see Table 4): (a) larch
middle-aged, (b) larch thinned, (c) larch long rotation, (d)
secondary forest, (e) old-growth forest.

we conclude that forest types are ordered from low to high

biodiversity, as follows: a, b, e, c, d. With exception of stand
m 8, for all other stands of Table 3 and Tables A.1A.4, it
holds D < H p < B1 S m ; rm .

4.
3.2.

Concluding remarks and discussion

Calculation for the b-e forest types

The calculations are similar to the described above. Results are

presented in the Appendix A, and Table 4 summarizes the
information about biodiversity indices and uncertainty windows for all forest types: (a) larch middle-aged, (b) larch
thinned, (c) larch long rotation, (d) secondary forest and (e) oldgrowth forest. After displaying results in Fig. 5, and by using
the fact that the biodiversity indices Bk are ordering systems,

In the literature there are several biodiversity indices in use,

satisfying different criteria, e.g.: the Simpson index D and
the Shannon information index H p. This paper introduces
a new index as a quantitative indicator of biodiversity that is
sensitive to species richness and relative abundance of
species: B1 S; p or B2 S; p. The biodiversity index Bk S; p is
based on a well-defined geometrical relation between the
Simpson index D r2 and the volume of a S-dimensional

Table 4 Forest types: (a) larch middle-aged forest, (b) larch thinned, (c) larch long rotation, (d) secondary forest, (e) oldgrowth forest
Forest type
a
b
c
d
e

hri  12 Dr

hB1 i  12 DB1

DrDB1

hB2 i  12 DB2

DrDB2

0:663  0:084
0:584  0:056
0:508  0:027
0:435  0:046
0:498  0:036

1:968  0:432
2:52  0:36
2:66  0:23
3:51  0:41
2:54  0:30

0.15
0.082
0.026
0.076
0.043

4:85  2:39
7:12  1:76
7:61  1:37
13:5  3:0
7:16  1:55

0.81
0.40
0.15
0.57
0.22

657

ecological indicators 9 (2009) 651658

Table 5 Right values of H and D0 for correcting some typos in Ohsawas Table 12 (Ohsawa, 2005)
Stand
H
D0 1  D

28

30

34

Standard W

0.607

0.568

2.442
0.769

3.152
0.871

3.149
0.837

2.256
0.688

sphere of radius r. The index Bk S; p acts as an ordering

system that allows a mathematical and visual representation of the biodiversity value in a bidimensional plane and
also the comparison between several systems (e.g. forest
types).
Due to the relation Bk S; r ak SB1 S; r=a1 Sk , for k  2,
the index B1 S; r can be chosen as the basic biodiversity
index. Furthermore, the relation Bk S; r ak S=Dk=2 indicates that Bk S; r is a function of the inverse of the Simpson
index D.
At this point it is worth stablishing the relation of Bk S; r
with Renyi information (or entropy) of order q that is associated
to an arbitrary probability distribution p f p1 ; p2 ; . . . ; pS g.
After generalizing Eq. (1) and redefining r through the relation
P
rq : Sn1 pn q , where q is an arbitrary non-negative real
number, the Renyi information of order q is by definition (e.g.
Bogaert et al., 2005)
Iq p Iq p1 ; p2 ; . . . ; pS :

1
ln rq :
1q

W Eq. (7)
2.435
0.714

time. Spatial and temporal resolutions play a central role in

the biodiversity evaluation in a given area A. Biodiversity
comparison for two different times (t1  Dt and t2  Dt)
requires to use the same window Dt, identical area and
sub-areas.
Finally we note that in Table 12 of Ohsawas paper there
are some typos for the diversity index D0 1  D and the
Shannon entropy H. In Table 5 of this paper we give account of
the right values. The last two columns correspond to
Ohsawas column entitled Average number of individuals,
now calculated by using standard assumption that all 44
stands have equal probability Wm 1=44 and also by using
statistical weight Wm of Eq. (7). Number of specimens as
reported in Ohsawas Table 12 are assumed to be right values
for all stands and all species.

