Seed Transfer Across Europe What Do We Know?

Attachment 1: Second draft

Seed transfer across Europe – What do we know? Synthesis of the results of a questionnaire, the workshop “Seed transfer across Europe – what do we know?” and literature research

Realised by: Doris Krabel and Heino Wolf, SBS Graupa, Germany With contribution of partner AT, DE, DK, ES, F, FI, GB, IE, IT, NL, NO, PL, RO, S, SK

Introduction

Since the antiquities, seed transfer of fruit trees in particular has been very active: the current distribution across Europe of species like chestnut and walnut is mostly due to intensive transfer starting at the Roman time and later on encouraged for example by Charlemagne. Inquiries on the history of seed transfer showed that first confirmed information are coming from the end of the 15th century e.g. when seeds from forest trees from the Baltic region were brought to Portugal. In 1601 Robin introduced for the first time Robinia pseudoacacia in France. Other famous examples of seed trading in the following centuries were e.g. the transfer of Scots pine seeds (so called “Riga pine”) to France in 1770 by Barbey (Daszkiewicz 2002) or in 1828 the introduction of Douglas-fir to Europe by David Douglas (Göhre 1958).

However, in the 19th century large cone kilns were constructed and huge amount of seeds and seedlings with unknown origin were transferred all across Europe. In most cases this uncontrolled transfer was accompanied by negative results concerning growth, quality and survival of the trees. The examples presented in Fig. 1a and 1b show for Germany during the 18th and 19th century that the centre of seed production and distribution for the whole country was often concentrated in one location (here west lower Germany).

Langner (1935) is giving an excellent overview on the origin of Larch seeds and its importance on forestry in Saxony. He found out that up to 1850 larch cultivation was carried out in two periods. The first one was during the period between 1767 until 1796 where foresters used mainly local material and the second one from 1796 until 1850 when nearly without exception seed material from Tyrol has been imported. These stands completely fell victim to the common “Larch-decline” whereas trees from the first period developed positively. 1931 Zimmermann published detailed information about the historical transfer of seeds of Norway spruce for Saxony. His overview contains a table with the demand and the origin of seed material in the period between 1836 and 1924. The table shows the following consumption of seeds in Saxony: 1836 -1854 consumption of local seeds as well as seeds from Thuringia and Harz mountains 1855 – 1864 local seeds, seeds from Thuringia, Harz mountains, Bohemian Forrest and Bavarian region 1865 -1879 seeds from almost exclusively unknown origin which was delivered by the Appel Company and which presumably originates from Hungary, Austria Scandinavia and the Black Forrest

0aeb80cb26147ce36cf20dde3f95202c.doc 1 1880 – 1924 seeds from local origin as well as material from Thuringia, Harz mountains, from Swabian Mountains, Hungary, southern part of Austria, Sweden and Norway were cultivated. Learning from the failures, in the early 20th century, well-documented studies on the origin of seeds and its effect on productivity, wood quality, survival and plasticity were established. The results obtained in many national and international provenance experiments were used to provide guidelines for seed transfer zones and to identify populations that show positive characteristics.

Seed transfer in west Lower Saxony (~18th century)

Source: DKV 1967

Fig. 1a: Seed transfer during the 18th century in west Lower Saxony (Liesebach et al.2008) Historic seed transfer in Germany

1978 Sources: ~1900 Borgmann (1904)

so ~1900 1978 Immel (1967) so Müller (1988) 1880+ Rohmeder (1972) 1820+ 1426 1966/7 1966 Schwartz (1991) 1680 1880+

1967 1880+

so seed orchard

0aeb80cb26147ce36cf20dde3f95202c.doc 2 Fig. 1b: Comprehensive map of the historic seed transfer (19th and 20th century) in Germany (Liesebach et al. 2008)

Up to the second third of the 20th century the extent of the trading of Forest Reproductive Material was increasing constantly for most species (Fig. 2). So for example in the 1990ies up to 1t Douglas-fir seeds per year have been imported to France compared to 370kg in 2004. As an example figure 1 shows the drastic decline of seed consumption in Denmark between 2000 and 2006.

kg 300000

200000

100000

0 2000 2001 2002 2003 2004 2005 2006 2000 2001 2002 2003 2004 2005 2006 Year Conifer tax Import Own

Fig. 2: Consumption (import and use of seeds of own seed production) of seeds in Denmark from 2000 until 2006 (Hansen 2008).

