Frequency distributions of sward height under
sheep grazing
G. T. Barthram*, E. I. Duff†, D. A. Elston†, J. H. Griffiths*, T. G. Common‡ and C. A. Marriott§
*Macaulay Institute, Hartwood Research Station, Shotts, UK, †Biomathematics and Statistics Scotland, Macaulay
Institute, Craigiebuckler, Aberdeen, UK, ‡Macaulay Institute, Sourhope Research Station, Kelso, UK, and
§Macaulay Institute, Craigiebuckler, Aberdeen, UK
Abstract
Cattle and sheep can create and maintain a mixture of
relatively tall and short patches in grass swards through
selective grazing. In swards that are grazed by cattle this
heterogeneous structure can result in the frequencies of
height measurements having a skewed distribution that
has variously been better described by the double-
normal distribution the gamma distribution and the
Weibull distribution than by the more common normal
distribution. The fit of these statistical distributions, and
the adequacy of the potentially useful log-normal
distribution, to sward height frequencies were tested
in sown temperate swards grazed by sheep and
compared within a single sward. It was concluded that
the single-normal and Weibull distributions were in-
adequate and that overall the log-normal and gamma
distributions had the best fit to the measurements.
Keywords: sward height, frequency distribution,
double-normal distribution, gamma distribution,
Weibull distribution, log-normal distribution
Introduction
Cattle and sheep can create and maintain a mixture of
relatively tall and short patches in grass swards through
selective grazing (Edwards et al., 1996). This is one
result of their tendency to forage on short vegetative
material and to avoid tall stemmy herbage where leaf
material is less accessible (Devries and Daleboudt, 1994;
Laca et al., 1994) and areas contaminated by faeces. The
opportunities for exercising selectivity in diet selection
are likely to be higher when stock densities are low and,
therefore, it can be hypothesized that there will be a
Correspondence to: E. I. Duff, Biomathematics and Statistics
Scotland, Macaulay Institute, Craigiebuckler, Aberdeen
AB15 8QH, UK.
E-mail: e.duff@bioss.ac.uk
Received 1 July 2004; revised 10 October 2004
4
2005 Blackwell Publishing Ltd. Grass and Forage Science, 60, 4–16
greater variability in sward height or biomass at low
stock densities. For predictive and modelling purposes it
is often desirable to know the distribution of sward
heights or herbage biomass in grazed systems. Although
a normal distribution is frequently assumed, sward data
may be skewed, bimodal or have a plateau shape and so
other distributions may be more appropriate. For
example, a positively skewed or bimodal distribution
may occur under low stocking densities where a broad
range of taller heights are created as a result of different
frequencies and extents of consecutive defoliation
events. Distributions that are appropriate for describing
positively skewed data include the double-normal, log-
normal, gamma and Weibull distributions. Gibb and
Ridout (1986; 1988) found that the frequencies of
height in swards continuously stocked by cattle had a
skewed distribution that was better described by a
double-normal distribution than by the more common
normal distribution. Since the five-parameter double-
normal distribution consists of a mixture of two
overlapping normal distributions, and an estimate of
the proportion of the data belonging to each distribu-
tion, this distribution has been used as a tool to quantify
the proportion of swards that are occupied by ‘tall’ and
‘short’ vegetation and to characterize these fractions
(Fisher et al., 1995; Gibb et al., 1999; Morris et al.,
1999). However, evidence of bimodality is not always
strong and the gamma distribution has also been
suggested as a good descriptor of frequency distributions
of herbage mass under rotational cattle grazing (Shiy-
omi et al., 1983; 1984; 1998; Tsutsumi et al., 2002;
Sanders et al., 2003) and for herbage height under
rotational cattle grazing (Sanders et al., 2003). The
three-parameter Weibull distribution has also been
shown to be a better fit than the normal distribution
to height and mass data in semi-arid short-grass steppe
grazed by cattle (Remington et al., 1992). Swards grazed
by sheep can also develop tall and short patches
(Edwards et al., 1996; Morris et al., 1999). Sheep and
cattle have different grazing habits and forage require-
ments and, consequently, can have different impacts on
 Frequency distributions of sward heights under grazing
sward structure (Grant et al., 1985; Hodgson et al., 1991;
Armstrong et al., 1997; Morris et al., 1999). From
published studies it is not clear which distribution
would best describe the frequency distribution of sward
height in pastures grazed by sheep, particularly in
swards that are grazed at low stocking density. We
determined which of five distributions, normal, double-
normal, log-normal, gamma and Weibull, gave the best
fit to the sward height frequencies of swards grazed by
sheep. All the distributions above, excluding the normal
distribution, are appropriate for fitting to data showing
some degree of positive skewness and, in addition, the
log-normal, gamma and Weibull distributions all
naturally fit to positive (above zero) data. The study
not only focused on swards grazed at a low stocking
density over two growing seasons at two sites but also
included measurements on a single date from more
intensively grazed swards at one site for comparative
purposes.
Materials and methods
Swards
Measurements of sward height were made at the
Macaulay Institute’s Hartwood (351¢W, 5549¢N) and
Sourhope Research (214¢W, 5529¢N) Stations in
Scotland. The two sites have different climatic and
edaphic conditions (Barthram et al., 2002). The Sour-
hope site has a slightly lower annual rainfall than the
Hartwood site. The soil at the Hartwood site is a poorly
draining non-calcareous gley while that at the Sour-
hope site is a freely draining brown-forest soil. Pasture
growth rate during dry periods appears to be reduced
more at Sourhope than at Hartwood. At each site,
measurements were made monthly during 2001 and
2002 in two replicate paddocks (each of 0Æ45 ha) that
had been grazed to maintain an average sward surface
height of 8 cm during the whole of each growing season
since 1990 and had not received any fertilizer during
this time. In addition, a limited set of height measure-
ments were also collected on shorter swards at one site,
Hartwood, in September 2003. The four paddocks
used had been maintained with average sward
heights of 4 cm since 1990. Two had received
140 kg N ha
1 year
1 as compound fertilizer (paddocks
2 and 9) and the remaining two had not. Sward height
in all paddocks was controlled by altering the number of
grazing Scottish Blackface ewes (and their lambs until
weaning) in response to weekly estimates of mean
sward height. As a result of differences in pasture
growth, sheep density varied during the season, e.g.
from 4 to 17 ewes ha
1 on swards maintained at 8 cm,
and between treatments and sites. At both sites, the
number of sheep carried on the 8-cm swards from the

