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Conservation of Norway
spruce genetic resources in Slovakia
Ladislav Paule
Faculty of Forestry, Technical
University, SKB96053 Zvolen, Slovakia
Introduction
The forest land in Slovakia is
about 1.94 million hectares, out of which 90% belongs to the
Carpathians. In the Polish Carpathians, the forests cover about 480
000 ha, in the Ukrainian Carpathians about 1 500 000 ha
and approximately 5 500 000 ha belong to the Carpathians
mountain range in Romania. This means that the Carpathian forests
cover about 9 million hectares in total.
The present distribution of
trees in European forests has been influenced by several factors:
- the natural migration of
tree species in the post-glacial period (different refugia and
geographical conditions B mountain ranges B influence
migration),
- long-lasting human
activity before the period of modern forestry,
- a short period of modern
forestry (in Europe about 100B200 years),
- the influence of human
activities and human-caused processes during the period of
industrialization.
Occurrence, origin and
distribution of Norway spruce in Slovakia
The forest stand area of
Slovakia covers in total 1 904 339 ha (Table 1). The proportion
of Norway spruce is 27.5% and that of conifers is 43.1%. The most
common broadleaved species in Slovakian forests (European beech)
covers in total 29.8%. The present proportion of Norway spruce is
higher than the original one. It is estimated (Han…inský 1972)
that originally Norway spruce covered about 10% and during the last
century it replaced the silver fir (e.g. 15% in 1920 and 5% at
present).
Natural distribution of the
Norway spruce forests in Slovakia is in altitudes between 600 m
and the upper tree limit, which is usually in the altitudes of
1400-1500 m. The upper tree limit is usually created by the mixed
Norway spruce and larch stands (e.g. the southern slopes of the High
Tatras or Norway spruce with the admixture of rowan or stone pine).
In all mountain ranges the Norway spruce creates the upper tree
limit except in the Eastern Beskydes where in the Carpathian
disjunction the Norway spruce is missing and the beech reaches the
highest altitudes.
Indigenous occurrence of the
Norway spruce populations is in all mountain ranges of central and
northern Slovakia. It is missing in the Little Carpathians and
Štiavnické vrchy and in eastern Slovakia. In eastern Slovakia the
Norway spruce's natural range is disrupted by the Carpathian
disjunction, i.e. from Košice toward the Ukrainian border and
further into CarpathoBUkraine a 200-km zone is without spruce. From
the northern side of the Carpathians, the limit of natural
occurrence of Norway spruce reaches Tarnawa, i.e. more or less the
border of Slovakia.
One of the classifications of
forest vegetation cover was elaborated on by Prof. A. Zlatník who,
on the basis of an intensive investigation of the Carpathian forests
in Slovakia and in Carpatho-Ukraine, described the natural character
of forest cover in eight vegetation levels, which could be defined
as the climax geobiocenoses which are conditioned by certain
geography, macro- and mesoclimate in given altitudes. The forest
vegetation zones are usually named according to prevailing tree
species and correspond with the first five altitudinal geographic
zones usually applied in geobotany (planar, colline, submontane,
montane, oreal, subalpine, alpine and nival):
- Oak forest vegetation
zone
- Oak-beech forest
vegetation zone
- Beech-oak forest
vegetation zone
- Beech forest vegetation
zone
- Beech-fir forest
vegetation zone
- Beech-fir-spruce forest
vegetation zone
- Spruce forest vegetation
zone
- Mountain pine forest
vegetation zone.
The principal characteristics
of forest vegetation levels are as follows (Table 2):
- increasing intensity of
solar radiation
- decreasing temperature
(for each 100 m, 0.54EC)
- increasing number of
frost days
- increasing difference
between air and soil temperatures
- decreasing vegetation
period
- increasing precipitation
total (but only up to a certain altitude over the upper
vegetation limit)
- increasing proportion of
snow precipitation
- increasing number of
foggy days
- increasing evaporation
- rapidly decreasing air
humidity with temperature decrease.