Movement of Forest Reproductive Material (FRM) from the collection site to another environment within the species range for reforestation or restoration may increase the risk of maladaptation (Campbell 1979, Bower and Aitken 2008). An increased rate of mortality or bad growth characteristics resulting from maladaptation could reduce the success of restoration projects, and gene flow from maladapted planted trees into adjacent native populations could negatively affect adaptation to local conditions (McKay et al. 2005). On the other hand, planting of individuals adapted to environmental conditions, e.g. more advantageous climatic conditions, could be a method to facilitate migration and provide a source of genotypes well-adapted to the local environment (Bower and Aitken 2008). Regarding failures and success, seed transfer should be guided by natural levels of genetic variation and local adaptation in quantitative traits specific to the individual species (Morgenstern 1996, Hufford and Mazer 2003, McKay et al. 2005).

From an economical point of view, breeders (and foresters) will usually have to balance between on one side the (hypothetical) benefits of using local material well-adapted to local environmental conditions but in many respects sub-optimum for many properties of economical importance (stem form, wood properties, etc.) and on the other hand, the risk to use less well-adapted foreign material but with better properties.

0aeb80cb26147ce36cf20dde3f95202c.doc 3 The question of the local optimality is raised in many instances. Many examples in the literature exist showing that local populations are not necessarily better adapted than foreign populations and that is particularly true in Europe where species distribution has been largely disturbed by quaternary glaciations. For most coniferous species native from Europe for example, populations from Central Europe proved better adapted than specialised populations from the western edges of their distribution ranges. ,A comprehensive study for hybrid larch (Larix x eurolepis Henry) has shown that firstly the local variety is not necessarily the best one and that secondly the performance of the tested varieties were more stable from site to site than expected (Philippe et al. 2002). The conclusions the authors were drawing out of these results were that variety transfers are possible and even desirable in that they increase the potential lists of FRMs useful for a given region. Nevertheless the potential impacts of climate change emphasises the importance of adaptation of populations/ individuals to their local environment. For plant material suffering from diseases and pests, minimizing maladaptation may mean the differences between establishing or maintaining viable populations (Bower and Aitken 2008).

Material and Methods In January 2008 a questionnaire (http://treebreedex.eu/IMG/doc/Ac3surveyseedtransfer.doc, 17-09-2009) dealing with the situation on “Seed transfer across Europe” has been answered by 19 TREEBREEDEX-partners. The aim of this survey was to collect data about the actual import/ export situation of forest reproductive material (FRM) of known and unknown provenances in Europe and the development of the current needs over the last 10 years. Additionally the results of the questionnaire should provide information as a basis for the evaluation of the impacts of this transfer on the stability and the yield of forests in Europe. These information have been completed by selected examples presented by the TREEBREEDEX partners on a workshop held in Pirna (Germany) in January 2008 on the subject “Seed transfer across Europe – What do we know?” (http://treebreedex.eu/spip.php? rubrique55, 17-08-2009) and finally a number of reviewed publications on the effect of seed transfer have been studied in order to insert the results into an international context. When talking about seed transfer today it has to be stated that according to the EU-regulations on the Marketing of Forest Reproductive Material (Council directive 1999/105/EC) there exists no import or export within the EU but only commercial transfer. Import/ export is meant for the commercial transfer of FRM from Non-EU-members and vice versa. In the following, the terms are used for both processes in order to classify the streams of FRM between the individual countries. Keeping in mind that for tree breeders and foresters the knowledge about the movement of genetic resources (FRM) is of more importance than the knowledge about the trading.

Results

Flow of Forest Reproductive Material First of all, the results of the questionnaire (http://treebreedex.eu/IMG/doc/Ac3surveyseedtransfer.doc, 17-09-2009) show that the information about seed transfer regarding the individual TREEBREEDEX partner countries is quite heterogeneous. These differences are on one side related to an insufficient documentation and the lack of standardized rules of documentation for the whole European

0aeb80cb26147ce36cf20dde3f95202c.doc 4 Union and on the other side, a lack of transparency. The latter concerns especially the trading of FRM between private companies and private forest owners.