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
5
start of the growing season in April until the end of
September was around 0Æ40 of that carried on fertilized
swards maintained at 4 cm, and it fell to around 0Æ25
from then until mid-November.
In 1990, the species composition of all swards was
around 0Æ60 Lolium perenne, 0Æ10 Trifolium repens, and
0Æ30 unsown species (these differed between sites, with
Agrostis capillaris dominant at Sourhope and Poa species
dominant at Hartwood) (Marriott et al., 2002). By 2001,
the proportion of L. perenne in swards maintained at
8 cm had fallen to around 0Æ40 at Hartwood and 0Æ25 at
Sourhope, with corresponding increases in unsown
grass species. There had been little change in the overall
proportion of sown species in swards maintained at
4 cm.
Sward height
Sward height measurements were made at approxi-
mately 300 locations spread across the whole of each
paddock. Measurements were made using the HFRO
sward stick (Bircham, 1981; Barthram, 1985). This
consists of a 2 cm · 1 cm ‘window’ that is lowered
until it makes contact with a green leaf. The height of
the window above ground level, measured to the
nearest 0Æ5 cm, is the height of the sward at that
location. There were eight locations (all in paddock 11
at the Sourhope site) where a height of 0 cm was
recorded. As the true height of these leaves was in the
range 0–0Æ25 cm (measurements were recorded to
the nearest 0Æ5 cm), the zero values were converted
to the midpoint of this range, 0Æ125 cm. This was to
avoid computational problems caused by zeros when
fitting the distributions.
Statistical analysis
In total, sixty sets of height frequency data were
generated, covering not only site but also temporal
variability. Single-normal, double-normal, log-normal,
gamma and Weibull distributions were fitted separately
to each set of measurements. The single-normal distri-
bution (the classic Normal distribution is referred to as
the single-normal distribution in the paper to avoid
confusion with the other forms of the Normal distri-
bution) is defined by its mean l and variance, r2. It has
the following density function (Payne, 2002):
 
1 1 x
l2
f ðxÞ ¼ pffiffiffiffiffiffi exp
¼ Uðx; l; rÞ
r 2p 2 r
The double-normal distribution can be used when an
observation may come from either of two single-normal
distributions with different means. It can occur when a
proportion p of the population is normally distributed
with mean l1 and variance r21 and a proportion (1
p)
 6 G. T. Barthram et al.
is normally distributed with mean l2 and variance r22 .
The distribution is thus described by five parameters.
The density function is:
f ðxÞ ¼ pUðx; l1 ; r1 Þ þ ð1
pÞUðx; l2 ; r2 Þ
with mean pl1 + (1
p)l2 and variance
pr21 þ ð1
pÞr22 þ pð1
pÞðl1
l2 Þ2 .
The log-normal distribution assumes that loge(X) has
a Normal distribution with mean l and variance r2, and
must have positive skewness. The density function is:
1
f ðxÞ ¼ UðlogðxÞ; l; rÞ
x Results
with mean exp (l + r2/2) and variance exp (2l + r2)
[ exp (r2)
1].
The gamma distribution is a general empirical distri-
bution described by two parameters, k (the shape
parameter) and b (the scale parameter), with mean
k/b and variance k/b2. The density function is:
Z1
e
bx
f ðxÞ ¼ bk xk
1 where CðkÞ ¼ x k
1 e
x dx; x > 0:
CðkÞ
0
The three-parameter Weibull distribution is a general-
ization of the exponential distribution. The density
function is:
c to the right when graphed as a frequency histogram).
f ðxÞ ¼ cbc ðx
aÞc
1 e
ðbðx
aÞÞ x > 0; b; c > 0;
which has a mean of a + (1/b)C((c + 1)/c) and a median
of a + (1/b)( log (2))1/c, and where c is the shape
parameter, b the scale parameter and a is a location
parameter (which allows the minimum theoretical
value of the distribution to take values other than zero).
For all five distributions, parameter estimates were
calculated using the method of maximum likelihood
(Genstat v. 6.2; VSN International Limited, Oxford,
UK), a technique which involves calculating values of
the parameters that give the highest likelihood for any
given set of data.
The Akaike information criterion (AIC) statistic
(Akaike, 1974), derived from the log-likelihood statistic,
was used to evaluate the fits of the different distri-
butions. The AIC statistic, defined as
2(maximized
log-likelihood) + 2(number of parameters), gives a
single quantitative measure, based on summarizing
the deviances of the fitted theoretical distribution from
the observed data. The AIC statistic was considered an
appropriate statistic as it is penalized by the number of
parameters, thus compensating for the greater flexi-
bility in fitting usually associated with higher numbers
of parameters (Ginzburg and Jensen, 2004). Lower AIC
values indicate a better fit between the observed data
and the theoretical distribution than higher AIC values.
In addition, to highlight any systematic differences
between the fitted theoretical density functions and the
frequency distribution of height observations over the

