Opportunities and risks for Douglas fir in a changing climate

Transcription

BERICHTE
FREIBURGER FORSTLICHE FORSCHUNG
HEFT 85
Opportunities and risks for Douglas fir
in a changing climate
- Abstracts October 18 – 20, 2010
ALBERT-LUDWIGS-UNIVERSITÄT FREIBURG
FAKULTÄT FÜR FORST- UND UMWELTWISSENSCHAFTEN
INSTITUT FÜR WALDWACHSTUM
FORSTLICHE VERSUCHS- UND FORSCHUNGSANSTALT
BADEN-WÜRTTEMBERG
ABTEILUNG WALDWACHSTUM
2010
ISSN 1436-1566
Herausgeber (editors):
Fakultät für Forst- und Umweltwissenschaften der Albert-Ludwigs-Universität Freiburg
Prof. Dr. Heinrich Spiecker, Institut für Waldwachstum
Forstliche Versuchs- und Forschungsanstalt Baden-Württemberg (FVA), Freiburg
PD Dr. Ulrich Kohnle, Dr. Kaisu Makkonen-Spiecker, Prof. Konstantin von Teuffel
Redaktion:
Dr. Kaisu Makkonen-Spiecker
Torge Brodersen
Umschlaggestaltung (cover design):
Bernhard Kunkler Design, Freiburg
Druck (print):
Eigenverlag der FVA, Freiburg
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Germany
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Content
III
Content
Content………………………………………………………………………………
Schedule
…………………………………………………………………………
III
IV
Preface ………………………….……………………………………………
VIII
Keynote I……………………………………………………………………
IX
Keynote II…………………………………………………………………………
XI
Session 1……………………………………………………………………………
1
Session 2………………………………………………………………………
5
Session 3…………………………………………………………………………
13
Session 4 ………………………………………………………………………
19
Session 5 ………………………………………………………………………
29
Session 6 ………………………………………………………………………
33
Session 7 ……………………………………………………………………………
45
Poster Sessions …………………………………………………………………
51
IV
Opportunities and risks for Douglas-Fir in a changing climate
OPPORTUNITIES AND RISKS FOR DOUGLAS-FIR
IN A CHANGING CLIMATE
October 18 - 20, 2010
Freiburg, Germany
Time
Sunday, October 17, 2010
17:00
19:00
Forest Research Institute, Günterstalstraße 61
Registration
Icebreaker
Time
Monday, October 18, 2010
08:00
08:30
09:00
09:40
10:2010:50
10:50
11:15
11:40
12:0513:35
13:35
14:15
14:40
15:05
15:3016:00
Forest Research Institute, Wonnhaldestr. 4
Registration (continued)
Welcome addresses (Moderator: Konstantin von Teuffel)
Max Reger; Konstantin von Teuffel
Keynote speech
The dynamics of Douglas-fir stands (Bruce Larson, Canada)
Keynote speech
Douglas-fir in France: history, recent economic development and overviews for the
future (Jean Louis Ferron, France)
Coffee break
SESSION 1 (Moderator: Konstantin von Teuffel)
Mapping Douglas-fir current and future abundance and potential site productivity in
western United States (Aaron R. Weiskittel, Nicholas L. Crookston, Gerald E.
Rehfeldt, Phillip J. Radtke, USA)
Going the distance: Testing seed transfer of coast Douglas-fir within its native range
(Peter Gould, Constance Harrington, Brad St. Clair, USA)
Sustainable cultivation of Douglas-fir on dry and acid sandy soils in the old-glacial
lowlands of eastern Germany – aspects of nutrient and humus dynamics (Thomas
König, Alexander Tischer, Franz Makeschin, Sven Martens, Germany)
Lunch
SESSION 2 (Moderator: Jürgen Bauhus)
Keynote speech
Douglas-fir and the management of forests as complex, adaptive systems (Klaus J.
Puettmann, USA)
Climatic and regional patterns in Douglas-fir climate-growth relationships in British
Columbia, Canada (Hardy P. Griesbauer, D. Scott Green, Canada)
Douglas-fir – a substitute species for Scots pine in dry inner-Alpine valleys? (Britta
Eilmann, Andreas Rigling, Switzerland)
Impact of Douglas-fir on the N cycle: Douglas-fir promotes nitrification? (Bernd
Zeller, Sitraka Andrianarisoa, Jean-Huques Jussy, Etienne Dambrine, Jacques
Ranger, France)
Coffee break
Schedule
16:00
16:40
17:05
17:30
17:5518:00
18:00
(1)
(2)
(3)
(4)
(5)
(6)
(7)
(8)
(9)
(10)
(11)
(12)
19:00
V
SESSION 3 (Moderator: Jürgen Bauhus)
Keynote speech
Tree-ring analysis of adaptation to drought in Douglas-fir (Philippe Rozenberg,
France)
Tree-ring growth of West-German Douglas-fir under changing climate conditions
(Stefanie Fischer, Burkhard Neuwirth, Jörg Löffler, Matthias Winiger, Germany)
Impact of water availability on wood density patterns in Douglas-fir tree rings (Bela
Johannes Bender, Heinrich Spiecker, Germany)
Findings from long-termed forest growth inventory data from 14 trial sites in the
II. International Douglas-fir progeny trials established in 1961 in north-western
Germany (Andreas Weller, Germany)
Break
POSTER SESSION
Bud burst and damage by spring frost of Douglas-fir seedlings in the south of
Sweden (Cecilia Malmqvist, Sweden)
Effects of surface humidity and light regime on germination success and seedling
development of Douglas-fir (Pseudotsuga menziesii Mirb. Franco) – a greenhouse
experiment (Franka Huth, Angelika Körner, Christine Lemke, Antje Karge, Jörg
Wollmerstädt, Sven Wagner, Martin Hartig, Dietrich Knoerzer, Germany)
Survival and growth of Douglas-fir seedlings of different provenances; results after
one growing season (Kristina Wallertz, Sweden)
Drought tolerance of native and non-native tree species in the Alps – results from a
large scale dendroecological study (Mathieu Lévesque, Britta Eilmann, Andreas
Rigling, Peter Brang, Switzerland)
Nutrient removal by wood harvesting in Douglas-fir stands for different silvicultural
treatments and varying harvesting intensities (Joachim Block, Julius Schuck, Thomas
Seifert, Germany)
The commercial development of Douglas-fir controlled mycorrhization in France: an
emerging tool for a new sylviculture (Jean Garbaye, Daniel Bouchard, Jean-Louis
Churin, François Le Tacon, Vincent Naudet, Bruno Robin, France)
Douglas-fir forests in Rhineland-Palatinate – structural diversity and future
perspectives (Ulrich Matthes, Germany)
Where has the Douglas-fir – Pseudotsuga menziesii – been first discovered, in
Canada or in the U.S.A.? (Rémy Claire, France)
Effects of climate change on growth and vitality of Douglas-fir plantations in
Caspian Forest (Iran)? (Farshad Yazdian, Iran)
Production and environmental functions of Douglas-fir on the School Training Forest
Kostelec nad Černými lesy territory (Vilém Podrázský, Jiří Remeš, Czech Republic)
Growth performance and reaction to biotic factors of Douglas-fir provenances in
northwest Germany (Mirko Liesebach, Germany)
Douglas-fir in beech forest ecosystems - diversity of two strict forest reserves
(Patricia Balcar, Germany)
Individual Program
VI
Time
08:30-
09:10
09:35
10:00
10:2510:55
10:55
11:20
11:45
12:1013:40
13:40
14:10
14:35
15:00
15:2515:55
Tuesday, October 19, 2010
Forest Research Institute, Wonnhaldestr. 4
SESSION 4 (Moderator: Philippe Rozenberg)
Keynote speech
Review of present and potential insect pests affecting Douglas-fir in Europe in a
context of global change (Alain Roques, France)
Vulnerability of Douglas-fir in a changing climate: study of decline in France after
the 2003 drought (Anne-Sophie Sergent, Philippe Rozenberg, Benoît Marçais, Yves
Lefévre, Jean-Charles Bastien Leopoldo Sanchez, Louis-Michel Nageleisen, Nathalie
Bréda, France)
Impact of the 2003 heat wave on Douglas-fir in France: comparison of dead and
surviving trees for juvenile and mature traits (Manuela Ruiz-Diaz, Sara Marin,
Alejandro Martinez-Meier, Leopoldo Sanchez, Guillermina Dalla-Salda, Philippe
Rozenberg, France)
Storm damage of Douglas-fir and Norway spruce in Southwest Germany: Stability
of Douglas-fir and the impact of silviculture on the vulnerability of conifers (Axel
Albrecht, Ulrich Kohnle, Marc Hanewinkel, Jürgen Bauhus, Germany)
Coffee break
SESSION 5 (Moderator: Philippe Rozenberg)
Interactions between Douglas-fir and European beech – what do growth parameters
indicate? (Martin Haßdenteufel, Rebekka Bögelein , Willy Werner, Frank M.
Thomas, Germany)
Growth, allocation of space and competition in mixed stands of Douglas-fir and
European beech (Lutz Hilbrig, Germany)
Douglas-fir growth response to variation in soil water holding capacity and
precipitation patterns in the Siskiyou Mountains of southwestern Oregon (Douglas A.
Maguire, David W. Hann, Douglas B. Mainwaring, USA)
Lunch
SESSION 6 (Moderator: Ulrich Kohnle)
Keynote speech
Douglas-fir - a look into the genetics (Monika Konnert, Germany)
Adaptation of Douglas-fir provenances to drought stress (Ingo Ensminger, Moritz
Heß, Thomas Müller, Henning Wildhagen, Karl Schmid, Germany)
Genetic value of a Douglas-fir natural regeneration in France (Alain Valadon, Ouzna
Boussaid, Jean-Charles Bastien, France)
Photoprotective isoprenoids as physiological markers for the adaption of
Douglas-fir to drought stress in context of climate change (Laura Junker, Henning
Wildhagen, Ingo Ensminger, Anita Rott, Jürgen Kreuzwieser, Germany)
Coffee break
Schedule
15:55
16:20
16:45
17:10
17:35
18:0018:05
18:0518:45
VII
SESSION 7 (Moderator: Ulrich Kohnle)
Genetic investigation on Douglas-fir seed stands by means of isozyme and
microsatellite markers (Barbara Fussi, Monika Konnert, Germany)
Ecophysiological isotope tools for characterising the drought sensitivity of Douglasfir (Pseudotsuga menziesii (Mirb.) Franco) (Jakob Sohrt, Kirstin Jansen, Arthur
Gessler, Germany)
Planted forests of Douglas-fir in New Zealand: challenges for the breeding
programme with climate change (Heidi Dungey, Charlie Low, Michal Watt, Ian
Hood, Jeff Stone, Mark Kimberley, New Zealand)
Root system response of naturally regenerated Douglas-fir (Pseudotsuga menziesii)
after complete overstory removal (Nathan Briggs, Christian Kühne, Jürgen Bauhus,
Germany)
Summary
Heinrich Spiecker, Germany
Break
POSTER SESSION (continued)
20:00
Joint Dinner
Time
08:0018:00
Wednesday, October 20, 2010
Fieldtrip to Kandern, Black Forest
Departure: Forest Research Institute, Wonnhaldestr. 4
PREFACE
Douglas-fir’s importance for forest management is not restricted to its sweeping natural North
American range, where Douglas-fir is among the most important tree species. The species has
been successfully introduced in many forests around the world including Europe and New
Zealand. Since the introduction of the first “exotic” Douglas-fir individuals in Europe in the
19th century, the species has received increasing interest as a forest tree. It has rapidly
developed into the most widespread non-indigenous tree species of major economic
importance in Germany and France. The reasons behind this amazing track record are the
species’ tremendous growth potential, its valuable timber, and robustness.
In spite of the species’ potential for timber production, (large-scale) cultivation of Douglas-fir
outside its natural range is subject to controversial discussion. In general, the introduction of
non-indigenous species is considered to be fraught with a variety of considerable risks.
Among these are for example concerns about detrimental effects on native ecosystems and
their biodiversity, or unpredictable problems with respect to pests and diseases. For example
in Germany, these concerns have resulted in rather severe restrictions on Douglas-fir
cultivation in the context of close-to-nature silviculture systems.
However, in the context of climate change Douglas-fir is again receiving increased attention
in Germany. Quite a number of tree species currently present in Central Europe display the
potential to cope successfully with increasingly warmer and drier climatic conditions. Among
these species, the economic potential of Douglas-fir is unrivalled. This renders the species a
major issue in the current discussions about forest management strategies mitigating climate
change effects.
This conference is intended to provide a platform for the exchange of state-of-the-art
knowledge on Douglas-fir in particular dealing with opportunities and risks related to
European climate change issues. This includes the species’ potential for adaptation, growth,
biotic disturbance factors, and effective silvicultural management strategies.
The conference proceedings consist of six keynote speech abstracts, 22 further abstracts of
oral presentations and 11 poster abstracts. The conference program is also included.
Freiburg, October 2010
Ulrich Kohnle
Kaisu Makkonen-Spiecker
Heinrich Spiecker
Konstantin von Teuffel
Keynote 1
IX
KEYNOTE 1
The dynamics of Douglas-fir stands
Bruce Larson
Forest Resources Management Department
University of British Columbia, Canada
Douglas-fir (Pseudotsuga menziesii) is one of the most important commercial tree species in
the world and has an extensive range in western North America. It is fairly long lived because
it has silvic characteristics that allow survival in areas that have an intense disturbance regime
of fire and wind. The disturbance regimes and growing characteristics of the coastal areas are
quite different from the more interior parts of North America and the silvic characteristics of
Douglas-fir in these two areas are sufficiently different enough that two different varieties of
the species are recognized. This paper will focus on the coastal variety (var. menziesii),
because the growing conditions of the coastal areas are more similar to those of Germany.
This paper will also be devoted to stand development pathways of stands less than 125 years.
Douglas-fir grows in a wide range of climatic conditions. The coastal region of the Pacific
Northwest has a maritime climate characterized by mild, wet winters, dry summers and a long
frost-free season. Precipitation, mostly as rain, is concentrated in the winter months. Douglasfir grows in mountainous areas (e.g. the Cascade Range and Sierra Nevada), where there is
more snow, but Douglas-fir is not found at the higher elevations with short frost-free periods.
Pure Douglas-fir stands are naturally found in the mid to drier areas of the rainfall range, but
pure plantations can be maintained throughout the rainfall range. In wetter areas, stands have
a large component of western hemlock (Tsuga heterohylla), and in the driest part of the range,
the stands are mixed with a variety of different conifer species depending on the latitude.
Douglas-fir is quite shade intolerant relative to the tree species that grow in association. This
leads to the assumption that Douglas-fir needs very large openings to regenerate. Historically,
it is found in areas naturally regenerated after widespread fires. One must be careful to
separate the autecological characteristics from the effects of competition. Growth in low light
levels can be very slow resulting in mortality from competition with both non-tree and other
tree species. Also, as the seedlings get larger the respiratory load may exceed photosynthetic
production. There is evidence that the interior variety of Douglas-fir is more tolerant of shade.
The degree of light interception by Douglas-fir crowns in the overstory is highly variable
depending on the age and condition of the trees leading to difficulty in establishing
management guidelines for regeneration of Douglas-fir in partially shaded conditions.
Other than the usual plethora of insect and disease agents, the main disturbances in Douglasfir stands are wind and fire. Douglas-fir stands are often referred to as ‘adapted to fire’. This is
primarily in reference to the thick bark characteristic that allows individual Douglas-fir to
survive low and medium intensity fires that are primarily ground fires. Thinner barked
species, such as western hemlock or grand fir (Abies grandis) are usually killed in these types
of fires. There are many examples of scattered large, old, Douglas-fir trees in stands 80 to 200
year old that naturally regenerated after stand replacing fires. These remnant trees survived
the fire and grow rapidly in the open conditions
Douglas-fir trees are quite resistant to windthrow compared to associated species. However,
in mixed species stands, if other species such as western hemlock are broken by the wind, the
Douglas-fir will be broken or uproot in subsequent gusts causing the stands to ‘unravel’. In
X
Germany, Douglas-fir mixed with beech may make very windfirm stands. In pure stands, the
height growth rate of Douglas-fir is often so great that if slender trees are widely spaced in a
thinning, the increased diameter growth is not sufficient to make the trees tapered enough to
be windfirm.
In mixed species stands, Douglas-fir is usually the tallest stratum and shade tolerant hemlock
and red cedar form the lower layers. Douglas-fir will make up the largest diameters of these
single cohort stands, which may have a ‘reverse J-shaped’ diameter distribution because of
the large number of smaller diameter hemlock. These layered stands have a diverse structure
providing habitat for diverse wildlife species. These irregular looking stands may have a
similar appearance to uneven age stands making them popular with hikers and walkers.
From a timber standpoint, Douglas-fir stands are very productive over longer periods.
Douglas-fir’s prolonged height growth rate means that even on average sites on Vancouver
Island MAI will not culminate until year 70 or 80. Stands can be grown at a variety of
spacings depending on the products desired. On many sites, Douglas-fir responds positively to
thinnings even at quite old ages depending on crown morphology.
The ability of Douglas-fir to survive and grow well over a broad range of conditions has led
to good growth in many different parts of the world, where it has been introduced; however
proper selection of provenance is important. Douglas-fir can survive windstorms better than
many other species and grows in fire prone regions, which may become more widespread in
various scenarios of climate change.
Keynote 2
XI
KEYNOTE 2
Douglas-fir in France: history, recent economic development,
overviews for the future
Jean Louis Ferron
France Douglas
Introduction
Two centuries ago, only the Western Coastal North America stands of Douglas-fir were
known all around the world. They covered a narrow stripe of more than 1.500 km from
British Colombia (North of Vancouver Island) to California, and from the Pacific Ocean to
the Cascade up to an elevation of 1.500 m. Douglas-fir was first described by Archibald
Menzies at the end of the 18th century. Thirty years later, a Scottish botanist and explorer
David Douglas wrote notes about the impressive development of theses stands to awaken the
interest of European scientists and politicians. The first seeds were exported to Europe – first
to England, then to France and Germany. The first tests of introduction - in parks and gardens
- occurred at the turn of the middle of the 19th century.
Reforestation
After this period, foresters started to get an interest in Douglas-fir, and engaged the first
programs of reforestation, in particular in France and Germany, between the years 1870 –
1914. After the First World War, however, the interest for Douglas-fir entered in sleep, in
France until 1960-1970. At that time with the support of the government policy, forests
owners were strongly stimulated to reforest abandoned agricultural lands and coppices.
Stimulated by the impressive results of the first stage of reforestation (second part of the 19th
century), Douglas-fir had gradually become the main species used in reforestation programs,
especially in the “Massif Central”, area located in center of France, characteristics of which
are the following ones: elevation between 600 et 1.200 m; rainfall more than 1.000 mm per
year; soils acid brown on granite source rocks.
XII
Today, Douglas-fir stands cover about 400.000 ha in France. They represent a standing
volume of 90 million m3 and an annual increment of 6.7 million m3 per year (16.7 m3 /ha /
year). The average age of Douglas-fir resources in France is comprised between 30 and 45
years. Harvesting has been increasing strongly during the last 15 years, being presently 2
million m3. Timber production has been following the same way and reaches today 800.000
m3, vs. 400.000 m3 at the end of the 1990’s.
volume (m3)
Evolution of sawn timber production between 1990 and
2008
900 000
800 000
700 000
600 000
500 000
400 000
300 000
200 000
100 000
0
1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 200 2 2003 2004 2005 2006 2007 2008
Evolution and trend of the french production of
Douglas-fir sawn timber in the future
volume (m3)
3 000 000
2 500 000
2 000 000
1 500 000
1 000 000
500 000
0
90
93
96
99
02
05
08
11
14
17
20
23
26
29
Overviews for the future
The French Douglas-fir production is now entering in a transition period. If the previous one
has been characterized by a production mainly oriented towards export, it is now necessary, in
comparison with the potential quantities concerned (3 million m3 per year for 20 years), to
develop also the domestic market. This will be the main task of the France Douglas
Organization for the next years. Therefore, there is a study with the aim to define and qualify
some specific Douglas-fir products well adapted to the construction market. This study
includes the analysis of the characteristics of the current supply, as well as of present and
future requirements of the construction market, in relationship with regulation and
standardization.