Table 1: List of main tree species transferred (inside/outside the respective country) in Europe.

Species Imported Exported Abies nordmanniana X X Larix decidua X X Picea abies X X Pseudotsuga menziesii X X Fagus sylvatica X X Prunus avium X X Quercus robur X X Abies procera X Pinus sylvestris X Alnus glutinosa X Betula pendula X Castanea sativa X Quercus rubra X Picea sitchensis X Pinus pinaster X Fraxinus excelsior X Tilia cordata X Quercus petraea X

Table 1 presents those tree species which are mainly imported and exported from the respective countries. In a number of cases, plants are simply imported for taking them in commissions work (e.g. Netherlands): they are listed as plant flow but they are more or less neutral in balance. Taking Slovakia as another example (Fig. 2 and 3): the main imported coniferous species is spruce (Picea abies) from Czech Republic - the plants are raised in Slovakia and returned back to Czech Republic. The same is true for the broadleaf species Quercus robur. Abies alba and Larix decidua are the main conifer species which are exported from Slovakia eg. to Germany resp. to Czech Republic. The broadleaves which are mainly exported are Quercus spec. as well as Fagus sylvatica. They are mainly transferred to Hungary and also Czech Republic.

0aeb80cb26147ce36cf20dde3f95202c.doc 5 300000

250000

200000

150000 Conife rs Broadle ves 100000 Total

0 Production in Import Export the country

Fig. 2: Slovakia - Survey on seed flow (in kg) across Europe (average 2005-2007)

150000000

100000000 Conifers Broadleves 50000000 Total

0 Production in Import Export the country

Fig. 3: Survey on the plant flow (total number) from and into Slovakia (average 2005-2007).

Another example for seed transfer was published recently concerning the import of Douglas fir seeds into eastern part of Germany (former GDR) between 1960 and 1990 (Schneck 2009). About 5800 kg Douglas fir seeds were imported from North America mostly via former FRG during this 30 years. The greater part of these seeds belonged from seed zones recommended for the use in eastern Germany according the results of provenance testing.

Reasons for the transfer of FRM

0aeb80cb26147ce36cf20dde3f95202c.doc 6 There are several reasons for shifting plant material from one country to another. The main reasons listed below reflect the answers of the survey among the TREEBREEDEX- partner institutions in 2008 (http://treebreedex.eu/IMG/doc/Ac3surveyseedtransfer.doc, 17-09-2009):

Reason 1: Better adapted material compared with local material (i.e.in France larch from sudetan mountains proved to be better adapted than alpine larch)

Reason 2: Material is “exotic” but does not rely on artificial stands (e.g. Douglas-fir, Sitka spruce all over Europe)

Reason 3: Lack of suitable plant material, because of an insufficient production of FRM (e.g. France has seed orchards for Sudetan larch but the production of seeds is insufficient; e.g. Denmark - there is a lack of FRM for Douglas-fir and Pinus sylvestris; e.g. Italy has a lack of FRM from Hybrid larch and Sitka spruce).

Reason 4: Some countries don’t have a breeding programme for the respective tree species: e.g. France for Sitka spruce or Sycamore maple; Poland for Wild cherry.

Reason 5: In many cases, seed dealers prefer getting seeds from eastern Europe to a cheaper price (e.g. Sycamore, Lime, Robinia)

Basis of choice

The decision for planting Forest Reproductive Material from another country or another region is mainly based on the following reasons: - Improved results from provenance and field tests as well as variety and clonal trials - best guess based on climatic similarities - the wish for diversification - decisions based on commercial reasons and finally best guess based on climatic similarities

Success and Failure

The analysis of significant successful cases (Table 2) of imported FRM shows for example for Sweden that seed sources of eastern European origin, mainly from Belarus have improved growth or resistance characteristics compared to domestic seed sources (e.g. Sitka spruce, European and Hybrid Larch). On the other hand seed sources of western and south-eastern European origin showed for Norway spruce reduced wood quality, higher number of stem cracks as well as a higher number of plants with top-drying disease. For Scots pine from Germany, plants from seeds, which were imported to Sweden during 1800 and 1900 have a reduced wood quality and rate of survival compared with plants from domestic seed sources.