Table 1. Norway spruce
area in Slovakia
|
|
Norway spruce |
Conifers |
Total |
|
Actual area (ha) |
523 692 (27.5%) |
820 770 (43.1%) |
1 904 339 |
|
Growing stock (1000 m3) |
126 391 (34.5%) |
183 068 (50.0%) |
365 946 |
|
Area by age classes (ha) |
|
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15+ |
53 446
45 981
34 724
33 711
53 120
53 832
53 802
64 956
50 077
27 871
17 820
10 874
7 858
5 761
9 839 |
|
Annual allowable cut (1000 m3) |
|
Final cutting
Thinnings
Total |
1 118
366
1 484 |
1 742
459
2 201 |
3 724
1 087
4 811 |
|
Share of Norway spruce composition (%) |
|
Natural
1950
Actual
Optimum |
7.7
27.3
27.5
23.6 |
23.6
42.8
43.1 |
|
Table 2. Principal
characteristics of forest vegetation zones
|
Vegetation zone |
Altitude(m a.s.l.)
|
Mean annual temp. (EC) |
Annual precip. (mm) |
Vegetation period (days) |
|
Oak |
< 350 |
> 8.0 |
< 600 |
> 165 |
|
Oak-beech |
350-400 |
7.5-8.0 |
600-650 |
160-165 |
|
Beech-oak |
400-550 |
6.5-7.5 |
650-700 |
150-160 |
|
Beech |
550-600 |
6.0-6.5 |
700-800 |
140-150 |
|
Fir-beech |
600-700 |
5.5-6.0 |
800-900 |
130-140 |
|
Spruce-beech |
700-900 |
4.5-5.5 |
900-1050 |
115-130 |
|
Beech-spruce |
900-1050 |
4.0-4.5 |
1050-1200 |
100-115 |
|
Spruce |
1050-1350 |
2.5-4.0 |
1200-1500 |
60-100 |
|
Mountain pine |
> 1350 |
< 2.5 |
> 1500 |
< 60 |
The ecological characteristics
of the forest vegetation zones are given in the paper of Van…ura
(this volume). The natural range of Norway spruce covers the
following forest vegetation zones: beechBfir, beechBfirBspruce and
spruce forest vegetation zone. In the first two forest vegetation
zones, the occurrence of mixed forests is prevalent (broadleaves
with spruce admixture or spruce with silver fir admixture), while in
the last forest vegetation zone the pure spruce forests prevail.
In Table 3 is shown the
proportion of the individual stand formation of spruce forests and
mixed forests with spruce: spruce-silver firBbeech and silver
fir-beech according to individual forest vegetation belts. From this
table it is clear that the proportion of the spruce forests within
the beechBfir, beechBfirBspruce and spruce forest vegetation belts
could be considered as more or less natural. The occurrence of
Norway spruce in the 1st to 4th vegetation belts is not natural and
partially also that within the beechBfir and beechBfirBspruce forest
vegetation belts could be considered as the replacements of silver
fir in the mixed stands.
Table 3. Proportion
(%) of individual forest formations (types) in forest vegetation
belts
|
Forest vegetation belt |
Spruce |
Spruce-fir-beech |
Beech-fir |
|
1 |
0.2 |
0 |
0 |
|
2 |
0.8 |
0.6 |
0.3 |
|
3 |
2.9 |
7.7 |
2.4 |
|
4 |
10.6 |
26.4 |
12.4 |
|
5 |
33.5 |
24.4 |
29 |
|
6 |
55.9 |
11.2 |
23.6 |
|
7 |
89.5 |
1.9 |
6.8 |
|
8 |
1.5 |
0 |
0.1 |
|
Total (ha) |
308 562 |
250 609 |
210 628 |
Conservation aims and current
state of conservation activities
In the forest practices of
Slovakia there are several methods of genepool conservation in
situ and ex situ.
In situ
1. National parks and
largeBscale protected areas
2. Gene reserves (bases)
3. Approved stands for seed
procurement (A and B category).