Keynote 2
XIII
This analysis will enable France Douglas’s members to define strategic choices about the
most relevant construction materials in relationship with the Douglas-fir's wood properties.
After that, it will be necessary to qualify all concerned products and to promote them. The
achievement of this strategic study is planned within two years.
Since 1993, France Douglas gathers producers, sawmill owners and other industrials, with
national federations, research centers in view to promote Douglas-fir production. France
Douglas is particularly involved in standardization, research; communication and assistance
of sawmill owners and industrials to develop Douglas-fir production.
To reach its target, France Douglas may rely on several advantages; in particular the context
(for instance increased demand of wood products by the construction industry; increasing
Douglas-fir resources; Douglas-fir wood properties), and the organization of Douglas-fir
production around France Douglas.
The climate change enforced by the high level of the industrial’s demand of Douglas-fir has
two main consequences: (1) some Douglas-fir stands are harvested too early compared to
their optimum maturity, and (2) the level of reforestation has strongly declined over the last
two decades.
25 000 000
20 000 000
15 000 000
10 000 000
5 000 000
0
19801981
19831984
19861987
19891990
19921993
19951996
19981999
20012002
20042005
20072008
Number of Douglas-fir seedlings sold in France between 1980 and 2008
To answer this challenge, there have being engaged studies on reforestation techniques,
including genetic improvement, in correlation with climate change and industrial requirement,
as well as on methods of silviculture, in correlation with the requirements of the sawmillers
and the results of the ongoing strategic analysis.
The results of these studies are expected to provide Douglas-fir producers with tools to boost
reforestation operations and promote silvicultural schemes adapted to the needs of the
industry.
For France, Douglas-fir production is a great opportunity to correct the national trade balance
that yearly imports more than 3 million m3 of coniferous timbers. So, Douglas-fir will be the
most important national emerging forest resource for the next decades.
Opportunities and risks for Douglas-fir in a changing climate
Session 1
1
Session 1
2
Mapping Douglas-fir current and future abundance and potential
site productivity in western United States
Aaron R. Weiskittel1 Nicholas L. Crookston2, Gerald E. Rehfeldt2, and Phillip J. Radtke3
1University of Maine, School of Forest Resources, Orono, ME
2USDA Forest Service, Rocky Mountain Research Station, Moscow, ID
3 Virginia Tech, Department of Forestry, Blacksburg, VA
Long-term forest planning requires knowledge of future species abundance and potential site
productivity in order to best match species planted to sites that are favorable for their growth
and survival. Douglas-fir is one of the primary commercial species throughout western United
States, but projected climate change is expected to significantly alter its abundance and
potential site productivity. The current distribution of Douglas-fir in western United States is
primarily driven by the amount of growing season precipitation as well as the interaction
between annual total precipitation and growing degree days. Based on projected climate
change scenarios, there is little anticipated change in the acreage potentially suitable for
Douglas-fir to grow by 2090, but the geographic range of its habitat may shift from the
Oregon and Washington Coast Range to higher elevations in the Cascades. Work is currently
focusing on matching the best Douglas-fir seed source to sites based on their expected future
climate.
In terms of site productivity, Douglas-fir productivity is driven primarily by the ratio of
annual moisture index to growing degree days and precipitation during the growing season to
the average difference between monthly maximum and minimum temperature. These
variables are able to explain over 72% of the original variation in Douglas-fir site index,
despite the wide range of conditions covered by the data. The predominant trend for future
Douglas-fir habitat is an increase in site index of 0 to 5 m by 2030. However, an increasing
proportion of the current prime habitat for the species will experience reductions in site index
of 10 to 15 m in 2060 and 2090. Increases in site index are more likely in the Idaho Central
Rockies and western Montana, while the reductions should occur in the Coast Range and
Cascades of Oregon and Washington.
This talk will explore the implications of this work for forest management activities, ongoing
research on developing growth and yield models that utilize these findings, and the potential
to conduct similar research in Germany.
Session 1
3
Going the distance: Testing seed transfer of coast Douglas-fir
within its native range
Peter Gould1, Constance Harrington1, and Brad St. Clair2
1
Pacific Northwest Research Station. Olympia, WA. USA.
2
Pacific Northwest Research Station. Corvallis, OR. USA.
Climate change in this century is predicted to increase temperatures and change the quantity
and distribution of precipitation within the native range of coast Douglas-fir. Despite these
changes, much of the species’ current range is expected to continue to be suitable for
Douglas-fir and its potential range will likely expand. However, Douglas-fir is a genetic
specialist and populations appear to be highly adapted to their local environment. A large
body of research has shown that differences in adaptive traits such as the timing of budburst
and budset occur over relatively short geographical distances and with small changes in the
local environment. Thus, local populations are at risk of maladaptation even where the
climate remains suitable for the species as a whole. The key challenge is to determine which
genotypes will be best adapted to the future climate. Long-term results from seed-transfer
studies are available for some species, but there is relatively little information for coast
Douglas-fir.
We initiated the Douglas-fir Seed-Source Movement Trial in 2008 to test seed transfers
among environments. The trial consists of nine test plantations in western Oregon and
Washington that contain trees from 120 half-sib families from California, Oregon, and
Washington. The trial is being intensively measured for traits including the timing of
budburst and budset, seasonal growth patterns, and annual growth increments. Automated
weather stations and other monitoring equipments have been installed at each site to collect
highly detailed information on the local environments. Long-term results from the study can
be used to help develop transfer functions to predict changes in productivity with changes in
genotype and local environment.
Short term results are also valuable particularly when viewed in mechanistic terms (i.e., how
genetics and the environment interact to influence productivity). For example, results from
an initial experiment with the plant material were used to develop a model to predict the
timing of budburst based on winter temperature and genetic variation in the chilling
requirement. The model indicates that genetic differences are important, but populations can
exhibit a considerable range in phenotypic plasticity depending on the winter environment.
This model and other short-term results can be used to develop a more complete
understanding of how specific traits are influenced by the environment and how climate
change will affect forest productivity.
4
Sustainable cultivation of Douglas-fir on dry and acid sandy soils in the
old-glacial lowlands of eastern Germany
- aspects of nutrient and humus dynamics
Thomas König, Alexander Tischer, Franz Makeschin, Sven Martens
Knowledge about site requirements of Douglas-fir in Germany on particularly acidic sandy
soils in regions of sub-continental climate is based on the experience of local foresters
mostly. Detailed research to determine limiting conditions is rare yet. Moreover, predicted
scenarios of climate change require tree species, which are suitable as silvicultural and
ecological alternatives in flexible management strategies.
A chronosequence of Douglas-fir stands (aged 12, 25, 34, 35, 63, 74 yrs.) was analysed. For
each stand, an adjacent stand of Scots pine (aged 80 to 120 yrs.) was used as reference. Later
on, the comparison was expanded by two stands of Sessile oak (aged 78, 136 yrs.), one of
Red oak (54 yrs.) and an additional one of Scots pine (106 yrs.). All test sites are
characterised as sandy soils (silt + clay 10 to 20 %) with annual precipitation and annual
mean temperature between 540-600 mm and 8.0 °C, respectively.
The authors predict an intact nutrient-cycle in older Douglas-fir stands as an indicator for the
suitability of Douglas-fir (as a climate-plastic tree) for cultivation in other parts of Germany,
where conditions like those on the investigated sites are likely occur in future times. The
suitability of cultivating Douglas-fir as economic alternative to Scots pine is of more current
interest in the investigation area. These investigations were focused on nutrient and humus
dynamics as important parameters to assess the status of soil fertility. Carbon and nitrogen
storage, pH, NH4Cl-extractable cations, Citrate-extractable Phosphorus, nutrient
concentrations in living and dead foliage, litter-fall and litter-decomposition as well as basalrespiration and concentration of microbial biomass were examined as important factors in
mineral soil and organic layer.
The authors conclude that even on the test sites, Douglas-fir is able to utilize the site potential
more efficiently than Scots pine. Neither negative effects on soil fertility nor the necessity for
compensation by fertilization were observed. Positive effects on nutrient status and organic
matter dynamic were recognised. For example enhanced pH and NH4Cl-extractable cations,
increased microbial biomass and basal-respiration in the uppermost part of the humus layer in
the oldest stands of Douglas-fir and Sessile oak were detected. Generally, the quality of
humus layer increased in the order Scots pine < Red oak < Douglas-fir ≤ Sessile oak.
Session 2
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Session 2
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KEYNOTE 3
Douglas-fir and the management of forests as complex, adaptive systems
Klaus J. Puettmann
Department of Forest Ecosystems and Society
Oregon State University
Global change consists of a variety of perturbations, including climate change, invasion of
exotic species, and atmospheric inputs. Accordingly, responses of forest managers to these
changes need to utilize a variety of approaches. To assess these approaches it is helpful to
group responses into practices that increase the resistance, resilience, and the adaptability of
forest ecosystems.
Treatments that increase the resistance of forests may provide the most benefit in the short
term, such as vegetation control practices that reduce impacts of invasive exotic species and
thinning to decrease competitive stresses and increase tree stability. Such practices can also
increase the resilience (ecological resilience, sensu Holling 1996) of forest ecosystems, if
they are specifically aimed to facilitate the forests’ ability to recover quickly after
disturbances. For example, the establishment of advanced regeneration will aid the forest’s
ability to continue carbon sequestration after a windstorm. Similarly vegetation control or
fertilization treatments may allow for crop tree establishment and growth when growing
conditions have been changed to a degree that a site is no longer favorable for tree growth.
However, all these practices will increase the cost and short-term environmental impacts of
forest management. In addition, they may have only limited value, if climate and other
ecological and environmental conditions continue to change in the future.
Shifting the emphasis to the ability of ecosystems to adapt to changing conditions by
managing forests as complex adaptive systems (CAS) has been suggested as a framework
that is especially useful in a changing and uncertain future (Puettmann et al. 2009). This
approach utilizes recent advancements in the complex systems and ecosystem sciences
(Levin 1998, 2005, Sole and Bascompte 2006, Norberg and Cummings 2008). Forest
ecosystems are a prime example of CASs and much can be learned from utilizing this
framework to manage forests for adaptability in light of climate change.
Complex adaptive systems consist of a diversity of agents that interact at local levels and
across a variety of hierarchical scales (Figure 1). These interactions include non-linear
relationships and positive and negative feedback loops, which at higher scales lead to
emergent properties; i.e., properties that cannot be predicted from information about the
individual components. An important feature of CAS is the bottom-up, decentralized control;
i.e., the multitude of agents and interactions at low-level, local scales are the drivers of
ecosystem behavior and thus critical for the ecosystem’s ability to adapt to changing
conditions. This implies that ecosystem responses to perturbations are determined by the
rules determining these local interactions and management practices have to be viewed in a
context, whether they can influence these rules and thus increase or decrease the adaptability
of ecosystems.
However, CAS theory and concepts are not yet developed to the stage, where they can be
directly applied to forestry settings. For one, because of the unpredictability of CAS behavior,
rather than focusing on minimizing uncertainty of future predictions, it may be more useful to
Session 2
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acknowledge the inherent uncertainty and developing plans for a range of possible future
scenarios (Biggs et al. 2010). While our understanding of the impact of forest management
practices on adaptability of forests is just developing, an emphasis on diversity and
redundancy at various spatial and temporal scales is likely a good starting point for
silvicultural prescriptions.
The presentation highlights these concepts using Douglas-fir forests as examples of CASs. A
special emphasis is placed on providing practical examples that link these conceptual ideas to
practical measures and practices. These include the impacts of thinning on ecosystem
resilience and “assisted migration” as a tool to increase the adaptability of forests to global
changes.
Figure 1: Simplified view of forest ecosystem as complex adaptive systems. The shaded box
highlights “lower level” components and interactions. These relationships organize
into emergent properties that interact through non-linear feedback loops to drive
system behavior.
References
BIGGS, R., M.W. DIEBEL, D. GILROY, A.M. KAMARAINEN, M.S. KORNIS, N.D. PRESTON, J. E. SCHMITZ, C. K. UEJIO, M.C.
VAN DE BOGERT, B.C. WEIDEL, P.C. WEST, D. P.M. ZAKS, AND S.R. CARPENTER. 2010. Preparing for the future: teaching
scenario planning at the graduate level. Frontiers in Ecology and the Environment 8:267-273.
HOLLING, C.S. 1996. Engineering resilience versus ecological resilience. P. 31-44 in Engineering within ecological
constraints, P. C. Schulze (ed). National Academy Press, Washington, DC.
LEVIN, S.A. 1998. Ecosystems and the biosphere as complex adaptive systems. Ecosystems 1:431-436.
LEVIN, S. A. 2005. Self-organization and the emergence of complexity in ecological systems. Bioscience 55:1075-1079.
NORBERG, J., AND G.S. CUMMING (eds.). 2008. Complexity Theory for a Sustainable Future. Columbia University Press,
New York.
PUETTMANN, K.J., K.D. COATES, AND C. MESSIER. 2009. A Critique of Silviculture: Managing for complexity. Island Press,
Washington,
8
SOLE, R. V. AND J. BASCOMPTE (2006). Complexity in Ecological Systems. Self-Organization in Complex Ecosystems.
Princeton University Press.
Session 2
9
Climatic and regional patterns in Douglas-fir climate-growth relationships
in British Columbia, Canada
Hardy P. Griesbauer, D. Scott Green1
University of Northern British Columbia
Prince George, BC, BC, Canada
Douglas-fir has been successfully introduced to many countries outside its natural North
American range and is now considered a tree species of global economic and ecological
importance. However, the continued successful management of Douglas-fir in introduced
areas such as Europe, New Zealand, and South America will require a better understanding of
its sensitivity to climate change as well as its capacity to adapt to new climatic conditions. To
address this knowledge gap, the authors present a novel approach using high-resolution
spatiotemporal climate data along with traditional dendroecological analyses to identify
spatial and climatic patterns in climate-growth relationships across the species range in the
Interior region of British Columbia.
Using this approach, the authors were able to identify a strong link between local climate
conditions and population sensitivity to climatic variability across a broad range of ecological
conditions. It was found that temporal Douglas-fir productivity is coherent over large regions,
and that local climatic and ecological conditions largely explain these spatial patterns. For
example, productivity in warm and dry low-elevations populations is primarily influenced by
annual precipitation, whereas growth in wetter and cooler high-elevation populations is more
limited by snowfall and cold temperatures. Perhaps most importantly, the results show that
while Douglas-fir productivity across most of its range is primarily sensitive to annual
precipitation, this sensitivity increases exponentially as sites become drier. Thus, while many
populations may become maladapted as a result of increased temperatures and their effects on
soil moisture, certain populations will likely express this maladaptation sooner or to a higher
degree than other populations, and this phenomenon may be related to strong adaptation to
local average climatic conditions.
These studies provide important information to manage Douglas-fir in Europe. The results
identify the key climate variables that need to be considered when regenerating and managing
Douglas-fir in this region. Because these results indicate that local climatic conditions can
predict population sensitivity to climate change, forest managers can use local climate data
for their region to ensure that Douglas-fir provenances are well suited to future climates. In
particular, these relationships can be used to highlight regions, where Douglas-fir may be
well buffered against climate change, or where local climates may become too warm and/or
dry for Douglas-fir to be productive. Furthermore, the authors’ approach allows forest
managers to use local climate conditions and geographical information systems to spatially
visualize sensitive in situ Douglas-fir populations in Europe that may be of management
concern.
10
Douglas-fir – a substitute species for Scots pine in dry inner-Alpine valleys?
Britta Eilmann, Andreas Rigling
WSL, Swiss Federal Research Institute
Birmensdorf, Switzerland
During centuries Scots pine (Pinus sylvestis L.) dominated the dry forest in Valais and other
inner-Alpine valleys. But since the 1990ies, the landscape is changing. Scots pine showed
increasing mortality rates, especially on dry sites and after drought years. Concurrently
pubescent oak is spreading. Therefore, pubescent oak was seen as number one candidate to
replace Scots pine under increasing drought and therefore as a grantee that the dry forests in
the inner-Alps can still provide ecosystem goods and services (esp. protection against natural
hazards) in the future. However, recent studies showed that also pubescent oak will soon
reach its physiological limits, if the frequency and severity of drought increases.
Unfortunately, no other native species with the potential to replace Scots pine and pubescent
oak under drier climatic conditions is available. Hence the forests on low elevation in the
Valais might be jeopardized. Consequently, non-native species need to be considered as
substitute species on the driest sites. Therefore, the authors analyze how non-native species
cope with the environmental conditions in Valais compared to Scots pine. This was done by
comparing the growth response to and the recovery after drought years in native - Scots pine
and European larch (Larix decidua Mill.) - and non-native species - Douglas-fir (Pseudotsuga
menziesii (Mirb.) Franco) and Black pine (Pinus nigra Arn.) in a 40 year old afforestation.
All species showed a growth depression in drought years. However, further analyses indicate
a clear advantage of non-native species over native species under dry conditions. Douglas-fir
showed the highest increment of all species studied. But the recovery after drought years was
delayed in Douglas-fir. In contrast, Black pine showed a lower level of increment, but at the
same time the fastest recovery after drought years. So, the question arise, if high increment
and a big crown system of Douglas-fir is really favourable in terms of sustainable growth in a
dry environment. However, black pine and Douglas-fir seemed to have a higher potential to
resist drier conditions and, hence, might be an alternative in future drought-prone protection
forests. This picture was supported by the results of mortality analysis showing no mortality
in black pine and Douglas-fir, but high rates of crown-dieback and mortality in Scots pine
and larch.
Session 2
11
Impact of Douglas-fir on the N cycle:
Douglas-fir promotes nitrification?
Bernd Zeller , Sitraka Andrianarisoa,
Jean-Huques Jussy,, Etienne Dambrine, Jacques Ranger
INRA Nancy, Biogeochemistry in Forest Ecosystems
Champenoux
Two Douglas-fir forests, Vauxrenard, (20, 40, 60-year-old stands) and Breuil (30-year-old
stand) in central France, had been intensively monitored over almost a decade. Both sites are
characterized by deep soils, with low pH, low base saturation and low contents of Ca and
Mg. The humus layer is thin and understory is only present in the oldest stand. The chemical
composition of the throughfall solution and the soil solution at different depths was
determined at monthly intervals. At Vauxrenard, net mineralization and net nitrification had
been monitored over 5 years. At Breuil, potential net and gross mineralization and
nitrification had been determined in the A horizon of a 30-year-old stand.
Nitrogen mineralization rate (net and gross) was high and mineralized nitrogen was mainly
nitrified. Root uptake of nitrogen was high but lower than mineralization. As the production
of nitrate exceeded the microbial immobilisation and root uptake, leaching losses of nitrate
were always high and especially in young stands. Additionally, former land use (arable fields
or meadows) boosted nitrate leaching. In addition, at Breuil where a common garden
experiment had been set up, Douglas-fir, beech, oak and Corsican Pine promote nitrification,
whereas spruce and Nordmann-fir reduce nitrification. Nitrification was positively related to
the number of nitrate oxidising bacteria (NOB). Nitrate concentration is negatively correlated
with the number of fungal species.
If soils from spruce and Nordmann-fir were transferred into the Douglas-fir stand, then the
nitrification increased within 18 months. The soil core exchange experiment showed that root
biomass and root colonization were species specific and were not simply dependant on the
NO3 availability in the soil. In contrast, root colonization was negatively related to soil
percent nitrification regardless of the tree species. In high nitrifying soils transferred into low
nitrifying stands, soil percent nitrification decreased gradually as root colonization increased.
