Plas ici y, In aspeci ic Va iabili y and Local
Adap a ion o Clima ic Ex eme E en s o
Eco ypes/P o enances o Key Plan Species
Disse a ion
zu E langung des akademischen G ades
D . e . na .
o geleg de
Fakul ä ü Biologie, Chemie und Geowissenscha en
de Uni e si ä Bay eu h
on
He n Daniel Thiel (M.Sc.)
geb. am 26.11.1977 in Ma k edwi z
Bay eu h, den 09. Mai 2012
Die o liegende A bei wu de in de Zei on Janua 2009 bis Ap il 2012 am Leh s uhl ü
Biogeog a ie de Uni e si ä Bay eu h un e de Be euung on He n P o . D . Ca l
Beie kuhnlein ange e ig .
Table o Con en
1.
Compendium ................................................................................................. 1
1.1.
Sho Summa y o he Thesis ..................................................................................... 1
1.2.
Ku ze Zusammen assung de Dok o a bei ................................................................ 3
2.
Backg ound o he Thesis ............................................................................ 6
2.1.
Global Clima e Change and Ex eme Wea he E en s .............................................. 6
2.2.
Clima e Impac on Ecosys em and Selec ed Species ................................................. 9
2.3.
Wi hin-species Di e si y as a Po en ial Tool in Clima e Change Adap a ion ......... 14
3.
Synopsis o he Thesis ................................................................................. 18
3.1.
Gene al Objec i es and App oach ............................................................................ 18
3.2.
Rema ks on Me hodological Challenges .................................................................. 21
3.3.
Main Findings and Conclusion ................................................................................ 23
4.
Re e ences o In oduc ion and Synopsis ................................................. 27
5.
Decla a ion o Own Con ibu ion ............................................................. 34
6.
Manusc ip s ................................................................................................. 39
6.1.
Manusc ip 1: ............................................................................................................ 39
Assis ed coloniza ion: a ques ion o ocal uni s and ecipien locali ies .............................. 39
6.2.
Manusc ip 2: ............................................................................................................ 56
Uni o m d ough and wa ming esponses in Pinus nig a p o enances despi e speci ic
o e all pe o mances ............................................................................................................ 56
6.3.
Manusc ip 3: ............................................................................................................ 78
Di e en eac ions o cen al and ma ginal p o enances o Fagus syl a ica o expe imen al
d ough ................................................................................................................................. 78
6.4.
Manusk ip 4: ......................................................................................................... 105
Eco ypes o Eu opean g ass species espond di e en ly o wa ming and ex eme d ough
105
6.5.
Manusk ip 5: ......................................................................................................... 128
Cold ha diness o Pinus nig a A nold as in luenced by geog aphic o igin, wa ming, and
ex eme summe d ough .................................................................................................... 128
6.6.
Manusk ip 6: ......................................................................................................... 153
La e os sensi i i y o ju enile Fagus syl a ica L. di e s be ween sou he n Ge many and
Bulga ia and depends on p eceding ai empe a u e .......................................................... 153
6.7.
Manusk ip 7: ......................................................................................................... 171
Eco ypic di e en ia ion and pas clima ic expe ience in luence he esponse o la e sp ing
os in ou common g ass species in Cen al Eu ope ....................................................... 171
6.8.
Manusc ip 8: .......................................................................................................... 189
In aspeci ic di e si y abe s chances o high yields unde di e en clima ic condi ions – a
modelled example om p o enance ial da a ................................................................... 189
7.
Acknowledgemen s ................................................................................... 208
8.
Appendix ................................................................................................... 209
9.
E klä ung .................................................................................................. 210
1 Compendium
1
1. Compendium
1.1. Sho Summa y o he Thesis
Clima e change, and especially an inc ease o magni ude and equency o clima ic ex eme
e en s such as d ough pe iods o hea wa es, will al e g owing condi ions o plan s in he
u u e. Pe sis en ecosys ems, wi h long-li ing o ganisms, such as o es o pe manen
g assland will be pa icula ly impac ed by his de elopmen . The eloci y o hese changes is
likely o occu a a pace, which species may no be able o keep ack wi h by na u al dispe sal
o gene ic adap a ion. Ag icul u e, o es y and ecosys em managemen mus de elop
coun e ac ing p ac ices o secu e he pe sis ence and unc ioning o hese ecosys ems and hus
hei p o ision o goods and se ices. The e o e i is impo an o de elop a be e
unde s anding how species and ecosys ems may espond o u u e clima ic s esso s. Impac
assessmen s, e.g. ia clima ic en elope modelling a e p one o misin e p e a ions o he
adap i e capaci y o species, as hey do no inco po a e he in aspeci ic gene ic and
pheno ypic di e ences ha exis wi hin he popula ions acc oss he dis ibu ion ange o a
species.
