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Norway spruce (Picea
abies) genetic resources and their conservation in Lithuania
J. Danusevicius
and R. Gabrilavicius
Lithuanian Forest Research
Institute, Girionys, LT-4312 Kauno, Lithuania
Summary
Lithuanian forests cover an
area of 1.86 million hectares. The total wooded area of the country
is 30.1%. Coniferous forests comprise 61.6%, and spruce stands
24.2%. Native spruce stands make up more than two-thirds of their
total area. Middle-aged and older stands are divided into three
selection groups. The best (1st selection groups) spruce stands
cover 41 000 ha. Genetic resources of spruce stands are
conserved in situ and ex situ. Permanent inventory and
assessment of genetic objects are carried out. Studies on phenotypic
and genotypic structures of populations are conducted; adaptation,
stability and selection effect of provenances and families, as well
as changeability of traits, peculiarities of reproductive biology
and hybridization are determined. Standards for conservation and
utilization of genetic resources are being prepared.
Introduction
The Lithuanian forests are
situated throughout the territory and cover an area of 1.86 million
hectares. The total wooded area of Lithuania is 30.1%, in which
coniferous forests comprise 61.6%, and spruce stands 24.2% (Table
1). During the last two decades, intensive reforestation has had a
tendency to increase the area of spruce stands. Mature spruce groves
are the most productive (285 m3/ha) of all stands growing
in Lithuania while they suffer the most from negative natural and
anthropogenic factors such as storms, droughts, insect pests, fungus
disease, frosts, animal damage and, finally, industrial pollution.
In Lithuania, spruce stands
are situated mostly in the western, northern and central parts of
the Republic. Scots pine forests take up a large area but there are
no clear big spruce forests. The forests of Rietavas-Taurage and
Birzai are somewhat larger. The characteristics of spruce groves are
presented in Table 2. The spruce groves of Piceetum myrtillosum
and Piceetum oxalidosum forest types prevail in Lithuania.
Mostly, mixed spruce stands with pine, birch and aspen are noted.
Mixed spruce stands with birch and aspen produce about 650 m3/ha
of timber at the cutting age. They distinguish themselves in better
stability and resistance to storms.
The area of spruce stands in
Lithuania has been decreasing continually (Fig. 1). True, it began
to increase during the last two decades because of young stands. At
present, young stands until 10 years of age make up 15.4% of the
total area of spruce stands. It is known how many disasters the
spruce groves suffer during their ontogenesis. Continually recurring
storms (in 1967, 1982 and 1993) destroyed most spruce stands.
Usually a big invasion of spruce bark beetles, which destroy what is
left after windfalls in spruce groves, follows the storms. The
latter process was particularly severe after the storm in 1992. One
of the main measures at present being used in Lithuania for cleaning
defaced and damaged spruce stands is to cut the damaged stands and
remove the stem together with spruce bark beetles. It is impossible
to determine the losses which we will suffer.
Another significant factor
destroying the spruce stands is industrial air pollutants. According
to the data of our specialists who are carrying out ecological
monitoring activities (Ozolincius 1994), defoliation of spruce
groves continually increases. For instance, in 1989 the spruce
stands damage class was 0.7 while in 1994 it was 1.1. General spruce
stand defoliation level at present comes to 21.7%.
[Table 1] not available
Table 2. Productivity
of various Norway spruce stands at 100 years of age and 0.7 of
stocking level
| Type
of forest |
Mean height (m) |
Mean diameter (cm) |
Mean volume (m3/ha) |
|
Piceetum myrtillosum |
24.1 |
27.5 |
313 |
|
Piceetum myrtilloso-oxalidosum |
25.7 |
29.0 |
333 |
|
Piceetum oxalidosum |
24.4 |
28.8 |
317 |
Fig. 1. Dynamics of
Norway spruce stand areas in Lithuania by years (not available)
At present the spruce stands
of natural origin in Lithuania make up more than two-thirds of total
spruce forest area. Annual spruce cutting woodlots consist of 2000
ha on average and the stands of natural origin are cut. Therefore,
the situation of the natural spruce grove genofund will change owing
to the artificial reforestation. The necessity arose to conserve the
valuable natural genofund of spruce as well as of other tree
species.
Main Norway spruce populations and their status
Six natural forest regions are
selected in Lithuania in which different populations of separate
tree species, among them Norway spruce, have formed. The main object
of forest tree selection is natural populations, because populations
are a primary link of genetic information. Valuable genotypes which
are adapted to the local conditions are best and genotype
conglomeration forms in natural stands, while the crossing of
separate individuals acts as natural selection under certain
environmental conditions.
Elucidation of population
bounds is a long and intricate process but it is possible to choose
and evaluate the populations most distant from each other and
separate due to the climatic differences and origins of specific
structural traits.
Diversity of genofund
decreases because the area of stands of artificial origin increases,
reforestation does not always use the seed of the best trees and
sometimes there are separate genotypes of low value. All potential
elements of a genofund participate in reproduction of new
generations in naturally formed autochthonic stands.