In low nitrifying soils transferred into high nitrifying stands, soil percent nitrification
increased as root colonization decreased. Root biomass and root colonization are mainly
related to tree species, while soil nitrification appears strongly influenced by root
colonization. Root colonization may be a common strategy of plant control over nitrification.
The authors showed that on acid soils, Douglas-fir increases nitrification rate and
subsequently leaching of nitrate and cations further resulting in an acidification of such
forests. Past land occupation and site fertility level influence the decoupling between nitrate
production and consumption by biota.
References
ANDRIANARISOA S., ZELLER B., POLY F., SIEGENFUHR H., BIENAIMÉ S., RANGER J., DAMBRINE E. (2010): Dynamic control of
nitrification by tree species (Ecosystems, in press)
12
BUÉE M., MAURICE, J-P., ZELLER, B., ANDRIANARISOA, S., RANGER, J., COURTECUISSE, R., MARÇAIS, B., LE TACON, F.
(2010): Influence of tree species on richness and diversity of epigeous fungal communities in a French temperate forest stand
(Fungal Ecology, in press)
JUSSY, J.H., COLIN-BELGRAND, M., RANGER, J. (2000): Production and root uptake of mineral nitrogen in a chronosequence
of Douglas-fir (Pseudotsuga menziesii) in the Beaujolais Mounts. For. Ecol. Manage., 128, 3 : 197-209.
JUSSY, J.H., KOERNER, W., DAMBRINE, E., DUPOUEY, J-L., BENOIT, M. (2002): Influence of former agricultural land use on
net nitrate production in forest soils. Eur. J. Soil Sci., 53, 3 : 367-374.
JUSSY J.H., COLIN-BELGRAND M., E. DAMBRINE. RANGER J., ZELLER B., BIENAIME S. (2004): N deposition, N transformation
and N leaching in acid forest soils. Biogeochemistry 69: 241-262
TRUM, F., (2004): Impact de l'essence sur la composition chimique des litières et de leurs percolats en conditions contrôlées.
In. Université Catholique de Louvain, Louvain-la-Neuve, p. 143.
ZELLER B., RECOUS S., KUNZE M., MOUKOUMI J., COLIN-BELGRAND M., BIENAIME S., RANGER J., DAMBRINE E. (2007):
Influence of tree species on gross and net N transformations in forest soils. Ann. For. Sci. 64: 151-158
Session 3
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Session 3
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KEYNOTE 4
Tree-ring analysis of adaptation to drought in Douglas-fir
P. Rozenberg1, L. Sanchez1, G. Dalla-Salda2, A. Martinez-Meier2,
A.S. Sergent1,4, M. Ruiz-Diaz1,3, S. Marin1, J. C. Bastien1
1
National Institute for Agronomical Research, Orléans, France
2
National Institute of Agronomical Tecnology, Bariloche, Patagonia, Argentina
3
Parque Tecnologico Misiones, Posadas, Misiones, Argentina
4
National Institute for Agronomical Research, Nancy, France
Adaptation is a process whereby an organism becomes better suited to its habitat. Considering
local adaptation, evolutionary adaptation and phenotypic plasticity are the two mechanisms
involved. Phenotypic plasticity is especially important for fixed and long-living organisms like
forest trees, above all when the environmental shift is shorter than one generation time. The ongoing climate change is characterized by a rapid temperature increase. A consequence of this
temperature increase is an augmentation of the atmospheric water demand during the growing
season. This augmentation may push forest trees beyond their adaptation limits, especially for
species with a high water needs growing in environments with already restricting water
conditions. This is the case for Douglas-fir in some regions of France: decline pictured by
abnormal ecophysiological reactions was reported in Douglas-fir, above all after the 2003 heat
wave, in different parts of France with symptoms ranging from needle coloration to individual
death.
The authors have shown that this decline was distributed among three distinct geographical
zones. In Bourgogne and in Massif-Central, decline started at the beginning of the 2000’s, with
2003 identified as a key-year in its apparition and in its development. However, in NorthWestern France, the decline was found to start as early as at the beginning of the 90’s, with a
several decline events succeeding up to day. Soil water deficit seems to be the main cause of
the most recent declines but not of the previous ones.
Because direct observation of tree reaction at the time of an unexpected climate event is
generally impossible, the authors used tree-rings as proxies for retrospective studies of tree
response to the 2003 heat wave. One of the important functions of wood is sap conduction.
Hydraulics of sap conduction is linked to duct internal dimension, itself interrelated to wood
anatomy and wood density. The authors have studied several components of Douglas-fir
reaction to the 2003 heat-wave. It was shown that the ring formed in 2003 is significantly
narrower than the previous and the following one, with a smaller, lower density latewood. It
was also shown that the response to the 2003 heat wave was significantly different among
clones and inherited. The authors demonstrated that dead and surviving trees after 2003 were
significantly different for traits measured in several rings formed prior to the heat wave. The
surviving trees had a higher proportion of higher density latewood. More recently, the authors
deepened the analysis of the difference between the dead and the surviving trees and confirmed
that the critical part of the ring was in the latewood. Above all, the wood trait better explaining
the difference between dead and surviving trees was found to be the quantity of high density
latewood cells.
Wood hydraulic properties involved in sap conduction could explain the relationship between
wood density and adaptation to drought. It was shown that on one hand, specific conductivity
and vulnerability to cavitation were related to ring-density traits, and that on the other hand,
Session 3
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these hydraulic properties were significantly different among Douglas-fir clones as well as
inherited. Complex relationships have been established among different parameters of
vulnerability to cavitation curves and several earlywood and latewood traits. According to
these results, the first steps of the cavitation process are related to latewood traits, while the
later steps are more related to earlywood traits.
The authors have also proved that within-ring variation of wood density (from earlywood to
latewood) is an expression of phenotypic plasticity: the relationship between within-ring
density and within-growing season weather variation is a norm of reaction. Douglas-fir trees
have the ability to form latewood: they radically decrease their tracheid lumen dimension as a
response to drought increase. Latewood formation is a fast adjustment to the conditions of the
second part of the growing season. It is a plastic reaction to the diminution of the water
availability and the increase of the water demand. It was also shown that this rapid response is
genetically variable and inherited and thus that it can be selected under the pressure of extreme
climate events. Among the wood traits exhibiting significant differences between the dead and
surviving trees after the 2003 heat wave, the authors have found contrasted result: for example,
the mean density of the highest density part of the ring (similar to latewood density) has been
found to be highly and significantly variable among provenances of very distinct origins. Other
traits showing significant differences between dead and surviving trees, the area of the highestdensity segment of the ring, has been found to be not significantly different among the same
provenances. Finally, at least one trait showing no difference between the dead and surviving
trees, the mean density of the low density part of the ring (similar to earlywood density), has
been found to be extremely variable among the same set of provenances. These results draw a
complex image of the adaptive value of distinct traits related with hydraulic wood properties.
In forest trees, telling the difference between evolutionary adaptation and phenotypic plasticity
for wood adaptive traits related to sap conduction is somewhat tricky: a new ring is
permanently added year after year to the trunk; few years after its formation, every ring
becomes progressively non-conductive. Thus the conductive part of the tree is constantly
modified along with tree growth. It is continually and strongly shaped by internal and external
factors, namely genetics, age, silviculture and environment, whereby weather variation during
the growing season seems to play a key-role. Thus the conducting phenotype of a given
individual may be gradually adjusted over time to match new water conditions. Hence the final
success or failure of a given individual in a given tree population submitted to a climatic shift is
the result of a complex process dynamically translating its fundamental genetic potential into
an adaptive plastic response.
16
Tree-ring growth of West-German Douglas-fir
under changing climate conditions
S. Fischer, B. Neuwirth, J. Löffler & M. Winiger
Institute of Geography, University of Bonn
Bonn, Germany
In terms of a changing climate there are several investigations dealing with suitable forest
management strategies, especially concerning the fitness of different tree species under diverse
site ecological features. Douglas-fir, introduced in Europe in the first half of the 19th century, is
a foreign tree species, which is discussed to be a part of forest management strategies in line
with changing climate conditions. Dendroecology providing at least an annual, retrospective
resolution, is a successful tool to deliver a contribution to actual discussions.
In this study, radial growth behaviour of Douglas-fir on drier and moister sites will be
compared. The investigation is based on analyses of pointer years (after Cropper 1979), mean
annual growth and linear trends using raw data as well as index data for different time periods
since the 1940´s. Afterwards these findings will be compared with growth behaviour of local
tree species.
For this investigation, taking place in the low mountain ranges of W-Germany an already
existing network of local tree species, like Fagus sylvatica, Quercus petraea, Picea abies and
Pinus sylvestris will be completed with data from Pseudotsuga menziesii. More than 400 trees
from sites representing a various ecological spectrum will be analysed. Detailed site
information like soil, hydrological attributes or vegetation type is collected for forest ecological
test areas in line with the biomonitoring of the LANUV (Landesamt für Natur, Umwelt und
Verbraucherschutz). All investigated trees are in adult state.
The presentation shows growth behaviour of the considerably fast-growing Douglas-fir with
special regard to rising temperatures in the last decades for two regions, national park Eifel and
national park Kellerwald, both in the low mountain ranges of West Germany.
Session 3
17
Impact of water availability on wood density patterns
in Douglas-fir tree rings
Bela Johannes Bender, Heinrich Spiecker
Institute for Forest Growth, University of Freiburg
Drought induced sensitivity of Douglas-fir provenances is linked to the structure of latewood
cells (Martinez-Meier et al. 2008). Less sensitive provenances show higher annual maximum
densities. In general, plant species with higher wood densities are less susceptible to drought
damages (Hacke et al. 2001). Furthermore, water shortage influences variations in
microstructures of annual increment zones as evidenced by false rings. In this study, the
authors analyse the relation between spatial variation of wood density in radial direction and
the temporal variation of the water regime assuming that water availability is the main driving
factor. The results could be used to characterize in a quantitative way the ability of trees to
react on dry conditions.
The authors analysed the microdensity profiles of four 75 year old dominant trees along eight
radii of each individual applying the H-F-Density method (Schinker et al. 2003). For analysing
the relation between water availability and wood formation it was focused on exceptionally dry
and humid summer periods respectively. Variations in water availability are calculated by a
simple drought index and the precipitation sums.
The deduced method may be used for estimating the plasticity and drought tolerance of
Douglas-fir provenances. Furthermore, it could improve retrospective analyses of intra-annual
growth processes and our understanding of growth-environment relationships. Further analyses
on Douglas fir provenances are object of current investigations.
References
HACKE, U.G.; SPERRY, J.S.; POCKMAN, W.T.; DAVIS, S.D.; MCCULLOH, K.A. (2001): Trends in wood density and structure are
linked to prevention of xylem implosion by negative pressure. Oecologia. 126. pp 457-461.
MARTINEZ-MEIER, A.; SANCHEZ, L.; PASTORINO, M.; GALLO, L.; ROZENBERG, P. (2008): What is hot in tree rings? The wood
dendity of surviving Douglas firs to the 2003 drought and heat wave. Forest Ecology and Management. 256. pp 837-843.
SCHINKER, M.G., HANSEN, N., AND SPIECKER, H.( 2003): High-frequency densitometry - a new method for the rapid evaluation
of wood density variations. IAWA Journal vol. 24 (3). pp 231-139.
18
Findings from long-termed forest growth inventory data
from 14 trial sites in the II. International Douglas-fir progeny trials
established in 1961 in north-western Germany
Andreas Weller
North-western German Forest Research Institute
Growth (i. e. stand top height (h100), diameter of the top height tree (dbh100) and total volume
production), and quality characteristics of Douglas-fir were evaluated from 36 indigenous
provenances and two German sources up to the age of 38 years. A visual assessment of the
categories branchiness and tree form characteristics of the future crop trees was available for
describing the qualitative development of the trees tested. The influence of future crop tree size
was used as a covariate in addition to provenance effects to determine the quality characteristic
branchiness. Based on the quality score of the future crop trees, the multi-nominal logistic
regression model estimated the probabilities of provenance-specific tree stem form, and of
branch scores in relation to provenance and to defined diameters at breast height of the future
crop trees.
The main results:
a)
The two provenances recommended for planting in north-western Germany, Ashford
and Gold Hill, originate from the west slopes of the Cascade mountains in central region of
Washington state (seed lot zone 422) and the Darrington region in northern Washington
(seed lot zone 403) respectively. They revealed an above-average, and more reliable yield
than that found for all other trials as well as good, above-average genetic traits of small
branches and straight stems.
b)
The Canadian provenances, South Wellington and Cameron Lake from the East Coast
of Vancouver Island (seed lot zone 1020), should be excluded as they display both poor
production and poor quality for branchiness and stem form. Given its generally slow
growth, the Fraser River provenance from the central interior of British Columbia (seed lot
zone 5010) also should be excluded.
c)
Based on the results of this investigation, the seed collection sites along the coastal
region of North Oregon, seed lot zone 052, and at Puget Sound in Washington State, seed
lot zone 430, in USA are recommended as seed collection areas. Here, the region is divided
into broad landscape units with uniformly good production and, simultaneously, high
quality stock in the indigenous ecotypes occurring.
Likewise, the author responded to the question of climatic variation of sampled data as to total
volume production in cubic metres of stem wood per hectare up to the age of 38 years. In the
process the trial sites were characterized by the use of annual variables of precipitation (Fvalue 0.9599, p > 0.05), and air temperature (F-value 1.485, p< 0.05).
Session 4
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Session 4
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KEYNOTE 5
Review of present and potential insect pests affecting Douglas-fir in Europe
in a context of global change
Alain Roques
INRA, Zoologie Forestière, Orléans, France
After more than a century of extensive plantations in Europe, Douglas-fir has recruited a
limited number of insect species compared to the importance of the associated entomofauna in
the native range (> 250 spp.). Less than 100 insect species, mostly belonging to the orders
Coleoptera (45 spp.) and Lepidoptera (34 spp.) have been observed to switch from indigenous
European trees to Douglas-fir. This limited recruitment has been related to the phylogenetic
distance of Douglas-fir with the European conifers. Most of the colonizers are polyphagous,
originally feeding on different conifer families and/or on angiosperms, and account for 42.5%
of the total whereas oligophagous species (feeding on different genera of Pinaceae) and
monophagous species account for 27.6% and 29.9% of the switches, respectively. The major
part of the monophagous insects originated from European fir and pines rather. The
entomofauna predominantly consists of external defoliators and xylophages rather than
endophagous insects. They rarely caused outbreaks, which were only observed for a lymantrid
defoliator moth, Orgyia antiqua L., in Poland and a geometrid moth, Pseudocoremia suavi, in
New Zealand during the 1970s-1980s. The 20 native European bark beetles observed, so far, on
Douglas-fir did not cause any severe damage even during the recent storms and droughts,
which hit Europe since 1999. In contrast, several alien insects introduced from North America
together with Douglas-fir are susceptible to induce significant damage after having invaded
most of Europe. In the absence of natural enemies and indigenous competitors, they tend to
occupy the entire niche, causing more damage than in the original range. They include woolly
aphids such as Adelges cooleyi and A. coweni, and seed insects such as the Douglas-fir seed
chalcid, Megastigmus spermotrophus, and the conifer seed bug, Leptoglossus occidentalis,
which may considerably decrease seed yield in seed orchards.
In the future, global warming may speed up the slow process of adaptation of native insects to
Douglas-fir. Damage by the pine processionary moth, Thaumetopoea pityocampa, is apparently
increasing since the moth is progressing in both latitude and altitude with global warming and
is thus reaching the bioclimatic range, where Douglas-fir has been extensively planted.
Experimental studies revealed that the moth larvae perform better on Douglas-fir, but female
moths still need to adapt to lay their eggs on Douglas-fir needles. If such an adaptation occurs,
processionnary moths may constitute serious problem for Douglas-fir stands. Besides that, the
present trends in introduction of alien arthropods in Europe (19.6 new species per year on the
average) are likely to result in the arrival of new species associated with Douglas-fir in the
native American range. Quarantine measures to prevent the introduction of Douglas-fir bark
beetles, and especially of the highly damaging Dendroctonus pseudotsugae, have to be
reinforced.
Session 4
21
Vulnerability of Douglas-fir in a changing climate: study of decline in France
after the extreme 2003’s drought
Anne-Sophie Sergent1*, 2, Philippe Rozenberg2, Benoît Marçais1+, Yves Lefévre1*, Jean-Charles
Bastien2, Leopoldo Sanchez2, Louis-Michel Nageleisen3 and Nathalie Bréda 1*
1
INRA UMR 1137* 1136+, National Institute of Agronomic Research, Nancy, France
2
INRA UR 588, National Institute of Agronomic Research, Orléans, France
3
DSF Forest Health Department, Paris, France
With 385.000 ha, Douglas-fir is an important species in France, especially in private forests. It
is the first exotic tree species planted in France. Some of the most important productive regions
in France have suffered an important decline and dieback after the 2003 drought and heat wave.
This decline caused early mortality and loss of production. The main aim of this study was to
answer foresters’ questions concerning the causes of decline. The authors analysed Douglas-fir
decline and dieback in France using two joint approaches. First a national study was performed
using the Forest Health Department (DSF) database. Second a regional dendroecological study
included soil and site description, dendrometric and biotic observations as well as soil water
balance calculation in all sampled stands.
Spatial and temporal Douglas-fir decline analysis at the national scale
The Forest Health Department database reported all mentions of biotic and abiotic problems on
Douglas-fir between 1989 and 2006 (as described by permanent forest health survey agents).
Comparison between mentions of decline and other types of observation allowed to map and
reconstruct the chronology of Douglas-fir decline in France and to identify vulnerability factors
at the national scale. Douglas-fir decline was mapped using reference events and decline events
with quadratic kernel estimator (grid 10km x 10km, window 90km).
This map shows three mains declining regions: two production regions (Burgundy and MidiPyrénées) and one region where Douglas-fir is less current - north-west of France (Normandy,
Pays de la Loire). Not all production regions are affected. This map raises one question: when
did this decline occur? To answer this question the rate of annual decline mentions was
calculated for each three “DSF regions” concerned by declines between 1989 and 2006.
Declines took place during different periods. In North-West DSF region mentions of decline
occurred during three periods (1990-1991, 1999, 2003-2005). In the two other regions the
declines are more recent (2001, 2003-2006 in south-west France and 2002-2006 in Massif
Central). The 2003 drought affected especially those regions. Preliminary results show
significantly higher soil water deficit (cumulated on three years) as computed by soil water
balance modelling for periods with declines. Soil water deficit seems to be a determinant factor
in recent declines in these two regions. In north-west France declines are older. Inappropriate
station condition could explain old and recurrent declines. To complete this result, factors of
vulnerability were identified using a logistical regression. A model was developed, including
topography, mean annual temperature, total annual rainfall and growing-season rainfall.
Thresholds of decline were identified using a regression tree. The main explicative trends
which were able to be extracted from these results were drawn at the national scale. Mains
trends can locally vary with local off-set factors. These local factors (soil water capacity,
silviculture, leaf area index) were not available. Their contribution was analysed in the second
part of this study.
22
Douglas-fir decline analysis at regional scale
A retrospective analysis of radial growth, microdensitometric profiles and daily soil water
balance calculation was performed. The purposes were to date growth reduction, to quantify
the impact of drought events on radial growth and tree ring properties, to search past growth
pattern related to actual tree health or tree mortality. Site conditions, silviculture and biotic
interactions were tested as vulnerability sources. We selected two regions severely impacted in
2003 according to the Forest Health Department database. These regions were the second
(Burgundy) and the fourth (Midi-Pyrénées) among Douglas-fir production regions. 30 plots
were selected in each region, 1768 trees were observed and 925 healthy or declining trees were
cored to the pith. Tree ring width and microdensity profiles were measured and related to
climatic data and drought indices. Growth reduction was related to local soil water deficit.