Ye , in aspeci ic a ia ion may exhibi po en ial ools o he de elopmen o clima e change
adap a ion s a egies. He e, I ocus on key ecosys ems in Cen al Eu ope. In pa icula he
selec i e use o plan p o enances o eco ypes may help o make ecosys ems clima e- esilien
wi hou a po en ially mo e p oblema ic in oduc ion o exo ic species. Especially p o enances
om wa me , d ough -p one egions, wi h a cu en clima e simila o he p ojec ed one o
Cen al Eu ope ecen ly came in o ocus as po en ial subs i u es o local p o enances, as hey
migh ha e de eloped local adap a ions o clima e condi ions a hei loca ion o o igin.
Insigh s abou he esponse o hese p o enances o changing a e ages and ex eme e en
egimes a e c ucial o a easonable use o wi hin-species di e si y in clima e change
adap a ion.
Fi s , he concep o assis ed coloniza ion o mig a ion o species o eco ypes and he ole i
can play as an adap a ion s a egy in ag icul u e, o es y o na u e conse a ion is in oduced
(Manusc ip 1). I is sugges ed ha a ocus should be laid on keys one species ha ensu e
ecosys em pe sis ence and unc ioning as hey go e n he habi a s uc u e and mic oclima e
o a si e. The assis ed coloniza ion o p e-adap ed eco ypes o keys one species om clima es
simila o u u e p ojec ions o he a ge si e is p oposed.

1 Compendium
2
Fu he mo e, p o enances o selec ed g assland and o es key-species we e exposed o
d ough and wa ming in wo expe imen s in Bay eu h and Landau, and hei ecological
esponses we e analysed. Resul s sugges ha local adap a ions o clima ic s esso s exis .
Howe e , he magni ude and di ec ion o esponses s ongly depend on species and clima ic
a iables. Fo g assland species, e.g. di e ences in d ough sensi i i y could be demons a ed
in some cases (Manusc ip 4). Fagus syl a ica exhibi ed di e ences be ween he p o enances
in esponse o d ough condi ions, as well (Manusc ip 3). I seems ha ma ginal p o enances,
om he d y ma gins o he dis ibu ion ange, show less inc emen educ ion due o he
d ough ea men . Ye , unde mo e a ou able condi ions o wa e supply hese p o enances
did no yield he same high inc emen a es han mo e cen al p o enances, indica ing a ade-
o be ween s abili y unde s ess and yield unde non-s ess condi ions. A pine species ha is
gene ally conside ed o be a he d ough - esis an , Pinus nig a, which is a po en ial subs i u e
o clima e- h ea ened coni e s on d y si es in Cen al Eu ope, did no show any di e ences in
esponse o d ough and wa ming (Manusc ip 2), maybe due o a weak selec i e p essu e as a
esul o high d ough - esis ance ac oss he whole dis ibu ion ange. The impac s o d ough
on inc emen became no isible be o e he second yea a e he ea men , s essing he need
o mo e long- ime expe imen s in clima e impac esea ch.
E en in a gene ally wa me en i onmen , cold ex emes in win e o sp ing a e expec ed s ill
o p e ail in he u u e. The e o e, he p o enances o he selec ed species we e es ed o hei
cold-ha diness and la e os esis ance (Manusc ip s 5-7). G ow h o he g assland species
and F. syl a ica we e nega i ely impac ed by a la e os e en and di e ences in la e- os
sensi i i y be ween p o enances o eco ypes we e iden i ied. The (sub-) medi e anean
species P. nig a showed di e ences be ween p o enances in hei win e cold ha diness.
Co ela ions be ween pe o mance unde cold s ess and win e condi ions o la e os
p oneness o he places o o igin could be es ablished o almos all species. Howe e ,
p eceding clima e expe ience, such as he wa ming o d ough ea men o he plan s al e ed
hei eac ion o cold ex emes compa ed o he con ol ea men , indica ing he complexi y o
he in e ac i e impac s o clima e ac o s on ecosys em and plan pe o mance.
