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Potential density and tree survival: an analysis based

on South African spacing studies
ab

Klaus von Gadow , Heyns Kotze , Thomas Seifert , Kai Staupendahl & Juan G lvarez
Gonzlez

Department of Forest and Wood Science, University of Stellenbosch, South Africa

University of Gttingen, Gttingen, Germany

Mondi, Hilton, South Africa

Argus Forstplanung, Worpswede, Germany

University of Santiago de Compostela, Santiago de Compostela, Spain

Published online: 03 Dec 2014.

To cite this article: Klaus von Gadow, Heyns Kotze, Thomas Seifert, Kai Staupendahl & Juan G lvarez Gonzlez (2014):
Potential density and tree survival: an analysis based on South African spacing studies, Southern Forests: a Journal of
Forest Science, DOI: 10.2989/20702620.2014.984151
To link to this article: http://dx.doi.org/10.2989/20702620.2014.984151

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Southern Forests 2014: 18

Printed in South Africa All rights reserved

SOUTHERN FORESTS

This is the final version of the article that is

published ahead of the print and online issue

ISSN 2070-2620 EISSN 2070-2639

http://dx.doi.org/10.2989/20702620.2014.984151

Potential density and tree survival: an analysis based on South African

spacing studies
Klaus von Gadow1,2, Heyns Kotze3, Thomas Seifert1, Kai Staupendahl4 and Juan G lvarez Gonzlez5
Department of Forest and Wood Science, University of Stellenbosch, South Africa
University of Gttingen, Gttingen, Germany
3 Mondi, Hilton, South Africa
4 Argus Forstplanung, Worpswede, Germany
5 University of Santiago de Compostela, Santiago de Compostela, Spain
* Corresponding author, e-mail: heyns.kotze@mondigroup.co.za

Downloaded by [Klaus v. Gadow] at 00:31 08 December 2014

Because of the high cost of maintaining a series of unthinned, densely stocked stands over long periods of time,
suitable data about potential forest density and tree survival for different planting espacements are difficult to
find. Direct assessment of the potential density (which is always preferable to speculation, however ingenious the
estimation may be) requires densely stocked unmanaged field studies that are remeasured regularly during long
observation periods. An example of such a study is the Correlated Curve Trend (CCT) series of spacing studies
established by OConnor in South Africa. This contribution presents results for unthinned Pinus patula stands
using an unusually large data set, specifically: (1) the potential density using the relationship between the quadratic
mean diameter and trees per hectare, which does not confirm Reinekes constant of 1.605; (2) the relationship
between average spacing and average tree diameter (known as Nilsons sparsity), which is found to be non-linear,
thus contradicting previous assumptions; and (3) the development of the ratio basal area/trees per hectare, which
appears to remain unchanged during the life of a planted forest, irrespective of the planting espacement. Finally,
we present a tree survival analysis, based on the Weibull distribution function, for the Nelshoogte replicated CCT
study, which has been observed for almost 40 years after planting and provides information about tree survival in
response to planting espacements ranging from 494 to 2 965 trees per hectare.
Keywords: CCT, Pinus patula, South Africa, stand density, survival

Introduction
The aim of the early field experiments, established during
the late-nineteenth and early-twentieth centuries, was to
estimate timber yields at different stages of forest development and thus create a scientific basis for the sustainable
management of timber resources. Resource degradation
and depletion can be prevented if it is possible to estimate
the effects of future harvest levels. Some of these experiments have been remeasured for over a century. They
provide information, not only about timber volumes, but
also about a variety of phenomena that were previously
considered unimportant, but later turned out to be crucial
for understanding the long-term dynamics of a forest
ecosystem. One of these phenomena, which has been of
considerable interest to scientists, is the potential density,
which has also been described as the maximum living
biomass or the potential density of woody tissue (using the
terminology of Luyssaert et al. 2008) of a community of
trees in response to a variety of site conditions. Among the
numerous studies that evaluate density-dependent mortality
or self-thinning for a given forest type and set of environmental conditions are those of Reineke (1933), Yoda et al.
(1963), Zeide (1987) and Hynynen (1993).
Considering the high cost of maintaining a series of
unthinned, densely stocked stands over long periods of