Pointer years were related to both management (thinning) and soil water deficit indices. The
2003 drought was the most severe according to soil water deficit calculation in Burgundy,
while recurrent drought years during the last decade were pointed out in Midi Pyrénées region.
The impact of either extreme drought or recurrent droughts on Douglas fir growth and health
will be compared. Main factors of vulnerability and threshold of irreversible dieback will be
presented and discussed.
Session 4
23
Impact of the 2003 heat wave on Douglas-fir in France:
comparison of dead and surviving trees for juvenile and mature traits
Manuela Ruiz-Diaz13, Sara Marin1, Alejandro Martinez-Meier2, Leopoldo Sanchez1,
Guillermina Dalla-Salda2 and Philippe Rozenberg1
1
2
National Institute of Agronomical Tecnology, Bariloche, Patagonia, Argentina
3
Parque Tecnologico Misiones, Posadas, Misiones, Argentina
The 2003 heat wave in Europe is considered a first worrying sign of the climate change
process, being for instance the most severe heat wave in France since the year 1370. From
partial foliage necrosis to individual death, different types of damages were reported as
consequence of this heat wave in many Douglas-fir populations. However, some Douglas-fir
trees neighboring the affected trees had no visible damage. In a previous study on the effects of
the 2003 heat wave in a Douglas-fir provenance test located in the Orléans forest, in the center
of France, it was shown that the wood of dead and surviving neighboring trees already showed
systematic differences across years before 2003.
In the present study, the same population of dead and surviving trees will be considered
whereby two new sets of traits were analyzed. The first set is composed of 17 juvenile traits
that were available in the INRA database, but not used in the previous study. These traits are
height, circumference, summer shoot phenology and architecture traits measured between 1976
and 1985 (when the trees were between one and 10 year-old).
The first study (already published) demonstrated that the trees surviving the 2003 heat and
drought wave have a significantly higher mean ring density, higher latewood density and
higher latewood proportion. These results, together with other published results, suggest that
this change is related to the sap conduction function of wood. The classical latewood variables
used in this study were based on the location of an arbitrary earlywood-latewood boundary.
Whether this boundary is the most relevant for studying the relationship between density traits
and the sap conduction function is unknown. Thus we wonder whether ring variables not based
on the standard earlywood-latewood boundary would better explain the difference between
dead and surviving trees. Hence the second set of traits was calculated, composed of original
ring variables. These variables are based on a moving density criterion dividing the
microdensity profiles into two parts, high density and low density segments. The annual rings
employed to calculate the second set of traits are from 1986 to 2002. The variables were
computed at the profile and at the ring level.
For each high and low density segment, the following variables were computed: mean density,
length, profile area and standard deviation. For the variables calculated at the profile level only,
the number of cross points between the density profile and the moving density criterion
threshold were also computed.
For the comparison of the dead and surviving trees, the randomized approach conducted in the
previous study was followed: in order to avoid the eventual effects of a non-random
distribution of dead trees across the trial, and to minimize the impact of micro-environmental
differences onto the dead versus surviving comparison, each dead tree was systematically
associated to an adjacent surviving tree with no apparent sign of damage. The authors assume,
therefore that each surviving tree shares the same physical environment as its associated dead
24
tree. Then, the difference between the value of the surviving tree and the value of its associated
dead tree was computed for all the study traits. In order to construct a reference distribution for
each estimator all the trees of the analyzed pairs were used as reference population. A
distribution of one thousand random pairs was subsequently formed from this reference
population, without replacement and with no consideration of health status, with differences for
all variables computed and stored. These distributions corresponded to the expected difference
between any two given trees, or a difference that is due purely to chance. Each observed
difference was compared to its corresponding distribution of randomly associated trees. A
correlation analysis (using Pearson’s linear coefficient of correlation) was also conducted
among all density variables.
No significant differences were observed between the dead and surviving trees for the first set
of traits, that is the traits measured at juvenile stage.
Conversely, significant differences between the dead and surviving trees were observed for
several density variables of the second set: at the profile level, the dead and surviving trees
were significantly different for the area, the mean density and the standard deviation of the high
density segment, as well as for the number of cross points. No significant differences were
observed for the variables of the low density segment. At the annual ring level, significant
differences were also observed between dead and surviving trees for the mean density, the area
and the length of the high density segment, mainly for the ring-years 1994-95 to 2002. Here
again, no significant differences were observed for the variables of the low density segment. At
the profile level, the optimum value for the moving density criterion differed among variables
and ranged from 400 g/dm3 (mean density) to 600 g/dm3 (area and number of cross point). At
the ring level the optimum value ranged from 500-600 g/dm3 (mean density) to 700 g/dm3
(area, length and standard deviation).
Our results confirm and specify the key-position of the higher-density part of the ring in the
resistance to drought. The trait exhibiting maximum differences between dead and surviving
trees is the area of a high density segment cut at a 600-700 g/dm3 density level. Since density is
strongly related with lumen proportion in a given wood volume, this variable can be
understood as the quantity of smaller-lumen cells in the ring: the higher this quantity, the
stronger the resistance to drought. The relationship between this trait and the surviving ability
of Douglas-fir trees to drought can be put side by side with the relationship recently found in
another Douglas-fir study between some parameters of resistance-to-cavitation curves and
latewood microdensity variables. More work is necessary to precisely investigate the role of
density in resistance to drought in Douglas-fir. Especially we want to underline the interest of
the scientific strategy aiming to compare the wood of dead and surviving trees after a severe
drought. This type of study should be repeated and extended to other sites, other Douglas-fir
populations and other climatic events.
Session 4
25
Storm damage of Douglas-fir and Norway spruce in Southwest Germany:
Stability of Douglas-fir and the impact of silviculture
on the vulnerability of conifers
Albrecht A1, Kohnle U1, Hanewinkel M1 and Bauhus, J2
1
Forest Research Institute Baden-Wuerttemberg, Germany
2
University of Freiburg, Institute of Silviculture, Germany
Compared to other natural risks, winter storms cause the most extensive damage in forests of
Central Europe (Schelhaas et al. 2003), and conifers are especially vulnerable to storm damage
(König 1995, Mayer et al. 2005). In order to adapt forest management to storm risk it appears
reasonable to take risk factors into consideration that indicate storm damage resulting not only
from one storm event but rather from several different storms. Therefore we investigated
correlations between risk factors and multi-event storm damage in Southwest Germany
(Albrecht 2009).
Material and Methods
The base of this investigation is the comprehensive data set of the long-term experimental
growth and yield plots of the Forest Research Station of Baden-Württemberg. The data
describe damage caused by the major storm events in 1990, 1999 and also from medium severe
storms in 1967 and 1984 on 800 plots of Norway spruce and Douglas-fir. Within the 650,000
tree observations we found 17,500 damaged trees, either as windthrow or stem breakage. A
special feature of the analyzed data is the high proportion of Douglas-fir, a tree species whose
empirical storm risk in Central Europe is not yet well known. Another characteristic is that the
silvicultural interventions for the long-term experimental plots are well documented. This
documentation allows for a quantitative analysis of the interventions’ impact on storm damage.
In many previous storm damage analyses of a comparably large database this impact could not
be quantified reliably.
Data mining methods and statistical empirical modelling techniques were applied to the
experimental plot data. The data mining techniques employed CART (classification and
regression trees) algorithms and were especially chosen in order to pre-screen the vast data.
Being able to accommodate correlated observations and missing values, these methods were
used to pre-select potential predictor variables on the single-tree and the stand level
independently. The applied statistical models were based on common generalized linear
models, but were extended to take the spatial clustering of the experimental plots into account.
Special variance components, the so-called random effects, were added to the conventional
regression techniques, rendering them “mixed models”.
For the empirical modelling we applied a four-step modelling technique: In the first step, the
general stand-level occurrence of storm damage was modelled (binary data). Therefore, stands
that had experienced storm damage were coded as damage events, and stands without damage
were coded as non-events. In the second step, total damage of stands was modelled, coding
stands with storm-damaged basal area larger than 75% as events and stands with damage below
75% as non-events. These two modelling steps were performed for each of the six main tree
species in the data set independently. The remaining plots with damage between 0.1 and 75%
then proceeded to modelling step 3. In this step, the response variable was coded as the
proportion of storm-damaged basal area. After these three steps of stand-level modelling, the
fourth step used the estimated stand-level storm damage of step 3 as an offset (e.g. value 0.32
26
in fig. 1), and distributed this estimate among the single trees in the stands, taking single tree
information into consideration. The modelling technique is visualized in fig. 1.
Fig. 1: Flowchart scheme of the consecutive modeling steps.
Our findings corroborate that tree species was the most important attribute to explain the plots’
vulnerability to storm damage. Surprisingly, storm damage in the Douglas-fir plots was higher
than in Norway spruce. Even after accounting for site conditions and stand height, Douglas-fir
could not be judged as more stable than Norway spruce. In contrast, other coniferous species
like Silver fir, Scots pine or European and Japanese larch exhibited significantly less damage
than Norway spruce or Douglas-fir. Other studies have also pointed out the increased risk of
Norway spruce compared to other coniferous species (Aldinger et al. 1996, Hanewinkel et al.
2008). However, the elevated risk of Douglas-fir is partly contradicting findings of Schütz
(2006) who found a stabilizing effect of Douglas-fir in Norway spruce stands. Only Lohmander
and Helles (1987) also found an elevated storm risk of Douglas-fir compared to Norway
spruce. A considerable proportion of the Douglas-fir plots in our study were found on clay
soils. This may have contributed to a site-driven higher disposition to storm damage. In
general, Douglas-fir tends to form a heart root system (Groth 1927, Nicoll et al. 2006,
Studholme 1995) which indicates more stable anchoring than the shallow rooting of Norway
spruce. However, some practical experience exists that also Douglas-fir roots penetrate the soil
less deeply when the rooting depth is restricted for example by clay (Oeschger 1968). In
conclusion and based on the findings of our study, Douglas-fir cannot be considered more
stable to storm damage than Norway spruce. However, it is not clear to date, whether Douglasfir’s high storm risk is due to the species’ characteristics or to the spatial distribution and site
characteristics of the analyzed data set. An additional study with primary orientation on this
question is needed.
After the risk factor tree species, stand height was the second most important attribute to
characterize storm damage, indicating higher risk with increasing stand height. In the first three
Session 4
27
modeling steps, this predictor variable was selected as significant. The high importance of
stand or tree height was also found by other authors and is plausible in general, since the
mechanical stability of stands and trees decreases with increasing height due to higher leverage
(Dobbertin 2002, Lohmander and Helles 1987), and since wind speeds naturally increase overproportionally above the ground (Mitscherlich 1971).
Variables describing silvicultural interventions appeared third in rank; thus, interventions and
selective removals/thinnings were more important to explaining storm damage on the
experimental plots than e. g. orographic exposure or soil characteristics. Especially removing
dominant trees destabilized stands more than thinnings from below. This strong effect was
indicated by the predictor variable ‘thinning quotient’ which was calculated as the mean
quadratic diameter of removed trees divided by the mean quadratic diameter of all trees prior to
intervention. Higher values of this quotient – which indicate the removal of dominant trees –
were found to coincide with higher damage. Variables coding the amount of removed timber
were also selected in some sub-models, but were not as strong in their overall impact as the
thinning quotient. The temporary destabilization caused by silvicultural interventions is
confirmed by numerous other studies (e.g. Cremer et al. 1982, Nielsen 1995) and can be
explained by the temporary increase of canopy roughness, leading to a reduced mechanical
support of tree neighbors. However, the degree of destabilization as described by the thinning
quotient in our study has not been quantified before.
Two literature reviews come to contradictory rankings concerning the importance of risk
factors (Colin et al. 2009, Dhôte 2005): meteorological and site factors are considered more
important than forest attributes or silvicultural interventions. It is possible, that the plots
analyzed in our study do not capture the variability in site conditions which exists in the entire
forest of Southwest Germany. In combination with the lack of significant information about
gust speed, this lack in site variability may explain this contradictory ranking of risk factors.
In the consequence for forest management we can reason that storm damage can be reduced by
decreasing the proportions of Norway spruce and Douglas-fir. Pure stands of these species
should generally be avoided. Limiting the maximum height of stands seems to be another
highly effective means in order to reduce and avoid damage. Therefore, rotation age and target
diameters need to be adapted accordingly. Another recommendation for reducing storm risk is
to concentrate thinnings from above and heavy thinnings during the early stages of stand
development and to avoid them at older ages.
Key words: storm risk, long-term experimental plots, silvicultural intervention, thinning, site
factors, Baden-Wuerttemberg, Douglas-fir, conifers
The list of references can be ordered in: Axel.Albrecht@forst.bwl.de
28
Session 5
29
Session 5
30
Interactions between Douglas-fir and European beech
– what do growth parameters indicate?
Haßdenteufel M, Bögelein R. , Werner W. & Thomas F.M.
University of Trier Department of Geobotany
It is assumed that beech and Douglas-fir have similar ecological demands. At a study site near
Merzalben (Pfälzerwald, Rhineland-Palatinate) direct interactions between Douglas-fir and
beech are investigated. The site covers a pure stand area of each species (intra-specific
competition), as well as a transitional zone, where Douglas-fir and beech individuals are mixed
and interact (inter-specific competition). Douglas-fir was planted about 50 years ago, beech arising from natural rejuvenation - is aged about 55 years. Every competitive situation is
studied similarly, in order to work out differences of inter- and intra-specific competition.
Particular attention is given to various growth parameters.
Allocation behaviour of beech in mixed stand differs from pure stand, concerning tree height
and stem diameter. Beech individuals' height is increased and stem thickness is decreased under
interaction, compared to pure stand. This indicates strong competition for light in the transition
zone. Light interception by Douglas-fir is higher than by beech. Therefore beech trees have to
allocate more biomass into height growth than in crown width; to keep up with Douglas-fir and
to avoid shading; beech grows in height at the expense of radial increment.
The shading effect of Douglas-fir is also shown by the specific leaf area (SLA) of beech leaves.
Leaf samples were yielded in the crown using a hydraulic lift. Within the upper crown of the
mixed stand SLA of beech is significantly higher than in pure stand. Leaves from the mixed
stand show stronger shadow leaf character, distinguished by ecophysiological measurement
with porometer as well as by SLA. Beech seems to be shaded and to form leaves which are
more efficient under diffuse light conditions. The light efficiency was characterised by light
response curves and A/ci-curves measured in the crown. In addition, differing vertical crown
gradients of SLA were found under diverse species interactions.
In addition, displacement effects in the upper rooting zone were found. Compared to pure stand
root mass, root surface and the number of root tips of Douglas-fir are increased in the mixed
zone. Concerning beech, those parameters are decreased. This negative effect of interaction
indicates competitive exclusion at higher expense of beech. Furthermore, a shift in root
distribution was observed. Under conditions of inter-specific competition beech roots are
forced into deeper soil regions, while Douglas-fir captures the upper soil area of highest
nutrient and water availability.
Beech is under strong competitive pressure exerted by Douglas-fir. Superiority of Douglas-fir
under current climate conditions at this specific site is indicated. Current studies concerning
water-use-efficiency of Douglas-fir and beech could give information about the species'
success in competition under expected future climate conditions
Session 5
31
Growth, allocation of space and competition
in mixed stands of Douglas-fir and European beech
Lutz Hilbrig, Germany
Federal Research Institute for Rural Areas, Forestry and Fisheries; Institute of Forest Ecology
and Forest Inventory, Eberswalde, Germany
Douglas-fir (Pseudotsuga menziesii) already plays an important role as an economic tree
species in the current forest practice of Germany. In future, it is expected that silvicultural
interest in this species will grow. However, pure stands of Douglas-fir are mostly not accepted.
European beech (Fagus sylvatica) could be one option for mixed stands together with Douglasfir. As it is the most competitive endemic broadleaved species, beech lets us expect a
successful co-existence with Douglas-fir.
In this context, some questions under the keywords „growth“ and „competition“ were
investigated. For this project, long term experimental mixed stands of Douglas-fir and
European beech in the lowlands of Lower Saxony were used. Investigations were done in the
aboveground and additional on the belowground stand compartments. Principally, mixed stands
of Douglas-fir and European beech up to an age of about 100 years seem to be promising. At
the given site conditions, both species are able to perform growth on high levels. Mixed stands
give higher results than pure stands of beech, however less than pure stands of Douglas-fir.
The range of diameter and height-distributions showed noticeable differentiations within both
species, however, with wide overlaps between the species. Looking at vertical canopy layers,
there is dominance of Douglas-fir in the upper layer and a dominance of beech in a middlelayer. It is notable that growth of diameter is looking similar. Douglas-fir, which is mostly
dominant, only achieved an equal or even smaller diameter growth in the middle-layer than
beech. That could be a first sign about competitions between both species. Looking at the treeheight, height of maximum crown-diameter, and height of crown-base, it became clear that
photosynthetically effective crown parts of Douglas-fir are growing above those of beech,
never reverse. According to that, dominance of Douglas-fir is a result from a higher increment
in height.
Appropriate competition-indexes are always showing a dominance of Douglas-fir. This applies
to absolute as well as to relative rates. The absolute competitive pressure on beech is much
higher than on Douglas-fir. Differentiated in intra-and interspecific competition, beeches are
prone to higher interspecific and Douglas-fir more intra-specific competition.
A second subject of this investigation was growth and structure of fine roots. Layering like we
had found for canopies are also given in vertical distribution of fine root biomasses. Douglasfir tends to subdue organic and upper mineral layers down to 10cm. However, beeches tend to
establish fine-roots within the deeper soil layers. Here both species seem to avoid each other.
Another aspect of competition is the horizontal range of fine roots surrounding one central tree.
In this case, fine roots of both species do not avoid, they are interlocking. Both species are able
to establish fine-roots near to the tree trunks of both competing species. A difference was
shown in the radii in which the central trees are dominant. This “zone of dominance” is greater
for beech than for Douglas-fir.
32
Douglas-fir growth response to variation in soil water holding capacity and
local climatic patterns
in the Siskiyou Mountains of southwestern Oregon
Douglas A. Maguire, David W. Hann1, Douglas B. Mainwaring
Department of Forest Engineering, Resources and
Management, Oregon State University
The Siskiyou Mountains of southwestern Oregon and northern California, USA, are unusually
complex with respect to topography, geology, climate, and vegetation composition (Whittaker
1960). Douglas-fir has relatively wide ecological amplitude across this complex landscape,
and its productivity in the region is probably more variable than in any other region of
comparable geographic extent. A large proportion of this variability in Douglas-fir production
can be attributed to the wide range in soil water holding capacity, total and seasonal
precipitation, and constraints imposed by soil moisture limitations, high late-summer vapor
pressure deficits, and growing season length. Productivity has been shown partly predictable
from basic ecophysiological principles (Swenson et al. 2005), with soil water holding capacity,
available soil water, and vapor pressure deficits playing a major role (Coops and Waring
2001a, b; Waring et al. 2008). Improved spatial analysis of local climatic diversity and detailed
field description of soils may provide significant insight into possible Douglas-fir responses to
long-term changes in climate.
Five-year stem volume growth was measured on 148 Douglas-fir plots in the Siskiyou
Mountains of southwestern Oregon. Soil water holding capacity was estimated at each site
from detailed soil profiles, and weather variables were compiled from the DAYMET U.S. Data
Center. Soil water holding capacity ranged from 101 to 411 cm with an average of 191 cm.
Total annual precipitation ranged from 76 to 208 cm, with an average of 122 cm, but growing
season precipitation ranged from only 10 to 22 cm with an average of 15 cm. Corresponding
gross periodic annual stem volume increments ranged from 1.3 to 33.7 m3ha-1yr-1 with a mean
of 13.6 m3ha-1yr-1.