Ex situ
1. Reproduction plantations ex
situ
2. Genebanks (seed)
3. Clonal archives
4. Seed orchards
5. Experimental trials.
The first regulation aimed at
the seed procurement only from approved stands originated in the
territory of Slovakia in 1938 and in the Czech Republic in 1939. In
the sense of this law, the seed of conifers (Norway spruce, silver
fir, Scots pine and European larch) could be procured only in
approved forest stands of A and B category and seed transfer was
allowed only within silvicultural zones. The silvicultural zones
were defined according to their geographical distribution and the
length of the vegetation period:
- subalpine regions with
the vegetation period shorter than 100 days,
- mountains with cooler
climate with the vegetation period 100-129 days,
- mountains with mild
climate and vegetation period 130-165 days,
- lowlands and hills with
plain climate and vegetation period longer than 165 days.
Subsequently, this regulation
was renewed in 1965 and since 1985 and 1988 there are new
regulations valid for the Slovak and Czech Republics. It has to be
noted that these two different regulations for both republics of
former Czechoslovakia were necessary due to different ecological
conditions and history of the forest stands.
Following these regulations
there are two different categories of approved stands, A and B, and
a new term was introduced for the territory of SlovakiaC`seed zone'.
The seed zone is defined as the zone with similar conditions within
which seed transfer is allowed. Exceptionally, seed transfer is
allowed between individual zones (as stated in the regulation).
For Norway spruce there are
seven seed zones in Slovakia (Fig. 1), five of which are within the
natural range and two are outside the natural range. Vertical
transfer of seed is allowed for altitudes up to 1200 " 200 m
and over 1200 " 100 m in the Tatra seed zone and in other zones
over 1200 " 150 m from the altitude of forest stands.
It has to be noted that there
are two seed zones into which it is prohibited to import seed: the
Tatra seed zone (Tatra National Park) and the Beskydy-Orava seed
zone which is adjacent to the Polish Beskydes (Istebna and Rycerka)
and also to the Moravian Beskydes.
At present there are in total
6075.25 ha of approved Norway spruce stands of the A category and
16 328.81 ha of stands of the B category, selected 345
phenotypically superior plus trees (in total 3171) and established
4.50 ha of clonal seed orchards and 2 ha of seedling seed
orchard (Stará Lesná) (in total 202.82 ha of seed orchards of all
species). The highest proportion of the Norway spruce approved
stands is in Northern Slovakia (2958 ha of A category and 7907 ha of
B category) and in Central Slovakia (1668 ha or 2887,
respectively), while in the Tatra National Park alone there are 722
ha of the approved stands of the A category and 1554 ha of stands of
the B category.
Fig. 1. Natural
Norway spruce zones of the Slovak Republic
The gene reserves (bases) are,
in the forestry practice of Slovakia, a new item aimed at genepool
conservation in situ. The gene bases are in general units of
the gene conservation practices established in the regions of the
most valuable genepool and the size is usually several hundreds of
hectares (200-2000 ha). In comparison, the size of approved stands
is equal to the size of a stand as a forest management unit, which
is usually a maximum of 10-15 ha. The gene bases usually include the
forest stands of different age classes with the aim to maintain the
dynamic aspect of genepool conservation. From the forest management
point of view, the gene bases are forest stands with a special
management regime, in which natural regeneration is the principal
method of regeneration allowed (the artificial regeneration with
plants originating from the local seed sources is allowed). At
present 44 gene bases are established in Slovakia (in total
12 673 ha) of which 3154.46 ha are declared to conserve the
genepool of Norway spruce. From those seed orchards should be
mentioned those which are aimed at the gene conservation of Beskydy
spruce (region adjacent to Polish Beskydes), those in the High
Tatras and in the Low Tatras.
Seed stands are special units
established mainly for genepool conservation in situ or ex
situ for the purposes of conservation and future procurement of
seed. In general the seed stands are progenies (from natural
regeneration or artificial regeneration as open-pollinated
progenies) of the approved stands of A category.
In total 127 seed stands are
established in Slovakia (not identical with seed stand according to
the OECD/EEC regulations) with the area of 635.5 ha and of this
number 37 seed stands of 196.5 ha are established for Norway spruce.
Forty percent of Norway spruce seed stands are established ex
situ and 60% in situ. From those established in situ
40% again originate from natural regeneration and the remaining 60%
originate from artificial regeneration.