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
range of the data, we constructed charts showing the
differences between the proportions of observed heights
and the theoretical density functions that fell
within intervals of the observed height data. One
centimetre intervals were used for the 8-cm swards and
0Æ5 cm intervals for the 4-cm swards. These charts
allowed an examination of any pattern in the deviances,
whereas the AIC statistic, described above, is a measure
based on summarizing deviances. Each chart was based
on the combined data from the two paddocks at each site.
The mean heights in the swards with target heights of
8 cm varied around the target height (Figure 1, row 1,
parameter l). The standard deviations changed with
time of year, being greatest in midsummer to late
summer (June–September) and with greater maxima at
Sourhope than Hartwood (Figure 1, row 1, parameter
r). The 4-cm swards were nearer target and less variable
(Table 1, single-normal distribution) than the 8-cm
swards.
Preliminary visualization of sward height data using
frequency histograms showed that most datasets were
positively skewed to some extent (i.e. data showed a tail
Typical examples are shown in Figures 2 and 3. Con-
sequently the single-normal distribution had the poor-
est fit (highest AIC statistic) to the data (in fifty-three of
fifty-six instances in the 8-cm swards, and in four of
four instances in the 4-cm swards; Tables 2 and 3). It
tended to fit comparatively well at the beginning of the
seasons in the 8-cm swards before grazing commenced
after the winter rests, but never had the outright best fit
of the five distributions. In the 8-cm swards the Weibull
distribution had the best fit (lowest AIC statistic) of the
tested distributions to the data on five occasions. The
double-normal distribution was best on seven occa-
sions, but the gamma was best on eighteen and the log-
normal best on twenty-eight occasions (note that there
were three instances when two distributions were
equally the best). In the 4-cm swards the log-normal
and gamma distributions each fitted best in two swards,
with the double-normal being joint-best with the
gamma distribution in one sward.
The mean improvement in fit of the distributions,
compared with the fit of the single-normal distribution,
was examined. The mean difference between the
single-normal and double-normal distributions was 79
AIC units, and this improvement in fit was smaller in
the 4-cm swards (43 AIC units). In the 8-cm swards, the
log-normal (mean difference 81 AIC units) and the
gamma distributions (mean difference 82 AIC units)
were, on average, marginally better fits to the data
than the double-normal distribution. The Weibull
 Frequency distributions of sward heights under grazing 7

Figure 1 Parameter values fitted to sward height frequency distributions in 8-cm swards. Row 1, parameters l and r of single-
normal distributions; row 2, parameters l1, l2, r1 and r2 of double-normal distributions; row 3, parameter p (proportion ‘tall’)
of double-normal distributions; row 4, parameters l and r of log-normal distributions; row 5, parameters k and b of gamma
distributions; row 6, parameters c, b and a of Weibull distributions. Measurements were made during 2001 and 2002.

distribution was marginally worse than the double-
normal with a mean difference of 78 AIC units. The best
average improvement over the single-normal distri-
bution in the 4-cm swards was for the log-normal
distribution with a mean improvement of 52 AIC units.
Figure 4 shows examples of distributions from the 8-cm
swards where the double-normal, the log-normal, the
gamma and the Weibull distributions each had low AIC
statistics.
In the 8-cm swards, the parameters of the distribu-
tions varied over time and between paddocks (Figure 1,
Tables 1 and 4). The parameters of the log-normal
distribution appear to be the most stable although this is
likely to be mainly an effect of the log-scaling. The

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 8 G. T. Barthram et al.

Table 1 Parameter estimates for five theoretical distributions A B

fitted to sward height distributions, and coefficients of variation
(CV), from swards with 4-cm target heights at the Hartwood 50
site. 40
1 30
Paddock no. 20
10
Distribution 2 6 7 9 0

Single-normal
50
l 4Æ10 3Æ72 3Æ54 3Æ98
40
r 1Æ45 1Æ56 1Æ75 1Æ29 30
Double-normal 2 20
p 0Æ273 0Æ181 0Æ173 0Æ030 10
l1 5Æ49 5Æ65 5Æ97 7Æ71 0
r1 1Æ53 1Æ94 2Æ25 0Æ38
l2 3Æ58 3Æ29 3Æ03 3Æ84 50
40

Frequency
r2 1Æ00 1Æ07 1Æ07 1Æ10
Log-normal 3 30
20
l 1Æ35 1Æ23 1Æ15 1Æ33
10
r 0Æ363 0Æ408 0Æ466 0Æ330
0
Gamma
k 8Æ09 6Æ21 4Æ76 9Æ62 50
b 1Æ97 1Æ67 1Æ34 2Æ42 Frequency 40
Weibull 4 30
c 2Æ33 1Æ85 1Æ55 2Æ19 20
b 0Æ277 0Æ323 0Æ345 0Æ328 10
a 0Æ897 0Æ968 0Æ931 1Æ270 0

CV (%) 35Æ3 42Æ1 49Æ6 32Æ5 50

40
Frequency

Parameter p is the proportion that is ‘tall’.