Predictions of Douglas-fir productivity varied across a hierarchy of models that incorporated
soils and weather variables at different resolutions. The simplest were regression models that
included yearly averages, and the most complex were hybrid models that attempted to simulate
ecophysiological processes on a daily time step. Douglas-fir productivity is driven by a
complex interaction of initial stand structure, soil moisture holding capacity, seasonal
distribution of precipitation, evapotranspirational demand, and other climatic variables. As
these factors fluctuate or change over time, Douglas-fir productivity may change in the short
term, natural selection may force genetic change in the long term, and extreme conditions may
drive a shift to other species in the still longer term, depending on the speed and consistency of
climate change.
Session 6
33
Session 6
34
KEYNOTE 6
Douglas-fir – a look into the genetics
Monika Konnert
Bavarian Institution for Forest Seeding and Planting
Teisendorf, Germany
Douglas-fir has one of the widest natural ranges of any tree species and the largest south-tonorth distribution of any commercial conifer in North America, extending from 19oN in
Mexico to 55oN in Western Canada. In Western Oregon and Washington it occurs from sea
level to 1700 m (Hermann and Lavender 1990). Within this large geographic area contrasting
climatic conditions are found. Douglas-fir populations seem to have a great adaptive potential
and are generally regarded as being closely adapted to the different environments. Winter
temperatures and frost dates are of great importance to the adaptation of Douglas-fir, whereas
summer drought is of less importance (St. Clair et al. 2005).
Two taxonomic varieties are recognized: the coastal form (P. menziesii var. menziesii or
viridis) found along the North American Pacific Coast and the interior form (P. menziesii var.
glauca) found inland in the mountains from British Columbia to Central Mexico. In Europe,
the caesia type is widely recognized as an intermediate type between the coastal (viridis) type
and the interior type (glauca) (Aas 2008, Schober 1954). There is no reproductive barrier
between the different types.
The varieties differ in a number of important traits: The coastal variety grows faster and gets
considerably larger then the interior variety, which tend to be more shade tolerant and more
cold hardy. Within the coastal variety trees from the coastal areas are less cold hardy than trees
from the western slope of the Cascades (Aitken at al. 1996). Seedling tests in common garden
experiments showed that variation in bud-burst, emergence and growth is strongly related to
elevation and cool-season temperatures. Seedlings from the east side of the Washington
Cascades were considerably smaller, set bud later and burst bud earlier than populations from
the west side (St.Clair et al. 2005).
Beside the high phenotypic diversity Douglas-fir shows high genetic variation. Studies using
allozymes (e.g. Merkle and Adams, 1987, Li and Adams 1989, Yeh and O’Malley 1980, ElKassaby and Ritland 1996) have found that Douglas-fir has an enormous amount of genetic
diversity and that more than 95 % of it resides within populations and not between them. There
is a clear difference between the coastal and inland varieties in gene frequencies at specific
allozyme loci. Over 75 % of the variation in allozymes among populations is accounted for by
differences among the varieties. Allozymes also show variation within varieties of Douglas-fir
as well. Artificial populations of Douglas-fir in Europe do not appear differently from the
natural ones as long as a sufficient number of trees are involved in the original collection (e.g.
Prat et Arnal 1994). Numerous investigations of Douglas-fir stands in Europe have shown that
the regional specific structure fund in North America was not altered in the second generation
by the transfer and only a few rare gene variants were lost (e.g. Hofmann and Geburek 1995,
Klumpp 1999, Leinemann 1996, 1998).
Since 2002 highly variable SSR marker have been developed in Douglas-fir (Carlsson 2002,
Slavov et al. 2004). They are useful tools for DNA fingerprinting and parentage analysis.
Slavov et al. (2004) report that only three SSR were sufficient to achieve an average probability
Session 6
35
of exclusion from paternity of 0.998 in a seed orchard block with 59 parents. On the other part
SSR-markers can improve the results on genetic variation obtained with allozymes. A study on
the population structure in coastal Douglas-fir using allozymes and microsatellites (Krutovsky
et al. 2009) demonstrates this but shows also that at first these two marker types should always
be analyzed separately due to their different nature. Both types of markers confirmed the
relatively high genetic diversity within populations and low differentiation among them similar
to other conifers with large effective population sizes. Only for SSR- markers a positive and
significant correlation between genetic and geographic distances was found. It is suggested that
isolation by distance may play an important role in the coastal Douglas-fir differentiation at
neutral SSR-loci, while allozyme variation are under different mostly stabilizing forms of
selection.
Douglas-fir is a wind-dispersed and wind pollinated species with high gene flow (El-Kassaby
and Ritland 1986) and oucrossing rates exceeding 90 % (e.g. El-Kassaby et al.1981, Shaw and
Allard 1982, Neale and Adams 1985, Yeh and Morgan 1987). The effective pollen dispersal
distance was estimated to be around 30 m. Within 30 m mating success is only weakly related
to distance (Erickson and Adams 1988). The reproductive phenology significantly affects the
mating system and induces a larger part of crossing between the same phenological classes (ElKassaby and Ritland 1986). As flowering is a heritable parameter crossing between related
trees might represent a major part in the mating system in natural populations of Douglas-fir
and can explain the often higher fixation index observed in progenies in comparison with old
stands (Prat and Arnal 1993).
Douglas-fir has been grown in Europe for over 120 years and is the most important introduced
species in Germany. Beside site conditions and silvicultural treatment, the provenance is
crucial for a successful introduction of a species. Until the late sixties frequently nothing was
known about the provenances which were imported. In the mid-sixties seed zones for Douglasfir were established in the Pacific-Northwest. At the same time (1967-1968) an IUFRO
provenance field trial was started including more than 100 provenances from Oregon,
Washington and British-Columbia. The collection was distributed to 33 countries mainly in
Europe; a part of the numerous sites still exist. The results of this trial demonstrate the better
adaptation and growth of the coastal variety on nearly all European sites (Breidenstein et al.
1990, Kleinschmit et al. 1974, 1991, Ruetz 1981). In oceanic Europe provenances from low
elevation in Washington perform the best. In Central and Eastern Europe, the choice of
Douglas-fir reproductive material is restricted to the middle elevation zone of the Cascade
range in Northern Washington as provenances from high latitude and elevation have a low
growing rate. In Southern and Mediterranean Europe provenances from low elevation in
northern Oregon and southern Washington seem to be of high interest (Breidenstein et al.
1990). To refine provenance recommendations in 1971 and 1976 supplementary field tests
were installed in Bavaria, which included an elevational sampling up to 1000 m in areas on the
western Cascade slope as well as provenances from the Cascade crest. As a result new
guidelines for Douglas-fir seed import were established based on seed zones and elevation
(Ruetz and Foerst 1984, Ruetz 1985). Under climate change this field trials provide valuable
concrete data on population responses, which are not obtainable from theoretical models
(Matyas et al. 2010).
Douglas-fir has one of the largest breeding programs in the world, with more than 4 million
progeny from nearly 34.000 parents growing on almost 1.000 test sites in western North
America alone. Breeding programs began in the mid’s 1950s to improve growth, stem and
36
wood quality and are today well into the third generation of selection and testing. The average
volume gain in the second generation seed orchards is reported to be around 12 %, whereas the
top-crosses outperformed the controls by an average of 17 % (Stoehr et al. 2009). In Europe the
most detailed breeding program for Douglas-fir started in 1960 in France with the selection of
more than 1000 plus-trees followed by field tests, next selection steps and finally the
establishment of more than 60 ha seed orchards with the best performing clones (Heois 1995).
Comparison between two generations of seed orchards and their 49 wild populations indicate
that early selection and breeding of this highly polymorphic species does not reduce genetic
variation significantly (El-Kassaby and Ritland 1996), if the number of clones is not too low.
Similar results were obtained by comparing seeds from French seed orchards and from seed
stands in South Germany. This is a very important result as the maintenance of genetic
diversity in inbreeding and production populations has to be a priority of all breeding programs
especially under climate change where a broad adaptation basis is needed.
An important yet unanswered question is what specific genetic and epigenetic phenomena are
responsible for the variation observed in adaptive traits. Associations between polymorphism at
the DNA level and adaptive traits measured in common garden experiments should provide a
first step towards elucidating the genes responsible for adaptive variation. Furthermore, for a
spieces as Douglas-fir with high phenotypic and genetic variability and a large and continuous
natural distribution range, association mapping approaches are promising. Under this approach
single nucleotide variation (SNP) in expressional and functional candidate genes is associated
with variation in adaptive traits as cold hardiness or drought resistance in segregating and
natural populations. The number of studies in this field is steadily increasing. SNP markers in
numerous candidate genes were recently developed (Krutovsky and Neale 2005, Pande et al.
2007). The identification of more adaptive genes or gene groups will be the task of coming
decades.
Remark: The cited references represent only a small fraction of the literature published on the
subject. References can be requested from the author.
Session 7
37
Adaptation of Douglas-fir provenances to drought stress
Ingo Ensminger1,3, Moritz Heß1, Thomas Müller2, Henning Wildhagen1, Karl Schmid2
1
Forest Research Institute Baden-Württemberg, Department Forest Ecology
2
Institute of Plant Breeding, Seed Science and Population Genetics
University of Hohenheim
3
Department of Biology, University of Toronto, ON, Canada
Climate change presents itself as an increase in temperature, change in precipitation patterns as
well as an increase in frequency of extreme events such as drought, heat waves or storms.
Future growth and productivity of forests depend on the adaptability, vulnerability, resistance
or resilience of trees to future climatic conditions. However, the current understanding of
adaptation and acclimation potentials of trees to climatic change is rather incomplete.
Within their natural range, trees of the same species may grow under various different climatic
conditions and these environmental differences act as selective forces on the genetic and
physiological growth response of the different provenances. This applies in particular to
Douglas-fir, which – as a consequence of its wide geographical distribution in North America –
shows a wide range of ecological tolerance and adaptability. This offers the opportunity to
identify Douglas-fir provenances which are adapted or may acclimate to the climatic conditions
anticipated for various regions of Central Europe in the future.
The focus of our project is on the drought response of forest trees, since significant increases in
temperature and more frequent extreme summer drought events are predicted by regional
climate models for Central Europe and the Upper Rhine valley in particular (Christensen et al.
2007; Rennenberg et al. 2004).
Our project is part of a multidisciplinary research consortium which aims at understanding the
diversity of drought responses among Douglas-fir provenances from environmentally different
habitats and the molecular and physiological mechanisms underlying drought tolerant
phenotypes. The ultimate goal is to identify physiological and molecular markers for the
selection of Douglas-fir provenances that are adapted to future climatic conditions. This will be
achieved by linking allelic variation in candidate genes with phenotypic variation using an
association genetics approach. The research program of this consortium consists of four
independent but complementing projects dealing with i) molecular markers and the molecular
regulation of drought responses, ii) metabolites and stable isotopes as physiological markers
(see Sohrt, Jansen & Gessler in this volume), iii) growth dynamics and iv) regulation of
isoprenoid metabolism and emission patterns of volatile organic compounds (see Junker et al.
in this volume). In this joint approach, the diversity of drought responses of Douglas-fir is
analysed on different temporal scales and different levels of biological organisation ranging
from single genes to whole trees and stands.
These responses will be investigated in experiments under controlled conditions as well as in
field studies. The field experiments are conducted on two sites of the International Douglas-fir
provenance trial. The selected sites (Wiesloch and Schluchsee) are characterised by contrasting
climatic conditions with temperature decreasing and precipitation increasing from Wiesloch to
Schluchsee. At the two sites, four identical Douglas-fir provenances are studied. Their sites of
origin in North America reflect a climatic gradient from dry interior sites to moist coastal sites.
38
Using the selected provenances and the two sites, we have the unique opportunity to study
differences in the adaptation to local climate and the adaptability to climatic changes in 50 year
old trees.
Here we present first results on the molecular and genetic basis of drought responsive
mechanisms of Douglas-fir. The first step to identify drought-stress candidate genes from the
whole set of transcribed genes, and to determine their allelic variation is to establish a
comprehensive reference database of gene sequences. Since conifer genomes are very large
(Ahuja & Neale 2005), we sequenced the Douglas-fir transcriptome instead of the whole
genome. For this purpose, we constructed normalized cDNA libraries of wood and needle
tissues of Coastal and interior Douglas-fir exposed to artificial drought stress. In a glasshouse
experiment, the two varieties were represented by eight provenances (see fig. 1) with a least
five plants per provenance and drought treatment (control, mild and severe drought stress). The
normalized cDNA libraries were sequenced by pyro-sequencing. Most of the libraries
contained approximately 300000 reads, and the average read length was approximately 300
basepairs. This demonstrates that the sequencing was successful. The reads were assembled to
so called “contigs” by merging two or more reads with overlapping sequences using the
program MIRA (Mimicking Intelligent Read Assembly; Chevreux et al., 1999). Next, the
contigs will be compared to publicly available sequences of other plant and tree species in
order to assign these contigs to a biological function or process. While sequencing of
normalized cDNA libraries is the method of choice to generate a reference database of gene
sequences, its application to the transcriptome-wide analysis of gene expression patterns is
limited. Therefore, our approach to select candidate genes and determine allelic variation in
these genes will be complemented by shortread-sequencing of non-normalized cDNA libraries.
These libraries will be constructed from plants exposed to artificial drought stress conditions in
glasshouse experiments and from trees sampled in field experiments.
In summary, our approach to assess the diversity of responses of Douglas-fir populations
towards drought stress by a combined analysis of phenotype characteristics (eco)physiological,
molecular and genetic traits will provide validated information for the selection of provenances
suitable to grow under the anticipated future climatic conditions.
Session 6
39
Canada
Mexico
Figure 1: Map of origin of the provenances selected for glasshouse experiments and
characteristics of the environmental conditions of their native habitats. (Map modified
according to Aas, 2008).
References
AAS G (2008): Die Douglasie (Pseudotsuga menziesii) in Nordamerika: Verbreitung, Variabilität und Ökologie. LWF – Wissen
59: 7-11.
AHUJA MR, NEALE DB (2005): Evolution of genome size in conifers. Silvae Genetica 54: 126-137.
CHEVREUX B, WETTER T, SUHAI S (1999): Genome sequence assembly using trace signals and additional sequence information.
Computer Science and Biology: Proceedings of the German Conference on Bioinformatics 99: 45-56.
CHRISTENSEN JH, HEWITSON B, BUSUIOC A, CHEN A, GAO X, HELD I, JONES R, KOLLI RK, KWON W-T, LAPRISE R, ET AL.
(2007): Regional climate projections. In: Solomon S, Qin D, Manning M, Chen Z, Marquis M, Averyt KB, Tignor M, Miller
HL. (Eds) Climate Change 2007: The physical science basis. Contribution of working group I to the fourth assessment report
of the Intergovernmental Panel on Climate Change. Cambridge University Press, Cambridge, UK: pp. 847-940.
RENNENBERG H, SEILER W, MATYSSEK R, GESSLER A, KREUZWIESER J (2004): Die Buche (Fagus sylvatica L.) – ein Waldbaum
ohne Zukunft im südlichen Mitteleuropa? Allgemeine Forst- und Jagdzeitung 175: 210-224.
40
Genetic value of a Douglas-fir natural regeneration in France
1
Alain Valadon1, Ouzna Boussaid12, Jean-Charles Bastien2
Genetic Conservatory of Forest Trees, National Forest Service, Orléans, France
2
Many Douglas-fir artificial stands planted for wood production in Europe will have to be
harvested and renewed in the next future. Natural regeneration is among the possible technical
choices. But most of the time information lacks about the origin and the genetic basis of the
adult stands. Forest managers sometimes question the genetic value of the natural regeneration.
In the context of the ongoing climate change, the adaptive potential of naturally regenerated
Douglas-fir populations is directly related with their existing genetic diversity.
We estimated the genetic diversity using molecular markers in two Douglas-fir populations: a
75-year-old adult population (158 trees) and a one-to-six year-old natural regeneration (463
seedlings) originating from the adult population. We used five chloroplastic and three nuclear
microsatellites markers. Our results show that there is a high genetic diversity in the adult
population, with no spatial structure. The allelic diversity is higher still in the population of
natural seedlings, with a significant spatial correlation between genetic and geographic distance
up to a distance of 60 m. The diversity of the natural regeneration is influenced by the
neighbouring parent trees as well as by annual gene flow entering the stand. These first results
highlight the combined roles of the origin of the parents and of the process of natural
regeneration on the establishment of future genetic diversity.
Session 6
41
Photoprotective isoprenoids as physiological markers for the adaption of
Douglas-fir to drought stress in context of climate change
Laura Verena Junker1, Anita Rott2, Henning Wildhagen1,
Ingo Ensminger1,3, Jürgen Kreuzwieser2
1
Forest Research Institute Baden-Württemberg, Freiburg, Germany
2
University of Freiburg, Germany
3
University of Toronto at Mississauga, Department of Biology
Mississauga, Ontario, Canada
The aim of this project is to determine the diversity of isoprenoid-related mechanisms of
drought tolerance in Douglas-fir provenances which differ in their geological origin and
therefore probably also in their drought responses. For this purpose, we will develop and link
genotypic and phenotypic markers for the adaption to drought stress and provide tools for
marker assisted tree improvement to select high-yielding Douglas-fir provenances.
Plants are always in danger of photooxidative damage, as excess light energy that cannot be
used for photosynthesis may impair the photosystems and produce reactive oxygen species
(ROS). Drought enhances the risk of photooxidative damage because it induces stomatal
closure and reduces photosynthetic CO2 assimiliation (Niyogi 2003).
Essential isoprenoids are comprised of different pigments and antioxidants and act as
photoprotective components by allowing non photochemical quenching (NPQ) of excess light
energy (Fig. 1). Under normal conditions, accessory pigments like carotenes enhance
photosynthesis efficiency by uptaking light of wavelengths that are not usable by chlorophyll
pigments; under stress conditions, they also absorb excess light energy from chlorophylls and
dissipate it as heat. Xanthophylls can also dissipate excess energy in terms of heat, while
tocopherols act as antioxidants that can scavenge ROS (Peñuelas and Munné-Bosch 2005).
Along with photoprotective compounds, isoprenoids also include hormones and membrane
components which can indirectly influence stress tolerance of plants (Fig. 1).
In addition to the essential isoprenoids, many plant species including Douglas-fir also produce
nonessential isoprenoids. These include volatile organic compounds (VOC) such as isoprene,
monoterpenes and sesquiterpenes (Jirovetz et al. 2000). The role of VOCs is still unclear
although they have been attributed to actions related to thermal stresses such as protection of
thermal damage, antioxidant properties and the uptake of excess energy during biosynthesis
(Fig. 1; Owen and Penuelas 2005).
42
Fig. 1: Isoprenoid compounds in plants serve different functions. Essential isoprenoids comprise
pigments and antioxidants which protect plants against photooxidative damage. Nonessential
isoprenoids seem to protect plants often in indirect ways when they get emitted into the atmosphere.
Previous studies demonstrated that isoprenoid content and composition in plants can vary
considerably between years, seasons, times of day and areas, and seem to be strongly related to
site-specific temperature regimes (Gambliel and Cates 1995). The biosynthesis of essential and
volatile isoprenoids is strongly linked because they share the same biochemical precursors
(Owen and Penuelas 2005). Therefore changing patterns of essential isoprenoid expression
should be consistent with volatile isoprenoid emission patterns.
We hypothesize that the adaptation of Douglas-fir to drought stress is mediated by patterns of
biosynthesis which can be traced via the abundance of essential and non-essential isoprenoids.
For this purpose we study field- and greenhouse-grown plants of four provenances which have
evolutionarily adapted to contrasting environments.
Field campaigns started in 2010 and will be continued in 2011 at two field sites in southwestern
Germany where growth traits of fifty-year-old Douglas-firs were regularly monitored (Kenk
and Thren, 1984). As the field sites differ in altitude, precipitation sums and mean annual
temperature, differences in summer drought periods are expected. To evaluate the trees
capability of photosynthesis under normal and drought stressed conditions, gas exchange
measurements of needles are conducted in May and July. These measurements will be taken
along with chlorophyll fluorescence measurements that indicate the amount of NPQ and
sampling of emitted VOCs. The content of essential and nonessential isoprenoids will be
determined by chromatographic methods in concurrently sampled needles. By this approach the
interrelation between photosynthesis efficiency, isoprenoid related NPQ mechanisms and VOC
emission will be investigated.