The unce ain y o clima e p ojec ions and he mul i ude o changing clima ic s esso s,
hough, make he p ospec o an easy and apid success in he sea ch o single “bes -adap ed”
p o enances e y ques ionable. In economics he po olio e ec shows ha a di e si ica ion
o in es men s dec eases he isk o a o al loss o p o i s. Hence, in a modelling p ocedu e
based on he inc emen da a om he abo e men ioned expe imen i was es ed i a “po olio
1 Compendium
3
in es men ” in se e al p o enances in one s and dec eases he isk o yield losses (Manusc ip
8). Resul s indica e ha he highe he numbe o p o enances he highe he chance o a
“bes -pe o me ” o be included in he se . So he likelihood o highe yields, unde di e en
clima ic condi ions inc eases, ye he isk o low yields s ays s able.
Gene ally, i seems ha he selec i e use o plan species and eco ypes in clima e change
adap a ion can be a easible ool o main ain ecosys em unc ionali y and p oduc i i y.
Howe e , he unce ain p ojec ions, he mul i ude o clima ic s esso s and hei in e play wi h
o he en i onmen al ac o s and he po en ial impac s o assis ed coloniza ion o eco ypes on
he gene ic di e si y wi hin species and popula ions equi e u he esea ch.
1.2. Ku ze Zusammen assung de Dok o a bei
De Klimawandel, und o allem Ve ände ungen in Au e enswah scheinlichkei und
In ensi ä on Ex eme eignissen, wie Dü en ode Hi zewellen, we den die
Wuchsbedingungen ü P lanzen kün ig s a k ände n. Hochs e e Ökosys eme, wie ex ensi
genu z es G ünland ode Wälde mi langlebigen O ganismen we den besonde s be o en on
diesen Ve ände ungen. Die Geschwindigkei mi de sich diese Wandel ollzieh , mach es
ü iele A en schwe bis unmöglich du ch A eal e schiebung ode gene ische Anpassung
Sch i zuhal en.
Vo diesem Hin e g und muss die Land- und Fo s wi scha Gegenmaßnahmen en wickeln,
die die Funk ion diese Ökosys eme e hal en und die E äge und die Be ei s ellung on
ökosys ema en Diens leis ungen siche n. Das Wissen um die Auswi kungen klima ische
S ess ak o en au A en und Ökosys eme is dahe une lässlich. Die Einschä zung solche
Auswi kungen, z.B. du ch Klimahüllenmodellie ung, be ücksich ig die inne a liche
gene ische Viel al und phäno ypische Plas izi ä , die inne halb und zwischen Popula ionen
exis ie , in de Regel nich , was zu Fehleinschä zungen de Anpassungs ähigkei üh en kann.
Diese inne a liche Va ia ion kann jedoch ein wich iges We kzeug da s ellen, wenn es um die
Klimaanpassung on Ökosys emen in Mi eleu opa geh . Diese S udie konzen ie sich au
Schlüsselökosys eme in Mi eleu opa. Die selek i e Nu zung on He kün en ode Öko ypen
wich ige Schlüssela en kann e en uell dazu bei agen Ökosys eme esilien e gegenübe
nega i en Klimaein lüssen zu machen, ohne dabei die o p oblema ische Ein üh ung on
exo ischen P lanzena en in Kau nehmen zu müssen. Dabei ge a en besonde s He kün e aus
wä me en und ockene en Gegenden, mi Klimabedingungen ähnlich zu den ü Mi eleu opa
1 Compendium
4
p ognos izie en in den Fokus, da diese ehe solche Bedingungen angepass sein könn en. Es
is jedoch wich ig zu wissen, wie e schieden He kün e au sich e ände nde Mi elwe e
und Klimaex eme eagie en, um dieses Mi el e nün ig einse zen zu können.
In de o liegenden A bei wi d das Konzep de Assis ed Coloniza ion o ges ell und
besch ieben welche Rolle es in Land- und Fo s wi scha und im Na u schu z spielen könn e
(Manusk ip 1). Es wi d da geleg . Dass dabei o allem Schlüssela en im Fokus s ehen
soll en, da sie o mals Habi a s uk u en und Mik oklima und dami auch Bes and und
Funk ion on Ökosys emen bes immen. Die geziel e Ein uh on angepass en Öko ypen aus
Regionen in denen heu e Klimabedingungen he schen, wie sie ü die Ziel egionen
p ognos izie we den, wi d hie bei zu Diskussion ges ell .
