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Genetic Implications of
        Forest Management
      in the Mediterranean

             Aristotelis C. Papageorgiou

Department of Forestry, Environment and Natural Resources,
        Democritus University of Thrace, Greece



                     Solsona 2004
Presentation plan

ā€¢   Genetic diversity in Mediterranean forests
ā€¢   Factors influencing genetic diversity
ā€¢   Impact of forest management
ā€¢   SFM and genetic diversity
ā€¢   Research and policy priorities



April 4, 2004       MEDFOREX meeting             2
What is so special about the Med?

ā€¢ Biological, cultural, social, historical diversity
ā€¢ Biological diversity:
      ā€“ rich mosaic of changing ecosystems and land use
        patterns (from alpine to tropical)
      ā€“ Large amount of species (especially plants) ā€“ high
        endemism
      ā€“ Populations of species with a broader distribution
        are the most variable in terms of genetic diversity

April 4, 2004           MEDFOREX meeting                  3
Genetic diversity
ā€¢ Differentiation among individuals and groups
  of individuals (populations) exists.
ā€¢ Differentiation at the gene level can be
  inherited = genetic variation
ā€¢ Key for adaptation in changing
  environments over space and time
ā€¢ Connection between generations
ā€¢ Is usually measured in traits that are not
  important for forestry (e.g. DNA markers)
April 4, 2004       MEDFOREX meeting         4
Genetic system of a forest population




                                    Hattemer & Gillet 2000
April 4, 2004   MEDFOREX meeting                       5
Generations of forest trees




April 4, 2004            MEDFOREX meeting     6
Genetics of Mediterranean forests
ā€¢ Disjunct distribution of populations and species ā€“
  high differentiation
ā€¢ Usually high genetic variation within populations ā€“
  comparison with N/C Europe
ā€¢ Many small populations
ā€¢ Large distributions of species over different
  environments (e.g. Pinus species)
ā€¢ Unique alleles and races
ā€¢ Paradox of variation in several cases
April 4, 2004        MEDFOREX meeting                   7
Genetic diversity of forest species




                Petit et al. 2003

April 4, 2004                       MEDFOREX meeting   8
Reasons for high diversity
ā€¢ The relief of the Mediterranean basin




April 4, 2004           MEDFOREX meeting     9
Reasons for high diversity
                             ā€¢ The Mediterranean
                               climate
                             ā€¢ The existence of
                               tectonic microplates
                               and their move




April 4, 2004           MEDFOREX meeting              10
Reasons of high diversity
ā€¢ The glaciations era ā€“ refugia




April 4, 2004           MEDFOREX meeting    11
Human and forests
                                  ā€¢ Most ancient
                                    human cultures
                                  ā€¢ Centre of the
                                    ā€œknown worldā€
                                  ā€¢ Organized trade
                                    and land use
                                  ā€¢ First ā€œglobalizedā€
                                    economies

April 4, 2004       MEDFOREX meeting                 12
Direct use of plant species




April 4, 2004        MEDFOREX meeting     13
Direct use of plant species




April 4, 2004        MEDFOREX meeting     14
Human shaping nature




Agriculture, grazing,
fuelwood collection, etc.
shape the landscapeā€¦
April 4, 2004         MEDFOREX meeting   15
ā€œMosaicā€ - forest landscape




April 4, 2004      MEDFOREX meeting     16
Mediterranean forests
ā€¢ Forest includes terrestrial ecosystems in a broad sense
ā€¢ Natural forest ecosystems ā€“ mixed forests ā€“ complex age
structures ā€“ connection between generations
ā€¢ Mainly small scale activities ā€“ high land use diversity ā€“
fragmentation ā€“ mixed with other activities
ā€¢ Degradation ā€“ desertification ā€“ loss of ecosystem
productivity ā€“ land abandonment
ā€¢ High protective role against erosion, drought ā€“ social
importance for rural development
ā€¢ Recreation ā€“ quality of life ā€“ Non Wood Goods
ā€¢ Culture ā€“ spirituality

April 4, 2004            MEDFOREX meeting                     17
Human impact on gene diversity
ā€¢ Far-reaching / global
     ā€“ Emission of air pollutants
     ā€“ Greenhouse gases
     ā€“ Policy processes
ā€¢ Local
     ā€“ Forest destruction (land conversion)
     ā€“ Forest fragmentation
     ā€“ Forest management
           ā€¢ Introduction of new species & populations
           ā€¢ Silvicultural activities
           ā€¢ Other forms on management (e.g. grazing)

April 4, 2004                  MEDFOREX meeting          18
Destruction and fragmentation
ā€¢ Forest destruction (deforestation)
      ā€“ Loss of species with small scale (e.g. Abies
        nebrodensis, Quercus euboica, Cedrus brevifolia)
      ā€“ Loss of populations (differentiated ā€“ adapted)
ā€¢ Forest fragmentation
      ā€“ Reduction of effective population size ā€“ drift
        effects ā€“ genetic bottlenecks
      ā€“ Increase vulnerability of populations
      ā€“ Caused by development, agriculture, grazing

April 4, 2004           MEDFOREX meeting                 19
Genetic bottleneck

            1400
            1200
            1000
                                                 RECOVERY
                800
 N
                600
                400   CRASH
                200
                 0                  Bottleneck

                          TIME
April 4, 2004                 MEDFOREX meeting          20
Example: Pinus leucodermis in Italy
ā€¢ Only a few small
  populations
  remaining ā€“
  fragmented ā€“ no
  connection possible
ā€¢ Low genetic variation
  ā€“ large differentiation
  among populations

      Morgante & Vendramin 1991

April 4, 2004               MEDFOREX meeting   21
Artificial regeneration
ā€¢ Adaptation on the final site in question
      ā€“ Different adaptations at the original site
      ā€“ Adaptation in nurseries
ā€¢ Genetic variation (evolutionary adaptability) reduced
  due to drift effects
      ā€“ Harvest from small number of plants (founder)
      ā€“ Unknown material ā€“ provenance ignored
ā€¢ Possible impact on surrounding forests due to gene
  flow
ā€¢ However: development of new land races (e.g.
  Quercus rubra in France)
April 4, 2004               MEDFOREX meeting            22
Example: Cupressus sempervirens
ā€¢ Low genetic variation
  of planted stands
ā€¢ Canker attack in
  planted stands
ā€¢ Gene flow from
  planted stands in
  natural populations
          Papageorgiou et al. 1994