Acknowledgments
(9)

For a fixed probability distribution p f p1 ; p2 ; . . . ; pS g the

standard Shannon entropy is recovered from Eq. (9) by using
lHospitals rule in the case q ! 1. If q 2, one gets the socalled quadratic entropy I2 p1 ; p2 ; . . . ; pS ln D 2ln r,
where I2 p is a positive decreasing function of Simpson index
D (0 < D  1).
Clearly the biodiversity index Bk S; r can be written as



k
I2 p1 ; p2 ; . . . ; pS :
Bk S; r ak Sexp
2
Thus, for a fixed k (e.g. k 1), Bk S; r provides a measure of
species richness (due to factor ak S) and also a measure of the
systems quadratic entropy. Bk S; r grows exponentially with
the quadratic entropy, I2 p.
Now we consider the fact that in a given study area
the biodiversity varies across the space and the time t. For
dealing with this situation, consider a sample region A
A1 [ A2 [ . . . [ AM that is the union of M sub-areas, and
assume these sub-areas do not overlap. Represent the
position of Am s central point by the position rm and be DAm
the area (or volume) of sub-area Am . Thus, the biodiversity
index Bk S; p; rm ; t can be considered as a function of
position and time and we speak of biodiversity in area m
th at time t (in a temporal window t  Dt). This is a way to
describe where biodiversity is located (spatial distribution)
and also a method for evaluating how the biodiversity
changes with time.
In conclusion, monitoring the biodiversity status and
evolution requires not only extensive field work but also a
theoretical foundation as the one described in this present
paper. A previous observational design (selection of area A,
sub-areas fA1 ; A2 ; . . . ; AM g, temporal window Dt, and so on) is
needed in order to follow biodiversity values in space and

The authors wish to thank the reviewers for important

recommendations that substantially helped to improve this
paper. We also thank Prof. Martha R. Campos, Universidad
Nacional de Colombia, for her valuable comments about the
manuscript.

Table A.1 Larch thinned forest

Stand
m th

H
(bits)

Sm

rm

12
13
14
15
16
17
18
19

1.419
1.297
2.579
1.319
2.197
2.918
2.315
2.882

0.520
0.595
0.368
0.504
0.235
0.171
0.293
0.252

5
6
19
5
5
11
11
26

0.721
0.772
0.607
0.710
0.484
0.413
0.541
0.502

B1 S m ; rm B2 S m ; rm
1.413
1.418
2.98
1.435
2.10
3.41
2.61
4.17

2.14
2.14
9.08
2.21
4.75
12.1
7.07
17.7

Stand 1218, S 26, N 249. Row 19$ represents a virtual stand

arranged with the accumulated data of all other stands.

Table A.2 Larch long rotation forest

Stand
m th

H
(bits)

Sm

rm

20
21
22
23
24
25

2.274
2.369
2.907
2.326
2.127
3.097

0.301
0.272
0.183
0.227
0.327
0.205

11
8
12
6
9
24

0.549
0.522
0.428
0.476
0.572
0.453

B1 S m ; rm B2 S m ; rm
2.57
2.36
3.42
2.30
2.26
4.45

6.87
5.85
12.1
5.61
5.35
20.2

Stand 2024, S 24, N 200. Row 25$ represents a virtual stand

arranged with the accumulated data of all other stands.

658

ecological indicators 9 (2009) 651658

Appendix B. Supplementary data

Table A.3 Secondary forest

Stand
m th

H
(bits)

25
26
27
28
29
31
32
33
34
35

2.541
3.559
2.086
2.442
3.326
2.861
2.944
2.913
3.149
3.973

0.275
0.0930
0.407
0.231
0.108
0.193
0.191
0.168
0.163
0.0991

Sm
13
13
10
7
11
12
13
24
11
40

rm
0.525
0.305
0.638
0.480
0.329
0.440
0.437
0.409
0.404
0.315

B1 S m ; rm B2 S m ; rm
2.90
4.98
2.12
2.42
4.29
3.34
3.47
3.45
3.89
8.17

8.68
25.7
4.69
6.21
19.2
11.5
12.5
12.4
15.6
67.5

Stand 2529 and 3134, S 40, N 346. Row 35$ represents a

virtual stand arranged with the accumulated data of all other
stands.

Table A.4 Old growth forest

Stand
m th

H
(bits)

Sm

rm

37
38
39
40
42
43
44
44

1.549
2.835
2.321
2.959
2.000
2.325
2.252
4.076

0.438
0.184
0.296
0.178
0.313
0.278
0.222
0.107

4
9
8
11
5
9
5
35

0.661
0.429
0.544
0.421
0.559
0.527
0.471
0.327

B1 S m ; rm B2 S m ; rm
1.417
3.02
2.26
3.35
1.822
2.45
2.16
7.37

2.18
9.53
5.37
11.7
3.57
6.30
5.01
55.0

Stand 3740 and 4244, S 35, N 106. Row 44$ represents a

virtual stand arranged with the accumulated data of all other
stands.

Appendix A
Tables A.1A.4 present calculations of Shannon entropy,
Simpson index, radius of S m -dimensional sphere, indices
B1 S m ; p and B2 S m ; p for the set of stands and forest types,
where m designates Ohsawas stand m th (Ohsawa, 2005).

Supplementary data associated with this article can be

found, in the online version, at doi:10.1016/j.ecolind.2008.
07.007.

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