J. Theoret. Biol.

(1969) 22, 221-237

Measurement of Non-circular Home Range

R. I. JENNRICH

Department of Biomathematics, University of California,

Los Angeles, California, U.S.A.
AND

F. B. TURNER

Laboratory of Nuclear Medicine and Radiation Biology,

University of California, Los Angeles, California, U.S.A.

(Received 16 July 1968)

We examinethe use of conventional samplingdata to estimatethe area

extent of an animal’shome range. Popular methodsof estimation based
on polygons and capture radii are discussed. A simpleexampleshowsthey
are not directly comparable.Furthermore, whenhomerangeslack circular
symmetry, methodsbasedon capture radii suffer largebiases.On the other
hand, methodsbasedon polygons have samplesizebiases.A new method,
basedon the determinant of the covariance matrix of the capture points,
is introduced. We show that this method, under specifiedassumptions,is
free from both forms of bias.Tablesthat identify the biasesin the polygon
methodsare provided. An important use of the tables, that of making
comparisonsbetweenvalues obtained by the various methodsof estima-
tion, is illustrated. The statisticalstability of eachmethodis given by means
of a table or formula for its relative variance as a function of samplesize.
Finally, the methods are evaluated. For purposesof estimating home
range area, the determinant method is recommended,the old convex
polygon method is defended, and the use of methodsbasedon capture
radii is discouraged.

1. Introduction
Estimates of the area of home ranges and territories have been a regular
feature of studies of the natural history of terrestrial vertebrates. The
desirability of such data has been recommended in a number of outlines
for ecological life history studies (Fitch, 1949; Skutch, 1950; Cagle, 1953,
1956). It is evident that analyses of home range data, particularly inter-
specific comparisons, have been discouraged by differences in the techniques
utilized. Presumably all methods are devoted to the measurement of the
227
228 R. I. JENNRlCH AND F. B. TURNER

same variable, but the same data analyzed by different methods may yield
appreciably different results. Then, too, the applicability of various methods
must be considered. Analytical procedures assuming circular home ranges
will yield misleading estimates if used with data exhibiting significant linearity
(e.g. see Stumpf & Mohr, 1962).
In this paper we review three techniques which have been used to estimate
home ranges, with a view towards (1) examining their limitations and biases
and (2) developing a set of scaling factors which may be used to render
data analyzed by different methods comparable. We also propose a new
technique which is free from sample size bias and the assumption of circular
symmetry.

2. Three Earlier Procedures

One of the simplest methods of estimating home range (Southwood,
1966, p. 262) involves drawing the smallest convex polygon which contains
all of the capture points (pi, pz, . . . , p,) for an animal and taking
A, = (area of the convex polygon) (1)
as an index of the animal’s home range.
A related procedure is obtained by dropping the requirement that the
polygon be convex. This leads to minimum polygon indexes for home
range (Stickef, 1954; Southwood, 1966). To our knowledge, this procedure
has never been precisely defined. Stickel(l954) simply stated that the capture
points are connected and the area of the resulting polygon measured. The
difficulty is that the shape of the polygon and its area depend on the order in
which the capture points are connected. We can make this method precise
by ordering the capture points counterclockwise about their geometric
center and connecting them to obtain a closed polygon. Then
A, = (area of the minimum polygon) (2)
defines an index of home range. The required area may be computed by
the mapmaker’s formula:

A2 = &i (xiYi+l-xi+lYi). (3)

i=l

Here (Xi, yi) is the ith ordered capture point from a total of 12points and
(x”+~, y,,,) = (x1, vi). This formula may also be used in evaluating A,.
Its use is illustrated in Table 1.
Finally, several procedures have been defined which are based on recapture
radii (Hayne, 1949; Dice & Clark, 1953; Harrison, 1958; Calhoun & Casby,
 MEASUREMENT OF NON-CIRCULAR HOME RANGE 229
1958; White, 1964). Our discussion will be based on the remarks of Calhoun
and Casby, but the basic concepts are common to all of these papers.
Calhoun and Casby defined the capture radius, rir for the ith capture
- -
point,p, = (xi, vi), as the distance from pi to the geometric center, p = (x, y),
of the capture points. They computed the statistic,

(4)

and estimated home ranges by a formula of the form

A, = 67~‘. (5)
Actually, Calhoun and Casby use the value 9 in place of the value 6 in this
formula. The choice is somewhat arbitrary and will be discussed in section 4.