April 4, 2004                   MEDFOREX meeting   23
Natural regeneration
ā€¢ Connection between generations - maintenance of
  dynamics of genetic structures in life cycle of forest
  trees ā€“ evolution goes on
ā€¢ How many individuals participate in the next
  generation? Where?
      ā€“ Natural processes (very improbable in the Med)
      ā€“ Non-forestry activities (e.g. grazing)
      ā€“ Forestry operations (e.g. light felling)
ā€¢ Limited potential for changes of genetic structures
      ā€“ No safeguard against unintentional & random changes of
        genetic structures, losses of genetic variation


April 4, 2004             MEDFOREX meeting                   24
Silvicultural treatments
ā€¢ Low population densities
ā€¢ Reduced gene flow
      ā€“ Possible founder effects (a few seeder trees)
      ā€“ Increased inbreeding ā€“ low germinability
      ā€“ Inbreeding depression
ā€¢ Threshold values for
      ā€“ Population sizes
      ā€“ Population densities
ā€¢ Problem for scattered species
      ā€“ Particularly in species-rich mixed forests
      ā€“ Species with ā€œpeculiarā€ mating system (Taxus baccata)

April 4, 2004              MEDFOREX meeting                     25
Example: Abies cephalonica
ā€¢ Higher values of inbreeding levels in uneven
  aged forests with lower population densities




                Fady & Conkle 1993
April 4, 2004                        MEDFOREX meeting   26
Selective cutting
ā€¢ Selection against superior phenotypes
      ā€“ Selective logging with short cutting cycles
      ā€“ Exclusion of superior phenotypes from
        reproduction
ā€¢ Reduction of mating trees
      ā€“ effective population size
      ā€“ Inbreeding
ā€¢ Coppice forests
      ā€“ Clear cuts with a few remaining trees
April 4, 2004            MEDFOREX meeting             27
Is genetic diversity at stake?
ā€¢ Disturbance of human ā€“ nature balance
ā€¢ The dynamic systems are broken (genetic, ecological,
  nutrients, water, energy, etc.)
ā€¢ Ecosystem are not able to provide goods and services in the
  long term ā€“ desertification.
ā€¢ Main problems occur mainly from factors outside SFM
ā€¢ The maintenance / restoration of the ability of
  Mediterranean ecosystems to continue their
  dynamic function should become the target of
  any conservation effort in the region
April 4, 2004             MEDFOREX meeting                  28
Forest management




April 4, 2004        MEDFOREX meeting   29
Forest management in the Med
ā€¢ Management strategies & techniques imported from
  the central and northern counties of Europe -
  production of timber is priority in most cases
ā€¢ Apply on the more temperate forests of the
  Mediterranean region - ignore the non-productive
  terrestrial ecosystems (e.g. maquis)
ā€¢ Fail to capture the complexity of Mediterranean
  forests & land use systems
ā€¢ The spatial reference of SFM is the stand and not
  the broader landscape - more appropriate for the
  description of the human - nature dynamics
April 4, 2004       MEDFOREX meeting              30
Protection of forest biodiversity




     SFM as a tool to achieve biodiversity conservation
     Protects BD elements, including genes & provenances

April 4, 2004         MEDFOREX meeting               31
Forest protection in the Med
ā€¢ Protection concepts and strategies imported from
  northern counties with empty spaces and productive
  forests - human influence on BD is ignored
ā€¢ Based mainly on the ā€œset ā€“ asideā€ principle
ā€¢ Focus on absolute protection or special
  management of specific biological entities (e.g.
  genes) or spatial units ā€“ miss the broader picture
ā€¢ Restricted to reserves and networks
ā€¢ Selection of protected items ā€“ majority remains
  unprotected

April 4, 2004         MEDFOREX meeting             32
Forest conservation genetics
ā€¢ Forest genetic resources
      ā€“ Included in broader BD policies & plans
      ā€“ Most under-represented part of BD
      ā€“ Based in ā€œgenetic inventoriesā€ (assessments)
ā€¢ In situ
      ā€“ Actions of strict protection of rare or endangered entities
      ā€“ Gene reserves (special management allowed)
      ā€“ Conservation in managed ecosystems (Namkoong).
ā€¢ Ex situ
      ā€“ Protection of endangered or rare tree species
      ā€“ Storage of ā€œusefulā€ genes (gene banks)
April 4, 2004               MEDFOREX meeting                      33
An important gap
ā€¢ ā€œOther wooded landā€ ā€“ not protected, not managed
ā€¢ The classical protection / SFM approach
      ā€“ has a spatial gap of about 50%
      ā€“ cannot cope with the dynamic nature of Mediterranean
        forests
ā€¢ Conservation of forest genetic diversity cannot be
  seen separately from the general use and
  management of forest resources
ā€¢ Inventories possible in a few cases only (no time, no
  money for ā€œtotal inventoriesā€)

April 4, 2004             MEDFOREX meeting                     34
Linking SFM and BD
ā€¢ Environmental agencies consider BD as a priority in
  all management activities
ā€¢ Forestry parties see the economic aspect of forestry
  as a priority.
ā€¢ BD conservation and SFM can be linked:
      ā€“ Complexity of both concepts
      ā€“ Long term character
ā€¢ ā€œEcosystem approachā€ & ā€œclose to natureā€ forestry
ā€¢ Tools, such as SFM certification, ā€œcriteria &
  indicatorsā€, Special Management plans, NFP
April 4, 2004            MEDFOREX meeting            35
The challenge
ā€¢ Maintaining adequate levels of genetic diversity of
  tree (& other) species is crucial for adaptation and
  adaptability processes
ā€¢ Dynamic processes must maintain their role for the
  maintenance of productivity of goods and services
ā€¢ Genetic diversity is not just another goal of SFM,
  but the means to achieve it
ā€¢ We need: Management techniques that will
  prevent disturbances in critical ecological and
  genetic processes

April 4, 2004        MEDFOREX meeting                    36
The challenge




April 4, 2004     MEDFOREX meeting   37
Future research priorities
ā€¢ Develop inventory systems for genetic diversity
  status of terrestrial ecosystems (stand types)
      ā€“ Use results of existing genetic research projects
      ā€“ Plan future genetic research
      ā€“ Use models to simulate evolution under forest
        management scenarios
      ā€“ Connect gene diversity with stand & age structure,
        density, mating system, etc.
ā€¢ Develop criteria & indicators for genetic diversity
      ā€“ adjust existing efforts to the Mediterranean
      ā€“ introduce genetic criteria into existing SFM C&Is
        (certification standards, MCPFE criteria, etc.)
April 4, 2004              MEDFOREX meeting                  38
Future research priorities
ā€¢ Connect specific management techniques of
  Mediterranean forests with the previous steps
      ā€“ Give a ā€œgenetic gradeā€ for each technique
      ā€“ Evaluate human impact on genetic diversity
ā€¢ Set priorities for SFM considering genetic diversity
ā€¢ Extrapolate of appropriate strategies for most taxa
  from the results of studies of a few model cases
ā€¢ Identify genetic aspects that may become limiting
  for certain types of species and ecosystems
ā€¢ Monitoring and evaluation
April 4, 2004             MEDFOREX meeting               39
SFM for the Med