Wei e hin wu den He kün e ausgewähl e G ünland- und Bauma en in einem
Top expe imen küns liche Dü e und E wä mung ausgese z , um de en Reak ion da au zu
messen. Die E gebnisse zeigen, dass es lokale Anpassungen an klima ische S ess ak o en
gib , diese jedoch a spezi isch a iie en und s a k on de jeweiligen Klima a iable
abhängen. Un e schiede in de Dü e esis enz zwischen e schiedenen He kün en manche
G asa en (Manusk ip 4), sowie zwischen He kün en de Ro -Buche (Manusk ip 3) konn en
dabei nachgewiesen we den. Vo allem bei de Ro -Buche scheinen He kün e om ockenen
Rand des Ve b ei ungsgebie es wenige s a k au Dü e zu eagie en wie He kün e aus dem
Zen um des Ve b ei ungsgebie es. Jedoch konn en diese „Rand-He kün e“ un e güns igen,
aus eichend wasse e so g en Bedingungen auch nich so hohe E agsleis ungen e zielen.
Hie geh S abili ä un e S essbedingungen anscheinend au Kos en hohe E äge un e
güns igen Bedingungen. He kün e de Schwa z-Kie e , eine gene ell dü eangepass en A ,
die als mögliche E sa z ü ge äh de e Nadelbauma en au T ockens ando en in
Mi eleu opa gil , un e schieden sich nich in ih e Reak ion au T ockenhei und E wä mung
(Mausk ip 2), was du ch einen schwachen Selek ionsd uck au g und eine allgemeinen
hohen Dü e esis en im gesam en Ve b ei ungsgebie e klä we den könn e. Die
Zuwachsleis ung eagie e jedoch e s im zwei en Jah au das Dü ee eignis. Dies
un e s eich die No wendigkei on lang is ig angeleg en Klimaexpe imen en um die
Auswi kungen on ex emen ich ig beu eilen zu können.
In einem wei e en Sch i , wu den die He kün e de genann en A en au ih e F os hä e und
Spä os ole anz ge es e , da solche E eignisse auch in un e höhe en
Du chschni s empe a u en dennoch möglich sein we den (Manusk ip e 5-7).
Spä os e eignisse üh en zu einem ge inge en Wachs um bei den G asa en als auch bei de
1 Compendium
5
Ro -Buche und Un e schiede zwischen den He kün en in de Spä os ole anz konn en
au gezeig we den. Auch die He kün e de (sub-)medi e an e b ei e en Schwa z-Kie e
un e schieden sich in ih e F os hä e. Die Pe o mance un e Käl es ess ko elie e bei as
allen A en mi Minimum empe a u en in Win e ode F ühling in den He kun so en de
e schiedenen P o enienzen. In e essan e weise beein luss en die „Klimae ah ungen“ de
P lanzen, sp ich die o ausgehenden Tempe a u - und Dü emanipula ionen, die Reak ion au
F os , was die Komplexi ä des Zusammenspiels on e schiedenen Klima ak o en und de en
Auswi kung au A en und Ökosys eme deu lich mach .
Die Unsiche hei de Klimap ojek ionen und die Vielzahl sich e ände nde Klima ak o en
machen die Suche nach einzelnen bes angepass en He kün en jedoch nich seh
e olg e sp echend. In den Wi scha swissenscha en besch eib de Po olio-E ek die
Risikominimie ung du ch eine S euung de In es i ionen. In diese A bei , wu de in einem
Modell, basie end au den Zuwachsda en aus oben genann en Expe imen , ge es e , ob eine
„Po olio-In es i ion“ in meh e e He kün e inne halb eines Bes andes das Risiko on
g oßen Zuwachs e lus en un e S essbedingungen minimie en kann (Manusk ip 8). Die
E gebnisse deu en da au hin, dass mi s eigende Anzahl on He kün en die Chance s eig
eine „Supe -He kun “ im „Po olio“ zu haben. Das bedeu e , dass die Chance au hohe
E äge un e e schiedenen Klimabedingungen s eig mi s eigende Zahl He kün e, das
Risiko nied ige E äge jedoch gleich bleib .