30
5
20
shape parameters for the gamma and Weibull distribu- 10
tions (k and c respectively) for the most part followed 0
similar trends.
The AIC statistics were related to the distributions of 50
observed heights by comparing differences in AIC 40
Frequency

statistics in relation to the coefficient of variation. This 30

6 20
allowed a determination of whether the fits changed
with this measure of variability. When the coefficient of 10
0
variation was relatively high, the log-normal distri-
bution tended to have the lowest AIC statistics
50
(Table 2). Indeed, out of the twenty instances when 40
Frequency

the coefficient of variation was greater than 50%, the 30

log-normal distribution had the lowest AIC statistic on 7 20
eighteen occasions. The lower coefficients of variation 10
were mostly associated with the swards at the wetter 0
Hartwood site, with there being only one instance
where a sward from the Hartwood site had a coefficient 50
40
Frequency

greater than 55%. At both sites, elevated coefficients of

8 30
20
Figure 2 Frequency histograms of sward height measure- 10
ments made in an 8-cm sward (paddock 4 at Sourhope) during 0
2001 (column A) and 2002 (column B). Rows 1–8 show 0 10 20 30 40 0 10 20 30 40
histograms from measurements made monthly during March Height (cm) Height (cm)
to October of each year.

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 Frequency distributions of sward heights under grazing 9

(a) 60 overfitted the central portion of the distribution, as it

did at the Sourhope site. The log-normal and gamma
distributions showed least bias, but at the Sourhope site
Frequency
40 the gamma distribution tended to overfit both tails.
Overall, the log-normal distribution had a tighter
cluster of points around the 0% line. The data from
20
the 4-cm swards were consistent with these observa-
tions from the 8-cm swards.
0
Discussion
(b) 60
The distributions
Frequency

40 The parameters of the single-normal and double-

normal distributions, describing location and scale,
can be interpreted easily. However, a potential issue
20
in the fitting of these distributions to biological data is
that neither distribution can be constrained to fit to
0 positive data only. This issue is relevant to sward height
data where, although the data can never be negative, a
(c) 60 substantial proportion of recorded sward heights may
be relatively close to zero. In such cases, the mean
height may be low relative to the standard deviation,
Frequency

40 resulting in the left tail of the distribution containing

unrealistic negative values if a single-normal distri-
20 bution, or double-normal distribution, are fitted (e.g.
Figure 4). In an attempt to address this issue, the log-
normal, gamma and Weibull distributions can also be
0 considered. They are all appropriate for fitting to
frequency distributions showing positive skewness
(d) and, in addition, naturally fit to positive data, usually
60
requiring an additional parameter to achieve a fit to
negative data. The log-normal distribution is normally
Frequency

40 distributed on a log-transformed scale rather than the

20
0
0 10 20 30 40
Height (cm)
Figure 3 Frequency histograms of sward height measure-
ments made in 4-cm swards in (a) paddock 2, (b) paddock 6,
(c) paddock 7 and (d) paddock 9 at Hartwood.
variation were generally associated with the drier
midsummer months.
Figure 5 shows differences between the fitted density
functions and the observed frequency distribution of
the height measurements. In the 8-cm swards, the
single-normal and Weibull distributions overfitted the
left tail and had a bias that changed along the x-axis. At
the Hartwood site the double-normal distribution
measurement scale. As a consequence, it can only be
fitted to data exhibiting positive skewness and is not
appropriate where data show negative skewness or a
symmetrical distribution. When it is fitted to nearly
symmetrical data, the fit is poor as a result of this lack of
flexibility and this explains some of the poor fits in the
data of this paper, particularly in paddock no. 1 at the
Hartwood site in April 2002 (Table 2). However, when
data are positively skewed, this distribution is especially
suitable where the frequency histogram shows a long
right tail. The two-parameter gamma distribution as
used here is described in terms of shape and scale
(parameters k and b). When k is large (and b ¼ 1), the
gamma distribution closely approximates a normal
distribution, but with the advantage over the normal
distribution that it can be fitted to positively skewed
data and can be constrained to fit only to positive
observations. The three-parameter version of the
Weibull distribution, as fitted in this paper and by
Remington et al. (1992), is defined in terms of shape,
scale and location parameters (c, b and a). When c is

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 10 G. T. Barthram et al.

Table 2 Coefficients of variation (CV) of sward height data and Akaike information criteria (AIC) statistics for five theoretical
distributions fitted to sward height frequency distributions measured in paddocks with target sward heights of 8 cm.