3. Limitations of Earlier Procedures

Even if we define home range as simply the area an animal covers in its
day-to-day travels (Burt, 1943), we can identify three problems arising
from the use of polygon and capture radii indexes of home range size. First,
the indexes are not directly comparable. The co-ordinates (xi, ui) of the ten
points in Table 1 are capture loci for a lizard (Cnemidophorus tigris) obtained
with a grid of can traps. For these points the values of the three indexes
are A, = 42, A, = 24 and A, = 265, and graphical representations of
these areas are given in Fig. 1. While all three of these indexes measure
the extent of day-to-day movements, they obviously do not do so in directly
comparable terms. Second, the polygon indexes have a sample size bias.
Their values have a tendency to increase as the number of capture points
increases. (The actual increase in the expectations of A, and A,, under
specified sampling assumptions, are given in Table 2 under the headings c,
and rl,, respectively.) Third, the capture radii index fails to respond properly
to the shape of the home range. Circular home ranges tend to give smaller
estimates than long narrow ranges encompassing the same area. Actually,
A, does not measure the area an animal covers in its daily movements,
but rather how far from home it tends to be. (It is, in fact, a constant, times
its average squared distance from j7.) Some authors have observed circular
home ranges (Calhoun & Casby, 1958), but Stumpf & Mohr (1962) have
described obviously non-circular ranges for birds and mammals, and linear
home ranges have been observed in several species of lizards (Fitch, 1956,
1958). For comparative studies it is important that an index of home range
area evaluates circular and non-circular ranges without bias.
230 R. I. JENNRICH AND F. B. TURNER
1
TABLE
Capture points of a lizard (Cnemidophorus tigris) and the
computation of A,, A,, A, and A,

i Xl 4’1 xf 4‘; .U,Yl ~~-,?‘t+l

- .Yl,lJi -Yt?‘i+1.- x:+lr‘
t1 $69 88 4161 7744 6072 176 523
2 67 88 4489 7744 5896 176

7-i 63
65 88 4225
3969 1744
7744 5544
5720 -302
176 -32s
5 65 86 4225 7396 5504 -258

tl: 67
65 84
86 4489
4225 7056
7396 5460
5762 -374
124 -1176
8 71 86 5041 7396 6106 -830

tli 79
75 84
86 6241
5625 7396
7056 6636
6450 494
666 494
666
Totals 686 864 41290 74672 59236 48 84
Al = 4 x 84 = 42, AZ = : x 48 = 24.
47290 - 74672 -
s2 = 1 (686)2 + (864)2 I = 14.04, AU = 61-d : 265.
18 10 10
= 28 8 s = 2,8,
47290 10
- (686)’ I ., yy=- 74672 10
- (864)2 1
--4,3
, ISI = s,,syy - s& = 62.15.
1
A., = 6nlSlf = 149.

t Convex polygon vertices.

$ Co-ordinates are in grid units. One grid unit is 7.5 m.