Abschließend läss sich sagen, dass die selek i e Nu zung on He kün en ode Öko ypen
du chaus ein geeigne es Mi el zu Klimaanpassung sein kann. Die Unsiche hei en de
Klimap ognosen, die Vielzahl klima ische S ess ak o en und de en In e ak ion mi ande en
Umwel ak o en, sowie die Auswi kung on Assis ed Coloniza ion au die gene ische Viel al
inne halb on Popula ionen und A en, mach jedoch wei e e Fo schung no wendig.
2 Backg ound o he Thesis
12
p oduc i i y a e expec ed o be less se e e han in mo e con inen al o Medi e anean o es
ecosys ems, whe e g ow h is al eady wa e limi ed (Lindne e al. 2010). In hese wa e
limi ed egions, high empe a u es and d ough condi ions will mos likely educe o es
p oduc i i y and acili a e ee mo ali y (Allen e al. 2010).
In Wes e n and Cen al Eu ope especially na i e coni e species will su e unde inc eased
empe a u es along wi h educed p ecipi a ion in summe , and migh be eplaced by mo e
compe i i e deciduous ee species (Ma acchi e al. 2005; Kölling 2009).
On he o he hand, wa me win e empe a u es ha e been ound o educe he os ha dening
o ees, especially in he con inen al egions o Eu ope wi h ha sh os e en s du ing he cold
mon hs (Hanninen 2006; Lindne e al. 2010). Fu he mo e ees a e expec ed o become mo e
ulne able o la e os e en s in sp ing as wa me empe a u es igge ea lie lea lushing
(K ame e al. 2000).
Fu he mo e a wa me and d ie en i onmen may imp o e condi ions o he bi o e insec s
(Vanhanen e al. 2007; Wes ga h-Smi h e al. 2007) and o es pa hogens (Desp ez-Lous au
e al. 2007), esul ing in calami ies and he la ge-scale b eakdown o o es s ands.
Howe e , hese impac s o clima e change on o es ecosys ems a e s ongly species-
dependen acco ding o he dominan key ee species and can e en di e in se e i y wi hin
he dis ibu ion ange o one single species (Hlasny e al. 2011) as si e condi ions, pheno ypes,
and egional clima ic changes may di e (Lindne e al. 2010).
In he ollowing obse ed on p o ec ed impac s o clima e change and ex eme e en s on he
wo ee species, used in his s udy, will be p esen ed.
Fagus syl a ica
Fagus syl a ica is he na u ally dominan ee species in Cen al Eu ope, and he e o e o high
economic impo ance. The a ea s ocked wi h F. syl a ica cons an ly inc eases in Cen al
Eu ope/Ge many due o o es con e sion om coni e ous o mixed o deciduous s ands,
despi e he ac ha i is conside ed a “high- isk” species in e ms o clima e change
(Rennenbe g e al. 2004; Ohlemulle e al. 2006; Gessle e al. 2007; Sche e e al. 2011).
Pa icula ly due o i s d ough -suscep ibili y i dese es special a en ion in he ace o
changing g owing condi ions in he u u e (Fo elli e al. 2009). The sou he n edge o he
ecen dis ibu ion ange o F. syl a ica is mos p obably limi ed by d ough e en s (Jump e
al. 2006; Maxime and Hend ik 2011). In F ance, he obse ed ecen decline in beech o es

2 Backg ound o he Thesis
13
p oduc i i y could be linked o dec easing wa e a ailabili y in ea ly summe (Lebou geois e
al. 2005; Hewi e al. 2011). Fu he mo e dend oclima ological s udies in he Apennine
Moun ains in I aly e ealed a s ong co ela ion o basal a ea inc emen (BAI) o beech s ands
wi h wa e a ailabili y. Since he 1970s BAI has been declining due o wa e limi a ions
(Pio esan e al. 2008). Howe e , also in Cen al Eu ope d ough e en s nega i ely impac he
pe o mance o beech. The excep ional d ough pe iod and hea wa e o e cen al Eu ope in
2003 gene a ed dis inc nega i e e ec s on he egene a ion, g ow h, mo ali y and
pho osyn he ic ac i i y o F. syl a ica (Czajkowski e al. 2005; Leuzinge e al. 2005; Jung
2009; Be sch e al. 2011). Mo eo e , d ough pe iods inc ease he pa hogen- and ungi-
suscep ibili y o Eu opean beech, as shown by Jung (2009) o he pos -2003 yea s o a
Ba a ian F.syl a ica s and.
