Oecologia (1989) 78: 394-400
9 Springer-Verlag1989
Comparison of skewness coefficient, coefficient of variation,
and Gini coefficient as inequality measures within populations*
R.B. Bendel**, S.S. Higgins***, J.E. Teberg****, and D.A. Pyke*****
Program in Statistics, Washington State University, Pullman, WA 99164, USA
Summary. The moment skewness coefficient, coefficient of
variation and Gini coefficient are contrasted as statistical
measures of inequality among members of plant popula-
tions. Constructed examples, real data examples, and distri-
butional considerations are used to illustrate pertinent
properties of these statistics to assess inequality. All three
statistics possess some undesirable properties but these
properties are shown to be often unimportant with real
data. If the underlying distribution of the variable follows
the often assumed two-parameter lognormal model, it is
shown that all three statistics are likely to be highly and
positively correlated. In contrast, for distributions which
are not two-parameter lognormally distributed, and when
the distribution is not concentrated near zero, the coeffi-
cient of variation and Gini coefficient, which are sensitive
to small shifts in the mean, are often of little practical use
in ordering the equality of populations. The coefficent of
variation is more sensitive to individuals in the right-hand
tail of a distribution than is the Gini coefficient. Therefore,
the coefficient of variation may often be recommended over
the Gini coefficient if a measure of relative precision is
selected to assess inequality. The skewness coefficient is sug-
gested when the distribution is either three-parameter log-
normally distributed (or close to such), or when a measure
of relative precision is not indicated.
Key words: B r o m u st e c t o r u m - F e s t u c a i d a h o e n i s - Fre-
quency distribution - Plant populations - Size hierarchy
Plant ecologists have known for decades that in many plant
populations most of the population biomass and seed pro-
duction are concentrated in relatively few of the individuals
(Koyama and Kira 1956). A tremendous amount of evi-
dence has been collected to indicate that frequency distribu-
tions of individual plant biomass and seed production are
* Scientific paper no 7830. College of Agriculture and Home
Economics Research Center, Washington State University
** Current address and address f o r offprint requests: Center for
Environmental Health, University of Connecticut Storrs, CT
06268, USA
*** Current addresses: Department of Horticulture and Land-
scape Architecture, Washington State University, Pullman, WA
99164, USA
**** Lockheed Corporation, 1050 Flamingo Road, Suite 301, Las
Vegas, NV 89119, USA
***** Department of Range Science, Utah State University, Lo-
gan, UT 84322, USA
Oecologia
characteristically non-normal and skewed to the right. Fur-
ther, this skewness of individual biomass and seed produc-
tion is often exacerbated by time and by competition (Yoda
et al. 1957; Obeid et al. 1967).
Characterization of the tendency toward an increasingly
skewed or "unequal" distribution is important for several
reasons. If only a very few individuals are producing most
of the population's seed, then fitness is most likely concen-
trated in these few plants. The evolutionary consequences
of such a concentration of fitness are great (Levin and Wil-
son 1978, Solbrig 1981). Changes in the frequency distribu-
tions with density or with time have been used as indices
of the competitive ability of the species in question (Mack
and Harper 1977). Also, changes in the frequency distribu-
tion of plant biomass with time may be a characteristic
of the genotype under consideration (Grime and Hunt
1975).
Plant frequency distributions have been characterized
in several different ways. Many authors chose not to char-
acterize mathematically the inequality embedded in fre-
quency distributions, presenting instead, histograms to rep-
resent graphically the inequality within and among popula-
tions (Mack and Harper 1977). They spoke in general terms
of "skewness", and restricted statistical analysis to tests
of normality (Obeid et al. 1967). The coefficient of variation
(CV) has been used in plant population ecology for over
30 years as a measure of relative precision and inequality
(Kira et al. 1953; Edmeades and Daynard 1979). Others
reported inequality among plant distributions as the range
and/or CVs of weights (Stern 1965; Mack and Pyke 1983),
and some recommended use of the mode relative to the
mean (Naylor 1976). The moment skewness coefficient, de-
fined below, was computed by Ford (1975) in reference
to Harper's (1967) "hierarchy of exploitation" or Hutch-
ings and Budd's (1981) "dominance hierarchy", and this
statistic has since been used extensively (Rabinowitz 1979;
Dirzo and Harper 1980; Bookman and Mack 1983; Turner
and Rabinowitz 1983; Cannell et al. 1984; Higgins etal.
1984; Higgins and Mack 1987).
Recently, Weiner and Solbrig (1984) proposed a defini-
tion of "size hierarchy" and recommended the use of the
Gini coefficient as a statistic that reflected ecologists' con-
notation of inequality among members of a plant popula-
tion. Weiner and Solbrig (1984) proposed that frequency
distributions of plant size be characterized on the basis of
three intuitive attributes of "size hierarchy", 1) large varia-
tion, 2) few large and many small individuals, and 3) the
large individuals are dominant. Weiner and Solbrig (1984)
claim that the skewness coefficient measures only the sec-

