General paper

THE USE AND IMPORTANCE OF HYBRID

INTENSIVE FORESTRY IN SOUTH AFRICA
N.P. Denison and J.E. Kietzka
(Mondi Forests - A Division of Mondi Paper Company Ltd)
(Paper presented at the IUFRO Conference on "Breeding Tropical Trees" held at Cali, Colombia)

SYNOPSIS
Interspecific hybrids are becoming increasingly important to the South African forest industry. Presently,
hybrids of the subtropical eucalypts are the most prominent. Many thousands of hectares of selected clonal
plantings have been established. The most common hybrid combinations are Eucalyptus grandis crossed
with either E. camaldulensis, E. urophylla or E. tereticornis.
These hybrids have made it possible to extend tree planting to areas traditionally considered off-site
for plantation forestry. On these marginal sites, the hybrids growth and survival outperformed the pure
species, and they are consistently more resistant to diseases, pests, cold, heat and drought. Through
developing hybrid combinations, land closer to processing sites can now be planted.
The wood and growth properties of the hybrids are normally intermediate between the parent species,
but superior growth to both parent species is common. Great care must be exercised in selectingthe parents
for the hybrid combinations, which must be based on performance of both parents from breeding trials.
To ensure uniform and consistent end products, wood properties should be considered as it is essential to
ensure that the density range of the hybrid is within acceptable limits. The basic wood density of the hybrid
is generally higher than that of pure E. grandis, and is influenced by age and site, which in turn will affect
the harvesting age.
The authors predict an increased operational use ofhybrids for the cold-tolerant eucalypts and for Pinus
species as vegetative propagation technology improves. Hybrids used commercially must be carefully
screened over a range of growing sites to ensure that the selected clones are well adapted for site and
product. Mondi is presently screening in excess of 1 000 hybrid clones of which only 1 to 2 % will be
commercialised.

INTRODUCTION therefore occured largely on more marginal sites.