ā€¢ Develop management plans for the
  Mediterranean beyond the classical forestry
  approach, including broader ecosystems and
  activities, based on the maintenance of
  genetic diversity
ā€¢ Protecting biodiversity through management
  and planning, following rules that need minor
  assessments
April 4, 2004       MEDFOREX meeting          40
Thank you for your attention




April 4, 2004         MEDFOREX meeting     41

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Ī‘Ļ€ĪæĻ„ĪµĪ»ĪµĪÆ Ī· Ī”ĪæĪ“ĻŒĻ€Ī· ĻƒĪ·Ī¼ĪµĪÆĪæ ĻƒĻ…Ī½Ī¬Ī½Ļ„Ī·ĻƒĪ·Ļ‚ Ļ„Ī·Ļ‚ ĪµĻ…ĻĻ‰Ļ€Ī±ĻŠĪŗĪ®Ļ‚ (Ī“Ī±ĻƒĪ¹ĪŗĪ®Ļ‚) ĪŗĪ±Ī¹ Ļ„Ī·Ļ‚ Ī±Ī½Ī±Ļ„ĪæĪ»Ī¹Īŗ...Ī‘Ļ€ĪæĻ„ĪµĪ»ĪµĪÆ Ī· Ī”ĪæĪ“ĻŒĻ€Ī· ĻƒĪ·Ī¼ĪµĪÆĪæ ĻƒĻ…Ī½Ī¬Ī½Ļ„Ī·ĻƒĪ·Ļ‚ Ļ„Ī·Ļ‚ ĪµĻ…ĻĻ‰Ļ€Ī±ĻŠĪŗĪ®Ļ‚ (Ī“Ī±ĻƒĪ¹ĪŗĪ®Ļ‚) ĪŗĪ±Ī¹ Ļ„Ī·Ļ‚ Ī±Ī½Ī±Ļ„ĪæĪ»Ī¹Īŗ...
Ī‘Ļ€ĪæĻ„ĪµĪ»ĪµĪÆ Ī· Ī”ĪæĪ“ĻŒĻ€Ī· ĻƒĪ·Ī¼ĪµĪÆĪæ ĻƒĻ…Ī½Ī¬Ī½Ļ„Ī·ĻƒĪ·Ļ‚ Ļ„Ī·Ļ‚ ĪµĻ…ĻĻ‰Ļ€Ī±ĻŠĪŗĪ®Ļ‚ (Ī“Ī±ĻƒĪ¹ĪŗĪ®Ļ‚) ĪŗĪ±Ī¹ Ļ„Ī·Ļ‚ Ī±Ī½Ī±Ļ„ĪæĪ»Ī¹Īŗ...
Aristotelis C. Papageorgiou
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Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½
Aristotelis C. Papageorgiou
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Ī•Ļ€ĪÆĪ“ĻĪ±ĻƒĪ· ĪµĪ½Ī“ĪæĻƒĻ…ĻƒĻ„Ī±Ī“Ī¹ĪŗĻŽĪ½ Ļ€Ī±ĻĪ±Ī³ĻŒĪ½Ļ„Ļ‰Ī½ ĻƒĻ„Ī· Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ī“ĪæĪ¼Ī® Ļ„Ļ‰Ī½ ĻƒĻ€ĻŒĻĻ‰Ī½ ĪµĪ½ĻŒĻ‚ Ī“Ī¬ĻƒĪæĻ…Ļ‚
Ī•Ļ€ĪÆĪ“ĻĪ±ĻƒĪ· ĪµĪ½Ī“ĪæĻƒĻ…ĻƒĻ„Ī±Ī“Ī¹ĪŗĻŽĪ½ Ļ€Ī±ĻĪ±Ī³ĻŒĪ½Ļ„Ļ‰Ī½ ĻƒĻ„Ī· Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ī“ĪæĪ¼Ī® Ļ„Ļ‰Ī½ ĻƒĻ€ĻŒĻĻ‰Ī½ ĪµĪ½ĻŒĻ‚ Ī“Ī¬ĻƒĪæĻ…Ļ‚Ī•Ļ€ĪÆĪ“ĻĪ±ĻƒĪ· ĪµĪ½Ī“ĪæĻƒĻ…ĻƒĻ„Ī±Ī“Ī¹ĪŗĻŽĪ½ Ļ€Ī±ĻĪ±Ī³ĻŒĪ½Ļ„Ļ‰Ī½ ĻƒĻ„Ī· Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ī“ĪæĪ¼Ī® Ļ„Ļ‰Ī½ ĻƒĻ€ĻŒĻĻ‰Ī½ ĪµĪ½ĻŒĻ‚ Ī“Ī¬ĻƒĪæĻ…Ļ‚
Ī•Ļ€ĪÆĪ“ĻĪ±ĻƒĪ· ĪµĪ½Ī“ĪæĻƒĻ…ĻƒĻ„Ī±Ī“Ī¹ĪŗĻŽĪ½ Ļ€Ī±ĻĪ±Ī³ĻŒĪ½Ļ„Ļ‰Ī½ ĻƒĻ„Ī· Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ī“ĪæĪ¼Ī® Ļ„Ļ‰Ī½ ĻƒĻ€ĻŒĻĻ‰Ī½ ĪµĪ½ĻŒĻ‚ Ī“Ī¬ĻƒĪæĻ…Ļ‚
Aristotelis C. Papageorgiou
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Aristotelis C. Papageorgiou
Ā 
Ī£Ļ…ĻƒĻ‡Ī­Ļ„Ī¹ĻƒĪ· Ļ„Ī·Ļ‚ Ļ‡Ļ‰ĻĪ¹ĪŗĪ®Ļ‚ Ī“Ī¹Ī¬Ļ„Ī±Ī¾Ī·Ļ‚ Ļ„Ļ‰Ī½ Ī“Ī­Ī½Ļ„ĻĻ‰Ī½ ĻƒĪµ Ī­Ī½Ī± Ī“Ī¬ĻƒĪæĻ‚ Ī¼Īµ Ļ€Ī±ĻĪ±Ī¼Ī­Ļ„ĻĪæĻ…Ļ‚ Ļ„ĪæĻ… Ī±Ī½...
Ī£Ļ…ĻƒĻ‡Ī­Ļ„Ī¹ĻƒĪ· Ļ„Ī·Ļ‚ Ļ‡Ļ‰ĻĪ¹ĪŗĪ®Ļ‚ Ī“Ī¹Ī¬Ļ„Ī±Ī¾Ī·Ļ‚ Ļ„Ļ‰Ī½ Ī“Ī­Ī½Ļ„ĻĻ‰Ī½ ĻƒĪµ Ī­Ī½Ī± Ī“Ī¬ĻƒĪæĻ‚ Ī¼Īµ Ļ€Ī±ĻĪ±Ī¼Ī­Ļ„ĻĪæĻ…Ļ‚ Ļ„ĪæĻ… Ī±Ī½...Ī£Ļ…ĻƒĻ‡Ī­Ļ„Ī¹ĻƒĪ· Ļ„Ī·Ļ‚ Ļ‡Ļ‰ĻĪ¹ĪŗĪ®Ļ‚ Ī“Ī¹Ī¬Ļ„Ī±Ī¾Ī·Ļ‚ Ļ„Ļ‰Ī½ Ī“Ī­Ī½Ļ„ĻĻ‰Ī½ ĻƒĪµ Ī­Ī½Ī± Ī“Ī¬ĻƒĪæĻ‚ Ī¼Īµ Ļ€Ī±ĻĪ±Ī¼Ī­Ļ„ĻĪæĻ…Ļ‚ Ļ„ĪæĻ… Ī±Ī½...
Ī£Ļ…ĻƒĻ‡Ī­Ļ„Ī¹ĻƒĪ· Ļ„Ī·Ļ‚ Ļ‡Ļ‰ĻĪ¹ĪŗĪ®Ļ‚ Ī“Ī¹Ī¬Ļ„Ī±Ī¾Ī·Ļ‚ Ļ„Ļ‰Ī½ Ī“Ī­Ī½Ļ„ĻĻ‰Ī½ ĻƒĪµ Ī­Ī½Ī± Ī“Ī¬ĻƒĪæĻ‚ Ī¼Īµ Ļ€Ī±ĻĪ±Ī¼Ī­Ļ„ĻĪæĻ…Ļ‚ Ļ„ĪæĻ… Ī±Ī½...