ond of their attributes. They have proposed use of the Gini
coefficient as a more intuitively satisfying measure of in-
equality within plant population frequency distributions.
Their argument has resulted in increasingly frequent appli-
cation of the Gini coefficient (Weiner 1985; Weiner and
Thomas 1986; Schmitt et al. 1986). However, use of a statis-
tic must also be based on its important mathematical char-
acteristics. To date, a concise comparison of the mathemati-
cal properties of the several commonly reported statistics
for assessing inequality in frequency distributions is lacking.
Our objective in this paper is to present characteristics
and behaviors of three common measures of inequality,
the skewness coefficient, g~, the coefficient of variation,
C V, and the Gini coefficient, G. Also, we will attempt to
make recommendations regarding circumstances in which
one measure of inequality may be preferred over another.
Below we define gl, C V, and G, and we discuss their
basic and pertinent properties. The results first present
Weiner and Solbrig's (1984) constructed examples and
other examples which illustrate and contrast properties of
these statistics. Then, results based on real data sets are
presented. The discussion is concerned with distributional
considerations and recommendations. The more mathemat-
ical properties of the statistics have been presented by Ben-
del and Carlin (1988).
Definitions and properties of the statistics
Moment Skewness, gl
The population skewness is defined by 7~ =r and is
estimated by gl =tc3/~c~/e where ~c2 and ~c3 are Fisher's ~c
statistics which estimate/tz and//3, the second and third
central moments, respectively (Cramrr 1951):
~c2=/~ Z(x-~)2,
n--1
To3=/~3- nZ(x-x)3
(n--l) 0 - - 2 ) '
Although K2 and *c3 are unbiased estimators of #2 and P3
respectively, gl is not generally an unbiased estimator of
7~. Note that g, is invariant to both location and scale;
that is, gl is invariant under the linear transformation, y =
a+bx. As a popular statistic, gl has been used as a test
for normality (Ho: 7~ = 0) and for descriptive purposes as
a measure of skewness. It is well known that gl > 0 typically
indicates skewness to the right with large values in the right-
hand tail of the distribution.
Coefficient of Variation, C V
The sample C V is defined as a proportion, C V - s / 2 where
s = ~c2
*/2, the sample standard deviation, and )2 is the sample
mean. The coefficient of variation is a well known measure
of relative precision which is invariant to scale changes (x --+
kx, k > 0), but is not invariant to location changes (x--+ x +
k, k + 0 ) .
Gini, d
The Gini coefficient for a random sample of size n is esti-
mated as (Sen 1973):
d ~Z]xl--x)[
2 n 2 9~ (1)
395
If all of the observations are the same ( 2 4 0), then G = 0.
At the other extreme, if all of the observations are zero
except for one, then G is a maximum, equal to 1 - 1 / n if
all but one of the observations are zero. The upper bound
of G is one.
For comparison it is important to realize that G is
bounded between 0 and I and is not a measure of skewness
but of relative precision, like C V (Kendall and Stuart 1963;
Bendel and Carlin 1988). Also, G is similar to C V but differ-
ent than gl in the sense that G is invariant to scale changes
but is not invariant to location changes.
The Gini coefficient is often defined in reference to the
Lorenz curve (Sen 1973; Weiner and Sobrig 1984) which
results from a plot of the cumulative proportion of the
population to the cumulative proportion of the variable
(e.g., biomass). For example, if a data set has 5 observa-
tions, all with a value of 5, G will be 0 and the Lorenz
curve will be a straight, diagonal line. Alternatively, if one
observation has a value of 25 (or any other nonzero value)
and the other four observations are 0, G will be 0.80 and
the Lorenz curve will account for 80% of the area under
the 45 degree line. Weiner and Solbrig (1984) provide a
more extensive illustration of the relationship between
and the Lorenz curve.
Materials and methods
Simple, constructed data sets were either taken from the
literature (Weiner and Solbrig 1984) or were generated to
demonstrate some of the properties ofga, C V, and G. The
first two statistics were computed by SAS (1985) subrou-
tines while G was computed directly according to Equation
I above.
Frequency distributions of individual plant biomass for
Festuca idahoenis seedlings grown for different lengths of
time and with different densities of competitors were used
as one of two real-plant data sets. The experiment that
produced this data set is described elsewhere (Higgins et al.
1984).
Frequency distributions of the number of seeds pro-
duced by individuals of Bromus tectorum provide the second
real-plant data set. Details of this experiment are provided
by Pyke (1987). The data set presented here includes plants
subjected to different grazing frequencies, with the grazing
treatment having been initiated at different plant ages. The
B. tectorum data set includes healthy plants as well as those
attacked by smut.
The comparisons within the real data sets were made
by computing the Pearson product moment correlation be-
tween each pair of the statistics, ga, C V, and G. To test
for a two-parameter lognormal distribution (minimum
value theoretically zero; Aitchinson and Brown 1957), the
logarithm of the observations was tested for normality us-
ing the SAS (1985) PROC U N I V A R I A T E program. For
B. tectorum the plants with zero seed production were as-
signed the value 0.5 since the logarithm of zero is not de-
fined. For F. idahoensis, the biomass of the dead plants
was assigned the mass of the smallest plant for the respec-
tive treatment ( < 0.001 g).
Results
Constructed examples
Table I contains constructed examples which convey impor-
tant properties ofg~, C V, and G. Recall that all three statis-
396