The forestry and forest products industry is one ofthe
fastest-growing sectors ofthe SouthAfrican economy.
Major expansion programmes have occurred in the
industry since 1983, resulting in an explosion of
afforestation programmes and land purchases. To
meet future demand, it has been predicted that wood
and fibre production would need to double by the year
2005.
This is a dynamic industry which uses modern
technology and is based on a renewable resource
which is dependant on only 1,2 % of South Africa's
land area. Of a total land area of 119 000 000 ha about
1400 000 ha is presently planted to exotic plantation.
It is doubtful whether the planted area will ever
reach 2 000 000 ha. Generally, South Africa can be
described as a dry country with only a small area
suitable for plantation forestry. However, the indus-
try brings economic activity and rural prosperity to a
belt stretching from the southern Cape, up through
the Natal Midlands and Zululand to the eastern and
north-eastern Transvaal.
Although excellent growth potential exists in the
prime areas of good rainfall, climate and soils, this
type ofland is scarce and also in demand for agricul-
tural crops and livestock. Forestry expansions have
Suid-Afrikaanse Bosboutydskrif- nr. 165, Junie 1993
Further constraints that influence profitable for-
estry and the processing industry are the high land
prices for good land, long haulage distances and
associated transport costs, and a permit system which
restricts land availability for forestry.
These then are the constraints and challenges
facing forestry management responsible for meeting
future sustainable wood and fibre requirements de-
manded for profitable forestry. The challenges con-
fronting the forestry researcher, tree breeder, silvi-
culturist and forester are enormous. Hybrids will
play an increasingly important role on the marginal
sites needed for forestry.
HYBRID FORESTRY· TECHNICAL CONSID·
ERATIONS
The potential for hybrid forestry in South Africa has
been recognised for many years. Progress has how-
ever been slow, owing to the technical difficulties
related to either obtaining hybrid seed in sufficient
utilisable quantities for commercial forestry or prob-
lems in multiplying selected material through veg-
etative propagation. Van der Sijde and Roelofsen
(1986) reported the volume production of the hybrid
between Pinus elliottii andP. caribaea v. hondurensis
55
at age 13 yars to be 2,5 times that of pure P. elliottii,
the Pinus species commonly planted in the coastal
area of Zululand. Du Plooy (1984) reported the pulp
properties of the 13-year-old hybrid trees to be com-
parable with those of P. elliottii. The solution to
meaningful operational use of selected hybrids is
therefore dependant on developing vegetative
propagation systems to increase selected hybrid com-
binations for commercial forestry. Burdon (1988)
clearly distinguishes between clonal forestry (CF)
and vegetative multiplication (VM). In essence, CF
infers a finite number ofselected clones whereas VM
refers to multiplication of unidentified genotypes
starting with limited amounts of seed. These two
systems will be used in South Africa to effectively
exploit the benefits of hybridisation. Burdon (1988)
further states that to be worthwhile, CF an VM must
be biologically safe, technically feasible and economi-
cally advantageous.
Hybrids generally have properties midway be-
tween the parent species. To be of economic value,
they must exhibit performance superior to the pure
species. Martin (1988) states that hybrid vigour
(heterosis) is always correlated with the environ-
ment in which the hybrid is grown. The heterosis is
stronger in zones which are marginal for the pure
species. The environmentally dependent hybrid vig-
our is the basis for the success ofthe eucalypt hybrid
programmes ofUAIC in the Congo, Aracruz in Brazil
and more recently Mondi's and HL&H's hybrid pro-
grammes in South Africa. The same considerations
apply to the successful hybrid programmes of P.
caribaea and P. tecunumannii in Queensland, Aus-
tralia. South Africa has not yet taken advantage of
the potential of hybridisation programmes in pines.
The relationship of hybrid vigour and adaptation to
new environments will be of significance in develop-
ing economic and viable forests in marginal areas of
South Africa. Pure species, if not too distinct from a
genetic point of view, will be used for the hybrids.
Hybrid seed is obtained through controlled polli-
nation, using promising genotypes of the pure spe-
cies. The progeny produced in the crosses is screened
in trials covering the major site categories. From
these trials, high performers are selected and screened
further for operational CF programmes (Denison
and Kietzka, 1992). Where vegetative propagation of
the selected genotypes is not technically feasible,
usually as a result of maturation problems, the VM
procedure, if practicable, may be followed. Hybrid