Aristotelis C. Papageorgiou
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ĪšĪ±Ļ„Ī±Ī½ĪæĪ¼Ī® Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī’Ī‘ Ī•Ī»Ī»Ī¬Ī“Ī±
ĪšĪ±Ļ„Ī±Ī½ĪæĪ¼Ī® Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī’Ī‘ Ī•Ī»Ī»Ī¬Ī“Ī±ĪšĪ±Ļ„Ī±Ī½ĪæĪ¼Ī® Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī’Ī‘ Ī•Ī»Ī»Ī¬Ī“Ī±
ĪšĪ±Ļ„Ī±Ī½ĪæĪ¼Ī® Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī’Ī‘ Ī•Ī»Ī»Ī¬Ī“Ī±
Aristotelis C. Papageorgiou
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Ī”Ī¹Ī±Ļ„Ī®ĻĪ·ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ ĻƒĪµ Ī“Ī¹Ī±Ļ‡ĪµĪ¹ĻĪ¹Ī¶ĻŒĪ¼ĪµĪ½Ī± Ī“Ī¬ĻƒĪ·
Ī”Ī¹Ī±Ļ„Ī®ĻĪ·ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ ĻƒĪµ Ī“Ī¹Ī±Ļ‡ĪµĪ¹ĻĪ¹Ī¶ĻŒĪ¼ĪµĪ½Ī± Ī“Ī¬ĻƒĪ·Ī”Ī¹Ī±Ļ„Ī®ĻĪ·ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ ĻƒĪµ Ī“Ī¹Ī±Ļ‡ĪµĪ¹ĻĪ¹Ī¶ĻŒĪ¼ĪµĪ½Ī± Ī“Ī¬ĻƒĪ·
Ī”Ī¹Ī±Ļ„Ī®ĻĪ·ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ ĻƒĪµ Ī“Ī¹Ī±Ļ‡ĪµĪ¹ĻĪ¹Ī¶ĻŒĪ¼ĪµĪ½Ī± Ī“Ī¬ĻƒĪ·
Aristotelis C. Papageorgiou
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Ī ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī·Ī½ Ī•Ī»Ī»Ī¬Ī“Ī± ĻƒĪµ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĻŒ DNA ĪŗĪ±Ī¹ Ī¼ĪæĻĻ†ĪæĪ»ĪæĪ³Ī¹ĪŗĪæĻĻ‚ Ļ‡Ī±ĻĪ±...
Ī ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī·Ī½ Ī•Ī»Ī»Ī¬Ī“Ī± ĻƒĪµ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĻŒ DNA ĪŗĪ±Ī¹ Ī¼ĪæĻĻ†ĪæĪ»ĪæĪ³Ī¹ĪŗĪæĻĻ‚ Ļ‡Ī±ĻĪ±...Ī ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī·Ī½ Ī•Ī»Ī»Ī¬Ī“Ī± ĻƒĪµ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĻŒ DNA ĪŗĪ±Ī¹ Ī¼ĪæĻĻ†ĪæĪ»ĪæĪ³Ī¹ĪŗĪæĻĻ‚ Ļ‡Ī±ĻĪ±...
Ī ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī·Ī½ Ī•Ī»Ī»Ī¬Ī“Ī± ĻƒĪµ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĻŒ DNA ĪŗĪ±Ī¹ Ī¼ĪæĻĻ†ĪæĪ»ĪæĪ³Ī¹ĪŗĪæĻĻ‚ Ļ‡Ī±ĻĪ±...
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Refugia and Post-Glacial Movement of Beech in Greece
Refugia and Post-Glacial Movement of Beech in GreeceRefugia and Post-Glacial Movement of Beech in Greece
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Ī‘Ļ€ĪæĻ„ĻĻ€Ļ‰ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ Ļ„ĪæĻ… ĪšĻ…Ļ€ĻĪ¹Ī±ĪŗĪæĻ ĪŗĪ­Ī“ĻĪæĻ… (Cedrus brevifolia)...
Ī‘Ļ€ĪæĻ„ĻĻ€Ļ‰ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ Ļ„ĪæĻ… ĪšĻ…Ļ€ĻĪ¹Ī±ĪŗĪæĻ ĪŗĪ­Ī“ĻĪæĻ… (Cedrus brevifolia)...Ī‘Ļ€ĪæĻ„ĻĻ€Ļ‰ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ Ļ„ĪæĻ… ĪšĻ…Ļ€ĻĪ¹Ī±ĪŗĪæĻ ĪŗĪ­Ī“ĻĪæĻ… (Cedrus brevifolia)...
Ī‘Ļ€ĪæĻ„ĻĻ€Ļ‰ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ Ļ„ĪæĻ… ĪšĻ…Ļ€ĻĪ¹Ī±ĪŗĪæĻ ĪŗĪ­Ī“ĻĪæĻ… (Cedrus brevifolia)...