Table 1. Comparison of skewness, gl, coefficient of variation, CV, and Gini, G, for constructed examples. The "values" in the table
represent the numerical values of the response variable, e.g., biomass or seed number. Sample A t , for example, contains 50 plants
which weigh 101 and 5 plants which weigh 102. The mean and standard deviation of these 55 plants are 101.09 and 0.29 respectively

Sample Values Frequency ~ s gl CV

A1 - 101 102 - 50 5 101.09 0.29 2.93 0.0029 0.00082

A2 - 1 2 - 50 5 1.09 0.29 2.93 0.27 0.076
A3 - 1 10 - 50 5 1.82 2.61 2.93 1.44 0.409
A4 - 0.5 9.5 - 50 5 1.32 2.61 2.93 1.98 0.564
A5 - 0.1 9.1 - 50 5 0.92 2.61 2.93 2.84 0.810
B1 0.1 1.1 2.1 20 0 10 0.77 0.96 0.74 1.25 0.58
B2 0.1 1.1 2.1 19 2 9 0.77 0.92 0.74 1.20 0.58
B3 0.1 1.1 2.1 14 12 4 0.77 0.71 0.60 0.93 0.48
C1 0.01 t.0l 2.01 13 1 2 0.32 0.70 2.08 2.18 0.81
C2 0.01 1.01 5.01 13 1 2 0.70 1.70 2.42 2.44 0.85
C3 0.0t 1.01 10.01 13 1 2 1.32 3.40 2.48 2.57 0.86
C4 0.01 1.01 20.01 13 1 2 2.57 6.81 2.50 2.65 0.87
D1 - 0 20 - 9 6 8.00 10.14 0.46 1.26 0.60
D2 - 0 20 - 11 4 5.33 9.15 1.18 1.72 0.73
D3 - 0 20 - 13 2 2.67 7.04 2.40 2.64 0.87
D4 - 0 20 - 14 1 1.33 5.16 3.87 3.87 0.93
E1 - 1 21 - 9 6 9.00 10.14 0.46 1.13 0.53
E2 - 1 21 - 11 4 6.33 9.15 1.18 1.45 0.62
E3 - 1 21 - 13 2 3.67 7.04 2.40 1.92 0.63
E4 - 1 21 - 14 1 2.33 5.16 3.87 2.21 0.53
F1 - 3 23 - 9 6 11.00 10.14 0.46 0.92 0.44
F2 - 3 23 - 11 4 8.33 9.15 1.18 1.10 0.47
F3 - 3 23 - 13 2 5.67 7.04 2.40 t .24 0.41
F4 - 3 23 - 14 t 4.33 5.16 3.87 1.19 0.29
GI - 5 25 - 9 6 13.00 10.14 0.46 0.78 0.37
G2 - 5 25 - 11 4 10.33 9.15 1.18 0,89 0.38
G3 - 5 25 - 13 2 7.67 7.04 2.40 0,92 0.30
G4 - 5 25 - 14 1 6.33 5.16 3.87 0,82 0.20
H1 0 1 10 9 5 1 t.00 2.54 3.64 2,54 0.80
H2 0 1 10 5 9 1 1.27 2.46 3.62 1.94 0.62
H3 0 1 10 5 5 5 3.67 4.65 0.76 1.27 0.61