On the one hand results of different provenance trials in Germany (Schneck 2007) showed for Scots pine which originated from several regions in Poland (north-eastern, north central, and south-western part) that they perform very well in Germany especially in the north-eastern lowlands. Based on these observations seed orchards were established with clones of plus

0aeb80cb26147ce36cf20dde3f95202c.doc 7 trees deriving of the best Polish provenances in Germany. Few of theses orchards are approved as “tested” (Schneck 2001). On the other hand there also exist examples for failure cases in connection with the import of Scots pine seeds to Germany: In consequence of a shortage of seeds and the common ability of such provenances to produce a good timber quality (stem form, thin branches) in a wider spacing, between 1960 and 1975 a large amount of Scots pine seeds was imported from Bulgaria and the western and north-western part of former Soviet Union to East Germany (former GDR). In the following time nearly all plantations consisting of such material failed.

Another example is the introduction of European larch to the central and south-eastern part of Germany in the 18th and 19th century. Münch (1933) and Langner (1935), who studied historical papers in the archives of private forest owners found out that a large number of the very well growing stands in these regions originated from seeds coming from Silesia (Sudetian mountains). Furthermore Münch and Langner reported that enormous amounts of larch seeds were imported from Tyrol in the 19th century. Nearly all plantations planted with this material failed some years after planting. This phenomenon was known as the so called “larch decline”. Significant failure cases had been also observed for example in France for Romanian Ash provenances. In the field tests plants from these provenances produced a high number of forked and crooked trees. Up to now it is not clear whether those provenances are hybrids of Fraxinus excelsior and Fraxinus angustifolia.

For most conifers in France, many populations from Central Europe proved better adapted in lowlands than local populations: that is the case for Scots pine (populations from Mazuria), for European larch (populations from Sudetan Mts), for Norway spruce (populations from Blizyn, Istebna, etc). Some material from foreign breeding programmes proved also to be of high value for reforestation in France. For example good results were obtained in France with Japanese larch from Danish seed orchard (FP615DK) and selected seed stands, in particular for stem form. Hybrid larch from seed orchards in Denmark and Netherlands, except for the hybrid variety from seed orchard FP203DK, showed excellent growth and stem form compared to plants from selected seed stands in France. Plants from FP203DK proved sensitive to stem cracks (drought or growth cracks).

0aeb80cb26147ce36cf20dde3f95202c.doc 8 Table 2: Examples of significant successful and failure cases of FRM transfer in Europe Country Successful cases Failure cases Denmark -Scots pine from Norway and some south- -Norway spruce from western Swedish provenance Arpathian Mountains and PL -Scots pine from many D- provenances France -Norway spruce from Central Europe (PL, -European larch from Baltic, Beskidy, CZ) Alpine sources for lowland -Sudetan larch from D, CZ, SK, PL plantations, Mariental and -Japanese larch from Danish seed orchard Berkel from D, D-837-03 (FP615DK) -Hybrid larch from DK-seed -Hybrid larch from Danish seed orchard orchard (FP203, FP211), D- (FP201DK) and Vaals/Esbeek-NL seed orchard Lolla, B-seed -Douglas fir from Weyerhaeuser -(Cascades orchard Halle and Twin) , Harbor-, Coos Bay seed orchards -Douglas-fir from -Scots pine from PL (314/1) Weyerhaeuser seed orchard -Poplar hybrids Balotino, I214, Dorskamp (Springfield) and Lux -Scots pine from Scandinavian populations -Ash from Romania Finland -Norway spruce from Estonia -Norway spruce from southern provenances -Scots pine from southern provenances -Larches from continental regions Ireland -Norway spruce from Denmark -Sitka Spruce from Alaska, -Sitka spruce from Oregon and Washington California -Japanese larch from Hokkaido (Japan) -Scots pine from Norway -Douglas fir from Oregon, Washington -Oak from France -Scots pine from Scotland -Oak from N, UK, D -Beech from UK Norway -Sitka spruce from Alaska -Norway spruce: some provenances from Central Europe Sweden -Norway spruce from Belarus -Norway spruce: seed -Pinus contorta of Canadian origin (Yukon sources of western and south- territory eastern European origin -Sibirian larch of western origin in Russia -Scots pine: from D imported during 1800-1900 Slovakia -European larch from Czech Republic (Sudetic type) Germany - Scots pine from ‘Ostpreussen’ , from north- - Scots pine from southern eastern, north-central, and south-western France and Bulgaria Poland - Douglas fir southern and - European Larch from Sudetian mountains some northern provenances (in parts of the lowlands) from the interior type - European Larch from the Alps (Tyrol) in the lowlands