Figu e 1: D ough damages in Fagus
syl a ica, Ca pinus be ulus and Que cus
obu du ing he 2003 d ough close o
Bay eu h (Bad Be neck). Pho o: C.
Beie kuhnlein
In he u u e, g ow h and dis ibu ion o F. syl a ica is expec ed o decline especially a lowe
ele a ions a he sou he n ange limi (Ma yas e al. 2009; Hlasny e al. 2011), bu also on
xe ic si es in Cen al Eu ope (Czucz e al. 2011). Unde d ough condi ions he egene a ion
o F. syl a ica will be h ea ened, especially in unde s o ey due o i s conse a i e shade-
ole an g ow h s a egy (Robson e al. 2009). Fu he mo e i is obse ed and p ojec ed ha F.
syl a ica loses i s compe i i e ad an age o less d ough -sensi i e species, like Que cus
pe aea, unde wa e limi ed condi ions, especially on he Sou he n and Sou h-Eas e n
dis ibu ion edges (Bonn 2000; Fo elli e al. 2001; Leuzinge e al. 2005; F ied ichs e al.
2009; Cla k e al. 2011; Scha nwebe e al. 2011), whe eas Bol e e al. (2010) showed ha on
he No he n ma gin o he species’ dis ibu ion ange clima e change may b ing compe i i e
ad an ages o F. syl a ica e sus coni e ous ees such as Picea abies.
One u he aspec ha can no be neglec ed is he ac ha wa me win e and sp ing
empe a u es may will lead o ea lie lea lushing in F. syl a ica and hus inc ease he isk o
la e os damage (K ame e al. 2000, K eyling e al. (2012).
2 Backg ound o he Thesis
14
Pinus nig a
Pinus nig a, wi h i s (sub-) Medi e anean dis ibu ion is conside ed e y d ough - ole an
(Isaje 2004; Hube 2011), and was he e o e iden i ied as a po en ial subs i u e o coni e
species, like Picea abies and Pinus syl es is ha a e h ea ened by he changing clima e in
Cen al Eu ope. P. nig a exhibi ed a signi ican ly lowe mo ali y a e a e wo na u al
d ough s han P. syl es is in Eas e n Spain (Ma inez-Vilal a and Pinol 2002). Lebou geois e
al. (1998) showed ha he d ough ole ance o P. nig a saplings is a ibu ed o hei capaci y
o e ec i ely coun e ac wa e s ess by s oma al con ol o anspi a ional wa e loss. Ye , a
ade-o be ween his su i al s a egy unde wa e limi ed condi ions, and a educ ion in
g ow h becomes e iden . Especially, la e-wood o ma ion was ound o be sensi i e o he
p ecipi a ion egime du ing summe mon hs (Biel e al. 2004; Ma in-Beni o e al. 2008;
Ma in-Beni o e al. 2010). In addi ion o an immedia e g ow h esponse o d ough
condi ions, delayed g ow h declines ha e been epo ed o P. nig a. P ecipi a ion and
empe a u e condi ions o he p e ious yea impac he g ow h, especially in he ea ly phase
o he g owing season (Lebou geois 2000; And eu e al. 2007; Ma in-Beni o e al. 2008).
Dend och onological da a sugges ha impac s o inc easing empe a u e on he pe o mance
o P. nig a will di e be ween geog aphic egions (Ma in-Beni o e al. 2010). Wa ming
s imula es g ow h in he mo e empe a e pa s o i s dis ibu ion, while i inc eases wa e
s ess in he Medi e anean egion and hus nega i ely e ec s he g ow h o his species.
2.3. Wi hin-species Di e si y as a Po en ial Tool in Clima e Change Adap a ion
Gi en he abo e men ioned eloci y o clima e change and he in ol ed nega i e impac s on
g assland and o es ecosys ems, na u e conse a ion, ag icul u e and sil icul u e ha e o
adop coun e ac ing p ac ices ha aim on suppo ing dispe sal and acili a ing adap abili y in
o de o main ain ecosys em unc ioning and hus he p o ision o ecosys em goods and
se ices. T adi ional assessmen s o clima e change impac on species wi h clima ic en elopes
(Thomas e al. 2004; Thuille e al. 2005; Kölling 2007) migh misin e p e he adap i e
capaci y o a species o changing condi ions. Wi hin-species di e si y is po en ially impo an
in his con ex .