time, suitable data about tree survival in response to density

and age are very hard to find. The majority of growth trials
in the Northern Hemisphere were designed to provide
data that were supposed to be useful for practical forestry.
Such studies usually lack unrealistic variants of extremely
low or high densities. It is also difficult to obtain data from
forests of low management intensity, because some
trees are usually harvested ahead of natural mortality. To
overcome the lack of direct observations, various indirect
methods have to be used to estimate the potential density.
Examples are the attempts by Sterba (1975), Clutter and
Jones (1980) and Gadow and Hui (1993). Obviously, direct
observation is always preferable to estimation, however
ingenious the speculation may be. Direct assessment of the
potential density, and developments towards reaching it,
requires densely stocked unmanaged field studies that are
remeasured regularly during long observation periods.
Two rare examples of such an expensive experiment
are the spruce growth series established in Denmark
(Skovsgaard 1997: p. 97 et sqq.) and the Correlated Curve
Trend (CCT) series of spacing studies established by
OConnor (1935) in South Africa. The CCT study is particularly interesting because it includes espacements ranging
from free growth to extremely dense. There are 27 CCT

Southern Forests is co-published by NISC (Pty) Ltd and Taylor & Francis

Downloaded by [Klaus v. Gadow] at 00:31 08 December 2014

experiments, typically they consist of 18 plots, covering

0.081 ha (0.2 acres) each. Detailed descriptions of the
design and analysis of the CCT studies were published
by Craib (1939), Marsh (1957), OConnor (1960), Burgers
(1976), Van Laar (1982), Gadow (1987), Bredenkamp
(1984) and Bredenkamp et al. (2000). Most of the 27
experiments were established between 1936 and 1938.
Based on previous studies of long-term experiments
in unmanaged forests growing at very high densities, we
conclude that the problem of estimating potential density is
not as trivial as it may appear when reading some of the
relevant literature. It has been shown that different growing
sites can produce quite different potential densities for
plantation-grown Pinus patula (Gadow 1987). However,
in many early studies, only stand averages were used.
Accordingly, the objective of this contribution is to estimate
the potential density of unthinned Pinus patula stands using
a more detailed and comprehensive data set. We also apply
new quantitative approaches, including Nilsons sparsity
(Nilson 2006; see also Hilmi 1957), which represents a
simpler and more logical relation because the variables
have the same dimension (average distance vs mean
diameter at breast height [dbh]) as the Reineke model
(number of trees vs mean dbh). In addition, we estimate
individual tree survival functions.
Materials and methods
This section introduces the spacing trials, the data set and
the methods used in our study.
Description of field trials
The South African CCT experiment is a classic spacing
trial designed to predict yields from plantations of various
species of pine and eucalypts for a wide range of densities,
varying between extremely dense (2 965 stems ha 1)
and free growing (124 stems ha1). The CCT experiment
typically consists of 18 plots, covering 0.081 ha each.
Nine of the 18 plots were left unthinned, the other nine
were subjected to various thinning regimes. The unthinned
experiment, known as the basic series, provides information about the growth of unthinned stands for a wide of
range of planting densities. In the other, known as the
thinned series, the response to various thinning regimes
may be assessed. Eight nominal stand densities, ranging
from 124 to 2 965 stems ha1, were established in plots 1 to
8 of the basic series. In order to avoid suppression by grass
and weeds, all plots were initially planted at 2 965 stems
ha 1 and then thinned in advance of competition until
the nominal stocking was achieved. In this study only the
results from the basic series are used. The treatment details
for plots 18 are presented in Table 1.
Data from the CCT experiments are suitable for analysing
tree survival in response to stand density and tree age,
especially since recently additional climate and soil data
have become available, allowing a more detailed analysis of
environmental effects.
In the late 1980s Bredenkamp (1990) developed and
implemented an alternative spacing experiment, known
as the Triple-S CCT trial. The Triple-S experiment is
essentially an unthinned spacing trial with six treatments.

Gadow, Kotze, Seifert, Staupendahl and lvarez Gonzlez

The planting densities range from 245 to 2 981 stems ha1.