1 _ +.--i-J
60 65 70 ii 80

FIG. 1. Four methods of estimating home range size from a dispersion of capture loci
(PI, pa,. . . , plO). Illustrated are (1) the convex polygon (Al = 42), (2) the minimum poly-
gon (AZ = 24), (3) a circle based on the mean recapture radius (A, = 265), and (4) an ellipse
based on the covariance matrix of the capture loci (AI = 149).
 MEASUREMENT OF NON-CIRCULAR HOME RANGE 231
TABLE 2
Biasesc, and d, from formulas (20) and (21) together with the relative variances
of Al and A2
n C” dn relvar (A,) relvar (Aa)
3 ox)43 0.043 1.052 1.052
4 0.092 0.088 0.506 0.525
5 0.135 0.122 0.368 0.381
6 0.171 0.146 0.262 0.279
I 0.196 0.158 O-217 0.244
8 0.231 0.178 0.161 0.190
9 0.257 0.188 0.153 0.171
10 0.284 0.201 0.130 0.157
11 0.301 0.203 0.127 0.150
12 0.325 0.211 0.109 0.132
13 0.348 0.220 0.108 0.126
14 0.363 0.221 0.101 0.117
15 0.390 0.228 0.091 0.105
16 0404 0.230 0.082 0.101
17 0.418 0.232 0.080 0.097
18 0.436 0.239 0.073 0.093
19 0.448 0.236 0.068 0.086
20 0.463 0.240 0.067 0.076
21 0.468 0.241 0.061 0.075
22 0.481 0.239 0.058 0.073
23 0.496 0.244 0.058 0.017
24 0.505 0.244 0.056 0.073
25 0.518 0.241 0.057 O-066

Each valuejlwas!obtained by computing the areas of 1000 polygons constructed from

capture points selected at random from a bivariate normal distribution with a = 1.

4. Definition of a New Index

Our new index is similar to A, but is designed to measure non-circular
as well as circular home ranges. It is given by the statistic
A;= 67cjSl*. (6)
Here:ISI is the:determinant of the capture point covariance matrix

(7)
defined by the equations

(8)
232 R. I. JENNRICH AND F. B. TURNER

Although it is not relevant to the material which follows, this statistic ma)
be given a geometric interpretation. Let Aij be the area of the triangle
defined by the capture points pi and pj, i # j, and the capture center p, then

A, = a,
where
6rc[2n(n - l)]”
CI,=
(n-2) *
Thus A, is a constant (approximately 6n,,/2) times the root mean square
of the areas Aij.

5. Definition of Home Range

So far we have discussedhome range without giving it a precisetheoretical
definition. We will first assume that the intensity with which an animal
utilizes each point in its habitat is expressed by a bivariate normal distri-
bution. Utilization distributions were first suggestedby Hayne (1949) and
given detailed expression by Calhoun & Casby (1958). The home range of
an individual is considered to be the area of the smallest sub-region which
accounts for a specifiedproportion, p, of its total utilization. Unlike Calhoun
and Casby we will not assumethat the utilization distribution has circular
symmetry. The value of p, which is somewhat arbitrary, will be specified
below. Let

be the mean vector and covariance matrix of the bivariate normal utilization
distribution. It is easily shown that the regions of most intense utilization
are bounded by concentric, constant density ellipsesof the form
(Z - p)C- ‘(2 - r*)’ = constant, (11)
where z denotes an arbitrary point on the ellipse, 2-i denotes the inverse
of C, and (Z-P)’ denotes the transpose of the vector Z--P. An ellipse of this
form which accounts for a proportion p of an animal’s total utilization of
its habitat has an area
a = 7Cln(l-p)-2jC~+. (12)
By settingp = 1-e-3 = O-950formula (12) simplifies to
a = 674X]“. (13)
We will adopt equation (13) as the definition of home range. It is the arca
 MEASUREMENT OF NON-CIRCULAR HOME RANGE 233
of the smallest region which accounts for 95% of an animal’s utilization of
its habitat. The elliptic area calculated from the points in Table 1 is illus-
trated in Fig. 1. If the 6, in (13), is replaced by 9 (the value used by Calhoun
& Casby, 1958), the area accounts for 99% of an animal’s utilization of its
environment.