0aeb80cb26147ce36cf20dde3f95202c.doc 9 Results presented by Chalupka ( 2008) showed for Pinus sylvestris that shifts of provenances in latitude greater than ± 4°N or ±2°C of mean annual temperature resulted in reductions in height of more than 15%. In contrast to height growth, the DBH (diameter at breath height) growth was modestly stimulated by transferring them to more northern locations. DBH growth was negatively affected by shifts to sites that were generally both warmer and further to south. Transfer of seeds to colder, more northern environments resulted in dramatic decline in tree survival. However, a modest shift in the opposite direction (to warmer, more southern sites) was slightly beneficial to survival (Chalupka 2008). Results from Sibirian larch provenance tests in Sweden showed that concerning autumn frost hardiness the latitude seems to have a strong influence. Northern provenances are generally less damaged. For spring frost the pattern is more unclear. Larix sukaczewii is less damaged than other Larch species. The eastern provenances are nearly all damaged to a high degree regardless of the latitude (Westin 2008). In this context the results of Bower and Aitken (2008) are also interesting. They suggest for Pinus albicaulis, a high elevation, five-needle pine, a seed movement from milder to colder climates to a maximum of 1.9 °C in mean annual temperature in the northern portion of the species range, and 1.0°C in the U.S. Rocky Mountains to avoid maladaptation.

Role of private forest owners

The lack of information concerning import, export and where about of FRM is one of the main problems in nearly each European country, especially when plant material is traded via private companies. By taking into consideration that climate is changing, one of the major tasks for tree breeders will be to make forest owners and private nurseries as well as other public groups aware of this subject. For example the forest owners should be convinced to use specific well-adapted plant material. Therefore it will be important to transform the results from breeding programmes etc. into an appropriate way, which makes the results understandable and finally acceptable for a wider public. One inspiring example might be Austria, which implemented an internet platform, which allows a direct dialogue between the tree breeder and the forest owner (Höbarth 2009, Schüler 2010).

Conclusions

According to “the Council Directive 1999/105/EC “it is necessary to remove any actual or potential barriers to trade which may hinder the free movement of forest reproductive material (FRM) within the Community...”. But taking into account that the natural demands of the plant material might be very different or even contrasting, which is confirmed by the results presented above, the fulfilling of the EC directive may cause a number of problems. So one should be very careful, evaluating first of all older provenance experiments to gain more knowledge and to become aware of the biological and economical consequences of free FRM transfer. (Chalupka 2008).

Recommendations for seed transfer across Europe should take into account the following aspects: - A seedling’s response to its planting environment is significantly influenced by its parents’ location (question of epigenic memory?) (Campbell 1991).

0aeb80cb26147ce36cf20dde3f95202c.doc 10 - less seed movement is possible at higher elevation - the risk of maladaptation increases when transferring seeds across more than one environmental condition - Seed orchard seeds are most safely used in breeding zones of the parents or in the area where the parents have been tested (Campbell 1991). - Seed transfer to a higher elevation usually increases the risk of maladaptation. - Latitude seems to have a stronger influence than longitude (difference to observations made in the Washington area see Campbell and Sugano 1993, Campbell 1986, Sorensen 1983 and Campbell 1991) - In harsher environments like it is reported for some areas in Finland, utilization areas are defined individually for each seed orchard; they are based on the estimated mean temperature sum - the success of seed transfer also depends on the species

Literature cited:

Bower A. D. and S. N. Aitken 2008. Ecological genetics and seed transfer guidelines for Pinus albicaulis (Pinaceae). American Journal of Botany 95: 66-76. Campbell R. K. 1986. Mapped genetic variation of Douglas-fir to guide seed transfer in southwest Oregon. Silvae Genetica 35: 2-3. Campbell R. K.1991. Soils, seed-zone maps, and physiography: guidelines for seed transfer of Douglas-fir in southwestern Oregon. Forest Science.37:973-986. Campbell and A. I. Sugano 1993. Genetic variation and seed zones of Douglas-Fir in the Siskiyou National Forest. Research Paper PNW-RP-461, Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, 1-22. Chalupka W., R. Rożkowski and M. Misiorny 2008. Effects of seed transfer in Europe: a case study of Scots pine (Pinus sylvestris L.). TREEBREEDEX Workshop: “Seed transfer: what we know?”, Jan. 14-15, 2008, Pirna, Germany Daszkiewicz P. 2002. Introduction of “Riga pine” to France during XVIII and XIX centuries: biological, historical, political and economical aspects. Doctoral thesis, Institute of Dendrology, Kórnik, Poland, 201 pp + 15 annexes [in Polish]. Göhre K.1958. Die Douglasie und ihr Holz. Akademie-Verlag GmbH, Berlin W 8, 595pp. Hufford K. M. and S. J. Mazer. 2003. Plant ecotypes: Genetic differentiation in the age of ecological restoration. Trends in Ecology & Evolution 18: 147–155. Höbarth M. 2009. Wahl von Forstpflanzen. Kärntner Bauer, 15. September 2009, Bauernjournal Forst, S. V. Langner W. 1935. Historical studies about the origin of seeds and their importance for forestry (in German). Tharandter Forstliches Jahrbuch 86: 1-120. Liesebach M., V. Schneck and H. Grotehusmann 2008. Seed transfer in Germany – Pinus sylvestris L. and Fraxinus excelsior L. TREEBREEDEX Workshop: “Seed transfer: what we know?” Jan. 14-15, 2008, Pirna, Germany. http:// treebreedex.eu/spip.php?rubrique55. McKay J.K., C.E. Christian, S. Harrison and K.J. Rice 2005. “How local is local?" A review of practical and conceptual issues in the genetics of restoration. Restoration Ecology 13: 432–440.

0aeb80cb26147ce36cf20dde3f95202c.doc 11 Morgenstern K. E. 1996. Geographic variation in forest trees. UBC Press, Vancouver, British Columbia, Canada. Münch, E. 1933. The mystery of larch as a question of provenance (in German). Tharandter Forstliches Jahrbuch 84: 438-531. Philippe, G., Curnel, Y., Jaques, D. Lee, S. and S. Matz, 2002. Performances of hybrid larch (Larix x eurolepis Henry) varieties across Europe : early results for survival, stem form and growth rate. Symposium on improvement of larch for better growth, stem form and wood quality, IUFRO Working Party S2.02-07, Sep. 16-21, 2002. Rohmeder E. 1972. Das Saatgut in der Forstwirtschaft. Verlag Paul Parey, Hamburg und Berlin, 273 pp. Schneck, D. 2009. Stands of Douglas fir in East Germany – where do they come from? AFZ- DerWald 64: 848-850. Schneck, V. 2001. Seed stands and seed orchards of Scots pine (in German). AFZ/Der Wald 56: 232-233. Schneck, V. 2007: Growth of different provenances of Scots pine – Results of provenance tests in the horth-eastern German lowlands (in German) im nordostdeutschen Tiefland. In: Landesforstanstalt Eberswalde (Hrsg.): Die Kiefer im nordostdeutschen Tiefland - Ökologie und Bewirtschaftung. Eberswalder Forstliche Schriftenreihe Band XXXII, 2007, 374-382. Schüler S. 2010. Online-Hilfe sucht beste Herkunft. Forstzeitung, Wien, 121 (2), S. 15. Sørensen F. 1992. Genetic variation and seed transfer guidelines for lodgepole pine in central Oregon. Research Paper PNW-RP-453. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 30 pp. Westin J. 2008. Seed transfer of Scots pine and Norway spruce in Sweden. TREEBREEDEX Workshop: “Seed transfer: what we know?” Jan. 14-15, 2008, Pirna, Germany. http://treebreedex.eu/spip.php?rubrique55 Zimmermann, H. 1931. Fichtensamenbeschaffung in Sachsen. Tharandter Forstliches Jahrbuch 82, 821-864.

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