Aristotelis C. Papageorgiou
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Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…
Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…
Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…
Aristotelis C. Papageorgiou
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Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĻĪæĻƒĪ±ĻĪ¼ĪæĪ³Ī® Ļ„Ļ‰Ī½ Ī“Ī±ĻƒĪæĻ€ĪæĪ½Ī¹ĪŗĻŽĪ½ ĪµĪ¹Ī“ĻŽĪ½ ĻƒĻ„Ī·Ī½ ĪŗĪ»Ī¹Ī¼Ī±Ļ„Ī¹ĪŗĪ® Ī±Ī»Ī»Ī±Ī³Ī®: Ī¼Ī±ĪøĪ®Ī¼Ī±Ļ„Ī± Ī±Ļ€ĻŒ...
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĻĪæĻƒĪ±ĻĪ¼ĪæĪ³Ī® Ļ„Ļ‰Ī½ Ī“Ī±ĻƒĪæĻ€ĪæĪ½Ī¹ĪŗĻŽĪ½ ĪµĪ¹Ī“ĻŽĪ½ ĻƒĻ„Ī·Ī½ ĪŗĪ»Ī¹Ī¼Ī±Ļ„Ī¹ĪŗĪ® Ī±Ī»Ī»Ī±Ī³Ī®: Ī¼Ī±ĪøĪ®Ī¼Ī±Ļ„Ī± Ī±Ļ€ĻŒ...Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĻĪæĻƒĪ±ĻĪ¼ĪæĪ³Ī® Ļ„Ļ‰Ī½ Ī“Ī±ĻƒĪæĻ€ĪæĪ½Ī¹ĪŗĻŽĪ½ ĪµĪ¹Ī“ĻŽĪ½ ĻƒĻ„Ī·Ī½ ĪŗĪ»Ī¹Ī¼Ī±Ļ„Ī¹ĪŗĪ® Ī±Ī»Ī»Ī±Ī³Ī®: Ī¼Ī±ĪøĪ®Ī¼Ī±Ļ„Ī± Ī±Ļ€ĻŒ...
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĻĪæĻƒĪ±ĻĪ¼ĪæĪ³Ī® Ļ„Ļ‰Ī½ Ī“Ī±ĻƒĪæĻ€ĪæĪ½Ī¹ĪŗĻŽĪ½ ĪµĪ¹Ī“ĻŽĪ½ ĻƒĻ„Ī·Ī½ ĪŗĪ»Ī¹Ī¼Ī±Ļ„Ī¹ĪŗĪ® Ī±Ī»Ī»Ī±Ī³Ī®: Ī¼Ī±ĪøĪ®Ī¼Ī±Ļ„Ī± Ī±Ļ€ĻŒ...
Aristotelis C. Papageorgiou
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Ī£ĻĪ³Ļ‡ĻĪæĪ½ĪµĻ‚ Ļ„Ī¬ĻƒĪµĪ¹Ļ‚ ĻƒĻ„Ī·Ī½ ĪŗĪæĪ¹Ī½ĪæĻ€ĪæĪÆĪ·ĻƒĪ· Ī­ĻĪµĻ…Ī½Ī±Ļ‚
Ī£ĻĪ³Ļ‡ĻĪæĪ½ĪµĻ‚ Ļ„Ī¬ĻƒĪµĪ¹Ļ‚ ĻƒĻ„Ī·Ī½ ĪŗĪæĪ¹Ī½ĪæĻ€ĪæĪÆĪ·ĻƒĪ· Ī­ĻĪµĻ…Ī½Ī±Ļ‚Ī£ĻĪ³Ļ‡ĻĪæĪ½ĪµĻ‚ Ļ„Ī¬ĻƒĪµĪ¹Ļ‚ ĻƒĻ„Ī·Ī½ ĪŗĪæĪ¹Ī½ĪæĻ€ĪæĪÆĪ·ĻƒĪ· Ī­ĻĪµĻ…Ī½Ī±Ļ‚
Ī£ĻĪ³Ļ‡ĻĪæĪ½ĪµĻ‚ Ļ„Ī¬ĻƒĪµĪ¹Ļ‚ ĻƒĻ„Ī·Ī½ ĪŗĪæĪ¹Ī½ĪæĻ€ĪæĪÆĪ·ĻƒĪ· Ī­ĻĪµĻ…Ī½Ī±Ļ‚
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Ī ĻĪæĻƒĪ­Ī³Ī³Ī¹ĻƒĪ· Ļ„ĪæĻ… ĪµĻ€Ī¹Ļ€Ī­Ī“ĪæĻ… ĪæĪ¼ĪæĪ¼ĪµĪ¹Ī¾ĪÆĪ±Ļ‚ ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ Ī¼ĻŒĪ½ĪæĪ¹ĪŗĻ‰Ī½ Ļ†Ļ…Ļ„ĻŽĪ½ Ī¼Ī­ĻƒĪ± Ī±Ļ€ĻŒ Ļ„Ī¹Ļ‚ ...
Ī ĻĪæĻƒĪ­Ī³Ī³Ī¹ĻƒĪ· Ļ„ĪæĻ… ĪµĻ€Ī¹Ļ€Ī­Ī“ĪæĻ… ĪæĪ¼ĪæĪ¼ĪµĪ¹Ī¾ĪÆĪ±Ļ‚ ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ Ī¼ĻŒĪ½ĪæĪ¹ĪŗĻ‰Ī½ Ļ†Ļ…Ļ„ĻŽĪ½ Ī¼Ī­ĻƒĪ± Ī±Ļ€ĻŒ Ļ„Ī¹Ļ‚ ...Ī ĻĪæĻƒĪ­Ī³Ī³Ī¹ĻƒĪ· Ļ„ĪæĻ… ĪµĻ€Ī¹Ļ€Ī­Ī“ĪæĻ… ĪæĪ¼ĪæĪ¼ĪµĪ¹Ī¾ĪÆĪ±Ļ‚ ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ Ī¼ĻŒĪ½ĪæĪ¹ĪŗĻ‰Ī½ Ļ†Ļ…Ļ„ĻŽĪ½ Ī¼Ī­ĻƒĪ± Ī±Ļ€ĻŒ Ļ„Ī¹Ļ‚ ...
Ī ĻĪæĻƒĪ­Ī³Ī³Ī¹ĻƒĪ· Ļ„ĪæĻ… ĪµĻ€Ī¹Ļ€Ī­Ī“ĪæĻ… ĪæĪ¼ĪæĪ¼ĪµĪ¹Ī¾ĪÆĪ±Ļ‚ ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ Ī¼ĻŒĪ½ĪæĪ¹ĪŗĻ‰Ī½ Ļ†Ļ…Ļ„ĻŽĪ½ Ī¼Ī­ĻƒĪ± Ī±Ļ€ĻŒ Ļ„Ī¹Ļ‚ ...
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Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ļ€Ī¹ĪøĪ±Ī½Ī® ĪµĪ¾Ī­Ī»Ī¹Ī¾Ī· Ļ„ĪµĻƒĻƒĪ¬ĻĻ‰Ī½ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĻŽĪ½ Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī”ĪæĪ“ĻŒĻ€Ī·