At present there are no big
spruce forests in Lithuania. They are situated in the western,
central and northern parts of the Republics and divided by larger or
smaller deciduous stands. Larger spruce groves remain in four parts
of the country and make up the following spruce accumulations:
southwestern Zemaitija, northeastern Zemaitija, forest of Birzai and
central southern part. However, these spruce stands are cut by
woodlots and damaged by windfalls. There are some spruce stands in
other parts of Lithuania as well but they do not form big forests.
Mostly, they are mixed with big forests of other tree species.
Therefore, it is necessary to conserve the genofund or remaining
spruce groves and to utilize them rationally in reforestation (Gabrilavi…ius
and Pliura 1993).
At present, the following best
Norway spruce populations are selected in Lithuania: Survilai (1st
forest natural region); Rokiskis (2nd forest natural region);
Ignalina (3rd forest natural region); Mociskis (4th forest natural
region); Sakiai, Kazlu Ruda, Punia (5th forest natural region);
Trakai (6th forest natural region). Investigated populations differ
significantly in most structural traits. Height differences of
separate populations make up 21.4%, and ones of crown width comprise
35.8%.
It is important to ascertain
the limits of population transfer, using them to establish new
spruce stands. The influence of environmental conditions on trait
variability must be studied, and the progenies transferred into new
ecological conditions; other studies include determining the
influence of genetic characteristics (Gabrilavi…ius 1992).
Ecological adaptability of
populations is the genetic result of different reactions of each
population to site fertility, humidity and micro- and macroclimatic
conditions so it is possible to get additional selection effect by
selecting specifically adapted populations.
Studies have evaluated
reliable growth and adaptability differences between some Lithuanian
Norway spruce populations (Danusevi…ius 1993; Gabrilavi…ius
1992). Adequate populations (their progenies) are being recommended
for reforestation in every forest natural region.
In the winter of 1993 the
storm turned a significant part of the spruce stands upside down. A
dry summer followed and also in 1994 was an unforeseen drought. For
this reason, the spruces became significantly weaker and the pest Ips
typographus has spread. At present, there are
60 000-70 000 ha of spruce stands being damaged by the
abovementioned pest; this area makes up 32-38% of middle-aged and
older stands. It is planned to clear-cut an area of
35 000-40 000 ha of the damaged spruce stands, which
represents more than a 2-year cutting rate.
Methods to conserve genetic resources
Primary evaluation of forest
genetic resources is carried out during forest inventory, when
middle-aged and older stands are divided into three selection
groups: Group ICstands of Ia and I bonitet, high productivity and
quality, healthy, normal thickness; Group IICstands of II-III
bonitet, average productivity and quality; Group IIICstands of IV
and lower bonitets, low productivity and quality. At the present
moment, Lithuanian middle-aged, premature and mature spruce stands
are of good selection value (Tables 3 and 4). Stands of the Group I
cover 41 000 ha, or 22% of the area of middle-aged and older spruce
stands (Kenstavi…ius and Brukas 1993). They make up the most
valuable genofund of Lithuanian spruce stands, and therefore more
attention is paid to their conservation and regeneration. In
Lithuania two methods of genetic resources conservation are applied:
In situ
C genetic reserves, when
highly valuable stands are singled out and conserved up to natural
maturity, allowing the removal of dying or damaged trees, to fight
against pests and to protect the main species from overtopping by
others; natural regeneration is used,
C plus (elite) individual
trees, when individual trees are supervised and kept until their
death, while progenies are transferred into archives or
plantations,
C seed orchards, when the
stand is left until maturity, and the seed crop is gathered from
standing trees. Regeneration is natural.
Ex situ
C clone archives, when 15
ramets from each clone are grown in special sites,
C seed clone orchards,
C test plantations from plus
(elite) tree seeds or genetic reserves and seed sites.
Table 3. Distribution
of Norway spruce stands according to the selection (genetic) groups
in Lithuania
| Age classes |
Total area
(ha) |
Distribution according to the selection (genetic) groups
(%) |
| |
|
I |
II |
III |
|
Middle-aged |
75.2 |
24 |
64 |
12 |
|
Premature |
17.9 |
14 |
58 |
28 |
|
Mature |
93.1 |
22 |
63 |
15 |
|
Total |
186.2 |
C |
C |
C |
|
Mean |
C |
22 |
63 |
15 |
Table 4. The amount of first
selection group stands of Norway spruce by forest natural regions in
Lithuania
|
No. |
Forest
natural region |
Area |
| |
|
ha |
% |
|
I. |
Lowland of Zemaitija |
3013 |
14.9 |
|
II. |
West Aukstaitija |
4800 |
23.7 |
|
III. |
East Aukstaitija |
965 |
4.8 |
|
IV. |
Littoral lowland |
1891 |
9.3 |
|
V. |
Suvalkija and Kaunas region |
7810 |
38.6 |
|
VI. |
Dzukija |
1764 |
8.7 |
| |
Total |
20243 |
100.0 |
Data presented in Table 5 are
from work carried out in the field of genetic resources
conservation. In agreement with the Law of Lithuanian Forests,
genetic reservations in the future will acquire the status of
genetic reserves, because in the first case man's interference is
prohibited. Genetic reserves comprise 0.5% of the total area of
middle-aged and older spruce stands (Baliuckas et al. 1994).