The Triple-S treatments are also shown in Table 1.
The data set
Relevant details of seven South African spacing trials for
Pinus patula are presented in Table 2. The altitudes range
from 840 m above sea level (Wilgeboom) to 1 650 m
(Jessievale), the mean annual temperatures from 15 C
(Jessievale) to 20 C (Entabeni), and the mean annual
precipitation from 943 mm (Weza) to 1 300 mm (Entabeni
and MacMac).
Soil depths and geology were not assessed in the
Weza experiment. With a depth of 6090 cm the soils are
moderately deep in the Jessievale and Wilgeboom experiments, and deep in the other experiments. Humic soils are
encountered in the MacMac, Nelshoogte and Tweefontein
experiments and red apedal dystrophic soils in the Entabeni
and Wilgeboom experiments, whereas the Jessievale
experiment is situated on non-red neocutanic soils on
granite. The depth-limiting material is saprolite, except
in the case of the Jessievale experiment where there is a
periodic wetness hazard.
Methods
Natural stem number decline
N
We define the rate of survival as the ratio 2 while mortality

N1
N1  N2
may be expressed by
. The natural decline of stem
N1

numbers per unit area is defined as the natural elimination of trees as affected by age and planting density. This
process may be described in different ways. One candidate
model is the following linear equation:
N2 a0 a1Age1 a2Age2 a3N1

(1)

where N1 is the observed number of trees per hectare at

the initial age Age1, N2 is the estimated number of trees per
hectare at the projection age Age2, and a0...a3 are empirical
parameters.
Reinekes limiting relationship
Populations of trees growing at high densities are subject
to density-dependent mortality or self-thinning. For a given
average tree size there is a limit to the number of trees
Table 1: Treatment details for the eight plots of the CCT basic
series and the six plots of the Triple-S experiment
CCT unthinned series
Nominal
Plot
density
(trees ha1)
1
2 965
2
1 483
3
988
4
741
5
494
6
371
7
247
8
124

Triple-S experiment

Plot Measured
Nominal
trees
size
density
(ha) (trees plot1) (trees ha1)
0.081
240
245
0.081
120
403
0.081
80
665
0.081
60
1 097
0.081
40
1 808
0.081
30
2 981
0.081
20
0.081
10

Plot size
(ha)
0.1022
0.06197
0.03759
0.02280
0.01383
0.00839

Measured
trees
(trees plot1)
25
25
25
25
25
25

Southern Forests 2014: 18

per hectare that may co-exist in an even-aged stand. The

relationship between the average tree size (increasing over
time) and the number of live trees per unit area (declining
over time) may be described by means of a limiting line.
A convenient model for estimating this limiting relationship
is the following:
Nmax a0Da1

(2)

Downloaded by [Klaus v. Gadow] at 00:31 08 December 2014

with N max maximum number of surviving trees per

hectare,, D quadratic mean diameter (cm), and a0 and a1
are empirical parameters. The parameters of Equation (2)
can be estimated from fully stocked, unthinned trials.
Reineke (1933) plotted the number of trees per unit area
of fully stocked stands over their average diameter and
considered the a1 parameter to be a constant equal to
1.605 for all species and locations (Oliver and Larson
1996: 353354; Zeide 2004).
Nilsons sparsity
In the case of a regular spatial distribution of the trees
within a forest, the average distance between the trees may
be estimated by the square root of the number of square
metres in a hectare (10 000) divided by the number of trees
per hectare. Nilson (2006) named this average distance
between the trees stand sparsity (L):
L

100
N

irrespective of the initial stand density. Thus, if G2/N2

G1/N1 for any interval from initial age 1 to final age 2, which
is equivalent to log(G2) log(N2) log(G1) log(N1), this
would imply that the death of trees and the associated lower
density is compensated by basal area growth.
Tree survival
The probability of survival, is a central term of survival
analysis 1 , which investigates the distribution of the
non-negative random variable T, which in our study
describes the forest age (Staupendahl 2011; Staupendahl
and Zucchini 2011). According to Klein and Moeschberger
(1997: 21), the pattern of T can usually be characterised by
four functions. The probability density function f(t) describes
the frequency distribution of the points in time, in which
trees die. In the case of continuously measured time, it is
defined by:

f(t )

lim

't o0

P(t d T  t  't )
, with t t 0
't

For small t, f(t)t may be thought of as the approximate

unconditional probability that tree death will occur at time t.
The cumulative distribution function F(t), as the integral of
the density function, gives the probability that a death has
occurred by time t:

F(t ) P(T d t )

L a bD

(4)

where L is the stand sparsity (m), D is the quadratic

mean diameter (cm) of the trees in a stand, and a and b
are empirical parameters. The advantage of this model,
when compared to the Yoda approach, is the fact that the
two variables have the same dimension. Furthermore, the
relationship is assumed to be linear and, if that is the case,
then Equation (4) would be useful for comparing speciesspecific relations in multi-species forests.
Change of G/N ratios
We define G as the basal area (m2 ha1) and N as number
of trees per hectare and assume that the ratio (G/N)
remains constant in unthinned Pinus patula stands,

f( x )dx

(6)

(3)

where N is the number of trees per hectare. Nilson (2006)

proposed to estimate the potential density using the
following relationship:

(5)

The survival function S(t) is the complement of F(t). Thus, it

gives the probability that a tree survives at least until time t:
S(t) P(T t) 1 F(t)

(7)

The probability that the event occurs at a certain time,

conditional on the tree having survived until that time,
is also important. The density of this conditional death
probability is called the risk or hazard function (hazard rate)
h(t), and is defined as:
h(t )

lim

't o0

P(t d T  t  't | T t t )
't

f(t )
S(t )

(8)

For small t, h(t)t gives an approximation of the conditional

probability of tree death. In the case of discrete time, which
is normally considered in forest research, the unconditional
probability that a planted tree dies at age i is given by:
f(ti) P(T ti) S(ti1) S(ti)

(9)

Table 2: Relevant details of the seven spacing trials for Pinus patula. masl Metres above sea level, MAT mean annual temperature,
MAP mean annual precipitation, SI mean height of dominant trees at age 20, N/A not available

Location
Weza
Entabeni
MacMac
Nelshoogte
Tweefontein
Jessievale
Wilgeboom

Trial type
CCT
CCT
CCT
CCT
Triple-S
Triple-S
Triple-S

Altitude
(masl)
1 250
850
1 300
1 400
1 200
1 650
840

MAT
(C)
15.8
20.0
17.0
17.0
17.0
15.0
18.5

MAP
(mm)
943
1 300
1 300
1 175
1 175
950
1 175

Soil group
N/A
Red apedal dystrophic soils
Humic soil with yellow subsoil
Undifferentiated humic soils
Humic soil with yellow subsoil
Non-red neocutanic soils
Red apedal dystrophic soils

Soil depth
class (cm)
N/A
150
150
120150
150
6090
6090

Lithology

SI20 (m)

Shale
Basalt
Oaktree
Granite
Dolomite
Granite
Granite

20.6
23.7
24.3
23.9
22.3
23.4
24.8

Gadow, Kotze, Seifert, Staupendahl and lvarez Gonzlez

where t i denotes the assessment age i and S(t i) gives

the discrete survival function, with i 1, 2,, t0 0 and
S(t 0) 1. Accordingly, the discrete time hazard rate
(Fahrmeir 2007: 26) gives the conditional probability that a
tree that survived until age i dies at this age:
f(t i )
S(t i 1)

ti | T t ti )

(10)

D 1

f(t )

D t

E E

t D
exp 
E

(11)

t D
F(t ) 1  exp 
E

S(t )

h(t )

(12)

t D
exp 
E

D
E

(13)

N2 41.87 11.15Age1 11.76Age2 0.937N1

Maximum density
Traditional approaches
Reinekes limiting relationship was fitted to the four
individual CCT experiments MacMac, Weza, Entabeni
and Nelshoogte. For these experiments, which had been
remeasured over a long period of time, a limiting relationship could be established. The parameter estimates for
the entire data set are a0 788446.6 and a1 2.00. The
limiting relationship is thus defined by the equation:
Nmax 788 446.6D2

(14)

Natural decline

(15)

The four parameters are highly significant. The adjusted

coefficient of determination is 0.972.