tics are i n v a r i a n t to scale changes b u t o n l y g l is i n v a r i a n t a p p l i c a t i o n m a y be e n v i s i o n e d as shifts in b i o m a s s f r o m

to a shift o f the distribution, i.e., a change in location.
This is i m p o r t a n t in application. W e i n e r and Solbrig's
(1984) three c o n s t r u c t e d e x a m p l e s ( A I A 3 in T a b l e 1) all
h a v e the s a m e skewness, g l - - 2 . 9 3 , b u t different and '~m o r e
r e a s o n a b l e " values for C V and G. S a m p l e s A 1 a n d A 2
are related by the linear t r a n s f o r m a t i o n y = x + 1 0 0 and
samples A 2 a n d A3 are related by the linear t r a n s f o r m a t i o n
y = - 8 + 9 x, so A 1, A 2 a n d A 3 m u s t h a v e the s a m e n u m e r -
ical v a l u e o f gl. F o r the s a m e reason, A 4 a n d A 5 also
h a v e gz = 2.93. T h e i m p o r t a n t p o i n t is t h a t g~ is a shape
p a r a m e t e r a n d does n o t d e p e n d directly o n the m e a n or
v a r i a n c e o f the distribution. In contrast, C V a n d G are
m e a s u r e s o f relative precision, i.e., they reflect the ratio
o f a m e a s u r e o f dispersion to the m e a n , so they c h a n g e
as the ratio changes.
Samples in Series B all h a v e the same m e a n (0.77) a n d
are p r e s e n t e d to d e m o n s t r a t e w h a t e c o n o m i s t s call the Pi-
g o u - D a l t o n c o n d i t i o n o f transfers (Sen 1973) or S c h u r - c o n -
vex f u n c t i o n s ( M a r s h a l l a n d O l k i n 1979). T o illustrate this
transfer principle, which is i m p o r t a n t in t h e o r y (Sen 1973),
suppose the units are in g r a m s so t h a t in P o p u l a t i o n B 1
there are 20 plants o f w e i g h t 0.1 g and 10 plants o f w e i g h t
2.1 g. T r a n s f e r i n g 1 g f r o m a h e a v y p l a n t (2.1 g) to a light
p l a n t (0.1 g) w o u l d give the B 2 distribution. A n ecological
y e a r to y e a r a m o n g m e m b e r s o f a p o p u l a t i o n o f p e r e n n i a l
plants at c a r r y i n g capacity. (In e c o n o m i c s , the idea w o u l d
be to transfer one unit o f w e a l t h f r o m a rich p e r s o n to
a p o o r p e r s o n w i t h o u t c h a n g i n g the total w e a l t h o f the
p o p u l a t i o n . ) Six transfers result in B 3. E a c h transfer neces-
sarily decreases the v a l u e o f s, ~ V a n d G ( M a r s h a l l a n d
O l k i n 1979). Since the skewness coefficient is n o t Schur-
c o n v e x it will n o t necessarily decrease w i t h each transfer.
Samples in Series C illustrate t h a t all three statistics
increase as the weight o f the largest plants increase f r o m
2.01 to 20.01. This increase in the v a l u e o f the statistics
seems r e a s o n a b l e for m a n y ecological applications. F o r
S a m p l e Series D suppose the n u m b e r o f seeds is the v a r i a b l e
o f interest. N o t e t h a t all three statistics increase as the fre-
q u e n c y o f the highest s e e d - p r o d u c i n g plants decreases. R e -
call t h a t the largest v a l u e o f G is 1 - 1 / n w h e n a l l plants
h a v e z e r o seeds except o n e plant, e.g., w i t h n - - 1 5 , G = 0 . 9 3
(sample D 4 , T a b l e 1). F o r p o p u l a t i o n s in Series E, the m e a n
is shifted only slightly, by 1.0, f r o m those o f Series D.
A g a i n , due to l o c a t i o n invariance, g l is the s a m e for each
c o r r e s p o n d i n g s a m p l e in the D group. T h e slight increase
in the m e a n does n o t c h a n g e the m o n o t o n i c a l l y increasing
n a t u r e o f C V for these p o p u l a t i o n s b u t it does affect G.
I n particular, ~ increases as the f r e q u e n c y in the tail gets
 397