Popula ions wi hin species o axa a e known o di e pheno ypically. P o enance- ials ha e
a long adi ion in o es y and ha e been conduc ed o mo e han a cen u y now (e.g.
onWuehlisch e al. 1995) E idence om hese ials sugges s ha di e en ia ion wi hin
2 Backg ound o he Thesis
15
species is dis inc , a leas in o es ees. Fo g assland species, only a ew expe imen s
conside ed wi hin-species a ia ion (Fe che and Sha e 1990; Ryse and Aeschlimann 1999).
These di e ences in pheno ypic exp ession could be unde lined on a geno ypic le el by
molecula me hods o o es ees (Mag i e al. 2006) and common g ass species (Michalski
e al. 2010). Gene ally his pheno ypic and gene ic a ia ion is exp essed in local adap a ion o
clima e condi ions o o he abio ic ac o s such as soil ype (e.g. Joshi e al. 2001; Hu o d and
Maze 2003; Sa olainen e al. 2007; Chen e al. 2010; O i and Kigel 2010).
Especially species wi h la ge dis ibu ion anges ha co e a b oad ange o clima ic
condi ions, such as F. syl a ica a e likely o display high le els o wi hin-species a ia ion
and adap a ion o local condi ions. In o es y, he in oduc ion o p o enances o eco ypes
om egions wi hin he dis ibu ion ange o he species wi h cu en clima ic condi ions
simila o he p ojec ed condi ions o he a ge a ea has he e o e been sugges ed as one
po en ial ool in clima e change adap a ion (Heme y 2008; Bol e e al. 2009; Bol e and Degen
2010). Fo g assland species his has been no ye discussed on a men ionable le el, ye , in
spi e o he de ici in s udies which examine local adap a ion and i s implica ion o clima e
adap a ion in g ass species, Macel e al. (2007) ound e idence o a local adap a ion o
clima ic ac o s in wo eco ypes o Holcus lana us.
F. syl a ica exhibi s a high gene ic di e si y wi hin in popula ions in Cen al Eu ope (Konne
1995; Vo nam e al. 2004). Looking a he dis ibu ion ange on a con inen al scale he gene ic
di e ences be ween popula ions become mo e dis inc (Comps e al. 1990; Mag i e al. 2006).
The gene ic composi ion and di e si y o popula ions de e mine hei pheno ypic plas ici y
and hus hei adap i e capaci y (Schabe g e al. 2008; Ma yas e al. 2009), so di e ences in
gene ic con igu a ion mos likely display di e ences in adap i e capaci ies be ween
popula ions. In se e al p o enance ials, dis inc esponses o p o enances o F. syl a ica o
clima ic s esso s, such as d ough , ha e been demons a ed (Sch aml and Rennenbe g 2000;
Peuke e al. 2006; Czajkowski and Bol e 2006). E idence o mac oclima ic adap a ion could
be de ec ed in a Eu opean-wide p o enance- ial ne wo k, whe e he pe o mance o di e en
p o enances was nega i ely co ela ed wi h clima ic dis ance be ween es -si e and o igin o
p o enance (Ma yas e al. 2009). Ye , also in he ield local adap a ions o d ough a e ound.
In he ex ao dina y d y yea 2003, beech popula ions in G eece only expe ienced mild
d ough s ess compa ed o Cen al Eu opean beech o es s (Fo elli e al. 2009), which
indica es an adap a ion o G eek popula ions o d ough condi ions. Especially hese ma ginal
popula ions, which ace mo e ad e se condi ions and a e hus unde s onge gene ic selec ion
2 Backg ound o he Thesis
16
(Wo emann e al. 2011), a e he e o e unde ocus in he sea ch o d ough - esis an eco ypes
(Rose e al. 2009).