D 1

t

E

with t 0. These functions are directly derivable from

one another. In this study, we will fit the survival function
(Equation 9) to observed surviving Pinus patula trees
n years after planting in the Nelshoogte spacing experiment. The parameters of the survival function will be
evaluated for five initial planting densities.

(16)

Not surprisingly, the value of a1 deviates considerably from

the constant 1.605 proposed by Reineke (1933). It could
be shown that the exponent may assume a wide range
of values, depending on tree species and site conditions
(Gadow 1986). A relationship between the coefficients and
the local climate and soil data (which are only available for
three of the four experiments) could not be established. The

Limiting relationship

Nilsons sparsity

3000

10

TREES PER HECTARE

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In other words, h(ti) indicates the proportion of trees in age

i 1 that die on average at age i. According to Pienaar and
Shiver (1981), Gadow (1987), Kouba (2002) and Dickel
et al. (2010), the Weibull distribution (Weibull 1951) was
chosen because it requires only two parameters while at
the same time being quite flexible. If T is Weibull distributed
with scale parameter and shape parameter , the density,
distribution, survival and hazard functions are given as
follows (Staupendahl 2011):

Natural decline of stem numbers

The decline of the number of trees per unit area is affected
by age and stand density. Figure 1 shows a dramatic
decline in the experiments planted at very high densities,
starting approximately 15 years after planting. In contrast,
the population in the very low density plots remains stable,
at least until 50 years. The following linear model, because
of its simplicity, was found to be appropriate for estimating
the natural decline of the number of trees in response
to an arbitrary initial age and stem number and a defined
projection age:

L (AVERAGE SPACING) (cm)

h(t i ) P(T

Results

2500

0.975 percentile

2000
1500
1000
500
0
0

10

20

30
40
AGE (y)

50

60 0

10

20

30 40
D (cm)

50

60

70

4
0.1 percentile

10

20

30 40 50
D (cm)

60

Figure 1: Left: Natural decline of stem number over age for different initial planting densities (N ha1), involving all available South
African Pinus patula spacing trials. Centre: Estimates of maximum density for all Pinus patula trials with the limiting line fitted using the
0.975 percentile values and the R library quantreg. Right: Estimates of maximum density for all Pinus patula trials with Nilsons sparsity using
the 0.10 percentile values and the R library quantreg

Southern Forests 2014: 18

the death of trees, not through competition, but through

reduced ability to survive pathogen attacks.
Shvidenko et al. (2006) describe the development of
the number of trees per hectare and the average tree
diameter for fully stocked Pinus sylvestris forests in the
forest tundra and northern taiga ecoregions in the European
part of Russia. These data show linearity up to the age
of 200 years. Looking at the Pinus patula CCT data, the
lower-bound relationships may be interpreted as linear (as
suggested by Nilson 2006), but only up to a mean diameter
of about 50 cm. This result is unexpected and contradicts
Nilsons original assumptions of complete linearity. For
this reason, we tried to express the lower bound using a
non-linear model, but further studies with different data sets
are required to arrive at satisfactory conclusions regarding
the limiting relationship over all tree sizes.

Triple-S studies Tweefontein, Wilgeboom and Jessievale

are still very young and not suitable for this part of the
analysis.
Nilsons sparsity
The parameters of Equation (4) can be estimated using
data from fully stocked, unthinned trials, such as the Pinus
patula CCT and Triple-S CCT trials.
The following relationship was estimated using the 0.025
percentile values and the R library quantreg:
L 0.273 0.101D

(17)

L ae(bD)

Change of G/N ratio

Based on the CCT data, we may assume that the ratio of
basal area and trees per hectare (G/N) remains virtually
constant in unthinned Pinus patula stands, irrespective of the
initial stand density. The following equation, involving a slope
parameter close to unity, was estimated for all the data:

(18)

The relationship appears to be linear up to the age when

the trees stop growing but mortality continues. After that,
the assumption of linearity is no longer valid. The trees start
dying, not as a result of competition-induced self-thinning,
but possibly because of other unknown factors. Increasing
age usually leads to failing defence mechanisms causing

G2
N2

MacMac a = 1.08; b = 0.039

1.112

G1
N1

(19)