N=157 i gl = 0 . 0 5 N= 156 gl = 0 . 6 0 gl = 0 . 5 8
15
2I A A

20
I0

:i
0
0 1.5 5.0 4.5 6.0 1.5 3.0 4.5 6.0
I0

O.
0 0.7 1.4 2,1 2.8
30 5C
gl = 0 . 7 1 gl = 1 . 3 5 gl = 1 , 9 3
A A
CV= 0.78 CAV= 0.97 CV = 1.15
>- 20
h 50 A A
(.3 G = 0.45 G = 0.51 G = 0.55
Z
IJJ N= 178
I0
20
0
IJJ
n*"
LI- 0 I-7 0 0
25 50 75 I00 15 50 45 60 0 15 3,0 45

gl = 0 . 6 6 gl = 2 . 0 5 gl = 2 . 5 5
,5f A
CV = O. ~'7 4(]
A
CV : 1.24
A
CV= 1.61
A A 60 A
G = 0.45 G = 0.60 G = 0.69

1
40
N = 169

20

V~ o 0
125 250 3,75 0 50 I00 150 200 40 80 120 160

BIOM A SS (rng)

Fig. 1. Histograms, skewness coefficients (gl), coefficients of variation (C V), and Gini coefficients ((~) for Festuca idahoensis biomass
sown with competitors at increasing (from left to right) density and harvested at one-month intervals (earliest harvest in the top row)
(Higgins et al. 1984). The mean biomass is indicated by a dashed vertical line

smaller except for Sample E4, where G=0.53. This is an problems as Weiner and Solbrig's (1984) examples illustrate
important distinction between C V and G. In contrasting (Table 1, A 1 - A 3).
Series D with Series E, C V measures a larger dispersion
relative to the mean as the frequency in the tail decreases
Contrasting gl, C V, and G with real data
whereas G does not increase as we change from E3 to E4.
Mathematically, G is a more robust measure of relative In practice it is important to know how gl, C V, and
dispersion than C V, and is not as sensitive to the tails. apply to real, not artificially constructed data sets, although
This effect is even more dramatic as the mean is shifted constructed data sets are useful for fixing ideas. We present
away from the origin as in the F and G series. Note that two examples. Figure 1 (Higgins et al. 1984) shows values
C V also begins to decrease in F as well as in G, just like for g~, ~ V, and G for F. idahoensis. The statistics gl, C V,
does. Hence, both C V and G are statistics which may and G are all highly correlated (r = 0.98 or higher) and all
not pick up tails to the right when the distribution is re- three statistics increase with increasing competition and
moved away from the origin. with increasing time for plant growth. The histograms and
Samples in Series H reflect two additional concepts. the high correlations among gl, C V, and G for F. idahoensis
Note that H1 and H2 have similar g~ values as they both clearly indicate that it would make little difference which
have a long tail to the right. In contrast, G=0.80 for H1 statistic was used to assess the "size inequality" of the data
whereas G=0.62 for H2. The reason G drops from 0.80 in Fig. 1. In contrast, Table 2 presents results of B. tectorum
to 0.62 is because the mode shifts from 0 to 1 as we change seed production with correlations, r = 0.77 between g~ and
from H 1 to H2. As noted above, G is very sensitive to CV, r = - 0 . 1 3 between g~ and G, and r=0.42 between
a slight change in the histogram when the change occurs C V and G. These correlations show that g~, CV, and
near zero. Sample H 3 together with H 2 illustrates a point do not behave similarly here, as they did with F. idahoensis.
made by Weiner and Solbrig (1984) which is given here Four of the 17 B. tectorum samples are presented in Fig. 2
for completeness. They noted that two different distribu- to illustrate why gl and G correlate poorly. In Fig. 2A,
tions with different "inequality" attributes i.e., different both gl and G are high whereas in Fig. 2B, g~ is high but
Lorenz curves, may have the same G value. It should be is relatively small. The long, narrow tail to the right
noted, moreover, that the skewness coefficient has this same accounts for the high skewness in both Fig. 2A, B. The
drawback (Mood 1950, pg. 96). Gini coefficient is lower in Fig. 2B because there is less
We have deliberately chosen many examples which high- relative precision (C V is lower) and, perhaps more impor-
light problems with C V and G. However, ga is not without tantly, the mode is shifted away from zero. The difference
398