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ļ€Ī¹ĪøĪ±Ī½Ī® ĪµĪ¾Ī­Ī»Ī¹Ī¾Ī· Ļ„ĪµĻƒĻƒĪ¬ĻĻ‰Ī½ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĻŽĪ½ Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī”ĪæĪ“ĻŒĻ€Ī·Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ļ€Ī¹ĪøĪ±Ī½Ī® ĪµĪ¾Ī­Ī»Ī¹Ī¾Ī· Ļ„ĪµĻƒĻƒĪ¬ĻĻ‰Ī½ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĻŽĪ½ Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī”ĪæĪ“ĻŒĻ€Ī·
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ļ€Ī¹ĪøĪ±Ī½Ī® ĪµĪ¾Ī­Ī»Ī¹Ī¾Ī· Ļ„ĪµĻƒĻƒĪ¬ĻĻ‰Ī½ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĻŽĪ½ Ļ„Ī·Ļ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ ĻƒĻ„Ī· Ī”ĪæĪ“ĻŒĻ€Ī·
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Ī‘Ļ€ĪæĻ„ĪµĪ»ĪµĪÆ Ī· Ī”ĪæĪ“ĻŒĻ€Ī· ĻƒĪ·Ī¼ĪµĪÆĪæ ĻƒĻ…Ī½Ī¬Ī½Ļ„Ī·ĻƒĪ·Ļ‚ Ļ„Ī·Ļ‚ ĪµĻ…ĻĻ‰Ļ€Ī±ĻŠĪŗĪ®Ļ‚ (Ī“Ī±ĻƒĪ¹ĪŗĪ®Ļ‚) ĪŗĪ±Ī¹ Ļ„Ī·Ļ‚ Ī±Ī½Ī±Ļ„ĪæĪ»Ī¹Īŗ...
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Ī‘Ļ€ĪæĻ„ĪµĪ»ĪµĪÆ Ī· Ī”ĪæĪ“ĻŒĻ€Ī· ĻƒĪ·Ī¼ĪµĪÆĪæ ĻƒĻ…Ī½Ī¬Ī½Ļ„Ī·ĻƒĪ·Ļ‚ Ļ„Ī·Ļ‚ ĪµĻ…ĻĻ‰Ļ€Ī±ĻŠĪŗĪ®Ļ‚ (Ī“Ī±ĻƒĪ¹ĪŗĪ®Ļ‚) ĪŗĪ±Ī¹ Ļ„Ī·Ļ‚ Ī±Ī½Ī±Ļ„ĪæĪ»Ī¹Īŗ...
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Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½
Ī“ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±, Ī²Ī¹ĪæĻ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī± ĪŗĪ±Ī¹ Ī“Ī¹Ī±Ļ‡ĪµĪÆĻĪ¹ĻƒĪ· Ī“Ī±ĻƒĪ¹ĪŗĻŽĪ½ ĪæĪ¹ĪŗĪæĻƒĻ…ĻƒĻ„Ī·Ī¼Ī¬Ļ„Ļ‰Ī½
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Ī•Ļ€ĪÆĪ“ĻĪ±ĻƒĪ· ĪµĪ½Ī“ĪæĻƒĻ…ĻƒĻ„Ī±Ī“Ī¹ĪŗĻŽĪ½ Ļ€Ī±ĻĪ±Ī³ĻŒĪ½Ļ„Ļ‰Ī½ ĻƒĻ„Ī· Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ® Ī“ĪæĪ¼Ī® Ļ„Ļ‰Ī½ ĻƒĻ€ĻŒĻĻ‰Ī½ ĪµĪ½ĻŒĻ‚ Ī“Ī¬ĻƒĪæĻ…Ļ‚
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Refugia and Post-Glacial Movement of Beech in Greece
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Ī‘Ļ€ĪæĻ„ĻĻ€Ļ‰ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ Ļ„ĪæĻ… ĪšĻ…Ļ€ĻĪ¹Ī±ĪŗĪæĻ ĪŗĪ­Ī“ĻĪæĻ… (Cedrus brevifolia)...
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Ī‘Ļ€ĪæĻ„ĻĻ€Ļ‰ĻƒĪ· Ļ„Ī·Ļ‚ Ī³ĪµĪ½ĪµĻ„Ī¹ĪŗĪ®Ļ‚ Ļ€ĪæĪ¹ĪŗĪ¹Ī»ĻŒĻ„Ī·Ļ„Ī±Ļ‚ Ļ„ĪæĻ… ĪšĻ…Ļ€ĻĪ¹Ī±ĪŗĪæĻ ĪŗĪ­Ī“ĻĪæĻ… (Cedrus brevifolia)...
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Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…
Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…
Ī£ĻĪ½ĪøĪµĻƒĪ· Ī±Ļ€Ī»ĪæĻ„ĻĻ€Ļ‰Ī½ Ļ‡Ī»Ļ‰ĻĪæĻ€Ī»Ī±ĻƒĻ„Ī¹ĪŗĪæĻ DNA ĻƒĪµ Ļ€Ī»Ī·ĪøĻ…ĻƒĪ¼ĪæĻĻ‚ ĪæĪ¾Ī¹Ī¬Ļ‚ Ļ„ĪæĻ… ĻŒĻĪæĻ…Ļ‚ Ī Ī±Ī³Ī³Ī±ĪÆĪæĻ…
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Genetic implications of forest management in the Mediterranean