Distribution

Date n CV (%) Single-normal Double-normal Log-normal Gamma Weibull

Hartwood paddock no. 1

May 2001 398 41Æ1 1887 1855 1861 1845 1850
June 2001 394 62Æ2 2220 2047 1969 2007 2002
July 2001 394 54Æ4 2183 2083 2051 2059 2075
August 2001 396 44Æ6 1929 1891 1882 1870 1880
September 2001 388 42Æ2 1884 1816 1812 1816 1824
October 2001 388 37Æ0 1918 1888 1886 1879 1892
March 2002 380 38Æ8 1778 1749 1754 1744 1745
April 2002 387 33Æ4 1989 1992 2062 2016 1989
May 2002 387 40Æ5 1970 1959 1968 1948 1943
June 2002 388 45Æ9 1762 1706 1707 1695 1714
July 2002 390 44Æ2 2007 1956 1961 1943 1969
August 2002 386 42Æ6 2125 2083 2083 2074 2082
September 2002 380 50Æ4 2126 2074 2062 2054 2059
October 2002 378 50Æ9 1949 1867 1856 1858 1866

Hartwood paddock no. 10

May 2001 358 43Æ3 2006 1964 1975 1957 1975
June 2001 353 54Æ8 1855 1734 1697 1718 1735
July 2001 356 48Æ2 1917 1838 1788 1806 1804
August 2001 346 45Æ4 2095 2035 2020 2022 2035
September 2001 336 50Æ1 1980 1882 1865 1873 1894
October 2001 340 50Æ4 1882 1797 1771 1780 1790
March 2002 340 45Æ4 1574 1518 1496 1500 1499
April 2002 336 35Æ4 1648 1632 1622 1618 1619
May 2002 334 38Æ0 1711 1695 1693 1682 1684
June 2002 344 41Æ4 1657 1613 1594 1595 1596
July 2002 338 40Æ6 1728 1712 1724 1708 1702
August 2002 330 36Æ0 1727 1711 1725 1710 1711
September 2002 321 41Æ0 1756 1740 1742 1728 1728
October 2002 296 43Æ2 1494 1479 1500 1474 1477

Sourhope paddock no. 4

May 2001 368† 36Æ6 2012 1991 1990 1980 1985
June 2001 62Æ7 2219 2041 2012 2045 2037
July 2001 60Æ0 2069 1909 1866 1899 1891
August 2001 44Æ9 2179 2120 2106 2106 2109
September 2001 54Æ3 2103 1991 1974 1983 1989
October 2001 43Æ8 1895 1816 1795 1807 1811
March 2002 40Æ4 1710 1686 1676 1666 1678
April 2002 33Æ2 1789 1753 1737 1740 1745
May 2002 39Æ6 1984 1942 1934 1934 1932
June 2002 63Æ1 2073 1873 1833 1875 1861
July 2002 67Æ4 2209 1908 1893 1957 1927
August 2002 57Æ0 2247 2104 2066 2095 2089
September 2002 53Æ2 2258 2112 2076 2107 2105
October 2002 48Æ6 2007 1941 1915 1918 1927

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 Frequency distributions of sward heights under grazing 11

Table 2 (Continued)

Sourhope paddock no. 11

May 2001 306† 38Æ5 1629 1546 1550 1559 1590
June 2001 60Æ7 1773 1606 1580 1613 1616
July 2001 60Æ5 1948 1747 1723 1770 1742
August 2001 67Æ4 1897 1705 1665 1704 1685
September 2001 51Æ4 1600 1513 1481 1496 1499
October 2001 38Æ2 1507 1457 1439 1447 1453
March 2002 42Æ4 1457 1402 1449 1413 1439
April 2002 33Æ5 1531 1520 1709 1587 1545
May 2002 47Æ5 1689 1613 1670 1631 1654
June 2002 56Æ3 1571 1437 1413 1439 1430
July 2002 57Æ6 1756 1626 1691 1648 1678
August 2002 44Æ1 1665 1581 1653 1611 1641
September 2002 42Æ0 1751 1662 1745 1698 1733
October 2002 43Æ6 1594 1562 1561 1551 1558

The number of height measurements is shown under n; †At Sourhope the same number of measurements was made on each date.
A CV (%) above 50% is shown in italics. The lowest AIC statistic on each occasion is shown in bold.

Table 3 Akaike information criteria

Distribution
(AIC) statistics for five theoretical distri- Paddock
butions fitted to sward height frequency no. n Single-normal Double-normal Log-normal Gamma Weibull
distributions measured in paddocks with
target sward heights of 4 cm at the 2 283 1016 995 996 990 999
Hartwood site. 6 305 1142 1095 1073 1079 1084
7 288 1144 1065 1046 1057 1056
9 305 1025 999 1003 999 1005

The number of height measurements is shown under n. The lowest AIC statistic for
each paddock is shown in bold.

approximately 3Æ5, the Weibull distribution approxi-
mates to a normal distribution, while lower values of c
are associated with positive skewness and higher values
with negative skewness. The b parameter is a scaling
factor, which is a measure of the variability in the data,
and the location parameter, a, indicates the minimum
theoretical value of the distribution. The three-param-
eter Weibull distribution is more appropriate than the
two-parameter version which, for the right-skewed
data, fitted a longer left tail than is justified by the data
because the location parameter (a) is effectively con-
strained to be zero. In cases where the minimum data
value is not close to zero, allowing the location
parameter to vary, allows for an improved fit in the
left tail of the distribution. The Weibull distribution has
the general advantage, over the normal distribution, in
that it can be fitted to data showing positive or negative
skewness and, when the location parameter is positive
or zero, does not produce unrealistic negative values.
Although the log-normal and gamma distributions
have the advantage that they naturally fit to positive
data, they, and the Weibull distribution, appear to be
very sensitive to data values very close to zero, with
changes to these data leading to quite large changes
in the parameter estimates (this, combined with
the effect of a symmetrical frequency distribution,
resulted in the poor fit of the log-normal distribution
in paddock no. 11 at the Sourhope site in April
2002). In this respect, parameter estimates for the
normal and double-normal distributions will be much
more stable. The double-normal distribution has the
additional advantage of being able to take account of
both positive and negative skewness and bimodality.
The above characteristics alone of the tested five
distributions do not suggest that any one distribution
should be an obvious choice for modelling sward
height frequency distributions.
Shiyomi et al. (1984) derived the gamma distribution
analytically for herbage mass by assuming that growth is
logistic, defoliation is constant per unit time and that the
herbage mass without grazing is normally distributed. By
analogy, this will also apply to sward height distributions.
There does not seem to have been an analytical deriva-
tion for sward height data of any of the other distributions

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 12 G. T. Barthram et al.