6. Making Home Range Estimates Comparable

Comparison of the home range estimates A,, AZ, A, and A, requires a
mathematically precise definition of home range. For this we adopt that
given by equation (13) of the previous section. We also assume that the
capture points pr, . . . , p,, constitute a random sample from the bivariate
normal distribution introduced there.
The indexes A,, A, and A, differ from A, in an important way. As esti-
mators of a they are invariant under linear transformations. As a conse-
quence their expected values depend on a, and possibly on n, but in no
other way on the underlying normal utilization distribution. Unfortunately
this is not true for A3.
It can be shown that the expected value of A, is given by
E(A,) = ba, (14)
where the bias,
b = -%$

Here, tr (C) denotes the trace, or sum of the diagonal elements, of C. Thus,
although the bias of A, is totally independent of n it is not independent
of C. When the bivariate normal utilization distribution has circular sym-
metry, b = 1 and the estimate A, is unbiased. Otherwise, its bias depends
on the eccentricity of the constant density ellipses. In particular, if the ratio
of the major to minor axes is 2, then b = 1a25 and, if the ratio is 5, then
b = 2.6. In every case, where circular symmetry is lacking, the expected
value of A, exceeds a, and in some cases substantially. In contrast, the
estimate A, is completely unbiased. That is
E(A) = a, (15)
regardless of the eccentricity of the constant density ellipses, their center,
or the sample size. The expectations of A, and A, have the form
E(A,) = c,n (16)
E(A,) = c&u. (17)
The biases c, and d,, given in Table 2 depend on n but, as pointed out above,
are completely independent of the form of the underlying normal utilization
234 R. I. JENNRICH AND F. B. TURNER

distribution. While it is easy to show that the expectations of A, and AZ

have the form given by equations (16) and (17), we used simulation to obtain
the values of c, and d,, given in Table 2. Each value was based on 1000
samples of size y1from a standard bivariate normal population (a total of
23000 pseudo-trapping experiments). Investigation of the samples suggests
that the tabled values are accurate to about two significant figures. The bias
of A1 is somewhat more dependent on the number of capture points than
that of Al.
Perhaps the most valuable use of Table 2 is that of making comparisons
between particular values of Al and AJ, estimates commonly encountered
in the literature. Comparisons involving A3 require the assumption of
circular symmetry. Under this assumption, -Al has the same expected value
as A,. Thus, if one author has reported a convex polygon area of 300 square
feet based on 15 captures of an animal, and another has reported a home
range of 870 square feet based on any number of capture radii and computed
by formula (5), it would be reasonable to use

-Al = 300/0.390 = 769 and A, = 870

Cl5

as comparable, in fact unbiased, estimates of home range in the two studies.

If this bias adjustment is made for the example given in section 3, the two
unbiased estimates are

-Al = 4210.284 = 148 and A, = 265.

Cl0
The disparity between the two estimates reflects the evident lack of circular
symmetry in the dispersion of the data (Fig. l), By contrast A, has the
value 149, which compares surprisingly well with the value 148 of Al. Since
different authors use different values for the constant in formula (5) a further
adjustment may have to be made when the constant used differs from the
value, 6, used in (5).
Table 2 also helps in combining convex polygon estimates based on
unequal numbers of capture points. Thus, if Ay), Ai” and Ai3’ are convex
polygon areas for three animals based, respectively, on 5, 10 and 15 capture
points, then

1 (18)

is an unbiased estimate of their (assumed) common home range area a.