In Slovakia there are in total
seven provenance trials (12 to 49 provenances) established from the
material of the IUFRO 1964/68 Norway spruce Inventory Provenance
Experiment (Holub…ík 1980) and four provenance trials being part
of the IUFRO 1972 provenance experiment with Polish Norway spruce
provenances. There exist only single-progeny tests with 42
open-pollinated progenies of a single stand from the Tatra National
Park (considered also as a seedling seed orchard).
In the seed bank established
in Liptovský Hrádok there are in total 75 seedlots with Norway
spruce (363.5 kg). They are stored with the aim to cover the seed
needs in the period of lower crops as well as for scientific and
genepool conservation purposes. In Slovakia all seed used in
forestry practice has to originate from approved stands only. Each
seedlot has a 10-digit code which should be maintained also in the
nurseries and planting stock. In the Semenoles, Liptovský Hrádok,
there are at present 332 units (8996.65 kg) of Norway spruce seed
from approved stands. At present, there are no pollen and tissue
banks established in Slovakia.
There are in Slovakia in total
292 state nature reserves and 74 protected sites with the total area
of 79 415 ha. The proportion of Norway spruce in the state
nature reserves is 37.74% and in protected sites 20.64%; almost 48%
of all state nature reserves are found in the spruce-fir-beech and
spruce forest vegetation zone (Vološ…uk 1993). In contrast to
many neighbouring countries there are in Slovakia several remnants
of spruce or mixed virgin forests with spruce occurrence. The best
known of them are Pod Chlebom (222.77 ha) and Rozsutec (650 ha) in
the Little Fatra, „ierny KameÁ (34.40 ha) in the Great Fatra,
Babia Hora (530.33 ha) and Pilsko (580 ha) in the West Beskydes and
Kotlov đµab (46.94 ha), and Podbanské (1800 ha) in the High
Tatras (Korpeµ 1995).
Threats to Norway spruce
genetic resources
Several factors affect
negatively the genetic resources of Norway spruce in Slovakia: air
pollution, game grazing, abiotic damages (e.g. drought) and
subsequent biotic damages (e.g. bark beetle), and improper forest
management practices not supporting natural regeneration.
High-elevation Norway spruce
populations are usually damaged by air pollution. Most affected are
the populations in the Beskydes and the High Tatras on the western
to northern slopes facing Ostrava and Katowice industrial regions
from which the longBdistance transported pollutants are the main
damage factor. Also other air pollution sources are negatively
influencing the health state of Norway spruce populations in the Spiš
region and the Low Tatras. Along with the decreased vitality and
damage of the assimilatory organs the air pollution affects the
regeneration processes and decreases flowering intensity and seed
production.
Browsing by excess populations
of larger ungulates (red deer) is the main factor negatively
influencing natural regeneration and the healthy state of
middle-aged Norway spruce stands. The worst situation is when the
combination of several damage factors (air pollution, insects, deer)
occurs.
The negative impact of forest
management practices (higher proportion of the artificial
regeneration, improper seed transfer, simplification of forest
management practices (e.g. replacement of mixed stands by
oneBspecies spruce stands, etc.) is also known from the territory of
Slovakia and owing to these activities the high proportion of very
valuable indigenous Norway spruce stands disappeared.
Relevant research activities
and needs; further development of conservation strategies
At present there is not very
much information about the genetic diversity of economically
important tree species in the Carpathians. However, there are tree
species in which the genetic structures and genetic diversities have
been systematically investigated, also in the Carpathians, during
the last decade. Among these species are the Norway spruce, silver
fir (Abies alba) and European beech (Fagus sylvatica).
Rather scarce information is available on the genetic structure and
diversity of other economically valuable tree species (e.g. oaks,
European larch, Scots pine, etc.).
In the abovementioned tree
species, research has been predominantly on the geographic variation
of allelic frequencies in natural populations, with an aim to
understanding the differences between individual regions and/or
populations and to understand the migration of individual tree
species from the different post-glacial refugia or selection
processes. Three examples of intensive investigation of genetic
diversity and differentiation in which our team has participated can
be given.
The first investigation of the
genetic structure of Norway spruce populations from Slovakia was
done by Paule et al. (1990). They investigated the genetic
structure of adjacent populations and found insufficient genetic
differences between adjacent populations but in some cases also
between the more distant populations.