A B C D
50
40
1 Frequency 30
20
10
0

50
40
Frequency

2 30
20
10
0

50
40
Frequency

3 30
20
10
0

50
40
Frequency

4 30
20
10
0

50
40
Frequency

5 30
20
10
0
0 5 10 15 20 25 30 0 5 10 15 20 25 30 0 5 10 15 20 25 30 0 5 10 15 20 25 30
Sward height (cm) Sward height (cm) Sward height (cm) Sward height (cm)

Figure 4 Illustrations of the fit of single-normal (row 1), double-normal (row 2), log-normal (row 3), gamma (row 4) and Weibull
(row 5) distributions to sward height frequency distributions. The sward height frequency distribution shown in column A is from
paddock no. 11 at the Sourhope site in May 2002, in column B is from paddock no. 4 at the Sourhope site in June 2002, in column C is
from paddock no. 1 at the Hartwood site in June 2002 and in column D is from paddock no. 10 at the Hartwood site in July 2002.

that were tested in this paper. Although the double-
normal distribution does not appear to have been
supported empirically, it is qualitatively satisfying be-
cause it is consistent with the observed patchy sward
height distribution (Gibb and Ridout, 1988).
Sward heterogeneity
Like all grazed swards the swards studied showed
structural heterogeneity. The variance (Morris et al.,
1999; Hutchings and Gordon, 2001), the standard
deviation (Correll et al., 2003) and the coefficient of
variation (Hirata, 2000) have all been used as measures
of sward heterogeneity. These measures correspond to
spatial variability rather than spatial pattern (see Adler
et al., 2001).
On average, the variance (r2 of the single-normal
distribution) of the heights in the 4-cm swards was less
than that of the 8-cm swards (2Æ3 cm2 vs. 13Æ6 cm2) and
with no overlap in the values. However, the coefficients
of variation of the 4-cm swards largely overlapped those
of the 8-cm swards (Tables 1 and 2). Correll et al. (2003)
observed a similar overlap in cattle-grazed swards
maintained at 5 and 10 cm. This result indicates that
the magnitude of the variability changed with the mean
height. It is noteworthy that the 4-cm fertilized swards
were less variable than the 4-cm unfertilized swards
(Table 1), perhaps as a consequence of the fertilizer

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 Frequency distributions of sward heights under grazing
Table 4 Average estimated parameter values for five
theoretical distributions fitted to sward height distributions,
and coefficients of variation (CV), from swards with 8-cm
target heights at Hartwood and Sourhope.
Site
Distribution Sourhope Hartwood
Single-normal
l 7Æ94 7Æ48
r 3Æ89 3Æ29
Double-normal
p 0Æ21 0Æ36
l1 13Æ1 10Æ7
r1 5Æ23 3Æ66
l2 6Æ74 5Æ97
r2 2Æ24 1Æ97
Log-normal
l 1Æ95 1Æ90
r 0Æ47 0Æ45
Gamma
k 5Æ18 5Æ52
b 0Æ66 0Æ75
Weibull
c 1Æ83 2Æ02
b 0Æ145 0Æ149
a 1Æ46 1Æ24
CV (%) 49Æ6 44Æ4
Parameter p estimates the proportion that is ‘tall’.
swamping the effects of patchy nutrient returns from
dung and urine.
Intensively grazed sheep and cattle swards in the
sourveld of South Africa can have both a lower variance
and a lower proportion of tall material than less
intensely grazed swards (Morris et al., 1999). In tem-
perate regions, similar responses to intensity of cattle
grazing have also been observed in sown swards in the
UK (Gibb and Ridout, 1988) and in Arrhenatherion
grassland in continental Europe (Correll et al., 2003).
The results from swards grazed at different stocking
densities of sheep in this study are consistent with these
observations. Sward height in the cattle studies was
measured using a rising plate meter, which sampled a
much larger area (710–930 cm2) than the sward stick
(2 cm2) used in this study. Due to its smaller sampling
area, the sward stick is likely to be more sensitive to the
small-scale horizontal variability in sward height found
in swards grazed by sheep. The differences between
sward heights (or grazing intensities) are compatible
with the observation that grazing livestock are more
selective and patch grazing is more prevalent under low
than high grazing intensities (Edwards et al., 1996).
However, in rotationally grazed cattle swards, the more