Alongside these physiological measurements, samplings were carried out in May, June, July
and September to obtain an overview of seasonal variation of diurnal patterns. Twigs of the sun
exposed crown are simultaneously sampled three times a day. These samples will also be
investigated via means of molecular genetic methods and isotope analysis. To differentiate
between effects of drought and heat, greenhouse experiments with one-year-old plants will be
carried out with selective applied drought and temperature stresses. The combination of
Session 6
43
different methods will give a better insight to the interconnections between isoprenoid
dependent processes and drought adaptation in Douglas-fir.
References
GAMBLIEL HA, CATES RG (1995): Terpene Changes due to Maturation and Canopy Level in Douglas-fir (Pseudotsuga
menziesii) Flush Needle Oil. Biochemical Systematics Ecology 23: 469-476.
JIROVETZ L, PUSCHMANN C, STOJANOVA A, METODIEV S, BUCHBAUER G (2000): Analysis of essential oil volatiles of Douglas
fir (Pseudotsuga menziesii) from Bulgaria. Flavour and Fragrance Journal 15: 434-437.
KENK G, THREN M (1984): Ergebnisse verschiedener Douglasienprovenienzversuche in Baden-Württemberg. Teil I: Der
internationale Douglasien-Provenienzversuch 1958. Allgemeine Forst- und Jagdzeitung 155, 7/8, 165-184.
NIYOGI KK (2000): Safety valves for photosynthesis. Current Opinion in Plant Biology 3: 455–460
OWEN S, PEÑUELAS J (2005): Opportunistic emissions of volatile isoprenoids. Trends in Plant Science 10: 420-426.
PEÑUELAS J, MUNNÉ-BOSCH S (2005): Isoprenoids: an evolutionary pool for photoprotection. Trends in Plant Science 10: 166169.
44
Session 7
45
Session 7
46
Genetic investigation on Douglas-fir seed stands
by means of isozyme and microsatellite markers
Barbara Fussi, Monika Konnert
Bavarian Office for Forest Seeding and Planting (ASP)
Teisendorf, Germany
In order to be successful in the establishment of Douglas-fir, one must select the best
provenance, maintain a broad genetic base and carry out proper silviculture. European
provenance trials indicated that the “green” or costal type (Pseudotsuga menziesii var.
menziesii) is more suitable for Bavaria compared to the “grey” or interior type (P. menziesii
subsp. glauca var. caesia). The goal of our study was to check the origin/provenance of
Bavarian Douglas-fir seed stands, the respective genetic variation of the stands and also the
natural regeneration. This is important because for many stands the origin of the reproductive
material as well as the number of seed trees harvested to establish Bavarian stands is unknown.
The proportion of empty seeds and low germination rate within Bavarian seed lots are not
uncommon and we assume inbreeding effects and inbreeding depression as possible causes.
In this study we present results based on isozymes and nuclear microsatellites. Over 150
Douglas-fir stands from Bavaria were analysed with isozymes and resulted in 2-3% pure grey
Douglas-fir stands, 25% mixed stands (grey and green) and the majority of the stands consisted
of pure green Douglas-fir. The genetic diversity revealed large differences between the stands
(vgam 4.0-12.0). The results using microsatellites pointed in the same direction. Special
attention using microsatellites was directed towards gene flow, because several stands consist
of spatially separated small groups of individuals (less than 50 individuals on less than 1 ha),
composed of different varieties. The results of Bavarian stands were compared to stands from
the area of origin of Douglas-fir and to high breeding seed orchards from France. The results
are integrated in provenance recommendations for Douglas-fir and are therefore highly relevant
for practice. Particularly with regard to the goal of increasing the stocking with Douglas-fir in
Bavarian Forests (from at present 0.7% up to 3.0%) mixed with different native tree species.
Session 7
47
Ecophysiological isotope tools for characterising the drought sensitivity of
Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)
Jakob Sohrt1, Kirstin Jansen2, Arthur Gessler1,2
1
ZBSA, University of Freiburg
2
Institute for Landscape Biogeochemistry
Leibniz-Institute for Agricultural Landscape Research
The stable isotope ratio of Carbon and/or Oxygen (δ13C resp. δ18O, d) in whole wood and in
cellulose is directly linked to environmental factors as well as to the physiological response of
the trees to these factors – at the time when the organic material was formed. Thus the isotopic
signature in tree rings can be used directly as a physiological archive. Physiological isotope
investigations combined with molecular biological methods can in future enable efficient and
time-saving tests for climatic sensitivity of provenances of a tree species.
Within this paper, we present a study of δ18O and δ13C in the late wood of different Douglas fir
provenances assessed retrospectively in an altitudinal transect (Oberrheinebene-Schwarzwald)
by isotope ratio mass spectrometry. Special attention has been payed to the physiological
reaction of the provenances in years with extreme atmospheric conditions (e.g. the extreme dry
year 2003) and in the following years. Isotope ratio measurements were conducted for tree
rings formed in the years 2002 until 2007.
The results of the isotope ratio measurements showed clear distinctions between different
Douglas-fir provenances in their physiological reaction to rigorous drought, as observed in
2003. Isotope ratio measurements revealed that some but not all provenances intensively closed
the stomata to reduce water consumption as a consequence of the drought in 2003.
48
Planted forests of Douglas-fir in New Zealand:
challenges for the breeding programme with climate change
Dungey, H.S2., Low, C.B2, Watt2, M.S. 2, Hood2, I.A., Stone2, J.K1. and Kimberley2
Scion. Te Papa Tipu Innovation Park, Rotorua, New Zealand
*Department of Botany and Plant Pathology, Oregon State University, Oregon, U.S.A.
Douglas-fir (Pseudotsuga menziesii) has been grown in New Zealand since 1859 and is now
the second most important planted exotic forest tree in this country. There are currently around
112,000 ha of this species planted, representing approximately 6% of New Zealand’s planted
forest estate. Douglas-fir is grown mainly in the South Island, in colder, wetter environments
but has enormous potential to expand on to higher-altitude sites. This is particularly the case
with the possibility of receiving carbon credit payments to inject cash flows well before
rotation age.
Douglas-fir breeding was initiated between 1957-59, when provenance trials were established
across a large number of sites in both the North and South Islands of New Zealand. Origin of
the material was from Washington and Oregon (1957), and from coastal California and Oregon
(1959). Following an analysis that showed that Oregon- and Californian-origin material was
growing faster than other provenances, further collections were made from these areas. As a
result, additional breeding trials were established from the central North Island to the south of
the South Island. A recent genetic analysis of these trials has revealed an important genotype ×
environment interaction. Selecting the best material using breeding values estimated from a
multi-site analysis will result in a considerable reduction in realised gain in certain local
environments. We believe this is driven primarily by climate, and the current distribution of
Swiss needle cast.
Swiss needle cast (SNC), caused by the pathogen Phaeocryptopus gaeumannii, was first
detected in New Zealand in 1959, and has since spread throughout the country. This needle
cast is very important to the economics of growing Douglas-fir as it has been shown to cause
approximately 32% reduction in tree growth (New Zealand-wide average). Recent work has
determined the most important driver limiting the distribution of SNC in New Zealand is winter
temperature. With climate change, SNC is likely to become even more severe in the North
Island, and the distribution is likely to extend further into the colder South Island. This has
very important implications for the Douglas-fir breeding programme.
This paper will discuss the current breeding programme, the extent of genotype × environment
interaction and origin of provenances growing best in each environment. Climate change
scenarios will be discussed with a view to general temperature changes across the country and
their likely impact on SNC distribution. Implications for the breeding programme and the
forest industry will be discussed.
Session 7
49
Root system response of naturally regenerated Douglas-fir
(Pseudotsuga menziesii)
after complete overstory removal
Nathan Briggs, Christian Kühne, Jürgen Bauhus
Institute of Silviculture, University of Freiburg, Germany
Observations by forest practitioners in Baden-Württemberg, Germany indicate that naturally
regenerated understory Douglas-fir trees of two-layered high forests are at great risk of
becoming uprooted as a result of storm events or high snow loads. The problem is of great
interest as Douglas-fir is commonly under-planted in older stands and additionally, shelterwood
systems and target diameter harvesting are becoming more popular as a means to naturally
regenerate Douglas-fir in long regeneration phases. We suspect the high susceptibility to
windthrow is a result of an inadequate structural root system and hypothesize that the restricted
development of Douglas-fir growing in these conditions may not be overcome in
developmental stages following overstory release. A retrospective approach was used to
investigate the effect of canopy removal on spatial and temporal expansion of structural roots
of young Douglas-fir that had established and grown for several years underneath a closed
canopy. For this purpose, the complete root systems of 6 Douglas-fir were excavated using
subterranean explosives. Two primary laterals and 1 primary vertical root per tree were
analyzed to investigate response in root elongation and radial increment. Additionally,
complete stem analyses were carried out to relate above- and below-ground responses after
overstory release. Individual compartments as well as comparison between compartments were
completed using 3 growth periods surrounding storm Thalia in 1995. Results indicate that it is
possible to efficiently and accurately investigate response of belowground compartments in
previously suppressed Douglas-fir. Contrary to previous studies, stem increment did not suffer
a growth delay while root compartments increased growth rates. However, annual shoot growth
did experience a 4-year delay before significant growth resumed. Response in root radial
increment was found to be stronger than annual root elongation and near the end of the poststorm period showed higher relative growth than all compartments analyzed. Root elongation
in the post-storm period showed significant difference from pre-storm, but data indicate that
real increases were made at the end of the post-storm period. Results suggest that belowground
compartments in previously suppressed Douglas-fir are capable of strong response to release,
and increase in size and complexity which further improves stability. However, individuals
may experience a delay before differences from pre-disturbance rates are meaningful and
statistically significant. These findings have several silvicultural implications and provide
information for researchers and practitioners working to develop silvicultural practices to retain
and utilize natural regeneration in two-layered Douglas-fir forest stands.
50
Poster Sessions
51
Poster Sessions
52
Bud burst and damage by spring frost of Douglas-fir seedlings
in the south of Sweden
Cecilia Malmqvist
Linnæus University
School of Engineering, Sweden
In the prospect of a change of climate, the question of using various tree species in the future
Swedish forestry has become more urgent. Douglas-fir has been used as a forest tree on a
small scale in Sweden for more than a hundred years. Nevertheless, the experience and
knowledge of the suitability and viability of different provenances is limited. One of the most
serious threats to Douglas-fir seedlings in Sweden is spring frost, especially in the interior
parts. Therefore it is desirable to be able to predict time of bud burst in proportion to
longitude, latitude, altitude, and distance from the sea along with spring weather conditions of
the locality.
The aim of the study was to investigate the time of bud burst and occurrence of damage from
spring frost in two plantations of Douglas-fir in the south of Sweden. In each of the two
localities six provenances, three coastal and three interior were planted in the spring of 2009.
In each site 20 plants of each provenance were marked and monitored each week during the
bud burst period. The development of the buds was documented and compared to the scheme
used for Norway spruce, the Krutzch index. Occurrence of damage from frost was also noted.
Since there was no temperature below zero during the measuring period, there were no frost
injuries on the buds. There were differences between the provenances both regarding the
starting point of the bud burst process and the length of the bud burst period. There were also
differences between the two sites; at the Tagel site the bud burst started earlier and ended
earlier compared to the Asa site.
Poster Sessions
53
Effects of surface humidity and light regime on germination success
and seedling development of Douglas-fir
(Pseudotsuga menziesii Mirb. Franco) – a greenhouse experiment
Franka Huth1, Angelika Körner1, Christine Lemke1, Antje Karge1, Jörg Wollmerstädt,
Sven Wagner1, Martin Hartig1, Dietrich Knoerzer2
1
Institute of Silviculture and Forest Protection; Dresden University of Technology
Tharandt, Germany
Introduction
In Europe previous interest in Douglas-fir (Pseudotsuga menziesii Mirb. Franco) was mainly
concentrated on aspects of growth and yield. This applies for both – forest enterprises and
research institutions. Regarding fructification of mature Douglas-fir trees, changed
regeneration concepts in silvicultural management and uncertainty of climate development
demand for more detailed knowledge of natural regeneration development in the future. Thus,
we purposed the following main objectives in the greenhouse experiment: (1) define a
comprehensible system of morphological characteristics for the different early seedling stages
of Douglas-fir with physiological conclusions, (2) document the temporal development of
germination rates and seedling development stages in dependence of humidity and light
regime, and (3) quantify growth parameters for further developed seedlings.
Material and Methods
Therefore the pot experiment under greenhouse conditions was established with a mixture of
needle litter (L - horizon) and partly decomposed organic matter (F - horizon) as surface
material from pure Norway spruce (Picea abies (L.) Karst.) stands. After studies from
Knoerzer (1999) and Cornett et al. (2000) this kind of substrate can be seen as favourable for
Douglas-fir seedlings during the first time of development. Douglas-fir seeds from
provenances Darrington (Washington, 48°15’N, 121°36’W) and Astoria (Oregon, 46°11’N,
123°29’W) were sown as a grid. Coordinates of seed positions were required to document
single plant development. A scale of fifteen different development stages and morphological
characteristics was laid down. The light regime was classified into 100%, 74%, 55% and 35%
relative radiation compared with open field conditions. As a second factor, humidity classes
with 100ml, 500ml and 1000ml watering per pot were chosen. These humidity classes were
kept constant by daily watering to balance the differences caused by transpiration and
evaporation. The number of successful germinated seedlings was counted all two days,
respectively all seven days. In addition, germination ability was controlled by tests confirmed
with ISTA (1993, ‘International Seed Testing Association’). For that optimal humidity
conditions in germinators were established in the greenhouse chamber. Furthermore, both
provenances of Douglas-fir seedlings were classified by the scale of morphological stages.
After a period of 2½ months all seedlings were harvested and measurements of shoot and root
parameters (e.g. length, dry matter) were done.
Results and Discussion
First, as result of controlled germination tests (in germinators), no effects were found for
germination rate or speed caused by light levels. Seedlings of both Douglas-fir provenances
have shown germination rates between 77% and 87%, if humidity was not limited. As one
general result of pot experiments the percentage of germination decreased by both increasing
light availability and decreasing humidity level. Consequently, highest germination rates with
72.1% (Darrington) and 33.5% (Astoria) were found on a high level of shade (35% RLI) and
54
water content (1000ml) for both provenances of Douglas-fir seedlings (General Linear Model,
ANOVA, p < 0.000). Table 1 shows that light level (a1) can explain as much as humidity (a2)
in the model of multiple linear regressions. Parameters to characterize germination speed (e.g.
R50%, PV and number of seedlings by a particular day) confirm the advantage of seeds
germinating under low light level and high humidity. In the pots, seedlings have had low
mean mortality rates (< 10%) for all combinations of light and humidity. But mortality was
tightly connected to the first development stage. In most cases successful growth of radicle
and fast contact with substrate surface decided about seedling survival in the first stage
(Hermann and Chilcote 1965, Gray and Spies 1997). In contrast to germination rate results,
mean classified development stage was lower under high shading at the end of experiment,
because after successful germination seedlings adjusted to photoautotrophic respiration (Lyr
et al. 1992).
For both provenances a characteristic light effect was shown by the differences in lengths of
hypocotyls (light 100% - 2.63cm and light 35% - 3.59cm). Hypocotyls decreased significantly
with increasing light availability (ANOVA, p ≤ 0.000). For example Lorences and Zarra
(1986) have proved that higher lengths of hypocotyls were connected with reduced cell wall
and cuticle thickness.
Table 1. Parameters of multiple linear regressions to quantify the influence of light and
humidity to relative number of Douglas-fir seedlings for the provenances Darrington and
Astoria (percentage of germination [%] = a0 + a1* light level [%] + a2 * humidity level [ml]).
These authors have estimated less seedling stability and higher risks of fungus attacks. In our
study Douglas-fir seedlings (stages 6 and 7) from Astoria have had a better growth compared
with seedlings from Darrington (Mann-Whitney-test, p ≤ 0.05). The first significant
differences in biomass production of seedlings influenced by light were provable in stage
eight (shoot length > 1cm to 2cm without hypocotyl) of seedling development. This
increasing light effect was documented for both provenances; whereas influences of humidity
levels on seedling growth were not significant in these higher development stages. Overall it
seems to be necessary to quantify environmental factors (e.g. light, humidity) with special
regard to single development stages of seedlings. This can be helpful to understand their
temporal, but decisive relevance for seedling survival rate and the creation of advantageous
establishment conditions. Some differences between the chosen Douglas-fir provenances were
found in this study, but the final interpretation of results is complex, because of variable
temperature conditions in the greenhouse.
References
CORNETT M.W., REICH P.B., PUETTMANN K.J., FRELICH L.E. 2000. Seedbed and moisture availability determine safe sites for
early Thuja occidentalis (Cupressaceae) regeneration. American Journal of Botany 87(12): 1807-1814.
HERMANN R.K., CHILCOTE W.W. 1965. Effect of Seedbeds on Germination and Survival of Douglas-Fir. Forest Research
Laboratory, Oregon State University, Research paper No.4, Corvallis. 28p.
GRAY A.N., SPIES T.A. 1997. Microsite controls on tree seedling establishment in conifer forest canopy gaps. Ecology 78(8):
2458-2473.
KNOERZER D. 1999. Zur Naturverjüngung der Douglasie im Schwarzwald: Inventur und Analyse von Umwelt- und
Konkurrenzfaktoren sowie eine naturschutzfachliche Bewertung. Dissertationes Botanicae, Band 306. Berlin-Stuttgart. J.
Cramer, Gebrüder Borntraeger Verlagsbuchhandlung. 283p.
Poster Sessions
LORENCES E.P., ZARRA I. 1986. Hypocotyl growth of Pinus pinaster seedlings. Changes in osmotic potential and cell wall
composition. Physiol. Plant. 67: 377-382.
LYR H., FIEDLER H.-J., TRANQUILLINI W. 1992. Physiologie und Ökologie der Gehölze. Jena-Stuttgart. Gustav Fischer
Verlag. 620p.
55
56
Survival and growth of Douglas-fir seedlings of different provenances;
results after 12 months
Kristina Wallertz
Swedish University of Agricultural Sciences
Asa Research station
Douglas-fir is a North American tree species that has been planted on a small scale for quite
some time in the south of Sweden. The knowledge in a larger, more practical scale is however
limited. The climate change, with warmer and maybe longer vegetation periods, has made
Douglas-fir to be a potential tree species for the future. The aim of this study was to give
some guidelines on the choice of provenances on various sites in southern Sweden.
Growth and survival of Douglas-fir seedlings was examined in a large survey study. Seeds
from six provenances, three coastal and three inland, was imported from Canada and sown at
a nursery in Falkenberg situated in southern Sweden. In total 8 sites with various site
conditions were included for the first year. Seedlings from each provenance were planted in
plots of 0.5 ha; therefore the planted area on each site was at least 3.5 ha. All sites were
scarified with a harrow or similar, and a majority of the plantations was conducted as a
mixture of Douglas-fir and hybrid larch (Larix x eurolepis), in rows. For each provenance, all
seedlings within five circle plots were marked so that they could easily be found for further
measurements.
There are several threats to Douglas-fir seedlings during the first years after plantation.
Browsing from moose and row deer, winter desiccation, spring frost and damage caused by
pine weevil are maybe the most important. The first inventory of the seedlings was conducted
in June 2010. The measurements included growth of the first year, occurrence of damage and
survival after the first winter.
Survival as well as the amount of seedlings that had no damage was significantly higher for
seedlings originating from inland provenances compared to those from the coastal region.