Table 2. Comparison of gi, C V, and G for Bromus tectorum. Sam- tribution. In testing for a two-parameter lognormal distri-
ple sizes, means, skewness coefficients, coefficients of variation, bution, the average value o f the K o l m o g o r o v - S m i r n o v D
and Gini coefficients are given for number of seeds produced by statistic ( n > 5 0 ) was 0.13 for F. idahoensis and 0.26 for
individuals of B. teetorum grazed at weekly intervals (W), bi-weekly B. tectorum indicating that the F. idahoensis was closer to
intervals (B), monthly intervals (M), or grazed only once (O), and being two-parameter lognormally distributed than was B.
having these grazing treatments initiated at 7, 30, 90 or 150 days
tectorum (the hypothesis of lognormality was rejected ( P <
of age (plus ungrazed controls). The first four rows contain the
data associated with Figure 2A-D, respectively. Correlation (r) be- 0.05) for all o f the samples except for one F. idahoensis
tween gl and (~V=0,77, between g~ and G = - 0 . 1 3 , between CV sample). Moreover, robustness with respect to the value
and G = 0.42 0.5 used in place o f zero as well as visual inspection helped
to confirm the empirical evidence that F. idahoensis was
Interval Age N Mean gl CV (~ closer to following a two-parameter lognormal distribution.
Hence, the empirical results and the distributional consider-
B 30 147 47.2 7.31 2.43 0.71 ations are fairly consistent in explaining why g l , C V and
B 150 259 35.2 7.85 1.86 0.55 correlated well with F. idahoensis but not with B. tec-
W 90 232 20.4 1.09 1.16 0.61 torum.
M 7 34 104.7 1.35 1.44 0.70
B 7 106 108.5 4.89 1.74 0.57
O 7 41 181.6 1.29 1.38 0.68
M 30 247 14.0 3.34 1.53 0.68 Discussion
O 30 217 36.6 5.32 1.82 0.66
W 30 133 69.5 4.49 1.34 0.54 gl is primarily a shape statistic which measures skewness
B 90 259 12.9 4.19 2.08 0.78 and is b o t h location and scale invariant. In contrast, C V
M 90 265 19.1 7.95 2.04 0.66 and G are b o t h measures o f relative precision and are invar-
O 90 264 25.9 7.48 1.70 0.64 iant to scale changes but not to location changes. Further-
M 150 221 30.6 3.50 1.75 0.70 more, C V is a more sensitive measure of the (right) tail
O 150 195 31.0 6.69 2.16 0.66 of the distribution whereas G is less sensitive or a m o r e
W 150 215 20.1 2.28 1.35 0.64 robust measure. This knowledge is crucial when a choice
CTL CTL 265 28.5 3.27 1.36 0.60 is to be m a d e among these statistics. W h e t h e r these statistics
CTL CTL 247 26.1 3.10 1.41 0.62
are effective depends u p o n the question of interest as well
as the underlying distribution.
In practice, the distribution o f plant biomass or seed
in the Gini coefficients between Fig. 2 A and Fig, 2B is simi-
production is never k n o w n exactly and varies according
lar to the difference observed in the constructed examples
to the response variable, the species, etc. The lognormal
H1 and H 2 in Table 1. As noted in Table 1, G is very
model, however, is frequently discussed in the literature
sensitive to this small shift in the mean. Figures 2C, D have
(Koch 1966) and provides an i m p o r t a n t p o p u l a t i o n model
relatively small values o f gl and C V with m o d e r a t e and
for p l a n t distributions. Table 3 (see also Bendel and Carlin
relatively large Gini coefficients, respectively.
1988) presents the p o p u l a t i o n parameters for the lognormal
distribution as well as some other well-known distributions
Distributional considerations
(which m a y or m a y not be c o m m o n in ecology but are
Differences in the results for F. idahoensis and B. tectorum useful for illustration). N o t e for the two-parameter lognor-
are partially explained by differences in the underlying dis- mal where the lower b o u n d is 0 = 0 , that 71, CV, and G