  • 1. Genetic Implications of Forest Management in the Mediterranean Aristotelis C. Papageorgiou Department of Forestry, Environment and Natural Resources, Democritus University of Thrace, Greece Solsona 2004
  • 2. Presentation plan ā€¢ Genetic diversity in Mediterranean forests ā€¢ Factors influencing genetic diversity ā€¢ Impact of forest management ā€¢ SFM and genetic diversity ā€¢ Research and policy priorities April 4, 2004 MEDFOREX meeting 2
  • 3. What is so special about the Med? ā€¢ Biological, cultural, social, historical diversity ā€¢ Biological diversity: ā€“ rich mosaic of changing ecosystems and land use patterns (from alpine to tropical) ā€“ Large amount of species (especially plants) ā€“ high endemism ā€“ Populations of species with a broader distribution are the most variable in terms of genetic diversity April 4, 2004 MEDFOREX meeting 3
  • 4. Genetic diversity ā€¢ Differentiation among individuals and groups of individuals (populations) exists. ā€¢ Differentiation at the gene level can be inherited = genetic variation ā€¢ Key for adaptation in changing environments over space and time ā€¢ Connection between generations ā€¢ Is usually measured in traits that are not important for forestry (e.g. DNA markers) April 4, 2004 MEDFOREX meeting 4
  • 5. Genetic system of a forest population Hattemer & Gillet 2000 April 4, 2004 MEDFOREX meeting 5
  • 6. Generations of forest trees April 4, 2004 MEDFOREX meeting 6
  • 7. Genetics of Mediterranean forests ā€¢ Disjunct distribution of populations and species ā€“ high differentiation ā€¢ Usually high genetic variation within populations ā€“ comparison with N/C Europe ā€¢ Many small populations ā€¢ Large distributions of species over different environments (e.g. Pinus species) ā€¢ Unique alleles and races ā€¢ Paradox of variation in several cases April 4, 2004 MEDFOREX meeting 7
  • 8. Genetic diversity of forest species Petit et al. 2003 April 4, 2004 MEDFOREX meeting 8
  • 9. Reasons for high diversity ā€¢ The relief of the Mediterranean basin April 4, 2004 MEDFOREX meeting 9
  • 10. Reasons for high diversity ā€¢ The Mediterranean climate ā€¢ The existence of tectonic microplates and their move April 4, 2004 MEDFOREX meeting 10
  • 11. Reasons of high diversity ā€¢ The glaciations era ā€“ refugia April 4, 2004 MEDFOREX meeting 11
  • 12. Human and forests ā€¢ Most ancient human cultures ā€¢ Centre of the ā€œknown worldā€ ā€¢ Organized trade and land use ā€¢ First ā€œglobalizedā€ economies April 4, 2004 MEDFOREX meeting 12
  • 13. Direct use of plant species April 4, 2004 MEDFOREX meeting 13
  • 14. Direct use of plant species April 4, 2004 MEDFOREX meeting 14
  • 15. Human shaping nature Agriculture, grazing, fuelwood collection, etc. shape the landscapeā€¦ April 4, 2004 MEDFOREX meeting 15
  • 16. ā€œMosaicā€ - forest landscape April 4, 2004 MEDFOREX meeting 16
  • 17. Mediterranean forests ā€¢ Forest includes terrestrial ecosystems in a broad sense ā€¢ Natural forest ecosystems ā€“ mixed forests ā€“ complex age structures ā€“ connection between generations ā€¢ Mainly small scale activities ā€“ high land use diversity ā€“ fragmentation ā€“ mixed with other activities ā€¢ Degradation ā€“ desertification ā€“ loss of ecosystem productivity ā€“ land abandonment ā€¢ High protective role against erosion, drought ā€“ social importance for rural development ā€¢ Recreation ā€“ quality of life ā€“ Non Wood Goods ā€¢ Culture ā€“ spirituality April 4, 2004 MEDFOREX meeting 17
  • 18. Human impact on gene diversity ā€¢ Far-reaching / global ā€“ Emission of air pollutants ā€“ Greenhouse gases ā€“ Policy processes ā€¢ Local ā€“ Forest destruction (land conversion) ā€“ Forest fragmentation ā€“ Forest management ā€¢ Introduction of new species & populations ā€¢ Silvicultural activities ā€¢ Other forms on management (e.g. grazing) April 4, 2004 MEDFOREX meeting 18
  • 19. Destruction and fragmentation ā€¢ Forest destruction (deforestation) ā€“ Loss of species with small scale (e.g. Abies nebrodensis, Quercus euboica, Cedrus brevifolia) ā€“ Loss of populations (differentiated ā€“ adapted) ā€¢ Forest fragmentation ā€“ Reduction of effective population size ā€“ drift effects ā€“ genetic bottlenecks ā€“ Increase vulnerability of populations ā€“ Caused by development, agriculture, grazing April 4, 2004 MEDFOREX meeting 19
  • 20. Genetic bottleneck 1400 1200 1000 RECOVERY 800 N 600 400 CRASH 200 0 Bottleneck TIME April 4, 2004 MEDFOREX meeting 20
  • 21. Example: Pinus leucodermis in Italy ā€¢ Only a few small populations remaining ā€“ fragmented ā€“ no connection possible ā€¢ Low genetic variation ā€“ large differentiation among populations Morgante & Vendramin 1991 April 4, 2004 MEDFOREX meeting 21
  • 22. Artificial regeneration ā€¢ Adaptation on the final site in question ā€“ Different adaptations at the original site ā€“ Adaptation in nurseries ā€¢ Genetic variation (evolutionary adaptability) reduced due to drift effects ā€“ Harvest from small number of plants (founder) ā€“ Unknown material ā€“ provenance ignored ā€¢ Possible impact on surrounding forests due to gene flow ā€¢ However: development of new land races (e.g. Quercus rubra in France) April 4, 2004 MEDFOREX meeting 22
  • 23. Example: Cupressus sempervirens ā€¢ Low genetic variation of planted stands ā€¢ Canker attack in planted stands ā€¢ Gene flow from planted stands in natural populations Papageorgiou et al. 1994 April 4, 2004 MEDFOREX meeting 23
  • 24. Natural regeneration ā€¢ Connection between generations - maintenance of dynamics of genetic structures in life cycle of forest trees ā€“ evolution goes on ā€¢ How many individuals participate in the next generation? Where? ā€“ Natural processes (very improbable in the Med) ā€“ Non-forestry activities (e.g. grazing) ā€“ Forestry operations (e.g. light felling) ā€¢ Limited potential for changes of genetic structures ā€“ No safeguard against unintentional & random changes of genetic structures, losses of genetic variation April 4, 2004 MEDFOREX meeting 24