Highest survival was found for the provenance “Larch Hill” and lowest for “Caycuse River”
(92 and 56%, respectively). Most of the damage was classified as “unknown”, the reason why
the damage had occurred could not be known for certain. No differences between the
provenances were found regarding growth.
Poster Sessions
57
Drought tolerance of native and non-native tree species in the Alps
– results from a large scale dendroecological study
Mathieu Lévesque , Britta Eilmann, Andreas Rigling, Peter Brang
Swiss Federal Institute for Forest, Snow and Landscape Research WSL
Birmensdorf, Switzerland
Climate change projections predict an increase in the frequency, duration, and severity of
drought periods for the coming years in the Alps. Such climate anomalies could potentially
affect species composition or long-term stability of forest ecosystems. To assure that forests
can provide ecosystem goods and services in a future drier climate, the use of non-native tree
species has been seen as an option to consider in extreme cases. Therefore, it is important to
understand the drought tolerance of native and non-native tree species in order to facilitate
long-term forest planning as well as decision support in current operational forest
management. In a preliminary study conducted on a dry site in Valais, Switzerland, the nonnative Douglas-fir (Pseudotsuga menziesii (Mirb) Franco) and European black pine (Pinus
nigra Arn.) showed to have higher resistance to drought than co-occurring native European
larch (Larix decidua Mill.) and Scots pine (Pinus sylvestris L.).
To verify this ranking in drought tolerance we analysed the growth response to drought of
native and non-native species under various site conditions in a large scale study using at least
80-year-old afforestations and plantations over the alpine arc. Several stands where native
(Abies alba Mill., Larix decidua, Picea abies (L.) Karst., Pinus sylvestris) and non-native
(Pinus nigra, Pseudotsuga menziesii) species co-occurred were selected according to a
moisture gradient. Species-specific responses to drought periods, particularly extreme dry
years such as those of 1976 and 2003, and subsequent recovery were assessed using
dendroecological methods.
Our poster will present and discuss preliminary results of this large scale study started in
spring 2010. Focus will be on Douglas-fir specific reaction to drought on contrasting dry and
mesic sites.
58
Nutrient removal by wood harvesting in Douglas-fir stands
for different silvicultural treatments and varying harvesting intensities
Joachim Block1, Julius Schuck1 and Thomas Seifert2
1
Forschungsanstalt für Waldökologie und Forstwirtschaft Rheinland Pfalz
2
Department for Forest and Wood Science, Stellenbosch University, South Africa
Conservation of the ecosystem nutrient potential is a basic precondition for sustainable forest
management. Especially on poor sites, nutrient export through harvest of wood or woody
biomass represents a relevant objective in nutrient balances of forest ecosystems.
Forest biomass and nutrient removal were calculated for stands of Douglas-fir [Pseudotsuga
menziesii (Mirbel) Franco] in comparison to Scots pine [Pinus silvestris L.], European beech
[Fagus sylvatica L.] and sessile oak [Quercus petraea (Matt.) Liebl.] on nutrient poor sandy
soils on Triassic sandstone in the Palatinate Forest. The estimation of biomass and nutrient
removal is based on scenario simulations with the growth simulator SILVA (SEIFERT et al.
2006). The growth models of SILVA were adjusted to reflect the tree growth of the different
stand types for the typical site condition of Triassic sandstone in the Palatinate Forest.
Furthermore, SILVA was extended to estimate biomass and nutrient content in several
aboveground tree compartments. The calculations were conducted for different silvicultural
treatments and for different harvesting intensities (BLOCK et al. 2007). The calculations were
based on a detailed assessment of aboveground biomass and nutrient (N, P, K, Ca, Mg)
distribution of 15 (Douglas-fir) up to 67 (beech) sample trees. The Douglas-fir sample trees
had a range of diameter at breast height (dbh) between 18 and 74cm taken from three 32, 71
and 92-year-old stands.
In the oldest Douglas-fir stand 90 percent of the biomass but only 40-60 percent of the
nutrient stock is distributed to the stem. The silvicultural treatment (thinning from below,
crown thinning and selective thinning with 100 elite trees) had only minor effects on the
nutrient removal. In contrast, raised harvesting intensity increased the nutrient export
significantly. Whole tree harvesting increased the nutrient removal in comparison to a low
intensity (harvest of stemwood of trees > 30cm dbh) by factor 3 (tab. 1). In relation to other
tree species growing on this site, wood harvesting of Douglas-fir resulted in a comparatively
small export of Ca, Mg and K - despite its much higher biomass extraction (tab. 2). This
surprising finding could be explained by very low levels of these nutrients in the heartwood of
Douglas-fir. Whereas the concentration of these nutrients in sapwood of Douglas-fir is as high
as in pine and European larch, heartwood concentration is in Douglas-fir 85 – 96% lower than
the respective concentration in the other species (fig. 1). Obviously, in Douglas-fir relocation
of these important nutrients before heartwood formation is much more efficient than in Scots
pine, European larch and sessile oak.
References
BLOCK, J., SEIFERT, TH., SCHUCK, J. (2007). Einfluss der waldbaulichen Behandlung und der Holznutzung auf den
Nährstoffhaushalt von Traubeneichenökosystemen. Mitteilungen aus der Forschungsanstalt für Waldökologie und
Forstwirtschaft Rheinland-Pfalz 63/07: 117-150
SEIFERT, TH., SCHUCK, J., BLOCK, J., PRETZSCH, H. (2006): Simulation von Biomasse und Nährstoffgehalt von Waldbäumen.
In: Deutscher Verband Forstlicher Forschungsanstalten, Sektion Ertragskunde: Beiträge zur Jahrestagung 29.- 31.05. 2006 in
Staufen: 209-224
Poster Sessions
Table 1:
59
Biomass and nutrient removal in Douglas-fir ecosystems for different harvesting
intensities
Harvesting intensity
Biomass
N
P
t/ha · year
K
Ca
Mg
kg/ha · year
whole tree 1)
8,81
12,15
0,93
5,72
7,50
0,96
very high 2)
7,53
9,66
0,66
4,00
3,66
0,57
conventional 3)
6,72
4,96
0,43
2,65
3,24
0,39
low 4)
5,83
4,15
0,31
1,94
2,76
0,30
1)
whole tree harvesting during the whole rotation period 2) stemwood with bark and whole tree havesting in
stand age 22-52 years 3) stemwood with bark, trees > 17 cm dbh 4) stemwood with bark, trees > 30 cm dbh
Table 2:
Biomass and nutrient removal for Douglas-fir in comparison to other stand types
(thinning from above, conventional harvest intensity)
Stand type
Biomass
Ca
K
t/ha · year
Mg
N
P
kg/ha · year
Douglas-fir
6,9
3,3
2,8
0,4
5,1
0,5
Pine with beech
3,5
3,7
2,0
0,7
2,5
0,2
Beech
4,2
4,1
3,8
1,1
4,6
0,3
Oak with beech
4,6
8,0
4,5
0,8
7,7
0,4
Figure 1: Box-plots of potassium concentration in sapwood (left figure) and heartwood (right
figure) of different tree species on Triassic sandstone in the Palatinate Forest
60
The commercial development of Douglas-fir
controlled mycorrhization in France:
an emerging tool for a new sylviculture
1
J. Garbaye, 1D. Bouchard, 1J.L. Churin, 1F. Le Tacon, 2V. Naudet, 3B. Robin
1
INRA, Interactions Arbres-Microorganismes
Champenoux, France
2
Pépinières Naudet, Leuglay, France
3
Pépinières Robin, Saint Bonnet en Champsaur, France
Douglas-fir has been the main reforestation tree species used in France for several decades,
with 14 millions plants sold in 1992 and 8.3 millions in 2009. As all trees belonging to the the
family Pinaceae, Douglas-fir forms obligate symbiotic organs called ectomycorrhizas that
associate fine roots and specialized soil fungi. The fungi are beneficial to tree growth and
fitness through improved nutrition, growth regulators and protection against pathogens.
However, it has been demonstrated experimentally that a given tree species, for instance
Douglas-fir, can harbour many different fungi, but that these fungal associates differ widely in
terms of growth stimulation. This functional diversity of the symbionts is the basis for
controlled mycorrhization, a forestry technique that consists in selecting particularly efficient
ectomycorrhizal fungal strains and inoculating them to the tree seedlings in the nursery,
resulting in planting stocks performing better after outplanting, long enough at least (aroung
10 years) to ensure significant benefits in terms of maintenance during the early stage of the
plantation.
In France, a research project for the controlled mycorrhization of Douglas-fir has been
developed in collaboration between private forest nurseries and INRA (National Institute for
Agronomy Research). It has resulted in the selection of the strain S238N (from the native area
of Douglas-fir in North America) of the ectomycorrhizal fungus Laccaria bicolor. Highperformance Douglas-fir planting stocks massively mycorrhizal with this fungal strain are
now available on the market, with a high level of colonization by the selected strain certified
by an INRA control, and have been used by forest owners since 1995. Both bare-root and
containerized seedlings are proposed. The use of these mycorrhiza-controlled planting stocks
changes the usual way of creating and managing Douglas-fir plantation forests because of
faster initial growth and reduced cleaning expenses because of quicker crown closer and of
the selective benefit of ectomycorhizal inoculation towards Douglas-fir (most of the
competing woody and herbaceous plants harbour another type of mycorrhizal symbiosis and
cannot associate with Laccaria bicolour). The number of such planting stocks sold yearly is
still limited (55,000 to 75,000) but is rapidly increasing.
The poster will detail the experimental strategy used and the different steps of the project that
led to the present commercial application: choice of candidate fungal strains, long-term
screening in the nursery and in a network of experimental plantations, final selection of strain
S238N, its qualities, the production of mycelial inoculant by the nurseries, the seedling
cultivation techniques, the principle of the INRA-nurseries commercial licence and of the
INRA control and label, and the perspectives of the process in the scope of the present
condition of Douglas-fir sylviculture in Europe.
Poster Sessions
61
Douglas-fir forests in Rhineland-Palatinate
– structural diversity and future perspectives
Ulrich Matthes
Research Institute for Forest Ecology and Forestry of Rhineland-Palatinate
Trippstadt, Germany
Rhineland-Palatinate is, with 6% of its forested area, among the 16 federal states of Germany
the state with the highest amount of Douglas-fir, as well as it has the biggest absolute area
(approx. 46.000 ha) occupied by this tree species. Douglas-fir is characteristic of many parts
of the forest landscapes in Rhineland-Palatinate. However, the amounts vary between less
than 3 % and more than 13 %. Likewise, the forest stand types and the mixture forms are
quite different and have a space-temporal dynamics in the landscapes.
A forest structure analysis delivered valuable insight referring to the amount of pure Douglafir stands as well as to the mixture types, within this tree species occurs. Regarding a strategy
of risk diversification stressed in the context of climate change, the question of the mixtureform and the form-index will be particularly interesting. On this way it can be estimated, for
example, whether the occurrence of potential abiotic or biotic impacts could lead to large gaps
in the forest landscape.Concerning climate change, the area of Douglas-fir is expected to
expand. As an expert survey has showed, in terms of rate the species shall take exactly that
surface which Norway spruce shall loose, due to its susceptibility to dry periods and insect
damages.
Whether Douglas-fir is, admittedly, the new "dry spruce", and how it shall cope with climatic
changes will depend on the tree suitability against the different climatic change projections.
Besides, the climatic parameters, like temperature and precipitation, other factors must be as
well considered, such as the soil water budget. In this context, also the different Douglas-fir
provenances and races are of major importance. Rhineland-Palatinate is at this level involved
in a national wide provenance trial.
From the nature conservationist’s point of view, Douglas-fir in Germany is under the status of
neophytic species, and its invasive potential is clearly a strong inconvenience. During the
presentation will be discussed, among other things, conditions, criteria or arguments relating
to the forest sites selection for the cultivation of Douglas-fir. Referring to some aspects of the
biodiversity and structural diversity of Douglas-fir forests, a few conclusions as well as open
research questions about the future contribution of Douglas-fir in Rhineland-Palatinate can be
drawn.
62
Effects of Climate change on growth and vitality of Douglas -Fir plantations
In Caspian Forest (Iran)
Farshad Yazdian
Chaloos Branch Islamic Azad University
Like other regions of the world climate change effects has happened in Iran forest. Reduced
rainfall or drought during the past 10 years on the other hand irregularities in the temperature
have caused to change Douglas trees in the growing factors. About 40 years ago, two hundred
Douglas trees planted in a region in Central of Caspian Forest. These trees planted between
Beech stand. About 20% of annual rainfall during the past 10 years has declined (from
1250mm in 1999 to 1050mm in 2010). Annual changes of temperature also in this period
have been quite sensible. The above changes reduced growth of Douglas trees. About 20% of
all trees have been completely dried during the past 10 years. Study showed that the annual
diameter growth has been reduced from 12mm (1999) to 7 mm (2010).Populations of pests
during the 5 years has been extremely rebellion. For example some insects like Lymantria
dispae also appeared on Beech trees, now appears on the Douglas trees too.Over 50% of
Douglas trees have dried branches at the top of crown. Presence of leaves on the branches has
declined from 7 years to 5 years. This study showed that climate change has been the main
reason for these problems.
Introduction
About 40 years ago, the number of 200 Douglas fir trees in the Caspian forests with wet
climate was planted Spacing is about 4 meter (Yazdian 2004). The trees cultivated in
managed forests of Farim Company with the aim of consistency check and finally to produce
wood. The seeds provenance is from the forests of west Canada. This stand located in altitude
800 meter on the North Slope.
Materials and Methods
To determine the qualitative and quantitative characteristics of this stand, all of trees have
been measured. Factors such as diameter at breast height, full height of the tree, height of tree
crown and diameter of tree crown have been measured. Crown condition of trees dried
branches and the diseases and pests were also evaluated. About growth were provided this
survey was conducted for last 10 years. (1999-2009) Climatology statistics received from the
nearest meteorological stations to the region and two factors such as annual precipitation and
annual mean temperate have been evaluated.
Results
There are about 200 Douglas trees in this stand. Their diameters are from 48 to 58 cm. Thus,
they are in two diameter classes 47.5- 52.5 and 52.5 – 57.5 about 20% of trees are dead now
and almost 50% of trees have dead branches in top of crown. Other results are shown in table.
Study on wood discs showed that the annual mean increment is evident in the past 10 years.
This process results in comparison with changes has been investigated for 10 years.
Correlation test between annual precipitation and annual mean increment are shown that
strong correlation exists. Results of Pearson and Non- parametric correlation are in table 3.
Poster Sessions
63
Regression model between the relevant factor was calculated, according to this model
R2=0.785 (figure 1)
Figure 1: regression line with annual precipitation and annual periodic increment. Other
results of this study identified some pests such as Lymanteria dispare during the past five
years that considered an important point.
There is a strong relationship between growth rate reduction and climate change. It should be
noted however that this trees have been planted in a non-native environment. As climate
change has been effected on Douglas fir trees more Beech Trees. (M.Mohajer2005)
References
M.MOHAJER, M 2005. Silviculture, University of Tehran, 387 p
YAZDIAN,F 2004. Forests of Iran, Research Institute of Forest and Rangelands.55p.
64
Production and environmental functions of Douglas-fir on the School
Training Forest Kostelec nad Černými lesy territory
Vilém Podrázský, Jiří Remeš
Czech Republic
Douglas-fir represents the most promising introduced tree species in the Europe, including
Czech countries, cultivated here on 4,800 ha, representing 0.2 % of the total forest land. On
the area of the School Training Forest Kostelec nad Černými lesy (CULS Prague) it is planted
since 1880ies, occupying 10.5 ha at present. Studies were provided in the stands, aiming the
evaluation of the growth and production potential and the soil effects of Douglas-fir in the
studied property.
The study sites are typical for relatively large areas of the central Bohemia region, the altitude
ranges between 350 – 450 m, the average annual temperature 7.5 – 8.5 ºC, precipitation 550 –
650 mm. Sites are acid to fresh, soil are Luvisols and Luvic Cambisols, 3rd – 4th vegetation
altitudinal zone. In the oldest stand (410 m a.s.l., 650 mm, 8 ºC, gleyic Luvisol, 97 years) the
standing volume was recorded between 830 – 1030 m per ha, depending on the Douglas-fir
share (14 – 30 % Nr. of trees, 32.4 – 42.4% basal area, 36.6 – 58.3 standing volume). Natural
regeneration occurs after chemical weed control reaching 16,000 – 31,000 pcs per ha.
Seedlings disappear in older age in the relatively dark stand.
Next stand represents change from site-natural hardwoods (61 years) to the monocultures of
Norway spruce (61 years) and Douglas-fir (45 years). Site is relatively acid (420 m a.s.l., 8.5
ºC, 550 – 650 mm, Luvisols). The standing volume was 266 m3ha-1 for hardwoods, 507 m3ha1
for spruce and 579 m3ha-1 for Douglas-fir. Average annual increment was evaluated as 4.43
m3ha-1.a-1 for hardwoods, 8.45 m3ha-1.a-1 for spruce and 12.87 m3ha-1.a-1 for Douglas-fir.
Humus forming effects were very favorable.
Last case documents the production and soil forming effects of the Douglas-fir compared to
Scots pine, Norway spruce and white birch (all 39 years) on afforested agricultural land (430
m a.s.l., 7.5 ºC, 600 mm, gleyic Luvisols). The standing volume reached: pine, spruce, birch
and DF 352, 349, 157 and 438 m3ha-1, the average annual increment 9.03, 8.96, 4.03 and
11.25 m3ha-1.a-1 respectively. Soil characteristics were the most favorable comparing to other
conifers: pH, soil adsorption complex characteristics, nutrient content, surface humus and
nitrogen dynamics. Douglas-fir confirmed its position as the most productive species with
favorable effects on the soil status.
Poster Sessions
65
Growth performance and reaction to biotic factors of
Douglas-fir provenances in northwest Germany
Mirko Liesebach
Johann Heinrich von Thünen-Institut (vTI)
Federal Research Institute for Rural Areas, Forestry and Fisheries
Institute of Forest Genetics
Großhansdorf, Germany
Seed of 54 provenances of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) from 8 states
of the USA and 2 provenances from Canada collected within the natural range of the species
was sown in 1962. In spring 1965, a field trial was established in the forest district of
Nordhorn, northwestern Germany, as part of the Emsland afforestation program to test the
suitability of Douglas-fir provenances on former agricultural land. In the field trial several
traits were measured or assessed regarding growth performance, and susceptibility to biotic
agents. Regarding theses characters the provenances show a great variation. Regarding
growth considerable changes in ranking were still observed between age 11 and age 33. At the
age of 33 the provenances from coastal Oregon and Washington hat the largest diameters.
These provenances were as well resistant to Rhabdocline needle cast disease. Provenances
from the southern interior were partly heavily attacked by the fungus. Some of them failed
totally at age 33. To assess the quality of provenances, especially for exotic tree species, older
aged provenance trials are essential.
BERICHTE FREIBURGER FORSTLICHE FORSCHUNG
Verzeichnis der bisher erschienenen Hefte
ISSN 1436 -1566
Heft
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Waldbau-Institut, Univ. Freiburg. 91 S.∗
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Ergebnisse FWJ 1997. FVA, Freiburg. 91 S.
4 (1999) LABUDDA, V. (Bearb.): Die Bestandesstruktur des Bannwaldes "Grubenhau" im Forstbezirk Langenau
bei Ulm (Baden-Württemberg). Ergebnisse der Forstlichen Grundaufnahmen 1986 und 1997. FVA,
Freiburg. 55 S.
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Ergebnisse der Forstlichen Grundaufnahme 1994. FVA, Freiburg. 31 S.