combinations are selected on the basis of high family
performance and breeding value. Seedlings of the
desired hybrid cross portraying juvenile qualities
and rooting ability will then be used. Many copies of
each seedling will be produced for operational
plantings. The normal procedure is to use either the
hedging process, serial reproductions or tissue cul-
ture. Promising results for multiplication on a re-
search scale have also been obtained with somatic
embryogenesis for both P. patula and E. grandis in
South Africa (Herman, personal communication).
The VM route is being pursued by Mondi in South
56
Africa for the pines as well as the difficult to root cold-
tolerant eucalypts. Burdon (1988) notes that hedging
appears to be effective technically, although few
hopes are held for it halting maturation indefinitely.
Even with the most rigorous hedging regime, matura-
tion is expected to proceed gradually, but it might
remain tolerable until the clones are made obsolete
by further genetic advances in the breeding pro-
gramme.
According to Martin (1988), hybridisation is an
efficient method of eliminating a number of sub-
lethal genes. If the production population of Fl hy-
brids is managed in well-planned breeding pro-
grammes, many of our marginal sites will be made
profitable. This is certainly the case on marginal sites
in coastal Zululand, where sites oflow rainfall, poor
soils or a combination ofboth have been transformed
into productive units. This is particularly meaning-
ful when marginal sites close to the processing cen-
tres can be productively utilised. If, on the other
hand, breeding programmes are not well managed,
hybrid breakdown will occur in later generations,
when sub-lethal and recessive genes will once again
be expressed. Breeding populations must be well
managed to ensure new hybrid combinations with
genetic gains in successive generations.
To produce hybrid combinations, breeding pro-
grammes for the pure species involved must be main-
tained. For the subtropical eucalypts, the hybrids
involved are E. grandis x E. urophylla (GXU), E.
grandis x E. camaldulensis (GXC) and E. grandis x
E. tereticornis (GXT). For the more temperate areas
the hybrids produced are E. grandis x E. nitens
(GXN), and E. grandis x E. macarthurii (GXM). For
the production of theE. grandis hybrids, information
and materials are obtained from the E. grandis
breeding and clonal programmes. From more than
3 000 families/clones, 30 individuals have been iden-
tified for the current hybrid programmes. The E.
urophylla base consists of 356 imported families
from Indonesia covering most of the natural range of
the species. At present, the E. camaldulensis (125
families) and E. tereticornis (60 families) base origi-
nates from third generation material from the Zim-
babwean breeding programme, in addition to im-
ported material on a provenance/family level from
Australia. We recognise that the base for E.
camaldulensis and E. tereticornis needs to be broad-
ened. This is to a degree compensated for through
good cooperation and exchange of base material
between the companies. For the temperate hybrids,
material of E. macarthurii and E. nitens is obtained
from the ICFR (Institute for Commercial Forestry
Research) cooperative breeding programmes which
have 445 E. macarthurii and 420 E. nitens local and
imported families in their programme.
It is equally important to maintain and manage
base populations that remain relatively undisturbed
and intact for insurance against known risk ele-
ments which may include market shifts or biological
risks in the form of pathogens, pests, climatic changes
or edaphic changes (Burdon, 1988). Namkoong (1988)
South African Forestry Journal- No. 165 June 1993
points out that geneticists, instead of managing ge-
netic resources, focus their attention on culling tree
populations for the best performers, with scant
thought to the natural structure or development of
forests. Because of the need for fast profits, forests
are viewed as machines for mass-producing low value
goods, and genetics is viewed as a means ofimproving
the efficiency of the machinery. Namkoong states
that forests are highly complex biological systems
that must be understood and managed primarily at
higher levels of organisation than tissues or single
cells. It is for this reason that CAMCORE (Central
American and Mexican Coniferous Resources Coop-
erative), an international organisation, has as one of
its main objectives the preservation of genetic re-
sources of tree species or provenances threatened
with extinction. All member organisations are obliged
to estabish gene preservation compartments from
seed of all families supplied to the cooperative mem-
ber. These plantings are in addition to the individual
family and provenance trials used for the genetic
improvement of the economically valuable species.
Martin (1988) noted that through hybridisation, it