It is not necessary to assume that the home ranges have the same shape.
 MEASUREMENT OF NON-CIRCULAR HOME RANGE 235

7. Statistical Stability of the Estimates

It is convenient to measure statistical stability in terms of relative variance

Assuming the capture points pi,. . . , pn are a random sample

from the bivariate normal distribution introduced in section 5, the relative
variance of A, is given by the formula

The relative variance of A, is given by

(20)

where
c = 2W (9-2[Cll
tr2(C) ’
As with its expected value, the relative variance of A, depends on the eccen-
tricity of the constant density ellipses. If the ellipses are circles, then C = 1,
its minimum value. If the ratio of the major and minor axis is 2, then
C = 1.36 and, if the ratio is 5, C = 1%. In every case the relative variance
of A, satisfies the inequality
1
~ I relvar(A,) 5 2 (21)
n-l n-l’
Like that of A,, the relative variances of A, and A, depend only on n. These
values, obtained by the same simulation used to obtain the values of their
expectations, are given in Table 2. In particular, when it = 10 and the
bivariate normal utilization distribution has circular symmetry, relvar
(A,) = 0.130, relvar (A,) = 0.157, relvar (A,) = 0.111, relvar (Ad) = O-125.
These relative variances do not differ greatly; that of A, is the greatest and
that of A, is the least. When circular symmetry fails, however, A3 becomes
biased and its relative variance increases. If the ratio of the major and minor
axes of the constant density ellipses is 2, for example, the relative variance
of A, increases to 0.151.
It is interesting to note the magnitude of these relative variances. The
relative standard deviation (the square root of the relative variance) of
A, is O-354 when n = 10. This means that a 35% change in A, from one
determination to the next would not be statistically surprising. This suggests
averaging several estimates. For example, if Ay), . . . , A$“) are determinant
236 R. I. JENNRICH AND F. B. TURNER

home range estimates [equation (6)] obtained from f?z animals and based
on nr,. . ., II,,, captures, respectively, then

f (n,-2)Ak”
(j = i=’ (22)
iEl(i1i-2)

is an unbiased estimate of the animals’ (assumed) common home range.

A simple unweighted average could also be used. As in equation (18) it
is not necessary to assume that the home ranges have the same shape.

8. What Estimate Should We Use?

Many results have been published using the indexes A, and A,, and indexes
similar to A,. The index A, appears to be new. The index A, has historical
prominence, graphical simplicity, and exhibits reasonably good statistical
stability. It is important to remember, however, that A, has a bias which
depends on n, and to make allowances for this when comparing (or com-
bining) these indices. This may be easily done by the use of Table 2.
While the bias of A, is less dependent on n than that of A,, its graphical
evaluation becomes unwieldy as the number of capture points increases.
Its algebraic evaluation, through the use of formula (3), is simple, but its
statistical stability is somewhat inferior to that of A,, A, and A,. Further-
more, as far as we can ascertain, A, has never been rigorously defined. These
considerations make us reluctant to recommend any polygon procedure
other than the convex polygon index A,.
The estimate A, seems to be an unfortunate historical accident, because.
as the estimate A, demonstrates, the assumption of circular symmetry
upon which A, is predicated is unnecessary. With the assumption of circular
symmetry, A, is statistically a little more stable than A,. However, this
advantage is far outweighed by the bias A, suffers when this assumption
fails. Calhoun & Casby (1958) made tests for circular symmetry. When no
apparent departures exist it seems reasonable to accept such results as
essentially unbiased, but for future work the authors recommend replacing
the recapture radii estimator, A3, by the determinant estimator, A,.
On balance we recommend the use of A, because of its lack of bias and
high statistical stability. When a quick graphical estimate is required, we
recommend A, because of its simplicity and historical prominence. When
comparing or combining these estimates Table 2 should be employed to make
corrections for sample size bias.
 MEASUREMENT OF NON-CIRCULAR HOME RANGE 237
The authors wish to thank Dr Ellen Coutlee of the University of California at
Riverside for reading the original manuscript and providing valuable references,
Mr Anthony Lynn of UCLA for writing the computer programs required for the
construction of Table 2, and the Health Sciences Computing Facility at UCLA
for the use of their computer.
This research was supported in part by National Institutes of Health grant FR-3
and by contract AT(O4-1) GEN-12 between the U.S. Atomic Energy Commission
and the University of California.

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