The investigation of the
geographic variation of Norway spruce in Eastern Europe, including
the Carpathians, was done by Bergmann and Gömöry (in preparation)
and another investigation of the differences between the Norway
spruce virgin forest and naturally and artificially regenerated
Norway spruce forest stands was done by Gömöry (1992). Results of
this investigation give insight into the structure of the natural
Norway spruce forests of Slovakia and show that naturally
regenerated forest stands do not significantly deviate from the
natural forest of virgin character. Significant deviations could be
expected only in the case of artificially regenerated forest stands
in which significant narrowing of the genetic diversity could also
occur (Gömöry 1992).
The practical gene
conservation in Slovakia is carried out in the Forest Research
Institute, Zvolen, Research Station Liptovský Hrádok, where the
central register of approved stands, seed stands and gene bases is
localized. Experts from this research station (Hoffmann, Piovar…i
and earlier also Chudík) are responsible for approval of forest
stands for seed procurement and also of the approval of genebases
(reserves). The coworkers of the Faculty of Forestry of the
Technical University in Zvolen (Paule, Gömöry) are involved mainly
in basic population genetic research.
Public awareness of the
importance of forest genetic resources
The public awareness of the
importance of forest genetic resources has been reflected in the
updating of laws on the approval of forest stands (1985) and gene
reserves (1991) as well as in the preparation of the Law on Plant
Varieties which is prepared for approval in the Parliament. This is
the first time in the legislation of this country that the Law on
Plant Varieties (common with agricultural crops) will be a part of
the legislation. It will be compatible with EC and OECD rules.
The next updating of law
regulations is under discussion and has considered recent ownership
changes in Slovakia.
Reconstruction of the forest
seed bank in Liptovský Hrádok is planned for the near future. It
will be aimed not only at the technical reconstruction and
replacement but also the new regulation on the content (the
selection of samples and their quantities) of the Seed Bank will be
updated.
The topics of gene
conservation are permanently a part of the research programme of the
Forestry Research Institute in Zvolen and their research stations in
Liptovský Hrádok and Banská Štiavnica. In Liptovský Hrádok
there is also a central database on gene resources (approved stands,
genebases, seed stands and seed orchards). A part of this research
station is Seed Control, an organ of the Ministry of Agriculture
which aims to control the forest seed procurement and business in
Slovakia.
The most urgent needs in
Norway spruce conservation
There are several urgent needs
for the conservation of the Norway spruce genepool in Slovakia. Among
them, there is the danger of air pollution in several regions with
very valuable Norway spruce populations (e.g. Spiš and Beskydy),
the grazing by ungulates (e.g. red deer) and the simplification of
forest management activities (e.g. replacement of mixed forest by
Norway spruce monocultures).
Last but not least there is
danger from forest owners themselves. Within the reprivatization
process of the Slovak forests the highest proportion of small
private forest owners is in the region of the best Norway spruce
populations of the Beskydy ecotype of Norway spruce, i.e. Orava and
Kysuce.
References
Gömöry, D. 1992.
Effect of stand origin on the genetic diversity of Norway spruce (Picea
abies Karst.) populations. Forest Ecol. Manage. 54:215B223.
Han…insky, L. 1972.
Lesne typy Slovenska. Priroda, Bratislava.
Holub…ík, M. 1980.
[Provenance research of Norway spruce (Picea abies Karst.) in
Slovakia]. Lesnícke Stúdie 33:1B95.
Korpeµ, Š. 1995. Die
Urwälder der Westkarpaten. Gustav Fischer Verlag, Stuttgart, 305
pp.
Paule, L., A.E. Szmidt
and R. Yazdani. 1990, Isozyme polymorphism of Norway spruce (Picea
abies Karst.) populations in Slovakia. I. Genetic structure of
adjacent populations. Acta Facultatis Foresatalis, Zvolen 32:57-70.
Vološ…uk, I. 1993. The virgin
forests and reserves in Slovakia. Pp. 69-74 in European
Forest Reserves Workshop. Pudoc Publ., Wageningen.
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