B
120 -
80
A gl = 7.31 gl = 7 . 8 5

6c CV
^ = 2.43 90 i CAV= 1.86

^
: o7, 8o- G = 0.55
40

>-
2c

0
-I .
300
. .
N

600
147

. . .
900 1200
30

0 i
0
i L __
200
C--L---
400
N = 259

600 800
0
Z ZO
LU I 0 0 D
D C
gl = 1 . 0 9 gl = 1 . 3 5
0
'" 80 A A
n,,. CV = 1.16 CV = 1 . 4 4 Fig. 2.A-D. Histograms, skewness
1.1.
60 h coefficients (gl), coefficients of
G = 0.61 G
A = 0.70
IO variation ((~ V), and Gini
40 N = 232 N=34 coefficients (O~) for Bromus
teetorurn samples selected from
20
Table 2. The mean number of
seeds produced is indicated by a
0 dashed vertical line
50 60 90 120 t25 250 ,575 500

SEEDS PER PLANT

 399

Table 3. Parametric relationships of 7~,C V, and G for common distributions a

Distribution Skewness - ?,, Coefficient of Variation-C V Gini-G

q exp (5 + 22/2) (2 ~(2/1~ ) - 1) exp(~ + 22/2)

Three parameter lognormal b r/(2 + exp (22))
0 + exp (5 + 22/2) 0 + exp (5 + 22/2)
Two parameter lognormal (0 = 0) r/(2 + exp (22)) r/ 2 ~ (;4~/2)- 1
Two parameter exponential c 2 ~r/(0 + er) o.s a/(o + ~)
One parameter exponential (0 = 0) 2 t 0.5
Two parameter uniform d 0 dl/3/(3 a) d/(3 a)
One parameter uniform (a = d) 0 1/3/3 1/3
Normal (u = E(x), ~r= STD(x)) 0 cr//~ ~/(1/'~r')
a q = (exp(22)_ 1)~ and 4 ( - ) represents a standard normal cdf
b Density with lower bound 0> 0 is f ( x ) = [(x--0) 1 / ~ 2]- 1 exp [-0.5 (log(x- 0)-~)2/22] where ~ = location parameter and 2 = shape
parameter, x > 0. Note (E(x) = exp (5 + 22/2) + 0 and STD (x) = r/exp (~ + 22/2)
c Density with lower bound 0 > 0 is f ( x ) = G- 1 exp [ - ( x - 0)/a], x > 0, er> 0. Note: STD (x) = er and E(x) = 0 +
a Uniform densityf(x)= (1/2 d) defined on the interval a _+d, 0 < d_< a