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
13
intensely the pasture is grazed, the more heterogeneous
the spatial pattern of herbage biomass can become
(Shiyomi et al., 1983), with the structure changing from
homogeneous to a random spatial pattern (Shiyomi
et al., 1984).
The results also suggest that there can be detailed
differences between sites (the Hartwood site was less
variable and had a greater proportion of tall material
than the Sourhope site although the maximum heights
tended to be greatest at the Sourhope site). The reasons
for the site differences were not examined but may be
related to differences in species composition, seasonal
growth patterns or diet selection.
Best fit
The superior fit of the double-normal, rather than the
single-normal, distribution to sward height data ob-
served by Gibb and Ridout (1988) has been referred to
many times as evidence of the heterogeneous nature of
swards grazed by cattle. However, its fit does not appear
to have been tested in swards grazed by sheep. Neither
has the Weibull distribution which, like the double-
normal distribution, has only been compared with the
single-normal distribution (Remington et al., 1992). The
authors are not aware of any previous instance where
the fit of the five distributions to observations from a
single sward, grazed by cattle or sheep, has been
compared.
In the sheep-grazed swards, as in swards grazed by
cattle, the frequency distribution of sward heights was
most poorly described by the single-normal distri-
bution. The fit was particularly poor during the
summer months when coefficients of variation in
sward height were high. This corresponds to periods
when plant growth rates are high, and small differ-
ences in defoliation frequency can generate larger
differences in sward height than at other times of the
year; excluding the inadequate single-normal distribu-
tion, which had the best fit? Considering only the AIC
statistics, the log-normal distribution commonly had
the best fit in the 8-cm swards, but the magnitude of
the differences in AIC statistics between the distri-
butions, when compared with the difference between
them and the single-normal distribution, was small.
Thus, no distribution had clearly and consistently the
best fit to the data based on the AIC statistics. From
inspection of the pattern of deviances over the range
of heights (Figure 5), the Weibull distribution can be
rejected because of bias in its fit although it offers
considerable flexibility in determining the shape of the
distributions, at the expense of an extra parameter.
Overall the log-normal distribution showed bias least
often, closely followed by the gamma distribution and
then by the double-normal distribution. There was no
 14 G. T. Barthram et al.

8 cm-Hartwood 8 cm-Sourhope 4 cm-Hartwood

5

Difference (%)
1 0

–5

5
Difference (%)

2 0

–5

5
Difference (%)

3 0

–5

5
Difference (%)

4
0

–5

5
Difference (%)

5 0

–5
0 20 40 60 80 100 0 20 40 60 80 100 0 20 40 60 80 100

Midpoint of theoretical interval Midpoint of theoretical interval Midpoint of theoretical interval

(%) (%) (%)

Figure 5 Differences between the percentage of the observed data and the percentage of the distributions that fell in
rounding intervals of the height data. The x-axis shows the midpoint (%) of the rounding intervals calculated for the relevant
distributions, and the y-axis shows the difference between the percentage of the observations and the percentage of the fitted
distributions lying in each interval. Points that lie on the horizontal 0% line show perfect correspondence between fitted and
observed distributions. A point below the line shows that there were fewer observed heights than in the fitted density function
(overfit) and a point above the line shows that there were more heights than the fitted density function (underfit). The
left-hand column shows combined data from the 8-cm swards at the Hartwood site, the central column from the 8-cm swards
at the Sourhope site and the right-hand column from the 4-cm swards at the Hartwood site. Rows one to five show
single-normal, double-normal, log-normal, gamma and Weibull distributions respectively.

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
 Frequency distributions of sward heights under grazing
strong evidence to suggest that any one of these
models was most appropriate for describing sward
height distributions at different times during the
season.
Grazing models
Simple models of the effects of a partial rejection of ‘tall’
patches by grazers (a relative preference for ‘short’
patches) have shown that, where there is a high
probability of ‘tall’ patches being eaten, a skewed but
unimodal distribution of patches develops. However,
where there is a low probability of ‘tall’ patches being
eaten, a bimodal distribution emerges (Parsons and
Dumont, 2003). Bimodal distributions have been
reported, mainly for cattle (Gibb and Ridout, 1986,
1988; Fisher et al., 1995; Gibb et al., 1999; Morris et al.,
1999) but without comparing with the fit of skewed
unimodal distributions. The swards in this study were
not unequivocally best described by either a unimodal
or bimodal distribution. They did contain visually
distinct ‘short’ and ‘tall’ patches that persisted for
many months (G. T. Barthram and C. A. Marriott,
unpubl. data), but this is not evidence for or against a
unimodal distribution as the distribution type is
unrelated to the spatial aggregation of the sward
height measurements. It is concluded that the data do
not provide unequivocal evidence for either bimodal
or unimodal distributions in these swards and so do
not imply support for either of the above modes of
grazing behaviour.
Conclusion
The single-normal distribution is inadequate for mod-
elling the sward height distribution of swards grazed by
sheep, and the Weibull distribution shows unacceptable
bias. The log-normal distribution shows bias least often,
closely followed by the gamma distribution and then by
the double-normal distribution. The log-normal and
gamma distributions have the advantage over the
double-normal that they cannot produce unrealistic
modelled values (negative sward heights). However,
the log-normal distribution has the disadvantage that it
does not fit well to near symmetrical frequency distri-
butions. The double-normal distribution has the
advantage that parameter estimates are likely to be
more stable than for these other two distributions when
there are data values that are close to zero.
Acknowledgments
Ian Sanders provided access to some references. This
work was funded by the Scottish Executive Environ-
ment and Rural Affairs Department.