is possible to recover interesting genes or gene groups
from another species for incorporation into the first
order species. It may then be necessary to apply back-
crosses to improve the genotype. These hybrid ma-
nipulations hold promise in South Africa for the
widely and diverse climates, altitudes and soils on
which we practice forestry. In manipulating groups
of genes for improved productivity, it is essential to
take wood properties into account. The object must be
to match the pure species or hybrid to site and to end
product. So often in the past, trees are grown for their
high yield potential and adaptation to a site without
taking cognisance of the wood properties.
POTENTIAL BENEFITS OF HYBRID FOR-
ESTRY
Hybrid forestry is in the process of revolutionising
forestry in South Africa. Future progress is however
largely dependant on successful vegetative propaga-
tion in the form of either clonal forestry (CF) or
vegetative multiplication (VM). Both systems will be
utilised for developing operational hybrid forestry.
One must remember that hybrids only have a role to
play if the benefits of creating and developing hy-
brids outweigh the additional costs and difficulties
encountered.
The main benefit of hybrid forestry will include:
1. Hybrid vigour (heterosis)
There are many references which refer to heterosis in
forest trees. (Campinhos et al., 1980; Zobel and
Talbert, 1984; Brandao et al., 1984; Van der Sijde and
Roelofsen, 1986; Martin, 1988; Van Wyk et aI., 1988).
This can be clearly demonstrated on many sites in
South Africa, particularly for several Eucalyptus
grandis hybrids established in marginal areas, con-
sidered off-site for the pure species (Figures 1 to 4).
Suid-Afrikaanse Bosboutydskrif- nr. 165, Junie 1993
The exploitation ofhybrid vigour for use on these site
is increasing, especially as better techniques are
developed for propagating the hybrid combinations
vegetatively. The authors believe that hybrid pro-
grammes for the pines and cold-tolerant eucalypts
will be equally dynamic as vegetative propagation
methods improve. Provided careful screening and
care is taken in selecting hybrid clones, meaningful
yield gains, often in excess of60 % are obtained when
comparing clones to orchard seed of E. grandis.
There are many instances where the pure species has
failed or remains stunted, while the hybrid is fully
adapted, healthy and vigorous on the same site.
There are also examples where the pure species, well
adapted to a prime site, always out-performs the best
hybrid combinations. Good examples can be found for
E. grandis on high potential forestry sites in the
Eastern Transvaallowveld. Zobel and Talbert (1984)
warn that one must be careful in the assessment of
hybrid vigour, because time and location are most
important. Many of the E. grandis hybrids express
heterosis early in life, but do not maintain the good
growth through to rotation (sprinters). Decisions on
the acceptance ofhybrid combinations for operation-
al programmes should not be made until at least half-
rotation age for the eucalypt hybrids.
2. Disease resistance
With the onset of clonal propagation of Eucalyptus
species in South Africa, concern has been expressed
that diseases might become more threatening. It is
certainly true that risks are greater in clonal
plantings. Careful attention must therefore be given
to utilising large numbers of clones of known
backgroud and susceptibility to disease. To achieve
the latter goal, risk assessment programmes have
been initiated in cooperation with the TPC (Tree
Pathology Cooperative) centered at the University of
the Orange Free State, and large scale selection and
screening procedures introduced.
As part of the overall programme, hybrids be-
tween various species ofeucalypts are being utilised.
Once again the question has been raised as to the
relative risk ofthese hybrids to disease. The potential
does exist that several eucalypt hybrids will be less
susceptible, and evidence at this stage of our pro-
grammes points in that direction. Information from
other parts ofthe world such as Brazil and the Congo
has suggested that hybrids of E. grandis and E.
urophylla are particularly resistant to cryphonectria
canker caused by Cryphonectria cubensis. This is
most encouraging given that this disease poses one of
the greatest threats to the local forestry industry.
However, preliminary screening programmes in
South Africa have shown that certain E. grandis x
E.urophylla hybrids are as susceptible as pure E.
grandis clones. The observation is perhaps not sur-
prising given the inherent susceptibility ofE. grandis.
One of the most serious problems associated with
the hybridisation programme is that very little is
known about the relative susceptibility of different
57
 A IC50 NYALAZI 38 MONTHS .. lel1 NYALAZI 38 MONTHS
PERFOTMANCE OF E.GRANDI8 COMPARED TO HYBRID CLDNE8 PERFOTIIANCE OF LGRANDIS COMPARED TO HYBRID CLONES