  • 25. Silvicultural treatments ā€¢ Low population densities ā€¢ Reduced gene flow ā€“ Possible founder effects (a few seeder trees) ā€“ Increased inbreeding ā€“ low germinability ā€“ Inbreeding depression ā€¢ Threshold values for ā€“ Population sizes ā€“ Population densities ā€¢ Problem for scattered species ā€“ Particularly in species-rich mixed forests ā€“ Species with ā€œpeculiarā€ mating system (Taxus baccata) April 4, 2004 MEDFOREX meeting 25
  • 26. Example: Abies cephalonica ā€¢ Higher values of inbreeding levels in uneven aged forests with lower population densities Fady & Conkle 1993 April 4, 2004 MEDFOREX meeting 26
  • 27. Selective cutting ā€¢ Selection against superior phenotypes ā€“ Selective logging with short cutting cycles ā€“ Exclusion of superior phenotypes from reproduction ā€¢ Reduction of mating trees ā€“ effective population size ā€“ Inbreeding ā€¢ Coppice forests ā€“ Clear cuts with a few remaining trees April 4, 2004 MEDFOREX meeting 27
  • 28. Is genetic diversity at stake? ā€¢ Disturbance of human ā€“ nature balance ā€¢ The dynamic systems are broken (genetic, ecological, nutrients, water, energy, etc.) ā€¢ Ecosystem are not able to provide goods and services in the long term ā€“ desertification. ā€¢ Main problems occur mainly from factors outside SFM ā€¢ The maintenance / restoration of the ability of Mediterranean ecosystems to continue their dynamic function should become the target of any conservation effort in the region April 4, 2004 MEDFOREX meeting 28
  • 29. Forest management April 4, 2004 MEDFOREX meeting 29
  • 30. Forest management in the Med ā€¢ Management strategies & techniques imported from the central and northern counties of Europe - production of timber is priority in most cases ā€¢ Apply on the more temperate forests of the Mediterranean region - ignore the non-productive terrestrial ecosystems (e.g. maquis) ā€¢ Fail to capture the complexity of Mediterranean forests & land use systems ā€¢ The spatial reference of SFM is the stand and not the broader landscape - more appropriate for the description of the human - nature dynamics April 4, 2004 MEDFOREX meeting 30
  • 31. Protection of forest biodiversity SFM as a tool to achieve biodiversity conservation Protects BD elements, including genes & provenances April 4, 2004 MEDFOREX meeting 31
  • 32. Forest protection in the Med ā€¢ Protection concepts and strategies imported from northern counties with empty spaces and productive forests - human influence on BD is ignored ā€¢ Based mainly on the ā€œset ā€“ asideā€ principle ā€¢ Focus on absolute protection or special management of specific biological entities (e.g. genes) or spatial units ā€“ miss the broader picture ā€¢ Restricted to reserves and networks ā€¢ Selection of protected items ā€“ majority remains unprotected April 4, 2004 MEDFOREX meeting 32
  • 33. Forest conservation genetics ā€¢ Forest genetic resources ā€“ Included in broader BD policies & plans ā€“ Most under-represented part of BD ā€“ Based in ā€œgenetic inventoriesā€ (assessments) ā€¢ In situ ā€“ Actions of strict protection of rare or endangered entities ā€“ Gene reserves (special management allowed) ā€“ Conservation in managed ecosystems (Namkoong). ā€¢ Ex situ ā€“ Protection of endangered or rare tree species ā€“ Storage of ā€œusefulā€ genes (gene banks) April 4, 2004 MEDFOREX meeting 33
  • 34. An important gap ā€¢ ā€œOther wooded landā€ ā€“ not protected, not managed ā€¢ The classical protection / SFM approach ā€“ has a spatial gap of about 50% ā€“ cannot cope with the dynamic nature of Mediterranean forests ā€¢ Conservation of forest genetic diversity cannot be seen separately from the general use and management of forest resources ā€¢ Inventories possible in a few cases only (no time, no money for ā€œtotal inventoriesā€) April 4, 2004 MEDFOREX meeting 34
  • 35. Linking SFM and BD ā€¢ Environmental agencies consider BD as a priority in all management activities ā€¢ Forestry parties see the economic aspect of forestry as a priority. ā€¢ BD conservation and SFM can be linked: ā€“ Complexity of both concepts ā€“ Long term character ā€¢ ā€œEcosystem approachā€ & ā€œclose to natureā€ forestry ā€¢ Tools, such as SFM certification, ā€œcriteria & indicatorsā€, Special Management plans, NFP April 4, 2004 MEDFOREX meeting 35
  • 36. The challenge ā€¢ Maintaining adequate levels of genetic diversity of tree (& other) species is crucial for adaptation and adaptability processes ā€¢ Dynamic processes must maintain their role for the maintenance of productivity of goods and services ā€¢ Genetic diversity is not just another goal of SFM, but the means to achieve it ā€¢ We need: Management techniques that will prevent disturbances in critical ecological and genetic processes April 4, 2004 MEDFOREX meeting 36
  • 37. The challenge April 4, 2004 MEDFOREX meeting 37
  • 38. Future research priorities ā€¢ Develop inventory systems for genetic diversity status of terrestrial ecosystems (stand types) ā€“ Use results of existing genetic research projects ā€“ Plan future genetic research ā€“ Use models to simulate evolution under forest management scenarios ā€“ Connect gene diversity with stand & age structure, density, mating system, etc. ā€¢ Develop criteria & indicators for genetic diversity ā€“ adjust existing efforts to the Mediterranean ā€“ introduce genetic criteria into existing SFM C&Is (certification standards, MCPFE criteria, etc.) April 4, 2004 MEDFOREX meeting 38
  • 39. Future research priorities ā€¢ Connect specific management techniques of Mediterranean forests with the previous steps ā€“ Give a ā€œgenetic gradeā€ for each technique ā€“ Evaluate human impact on genetic diversity ā€¢ Set priorities for SFM considering genetic diversity ā€¢ Extrapolate of appropriate strategies for most taxa from the results of studies of a few model cases ā€¢ Identify genetic aspects that may become limiting for certain types of species and ecosystems ā€¢ Monitoring and evaluation April 4, 2004 MEDFOREX meeting 39
  • 40. SFM for the Med ā€¢ Develop management plans for the Mediterranean beyond the classical forestry approach, including broader ecosystems and activities, based on the maintenance of genetic diversity ā€¢ Protecting biodiversity through management and planning, following rules that need minor assessments April 4, 2004 MEDFOREX meeting 40
  • 41. Thank you for your attention April 4, 2004 MEDFOREX meeting 41