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
15
References
Adler P.B., Raff D.A. and Lauenroth W.K. (2001) The
effect of grazing on the spatial heterogeneity of
vegetation. Oecologia, 128, 465–479.
Akaike H. (1974) A new look at statistical model
identification. IEEE Transactions of Automatic Control,
AU-19, 716–722.
Armstrong R.H., Grant S.A., Common T.G. and Beattie
M.M. (1997) Controlled grazing studies on Nardus
grassland: effects of between tussock sward height and
species of grazer on diet selection and intake. Grass and
Forage Science, 52, 219–231.
Barthram G.T. (1985) Experimental techniques: the HFRO
sward stick. HFRO Biennial Report 1984–85, pp. 29–30.
Barthram G.T., Marriott C.A., Common T.G. and Bolton
G.R. (2002) The long-term effects on upland sheep
production in the UK of a change to extensive
management. Grass and Forage Science, 57, 124–136.
Bircham J.S. (1981) Herbage growth and utilisation under
continuous stocking management. Ph.D. thesis, University of
Edinburgh, Edinburgh.
Correll O., Isselstein J. and Pavlu V. (2003) Studying
spatial and temporal dynamics of sward structure at low
stocking densities: the use of an extended rising-plate-
meter method. Grass and Forage Science, 58, 450–454.
Devries M.F.W. and Daleboudt C. (1994) Foraging strategy
of cattle in patchy grassland. Oecologia, 100, 98–106.
Edwards G.R., Parsons A.J., Newman J.A. and Wright I.A.
(1996) The spatial pattern of vegetation in cut and grazed
grass white clover pastures. Grass and Forage Science, 51,
219–231.
Fisher G.E.J., Roberts D.J. and Dowdeswell A.M. (1995)
The manipulation of grass swards for summer-calving
dairy cows. Grass and Forage Science, 50, 424–438.
Gibb M.J. and Ridout M.S. (1986) The fitting of frequency
distributions to height measurements on grazed swards.
Grass and Forage Science, 41, 247–249.
Gibb M.J. and Ridout M.S. (1988) Application of double
normal frequency distributions fitted to measurements of
sward height. Grass and Forage Science, 43, 131–136.
Gibb M.J., Huckle C.A., Nuthall R. and Rook A.J. (1999)
The effect of physiological state (lactating or dry) and
sward surface height on grazing behaviour and intake by
dairy cows. Applied Animal Behaviour Science, 63, 269–287.
Ginzburg L.R. and Jensen X.J. (2004) Rules of thumb for
judging ecological theories. Trends in Ecology and
Evolution, 19, 121–126.
Grant S.A., Suckling D.E., Smith H.K., Torvell L., Forbes
T.D.A. and Hodgson J. (1985) Comparative studies of diet
selection by sheep and cattle – the hill grasslands. Journal
of Ecology, 73, 987–1004.
Hirata M. (2000) Quantifying spatial heterogeneity in
herbage mass and consumption in pastures. Journal of
Range Management, 53, 315–321.
Hodgson J., Forbes T.D.A., Armstrong R.H., Beattie M.M.
and Hunter E.A. (1991) Comparative studies of the
ingestive behavior and herbage intake of sheep and cattle
grazing indigenous hill plant-communities. Journal of
Applied Ecology, 28, 205–227.
 16 G. T. Barthram et al.
Hutchings N.J. and Gordon I.J. (2001) A dynamic model of
herbivore–plant interactions on grasslands. Ecological
Modelling, 136, 209–222.
Laca E.A., Distel R.A., Griggs T.C. and Demment M.W.
(1994) Effects of canopy structure on patch depression by
grazers. Ecology, 75, 706–716.
Marriott C.A., Bolton G.R., Barthram G.T., Fisher J.M.
and Hood K. (2002) Early changes in species composition
of upland sown grassland under extensive grazing
management. Applied Vegetation Science, 5, 87–98.
Morris C.D., Derry J.F. and Hardy M.B. (1999) Effect of
cattle and sheep grazing on the structure of Highland
Sourveld swards in South Africa. Tropical Grasslands, 33,
111–121.
Parsons A.J. and Dumont B. (2003) Spatial heterogeneity
and grazing processes. Animal Research, 52, 161–179.
Payne R.W. (2002) The Guide to Genstat 6.1, Part 2: Statistics.
Oxford: VSN International Limited.
Remington K.K., Bonham C.D. and Reich R.M. (1992)
Blue Grama-Buffalograss responses to grazing – a
Weibull distribution. Journal of Range Management, 45,
272–276.

2005 Blackwell Publishing Ltd, Grass and Forage Science, 60, 4–16
Sanders I., Topp C.F.E., Swain D., Wilson L. and
McRoberts N. (2003) Modelling seasonal changes in
spatial heterogeneity of sward height in grass-clover and
grass only swards. In: Seventh BGS Research Conference,
Aberystwyth, 1–3 September 2003, pp. 41–42. Reading, UK:
British Grassland Society.
Shiyomi M., Akiyama T. and Takahashi S. (1983) A spatial
pattern model of plant biomass in grazing pasture I.
Journal of Japanese Society of Grassland Science, 28,
373–382.
Shiyomi M., Akiyama T. and Takahashi S. (1984) A spatial
pattern model of plant biomass in grazing pasture II.
Journal of Japanese Society of Grassland Science, 30, 40–48.
Shiyomi M., Okada M., Takahashi S. and Tang Y.H. (1998)
Spatial pattern changes in aboveground plant biomass in
a grazing pasture. Ecological Research, 13, 313–322.
Tsutsumi M., Shiyomi M., Sato S. and Sugawara K. (2002)
Use of gamma distribution in aboveground biomass of
plant species in grazing pasture. Grassland Science, 47,
615–618.