are all monotonically increasing functions of the lognormal
shape parameter, 2. Hence, if the response variable of inter-
est follows an approximate two-parameter ( 0 = 0 ) lognor-
mal distribution, then all three of the statistics are likely
to be positively correlated in different samples. If the re-
sponse variable follows a three-parameter lognormal (0 > 0)
or a non-lognormal distribution, the three statistics may
or may not be correlated, depending upon the distribution
of the response variable. Aitchison and Brown (1957) dis-
cuss the two and three-parameter lognormal distribution
extensively. The primary difference between the two- and
three-parameter lognormal distribution is the lower bound,
0, which is zero for a two-parameter and positive for a
three-parameter lognormal distribution. Since there is no
difference in shape between the two, there is no difference
in skewness. To illustrate the use of Table 3 for an exponen-
tial distribution, the population skewness coefficient is nu-
merically equal to two regardless of the lower bound, 0.
In contrast, the population Gini coefficient is one-half as
large as the CV, so that the sample Gini and sample C V
are likely to be correlated to each other in samples from
different two-parameter exponential distributions. Both the
sample Gini and sample C V would probably correlate low
with the sample skewness coefficient.
The Pigou-Dalton condition of transfer, illustrated in
Table 1 (series B), is considered an important criterion for
a statistic to possess in economics (Sen 1973). C V and
possess this property but g~ does not. Although the Pigou-
Dalton condition is important in assessing economic in-
equality, it does not follow that this condition should be
important in assessing inequality in ecology. For example,
in economics total wealth is finite and the units of wealth
may be transferred from rich to poor population members.
Finite wealth may be analogous to carrying capacity, but
conditions under which the wealth can be transferred from
" r i c h " to " p o o r " plants are more difficult to imagine. One
such case may be under frequency-dependent selection
where the few large plants in a population are selected
against as the frequency increases from generation to gener-
ation. Given the evolutionary significance of changes in
the large plants in the right-hand tails of the distribution,
one would probably be well advised to use C V or g~ rather
than G. The choice between C V and g~ would depend upon
other previously mentioned considerations as well as the
underlying distribution.
Clearly, interpretation of results which stem from com-
parisons of frequency distributions may be influenced by
the statistic chosen for the comparison. Because of this,
the researcher should have some knowledge of the fre-
quency distribution before the comparison is attempted.
For example, say that individual seed production is the
response variable, and that many plants produce no seeds.
In this case, seed distribution is near the origin and the
distinction between zero reproductive potential and one
seed may be more important than individuals in the right-
hand tail; G may be the appropriate statistic (contrast D 1
and D 4 with E l and E4, Table 1). Alternatively, if this
distinction is not important, and very few individuals occur
in the right-hand tail, C V may be more appropriate. Yet
another case may be when comparisons of biomass distribu-
tions are to be made over time, so that the location of
each sample shifts, e.g., as in a three-parameter lognormal
distribution illustrated by series D through G in Table 1.
Now, the skewness coefficient may be indicated.
The importance of the knowledge of the underlying dis-
tribution is further emphasized by contrasting the results
from F. idahonensis with those from B. tectorum. For F.
idahonensis, all three statistics were highly correlated be-
cause the distributions followed, at least approximately, the
two-parameter lognormal distribution. With such a high
correlation, conclusions drawn from the results of the F.
idahonensis experiment (Higgins et al. 1984) would be the
same regardless of which statistic was used. For B. tectorum,
gl and G were negatively correlated (albeit non-significant-
ly, P > 0.05); these distributions were not as well approxi-
mated by the two-parameter lognormal distribution. Con-
clusions drawn from the B. tectorum results depended upon
the statistic used. Using gl, comparisons of those popula-
tions grazed biweekly with the controls (Table 2) indicated
that biweekly grazing consistently increased skewness and
decreased the proportion of the population producing seed
regardless of when grazing was initiated. But this increased
inequality was greatest when grazing was initiated at 30
or 150 days. Comparisons of the same populations based
on C V also indicated that biweekly grazing increased in-
equality over controls, but the effect was less when grazing
was initiated either early or late in the population's develop-
ment (Table 2). In contrast, comparisons based on G sug-
gested that inequality decreased when grazing was initiated
early or late (7 or 150 days), but inequality increased over
controls when grazing was initiated at 30 or 90 days (Ta-
ble 2).
400
W i t h some d a t a sets, the choice o f a statistic is n o t
clear and none o f the three statistics m a y be appropriate.
Histograms are, however, indispensible and should always
be used to support conclusions based on a single s u m m a r y
statistic such as g l , C V, or G. W e recommend that ecolog-
ists first determine which characteristics o f distributions are
i m p o r t a n t in their own research. F o r example, in an evolu-
tionary study, the concentration o f seed production in very
few individuals in the right-hand tail m a y be the m o s t im-
p o r t a n t characteristic. Alternatively, one m a y choose to ex-
amine machanisms affecting the biomass distributions from
populations planted at different densities or grown under
different competition regimes. In short, ecologists need to
know more a b o u t what is i m p o r t a n t a b o u t frequency distri-
butions in p o p u l a t i o n ecology.
The skewness coefficient is suggested when the distribu-
tion is either three-parameter lognormally distributed (or
close to such), or when a measure o f relative precision is
not indicated. However, if preference is given to measures
of relative precision, we believe that generally, the C V
should be used. The C V is more sensitive (less robust) to
observations in the right-hand tail o f the distribution. Rec-
ognition o f this difference is especially i m p o r t a n t in light
o f the evolutionary significance associated with having pop-
ulation reproductive potential concentrated in just a few
individuals.
Acknowledgements. We would like to thank K. Rice, A. Marcus,
R. Mack, R. A. Black, and J. Weiner for encouragement and for
helpful comments throughout the course of this study, and P. Pe-
dersen for technical assistance.
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Received April 15, 1988