6 (1999) HANKE, U.; PISOKE, TH.: Der Sturmwurfbannwald "Teufelsries": Waldstrukturerhebung mit
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Kolloquium des Arbeitskreises "Waldböden" der DGB und der Sektion Wald und Wasser im
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Fachtagung an der Albert-Ludwigs-Universität Freiburg am 3. und 4. Dezember 1998. Institut für
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10 (1999) WILWERDING, A.; RÖSCH, CH.: Anbau, Bereitstellung und energetische Nutzung holzartiger Biomasse
von Kurzumtriebsflächen aus ökologischer und ökonomischer Sicht. FVA, Freiburg. (unveröffentl.)
11 (1999) HANKE, U.; AHRENS, W.: Bannwald "Großer Trauben" im Forstbezirk Pfullendorf (BadenWürttemberg). FVA, Freiburg. 33 S.
12 (1999) WEBER, J.; WOTKE, S.A.; BÜCKING, W.: Bannwald "Sommerberg" im Forstbezirk Eppingen (BadenWürttemberg). FVA, Freiburg. 55 S.
13 (1999) SEEMANN, D. (Bearb.): Die Massenvermehrung des Schwammspinners (Lymantria dispar L.) in
Baden-Württemberg 1993-1994. FVA, Freiburg. 235 S.
14 (1999) BRANDL, H.; HERCHER, W.; LÖBELL, E.; NAIN, W.; OLISCHLÄGER, G.; WICHT-LÜCKGE, G.: 20 Jahre
Testbetriebsnetz Kleinprivatwald in Baden-Württemberg: Betriebswirtschaftliche Ergebnisse 19791998. FVA, Freiburg. 122 S.
15 (1999) KEGLER, H.-H.: Der Bannwald „Hechtsgraben“ im Forstbezirk Kenzingen (Baden-Württemberg).
Ergebnisse der Forstlichen Grundaufnahmen 1981 und 1994. FVA, Freiburg. 39 S.
16 (1999) Die Bedeutung der forstlichen Vegetationskunde für den naturnahen Waldbau – Ansprüche,
Möglichkeiten und Grenzen. Standortskundlich-Waldbauliches Kolloquium der FVA am 13. April
17 (2000) EISFELD, D.; HUSS, J.; OESTEN, G.; UERPMANN, B.; VOLZ, K.-R. (Hrsg.): Wald und Schalenwild.
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1999. Institut für Forstökonomie, Univ. Freiburg. 177 S.*
18 (2000) ULLRICH, TH.: Der Bannwald „Eiberg“ im Forstbezirk Bad Wildbad (Baden-Württemberg).
Ergebnisse der Forstlichen Grundaufnahme 1998. FVA, Freiburg. 48 S.
19 (2000) RAU, H.; BRANDL, H.: Zwei Jahrhunderte Forstgeschichte Baden-Württemberg. Von der KöniglichWürttembergischen Forstdirektion zur Forstdirektion Stuttgart. FVA, Freiburg. 191 S.
20 (2000) WEBER, J.: Geostatistische Analyse der Struktur von Waldbeständen am Beispiel ausgewählter
Bannwälder in Baden-Württemberg. FVA, Freiburg. 133 S.
21 (2000) BRANDL, H.; BÜRGI, M.; JOHANN, E.; SCHUMACHER, W.; SCHMIDT, U.E.; VOLK, H.; VOLZ, K.-R.:
Beiträge zur Forstgeschichte. FVA, Freiburg. 87 S.
22 (2000) PISOKE, TH.: Die Waldstruktur im Bannwald Taubergießen. Eine Luftbildauswertung im
strukturreichen Auewald. FVA, Freiburg. 96 S.
23 (2000) GAERTIG, T.; SCHACK-KIRCHNER, H.; VOLKMANN, J.; WILPERT, K. V.: Physikalische und chemische
Bodeneigenschaften als prädisponierende Faktoren neuartiger Eichenschäden. FVA, Freiburg u. Inst.
für Bodenkunde, Univ. Freiburg. 95 S.
∗
Bestellung über Waldbau-Institut, Universität Freiburg, e-mail: waldbau@uni-freiburg.de
Euro
vergriffen
6,50
3,00
4,00
2,50
4,00
6,00
10,20
5,00
3,50
4,00
6,50
4,00
3,50
6,00
13,30
6,00
12,00
12,00
9,00
13,00
9,00
24 (2000) AHRENS, W. ; GERBOTH, G. ; LABUDDA, V. ; LUDEMANN, T. ; SCHOTTMÜLLER, E.: Bannwald
"Feldseewald " im Schwarzwald im Forstbezirk Kirchzarten (vorher Schluchsee). FVA, Freiburg.
130 S.
25 (2001) Wiederbewaldung von Sturmschadensflächen. Beiträge zu einem gemeinsamen Kolloquium von
Forstwissenschaftlicher Fakultät der Universität Freiburg und Forstlicher Versuchs- und
Forschungsanstalt Baden-Württemberg am 21. und 22. September 2000 in Freiburg. FVA, Freiburg
u. Waldbau-Institut, Univ. Freiburg. 148, XIII S.
26 (2001) SCHRÖTER, H.; BECKER, T.: Die Ausbreitung des Borkenkäferbefalls im Bereich von SturmwurfSukzessionsflächen. Eine Untersuchung in Sturmwurfbannwäldern Baden-Württembergs. FVA,
Freiburg. 79 S.
27 (2001) BARON, U.; BRANDL, H.; HERCHER, W.; NAIN, W.: Testbetriebsnetz der Forstwirtschaft in BadenWürttemberg. Betriebswirtschaftliche Ergebnisse der Waldbesitzarten im FWJ 1999. FVA, Freiburg.
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28 (2001) SEILER, W.: Erläuterungen zum Bannwald "Pfannenberg". Forstbezirk Kirchheim (Teck). FVA,
Freiburg. 46 S.
29 (2001) BÜCKING, W.; GEISCHER, B.; WINTERHOFF, W.; WOLF, T.: Die Bannwälder "Franzosenbusch" und
"Kartoffelacker" in der Schwetzinger Hardt. Baden-Württemberg, Forstbezirk Schwetzingen. FVA,
Freiburg. 147 S.
30 (2001) WOHLFAHRT, D.; RIEDEL, P.: Bannwald "Wilder See - Hornisgrinde". Forstbezirk Schönmünzach.
FVA, Freiburg. 68 S.
31 (2001) KELLER, F.; RIEDEL, P.: Bannwald "Zweribach". Forstbezirk St. Märgen. Erläuterungen zur
Forstlichen Grundaufnahme 1999. FVA, Freiburg. 64 S.
32 (2001) 10 Jahre Waldentwicklung nach Sturm "Wiebke". Untersuchungen in Fichten-Sturmwurfbannwäldern Baden-Württembergs. FVA, Freiburg. 205 S.
33 (2001) Chemische und physikalische Schlüsselprozesse der Speicher-, Regler- und Reaktorfunktionen von
Waldböden. Gemeinsamer Workshop des Arbeitskreises "Waldböden" der DBG u. der Sektion
"Wald und Wasser" im DVFF in Freiburg vom 5. bis 6. April 2001. FVA, Freiburg. 208 S.
34 (2001) BARON, U.; HERCHER, W.; NAIN, W.; PISTORIUS, T.: Testbetriebsnetze der Forstwirtschaft in BadenWürttemberg. Betriebswirtschaftliche Ergebnisse der Waldbesitzarten im FWJ 2000 (Sturmjahr
"Lothar"). FVA, Freiburg. 100 S.
35 (2001) BOLKENIUS, D.: Zur Wurzelausbildung von Fichte (Picea abies L. [Karst]) und Weißtanne (Abies alba
Mill.) in gleich- und ungleichaltrigen Beständen. FVA, Freiburg. 155 S.
36 (2002) ABEL, U.; RIEDEL, P.: Bannwald "Scheibenfelsen". Forstbezirk Kirchzarten. FVA, Freiburg. 47 S.
37 (2002) KOCH, B. (ed.): Spatial technologies for forest inventories. Contributions to the XXI IUFRO World
Congress. Institut für Fernerkundung u. Landschaftsinformationssysteme, Univ. Freiburg. 238 S.∗
38 (2002) LOCH, R.: Statistisch-ökologischer Vergleich der epigäischen Spinnentierfauna von Bann- und
Wirtschaftswäldern. Beitrag zur Erforschung der Biodiversität heimischer Wälder. FVA, Freiburg.
311 S.
39 (2002) Orkan Lothar: Katastrophe für die naturgemäße Waldwirtschaft ? Oder: Über den Umgang mit
Störungen im Nordschwarzwald. Tagungsband der ANW-Bundestagung vom 12.-14. Juni in BadenWürttemberg. FVA, Freiburg. 60 S.
40 (2002) AHRENS, W.; GERTZMANN, CH.; RIEDEL, P.: Bannwald "Hoher Ochsenkopf". Forstbezirk Forbach.
Erläuterungen zu den Forstlichen Grundaufnahmen 1985 und 1995. FVA, Freiburg. 75 S.
41 (2002) NOWACK, A.: Bannwald "Bruchsaler Bruch". Forstbezirk Bruchsal. Erläuterungen zur Forstlichen
Grundaufnahme 2000. FVA, Freiburg. 50 S.
42 (2002) BANTLE, P. (Bearb.): Bannwald "Kleiner Imberg". Forstbezirk Eberbach. Erläuterungen zur
Forstlichen Grundaufnahme 1999. FVA, Freiburg. 36 S.
43 (2002) Holzasche-Ausbringung im Wald, ein Kreislaufkonzept. FVA-Kolloquium in Freiburg vom
5. bis 6. März 2002. FVA, Freiburg. 166 S.
44 (2003) Großkronige Buchen – ein Konzept zur Wertholzerzeugung? Kolloquium 19. März 2002, Freiburg
im Breisgau. FVA, Freiburg. 80 S.
45 (2003) Vom Wert des Waldes jenseits von Aufwand und Ertrag. Kolloquium 17. September 2002, Freiburg
im Breisgau. FVA, Freiburg. 87 S.
46 (2003) GÜRTH, P. (Bearb.): Geschichte des Waldbaus in Baden-Württemberg im 19. und 20. Jahrhundert. Ein
Projekt der LFV Baden-Württemberg. FVA, Freiburg. 224 S.
47 (2003) HANSEN, J.; SPIECKER, H.; TEUFFEL, K. v. (eds.): The question of conversion of coniferous forests.
Abstracts. Internat. Conference 27 September – 02 October, 2003, Freiburg im Breisgau, Germany.
48 (2003) MÜLLER, U.; STREIN, M.; SUCHANT, R.: Wildtierkorridore in Baden-Württemberg. FVA, Freiburg.
45 S.
49 (2003) Boden und Wasservorsorge. Gemeinsames Kolloquium des Arbeitskreises "Waldböden" der DBG
u. der Sektion "Wald und Wasser" im DVFFA in Freiburg vom 13. bis 15. Mai 2003. FVA, Freiburg.
229 S.
∗
13,00
11,80
15,00
7,00
7,00
13,00
10,00
10,00
18,00
9,00
8,00
16,00
7,00
10,00
23,00
5,50
11,00
11,00
11,00
12,00
6,00
6,00
20,00
5,00
8,00
15,00
50 (2003) HANEWINKEL , M.; TEUFFEL, K. V. (Hrsg.): Waldwachstumsmodelle für Prognosen in der
Forsteinrichtung. Stand der Wissenschaft in Mitteleuropa. FVA, Freiburg. 102 S.
51 (2004) International Symposium on Contributions of Family-Farm Enterprises to Sustainable Rural
Development, 28 July – 01 August 2002, Gengenbach, Germany. Proceedings. FVA, Freiburg. 388 S.
52 (2004) HAUFFE, PH. : „Weiterentwicklung der automatisierten Rundholzvermessung hinsichtlich der
Erfassung und Quantifizierung äußerer und innerer Holzeigenschaften“. KAISER, R.: „Weiterentwicklung des Anforderungskataloges für die Werksvermessung von Stammholz“. FVA, Freiburg.
80 S.
53 (2004) KÖHL, M.; DEES, M.: Statistische Analyse von Kronenzustandsdaten aus der Interpretation von CIRLuftbildern. Univ. Freiburg. 29 S.*
54 (2004) HUSSENDÖRFER , E.; ALDINGER, E. (Hrsg.): Herkunftssicherung und Zertifizierung von forstlichem
Vermehrungsgut. FORUM Genetik-Wald-Forstwirtschaft. Arbeitstagung vom 11. – 13. Juni 2001 an
der FVA Freiburg. FVA, Freiburg. 159 S.
55 (2004) BARON, U.; FILLBRANDT, TH.; HARTEBRODT, C.; HERCHER, W.: Testbetriebsnetze der Forstwirtschaft
in Baden-Württemberg. Betriebswirtschaftliche Ergebnisse der Waldbesitzarten im FWJ 2001 und
56 (2004) Umweltüberwachung und Waldwachstum. Ehrenkolloquium zum 75. Geburtstag von Prof. Dr. HansUlrich Moosmayer am 23. September 2003. FVA, Freiburg. 97 S.
57 (2004) Impacts of the Drought and Heat in 2003 on Forests. Scientific Conference 17 – 19 November 2004
Freiburg, Germany. Abstracts. FVA, Freiburg. 70 S.
58 (2005) Waldnaturschutz. Forschung, Planung, praktische Umsetzung. Kolloquium der Abt. Landespflege der
FVA zur Standortsbestimmung im Bereich Wald und Naturschutz am 15. Juni 2004 in Freiburg.
59 (2005) HUG, R.; HEPP, R.; WILPERT, K. V. : 18 Jahre Depositionsmessnetz der Forstlichen Versuchs- und
Forschungsanstalt Baden-Württemberg. FVA, Freiburg. 95 S.
60 (2005) BAUHUS, J.; CSAPEK, G.: Beiträge zur Tagung 2004 der Sektion Waldbau DVFFA, 6 – 8 September in
Freiburg/Staufen. Waldbau-Institut, Univ. Freiburg. 147 S.*
61 (2005) Differenzialdiagnostische Untersuchungen zu Eichenschäden in Baden-Württemberg. Ansprache,
Faktoren, Schlussfolgerungen. FVA, Freiburg. 193, 66 S.
62 (2005) Wasservorsorge in bewaldeten Einzugsgebieten. Gemeinsames Kolloquium des Arbeitskreises
"Waldböden" der DBG u. der Sektion "Wald und Wasser" im DVFFA in Freiburg vom 02. bis
03. Juni 2005. FVA, Freiburg. 240 S.
63 (2005) VEIT, H.; BECKER, T.: Untersuchungen zu den Befallsbedingungen, zum Befallsablauf und zur
Abwehr des Befalls von Laub- und Nadelholz durch den sägehörnigen Werftkäfer (Hylecoetus
dermestoides Latr. (Coleoptera, Lymexylidae)). FVA, Freiburg. 111 S.
64 (2005) Menschen für Raufußhühner. Beiträge des Statuskolloquium der Arbeitsgruppe Raufußhühner in
Freiburg, Baden-Württemberg am 10. Mai 2005. FVA, Freiburg. 249 S.
65 (2006) HOHENLOHE-WALDENBURG, F.K. zu: Zur Waldgeschichte des Hohenloher Landes. FVA, Freiburg.
102 S.
66 (2006) HERBSTRITT, S.; KOHNLE, U.; ABETZ, P.; KENK, G. (eds.):The European stem number experiment in
Norway spruce (Picea abies (L.)Karst.); 3. report. IUFRO working party 1.05.05 "Thinning
Experiments". FVA, Freiburg. 132 S.
67 (2006) ORF, S.: Neue Entwicklungen im Forstrecht. Gesetzgebung, Verwaltung und insbes. Rechtsprechung.
68 (2006) BAUHUS, J.; SCHERMBECK, J. (eds.): Ecosystem goods and services from planted forests. Abstracts of
the scientific forum of the International Congress on Cultivated Forests, 3rd - 7th October 2006,
Bilbao, Spain. Waldbau-Institut, Univ. Freiburg. 236 S. + 1 CD ROM∗
69 (2006) IVITS, E.; HEMPHILL, S.; LANGAR, F.; KOCH, B: Optimisation of land cover and forest habitats
classification for the monitoring of habitat directive areas in Thuringia, Germany: advantages and
disadvantages of pixel- and object-based classification methods based on the Spot 5 sensor. Univ.
Freiburg. 86 S.*
70 (2007) Europäische Flußauen und Auewälder. Ergebnisse des Kolloquiums vom 26. bis 28.04.2006. FVA,
Freiburg. 205 S.
71 (2006) HARTEBRODT, C.; FILLBRANDT, T.; HERCHER, W.; HAAGER, H.; BITZ, S.; HOLTHAUSEN, N.: Wald Besitz - Ökonomie - 2006. FVA, Freiburg. 113 S.
72 (2006) BUB, G.; DELB, H.; SCHRÖTER, H.: Die Gradation des Schwammspinners (Lymantria Dispar L.)
2004/2005 in Baden-Württemberg. FVA, Freiburg. 64 S.
73 (2007) PISTORIUS, T.; ZELL, J.; HARTEBRODT, C.: Untersuchungen zur Rolle des Waldes und der
Forstwirtschaft im Kohlenstoffhaushalt des Landes Baden-Württemberg. FVA, Freiburg. 182 S.
74 (2007) Wald – Besitz – Ökonomie – 2007. FVA, Freiburg. 152 S.
75 (2007) WILPERT, K. V.; NIEDERBERGER, J.; PUHLMANN, H.: Fallstudien zur Bewertung und Entwicklung
forstbetrieblicher Optionen zur Sicherung der Wassergüte in bewaldeten Einzugsgebieten. Wirkung
von Bodenschutzkalkungen auf die Wasserqualität. FVA, Freiburg. 65 S.
∗
17,00
25,00
17,00
10,00
14,00
7,50
14,00
5,00
18,00
14,00
14,00
14,00
15,00
14,00
20,00
20,00
12,00
6,00
12,50
17,80
25,00
15,00
14,00
17,00
16,00
12,50
76 (2008) Forstwissenschaftliche Tagung 2008: Ressourcenknappheit und Klimaänderung, 24.-27. September
2008.Univ. Freiburg. 134 S.*
77 (2008) Der Buchdrucker (Ips typographus [L.]) im "Bannwald Napf". Rekonstruktion der
Populationsdynamik in den Jahren 1990 bis 2006. FVA, Freiburg u. Univ. Freiburg. 52 S.
78 (2008) Starkholz – Premiumprodukt oder Problemsortiment? Chance und Herausforderung für Produktion,
Bereitstellung u. Vermarktung. Fachtagung im Juli 2008. FVA, Freiburg. 120 S.
79 (2009) Walderschließung und Bodenschutz. Abschlusskolloquium in Freiburg vom 6. bis 7. März 2008.
80 (2009) BERNAUER, B.: Hiebsreifeanalyse von Waldbeständen. FVA, Freiburg. 122 S.
81 (2009) ALBRECHT, A.; KOHNLE, U.; BÄUERLE, H. et al.: Perspektiven einer nachhaltig
wertleistungsoptimierten Waldwirtschaft im waldreichen ländlichen Raum Ostalb. FVA, Freiburg.
144 S.
82 (2009) Bedeutung des Klimawandels für Bodenprozesse : gemeinsames Kolloquium des Arbeitskreises
"Waldböden" der DBG und der Sektion "Wald und Wasser" im DVFFA in Freiburg vom 24. bis 25.
April 2008. FVA, Freiburg. 226 S.
83 (2009) BÄUERLE, H.; HOLZMANN, M. (Red.): Aktivierung von Wertschöpfungspotentialen zur nachhaltigen
Nutzung und Verwendung von Nadel- und Laubstarkholz. Abschlussbericht zum BMBFVerbundprojekt Starkholz. FVA, Freiburg. 244 S.
84 (2010) HARTEBRODT, C. (Red.): Proceedings. International Workshop on Figures for Forests ; 24. 26.11.2008, Freiburg, Germany. FVA, Freiburg. 172 S.
∗
12,50
14,00
12,00
12,00
13,00
14,00
15,00
15,00
18,00

Opportunities and risks for Douglas fir in a changing climate

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