Nelshoogte a = 1.17; b = 0.032

10

L (average spacing) (m)

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The relationship between average spacing (L) and average

tree size (D) was originally assumed to be linear, but a
closer look at the observations, as shown in Figure 2,
reveals a non-linear relation. We used the following model:

0.025 percentile

0.025 percentile

Weza a = 1.31; b = 0.028

Entabeni a = 1.44; b = 0.026

10

0.025 percentile

0.025 percentile

10

20

30

40

50

60

70

10

20

30

40

50

60

70

D (cm)
Figure 2: Estimates of maximum density for the four Pinus patula CCT studies using the R library quantreg for estimating Nilsons sparsity
with 0.025 percentile values

Gadow, Kotze, Seifert, Staupendahl and lvarez Gonzlez

Tree survival
The results of the parameter estimates of the survival
function are presented in Table 3. Each of the parameters
1 and 1 of Equation (13) are linear functions of the
number of planted trees per hectare. The estimated linear
function of the shape parameter is 5.38 0.001012N0
(R2 0.8327). For the scale parameter it is 49.36
0.005206N0 (R2 0.9921), where N0 refers to the initial
number of planted trees per hectare. 2 and 2 are the
highly significant parameter values estimated from the two
linear relationships.
This relationship is potentially useful in that it allows one
to estimate the lifetime distribution for a range of planting
densities. A graph of the survival functions, and the
corresponding mortality and hazard functions, are presented
in Figure 4 for different initial espacements.
Figure 4 shows that the probability of tree survival is
rather different for the different planting espacements. As
expected, the probability of tree survival is increasing with

0.35
0.30
0.25
G2/N2

Downloaded by [Klaus v. Gadow] at 00:31 08 December 2014

The residual standard error is 0.012 on 1 095 degrees of

freedom and the parameter estimate is highly significant.
The adjusted R2 is 0.979. Figure 3 shows the relationship
between two ratios G1/N1 is the ratio of basal area (m ha1)
to trees per hectare at an arbitrary initial age and G2/N2 is
that ratio at the projection age.
Interestingly, the change of the G/N ratio appears to be
independent of stand density. This is, of course, only valid
in unthinned stands. Selective thinnings are likely to modify
the ratio. Pinus patula is sometimes grown in unthinned
pulpwood plantations and Equation (19) may be usefully
employed in situations where the initial ratio G1/N1 is known.
An estimate of either G2, using a basal area growth model,
or N2 using a tree survival model, may produce the ratio and
thus the other unknown quantity. However, a closer look at
Figure 3 shows that there is a slight bias in the linear model
at higher ratios which may reveal the same effect as was
evident in Nilsons model.

0.20
0.15

decreasing number of planted trees per unit area. The

peaks of tree mortality occur earlier, and are more lasting,
in the higher planting densities. The differences are notable
for the entire range of ages.
Discussion
Empirical research provides the essential basis for
understanding and modelling the dynamics of managed
and unmanaged forest ecosystems (Vanclay 1995; lvarezGonzlez et al. 1999; Pretzsch 2001; Garca 2003). The
extensive South African spacing experiments, maintained
for unusually long periods of time, therefore provide very
important information for the scientific analysis of maximum
forest density and tree survival.
Reinekes limiting line is an attempt to simplify the
relationship between average tree size and number of trees
per unit area. Yodas 3/2 power line of ecology represents
a similar attempt to explain self-thinning with a very simple
model. Both models have been questioned by a number of
authors, including Zeide (1985, 1987) and Gadow (1986).
This study has also shown that generalisations should be
treated with caution and simplification is only possible for
very specific conditions.
Nilson (2006; see also Hilmi 1957) argues that the most
simple and logical relation is expected between variables
of the same dimension, which is not the case in the
Reineke model (Equation 2). For this reason, and because
of its simpler form, Equation (4) was thought to be preferable when compared with Equation (2). A general discussion on the assumption of linearity of Nilsons sparsity is
not possible, because the CCT material does not cover
the seedling stage and stands older than 50 years. The
tables published by Shvidenko et al. (2006) for Russian
Pinus sylvestris forests on site quality II show that Nilsons
sparsity is linear up to the age of about 180 years.
However, the available material shows that the assumption of linearity cannot be confirmed for the Pinus patula
CCT studies. Considering the culmination of diameter
increment of Russian P. sylvestris and the much fastergrowing P. patula in South Africa one might expect that a
deviation from linearity, should it signify a general, speciesindependent pattern, could be induced at much higher
ages in P. sylvestris. It is known that stem diameter scales
well with crown extension and thus light competition and
that these relations are very species specific, at least for
conifers, which might affect the relationship.
Table 3: Weibull parameter estimates of Equation (13) for five
planting densities in the Nelshoogte CCT experiment. The subscript
1 refers to the first estimate based on the field observations,
and subscript 2 refers to the estimate obtained with the linear
equations. MEF Model efficiency