MAl (M3\HA\YR)
40 r - - - - - - - - - - - - - - - - - - - ,

315
15 GRANDIS

10

25
GRAND IS

.*....
SEEDLING
I GRANDIS
CL()NE.. ....
so

21
!
CLONE GRANDIS
SEEDUNO

~11i.
20
20
111
15
10
1IIIIIIIIt
10
FIGURE 2.
FIGURE 1.

.. IC38 DAHL 31 MONTHS

A CT07B SHIRE 80 MONTHS
PERFOTIIANCE OF E.GRANDIS COMPARED TO HYBRID CLDNES
PERFOTIIANCI OF I.GRANDI8 COMPARID TO HY8RID CLONE8

MAl (M3\HAIYR)
MAl (M3\HA\YR)
21 r - - - - - - - - - - - - - - - - - - - - - , 50 r - - - - - - - - - - - - - - - - - - - - ,

20 40
GRANDIS GRANDI GRANDIS
CLONE SEEDLIN CLONE
11 30

"'1
GRANDIS
SEEDLING
10 20

I 10

o o
FIGURE 3. FIGURE 4.

species being used in hybridisation. Trials have now
been established in which these different species are
being artificially inoculated with the most important
pathogens affecting eucalypts ofinterest in the local
industry. This information is already proving useful
in choosing species used in hybridisation (Wingfield,
personal communication).
Hybrids between E. nitens and E. grandis show
tremendous promise in various parts ofSouth Africa.
E. nitens is however highly susceptible to an impor-
tant leaf pathogen, Mycosphaerella molleriana In
contrast, E. grandis is relatively resistant to the
disease. Hybrids between these species range from
highly resistant to highly susceptible to the disease.
Careful selections are therefore underway to select
hybrids resistant to this disease. The same situation
is true of E. camaldulensis which is highly suscepti-
ble to Botryosphaeria die-back. Certain hybrids be-
tween this species andE. grandis are also susceptible
to this disease and material to be used for large-scale
planting must be carefully selected.
58
From the pathology standpoint, hybridisation be-
tween species offers some exciting possibilities to
avoid disease problems. Resistance in certain species
that are perhaps less desirable in their pure form can
be capitalised upon. However, in order to fully benefit
from these opportunities, it is necessary to have a
broad knowledge of diseases affecting the species of
interest. It is also essential that disease-resistant
hybrids are selected before large-scale planting is
undertaken.
3. Wood properties
The species used in the hybrid combinations with E.
grandis all have on average a higher basic wood
density than E. grandis. Indications are that the
density of the hybrid is intermediate between the
species used in the combination. As noted earlier
(Denison and Kietzka, 1992), density is the single
most important characteristic influencing pulp yield
and quality. In selecting hybrid combinations, den-
South African Forestry Journal- No. 165 June 1993
 DENSITY-AGE RELATIONSHIP OF E.GRANDIS CLONES
OAKlANDS TRIAL SITE ZULULAND
DENIlm':Ko/InI
IlOO,---------------,
CI\h. .- '" '"
400~
_ --i'- _--- ---
300 »-:
200
100
~4'----::-:---::-----------.J
30 eo 72
ME: lOONnll
1- -..cTIiD Q.DIiCI - - +'nWlL lIMN _11M ....,. . . . . . . "'" De.
BASED ON F2 E.GRANDIS CLONES
FIGURES.
sity parameters of the genotypes of the pure species
must be carefully managed, especially since basic
density is a highly heritable trait (Zobel and Van
Buijtenen, 1989). Furthermore, it should be recog-
nised that the interactions between site, tree age and
silviculture affect the basic wood density. The Mondi
pulp mill at Richards Bay has indicated a preference
for wood of a more homogeneous density ranging
between 0,48 to 0,52. To ensure that the wood of the
hybrids meet the above specifications one would need
to carefully select genotypes for hybrid crosses to
optimise the density range at felling age. It is clear
from Figures 5 and 6 that the hybrids would have to
be managed on a shorter rotation age. For each major
site and clone, an annual assessment of basic wood
density is determined. For other purposes, as is the
case for firewood and charcoal, selection for higher
density with improved calorific values would be pre-
ferred.
4. Nursery efficiency
In the majority of cases, subtropical hybrids of E.
grandis are easier and cheaper to produce than pure
E. grandis, For cutting production the hybrids cop-
pice will produce more cuttings at each harvest and
root more readily. In the multiplication hedges, hy-
brid mortality is on average 20 to 30 % less than the
pure species, and produces double the number of
suitable cuttings per harvest. With the selective
harvesting system, three to four cuttings will be
produced from an E. grandis stock every 10 to 14
days, whereas hybrids will produce an average of
seven cuttings. The hybrids root quickly, and require
about four weeks less time in the nursery when
compared to the pure species. The plants are more
robust and less sensitive to handling, heat and drought
when planted.
5. Adaptability
Experience in the Zululand coastal area has revealed
that, under certain site conditions, the hybrids of E.
grandis with either E. camaldulensis, E. tereticornis
and E. urophylla generally outperform their pure
Suid-Afrikaanse Bosboutydskrif- nr. 165, Junie 1993
DENSITY-AGE RELATIONSHIP OF E.HYBRIDS CLONES
.ZULULAND TRIAL SITES
DENIITY:Ko/InI
8OOr----------------,
~.,.:.:.-
550
~------------'
~ ~ ~ ~ ~ ~ ~ n M M
ME: lOONTHS
I 1- -.acrsD~ II&NIl +'nIML ...... ..,. fUll'" ..... "" Da.1
BASED ON F2 E.HYBRIOClONES
FIGURE 6.
species counterparts. This is especially true for E.
grandis, where the hybrids can tolerate harsher
sites. The site quality or site type is determined by