This year additional selection
is still foreseen, while damaged ones will be reassessed. Additional
seed reserves also will be singled out. At present, standards for
selection, management and utilization of the abovementioned
conservation objects of genetic resources are being specified and
therefore an international experience here would be of great help.
Some plus trees are dead or
damaged, and therefore in the near future their additional selection
and evaluation will be carried out. Clones of most of dead plus
trees are concentrated in archives, so that their genotypes are not
lost.
Table 5. The conserved
Norway spruce genetic resources in Lithuania
| No. |
Way of preservation |
Number of objects |
Area (ha) |
|
In situ |
|
1.
2.
3.
4. |
Strict genetic reserves
Genetic preserves
Seed stands
Plus trees |
1
51
21
589 |
429.0
542.0
149.6
C |
|
Ex situ |
|
5.
6.
7. |
Clonal archives
Clonal seed orchards
Progeny plantations
Progeny tests
Provenance tests |
5
27
18
13
5 |
15.5
339.1
45.2
22.0
23.2 |
Studies on genetic resources
and the need for their improvement
At present, studies on genetic
resource are conducted by the Lithuanian Forestry Research
Institute. Owing to long-term isolation from West European countries
and inadequate financing, investigations were carried out on a small
scale and slowly. The works that have been and are being carried out
can be divided into two groups: fundamental and applied (Table 6).
More detailed studies were
performed concerning phenotypic and genotypic structure of
populations, selection efficiency of provenances and families in
situ and ex situ, including adaptability and stability
indices; investigations of genotype by environmental interaction and
geographical changeability, as well as peculiarities of reproductive
biology in seed orchards and interspecific as well as intraspecific
hybridization.
In the field of applied
sciences, recommendations and standards for stand selective
evaluation, for selection of plus trees, genetic and seed reserves,
as well as for establishing seed orchards and clonal archives, and
for increasing seed crop productivity, have been prepared.
It is foreseen to continue the
inventory of forest genetic resources, studies and compiling of
catalogues, work on tree assessment and genotype identification, as
well as to work out criteria for the selection of plants under
conservation.
The quality and extent of
these works depends considerably upon international collaboration
which is carried out by the International Plant Genetic Resources
Institute, and upon our united efforts in the EUFORGEN programme, as
well as on financial support.
Table 6. Research
investigations in genetic resources estimation, preservation and
utilization
|
No. |
Basic research |
No. |
Applied research |
|
1. |
Analysis of the phenotypical and genotypical structure of
populations and provenances. |
1. |
Criteria for selection of the plus trees, stands and
selection groups. |
| |
|
|
|
|
2. |
Methods for selection of the plus trees and stands. |
2. |
Selection of genotypes for the higher generation seed
orchards; methods and technologies for establishment of the
seed orchards. |
| |
|
|
|
|
3. |
Efficiency of selection of families and provenances, grown in
situ and ex situ. |
|
|
|
4. |
Limits of provenance transfer. |
3. |
Selection of the genetic reserves and
seed reserves. |
|
5. |
Methods of the early prediction and
evolution of genotypes. |
4. |
Establishment of the clonal archives. |
|
6. |
Bioecological peculiarities of flowering and bearing in
seed orchards. |
5. |
Means to increase the forest seed production. |
|
7. |
Creation of new hybrids. |
|
|
| |
|
|
|
References
Baliuckas V. et al.
1994. Forest genetics and breeding in Lithuania. Information
publication, Girionys, 30 p.
Danusevi…ius
J. 1993. Growth peculiarities of Norway spruce provenances and
families in Lithuania. Pp. 38-43 in Proceedings of IUFRO (S
2.2-11) Symposium, Latvia, 1993.
Gabrilavi…ius
R. 1992. Adaptation of spruce populations and effectiveness of
selection. Exp. Biol. 3-4 (Vilnius "Academia"):35-36.
Gabrilavi…ius
R. and A. Pliura. 1993. Breeding of Norway spruce in Lithuania. Pp.
189-192 in Proceedings of IUFRO (S 2.2-11) Symposium, Latvia,
1993.
Kenstavi…ius
J. and A. Brukas. 1993. Lietuvos misku pakartotinio tvarkymo
augimvieciu pagrindu rezultatas ir naudojimo perspektyvos [Results
of rests on basis of sites and perspectives of their usage]. Kaunas,
527 p. [In Lithuanian].
Ozolincius, R. 1994.
Forest condition assessment in regional network of forest
monitoring. Annual report of Lithuanian Forest Research Institute,
Kaunas-Girionys.
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