0.10
0.05
0.00
0.00

0.05

0.10

0.15 0.20
G1/N1

0.25

0.30

Figure 3: Relationship between G1/N1, the ratio of basal area

(m ha1) to trees per hectare at an arbitrary initial age, and G2/N2,
which is that ratio at the projection age

Planted trees
per hectare
2 965
1 483
988
741
494

2.35
3.71
5.04
4.36
4.71

34.18
41.04
44.01
45.79
47.06

MEF

Bias

0.9646 0.0026
0.9382 0.0033
0.8574 0.0044
0.7098 0.0008
0.9115 0.0071

2.38
3.88
4.38
4.63
4.88

33.92
41.64
44.22
45.50
46.79

Southern Forests 2014: 18

2 965
1 483
998
741
494

MORTALITY

SURVIVAL

0.8
0.6
0.4

1.0

1.0

0.8

0.8

HAZARD

1.0

0.6
0.4

0.0

0.0

0.0
0

10

20

30

40

50

60

0.4
0.2

0.2

0.2

0.6

10

20

30

40

50

60

10

20

30

40

50

60

Downloaded by [Klaus v. Gadow] at 00:31 08 December 2014

AGE (y)
Figure 4: Estimated survival (left) and corresponding mortality density (centre) and hazard (right) functions for five different planting
espacements N0, expressed in terms of planted trees per hectare

Dickel et al. (2010) have shown that if there were N1 trees

at the initial time point, and given that the lifetime distribution of the trees is Weibull distributed, it follows that the
number of surviving trees at t time units later is binomially distributed with parameters N1 and = 1 F(t; , ),
where and can be estimated from the initial stems per
hectare planted, as outlined in the section on Results
Tree Survival2:

P(N2

N
n ) 1 S n (1  S )N1n
n

This may be explained as follows. Suppose we have an

initial population of six trees (a,b,c,d,e,f). Four of the six
initial trees survive at the end of a given period of time.
There are 15 possible combinations of four of the six initial
trees surviving: (a,b,c,d)(a,b,c,e)(a,b,c,f)(a,b,d,e)(a,b,d,f)
(a,b,e,f)(a,c,d,e) (a,c,d,f)(a,c,e,f)(a,d,e,f)(b,c,d,e)(b,c,d,f)
(b,c,e,f)(b,d,e,f)(c,d,e,f) which may be obtained by the
following expression:

N1

n

N1 !
n !(N1  n )!

6!
4!2!

65
2

15

There are 15 possible outcomes and the probability for an

individual tree, given that two trees die and four survive, is
equal to the product of the probabilities for each tree. If i is
the probability of survival of one tree, and four trees survive,
then 4 is the survival probability of a group of four trees.
The probability that two trees die is then (1 )(64).
Notes
Survival analysis is most often defined as a class of statistical methods for studying the occurrence and timing of
events most often death (e.g. Cox and Oakes 1984).
2 In case N is not the number of trees at the initial time
1
point, the left truncated Weibull distribution has to be used.
1

Acknowledgements We wish to express our gratitude to

all those who conceived and implemented the South African

spacing experiments. Special thanks are due to the institutions (in sequential order: South African Department of Forestry,
South African Forest Research Institute, Council for Scientific and
Industrial Research, South African Forestry Company Limited
and Komatiland Forests) and individuals who measured the trees
and made sure that these unique observations are available for
research.

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Received 30 March 2014, revised 1 July 2014, accepted 14 July 2014