the combination of soil, climatic and terrain condi-
tions that prevail in a particular area. In several
cases where the soil conditions are good, the climatic
conditions can become limiting and vice versa. In
general, the hybrids can tolerate the limiting factors
and appear to withstand stress more readily than the
pure species. For this reason site/species matching is
carefully planned according to a natural resource
data base and whenever limiting factors are recog-
nised, hybrids are planted in preference to the pure
species.
HYBRID APPLICATION
The planting of hybrids in South Africa is increasing
and will continue to develop if future wood require-
ment predictions are to be satisfied. The develop-
ment of hybrid forestry has been confined largely to
the warmer zones, where clonal forestry is feasible
from crosses with E. grandis. It is predicted that
hybrid forestry will expand to include combinations
incorporating cold-tolerant eucalypts and pines. This
will become reality as vegetative propagations sys-
tems improve to include vegetative multiplication on
an operational scale.
The hybrids will then play an even more important
role, since they will more readily adapt to marginal
sites in closer proximity to processing plants. Trans-
port and associated costs will force production for-
estry closer to the mills if good growth and quality is
obtainable.
Hybrids will help alleviate many of the rural
demands for fuel-wood and building material. Many
of the hybrid combinations adapt well to the harsh
conditions encountered in many rural areas. The
yield, form and wood properties of these hybrids
planted under adverse conditions significantly out-
perform the pure species, thus meriting their appli-
cation in these areas, especially considering social as
well as conservation implications. Indigenous trees
and shrubs are being depleted at an alarming rate in
many developing countries and communities to sup-
59
ply essential fuel and wood requirements.
Mondi Forests has developed an exciting project in
KwaZulu (Natal) called Khulanathi, which in Zulu
means "grow with us". Khulanathi is aimed at help-
ing subsistance farmers in rural KwaZulu to grow
plots of Eucalyptus trees on a commercial basis.
Rural land holders are encouraged to grow the trees
on under-utilised land. While Mondi secures an addi-
tional source of timber, it provides management,
technical expertise, a guaranteed market and supe-
rior clonal plants. On most ofthe areas, hybrid clones
adapt the best and will ensure an earlier financial
return for the participants. Almost 800 Zulu farmers
are currently involved in the project with woodlots
covering 1 600 ha. Mondi plan to develop a further 12
000 ha of woodlots in the future, which will supply
Mondi Richards Bay with 300 000 t of fibre per year
(Mondi World, September 1992). There are similar
schemes practiced by other organisations.
CONCLUSION
Hybrids are becoming increasingly important to the
South African forestry industry.
Hybrids have made it possible to extend tree
planting sites for plantation forestry to marginal
areas, often closer to processing sites.
If the parent populations of F 1 hybrids are man-
aged in well planned breeding programmes, many of
our marginal sites will be made profitable.
The wood and phenotypic properties ofthe hybrid
are normally intermediate between parent species,
and growth is frequently superior.
The basic wood density ofthe hybrid combinations
are generally higher than that of E. grandis.
The authors predict an increased operational use
60
of cold-tolerant eucalypt hybrids as well as pine
hybrids as vegetative propagation technology im-
proves.
REFERENCES
BRANDAO, L.G., IKEMORI, Y. CAMPINHOS, E., 1984. The
new eucalypt forest. Marcus Wallenberg Foundation Sympo-
sia Proceedings. Marcus Wallenberg Foundation, Falun,
Sweden.
BURDON, R.G., 1988. When is cloning on an operational
scaleappropriate. Proceedings "Breeding Tropical Trees",
IUFRO, Pataya, Thailand, 1988.
CAMPINHOS, E., and IKEMORI, Y., 1980. Mass production of
eucalyptus species by rooting cuttings. Silvicultura 8: 770-
775.
DENISON, N.P., and KIETZKA, J.E., 1992. The development of
vegetative propagation in Mondi for commercial afforesta-
tion programmes. IUFRO, Cali, Colombia, 1992.
DU PLOOY, A.B.J., 1984. Kraft pulp properties of 15-year-oldP.
elliottii x P. caribaea hybrids C.S.I.R. Contract Report CI
Hout 315.
MARTIN, B., 1988. The benefits of hybridisation. How do you
breed for them? Proceedings "Breeding Tropical Trees",
IUFRO, Pataya, Thailand, 1988.
MONDI WORLD, September, 1992. "Khulanathi Takes Root·,
P.O. Box 61587, Marshalltown 2107, South Africa.
NAMKOONG, G., 1988. Systems ofgene management. Proceed-
ings "Breeding Tropical 'I'rees", IUFRO, Pataya, Thailand,
1988.
VAN DER SIJDE, H.A., and ROELOFSON, J.W., 1986. The
potential of pine hybrids in South Africa. South African
Forestry Journal, March, 1986.
VAN WYK, G., SCHONAU, A.P.G., and SCHON, P.P., 1988.
Growth potential and adaptability of young eucalypt hybrids
in South Africa. Proceedings "Breeding Tropical Trees",
IUFRO, Pataya, Thailand, 1988.
ZOBEL, B.J., and TALBERT, J., 1984. Applied Forest Treee
Improvement. John Wiley & Sons, New York.
ZOBEL, B.J., and VAN BUIJTENEN, J.P., 1989. Wood Varia;
tion . Its Causes and Control. Springer-Verlag, New York.
South African Forestry Journal - No. 165 June 1993