Suppo ed by i s sca e ed dis ibu ion ange, P. nig a also shows s ong gene ic di e ences
be ween popula ions and subspecies (Jagielska e al. 2007; So o e al. 2010). This gene ic
di e en ia ion is supposed o ha e been enhanced by geog aphic isola ion du ing he
Pleis ocene (Aguinagalde e al. 1997). P o enance ials showed a non-uni o m pe o mance
o P.nig a p o enances om a ious geog aphic o igins (Va elides e al. 2001; Seho e al.
2010); howe e , di e ences in esponse o clima ic s esso s, such as d ough , could no be
p o en ye , as he high d ough ole ance ac oss popula ion migh p e en a s ong selec ion.
Conside ing he ou lined po en ial impac s o changing clima ic condi ions on unc ions and
se ices o g asslands and o es ecosys ems i is impo an o know, whe he speci ic
p o enances o eco ypes o key species a e mo e o less suscep ible o be e adap ed o
clima ic s esso s, such as d ough , hea o os . This knowledge can be c ucial o assess he
po en ial o selec i e ansplan ing o clima e- esis an p o enances o eco ypes o na i e o
exo ic species as a ool o coping and adap a ion s a egy in ag icul u e and o es y o
dampen he ha m ul impac s o such ex emes in he ace o clima e change (Manusc ip s 2,
3, 4, 5, 7, 8, 9). Unlike in economics, ecosys em managemen has ha dly in oduced isk
managemen in o decision making p ocesses un il oday, despi e he s ong isk o an
unce ain u u e in e ms o clima ic condi ions (Knoke e al. 2005; Hanewinkel e al. 2011).
In economics, he isk o a comple e loss o p o i s is educed by a di e si ica ion o
in es men s. This e ec is called he po olio e ec and was desc ibed by Ma kowi z (1952).
In ecology, a compa able concep , he insu ance hypo hesis, desc ibes a he posi i e e ec o
biodi e si y on ecosys em unc ioning and eliabili y, as he highe numbe o species, he
mo e likely he unc ion o a ailing species can be adop ed by o he species in he sys em
(Yachi and Lo eau 1999). The con e sion o monocul u es in o mixed o es s, i.e. an inc ease
o species di e si y as insu ance agains ad e se bio ic and abio ic impac shas become
popula o e he las decades (Knoke e al. 2005), ye he ole ha wi hin-species di e si y
could play in his con ex jus ecen ly came in o ocus o o es science and managemen .
Wi h espec o he desc ibed unce ain ies and he po en ial posi i e e ec s o biodi e si y on
isk aba emen , he mixing o p o enances has been sugges ed by se e al au ho s (Kols öm e
al. 2011; F asca ia-Lacos e and Fe nández-Manja és 2012). Howe e , e idence has o be
p o ided whe he an an h opogenic enhancemen o geno ypic di e si y and pheno ypic
plas ici y, e.g. by in e mixing local, highly-adap ed and e y plas ic p o enances om
2 Backg ound o he Thesis
17
di e en clima ic egions, may main ain high yields unde a ou able condi ions and secu ing
ecosys em unc ioning, pe sis ence and se ices unde ex eme condi ions (Manusc ip 8).

3 Synopsis o he Thesis
18
3. Synopsis o he Thesis
3.1. Gene al Objec i es and App oach
Conside ing he challenges ha ecosys em managemen , o es y, ag icul u e, and na u e
conse a ion will ha e o mee in he ace o clima e change, he o e all objec i e o his
hesis was o
• assess he po en ial o he selec i e use o wi hin-species a iabili y (p o enances,
eco ypes) as a ool o clima e change adap a ion and o
• iden i y p o enances o eco ypes o key g assland and o es species, which a e be e
adap ed o u u e clima e condi ions, especially o clima ic ex eme e en s, such as
p olonged d ough pe iods.
The i s Manusc ip (Manusc ip 1) in oduces he opic o Assis ed Coloniza ion o
Assis ed Mig a ion o species o eco ypes/p o enances and discusses i s po en ial as a ool in
clima e change adap a ion. This is inc easingly discussed as a p oac i e s a egy bu s ill he e
is insu icien knowledge abou he p ospec s o success and isk.
Then he aim was o depic possible u u e clima e condi ions o Bay eu h/Ge many wi h he
help o egional clima e models, such as REMO (B G 2009) based on he IPCC emission
scena io A1B (IPCC 2000). In a second s ep, egions wi hin he dis ibu ion ange o he
species es ed in his s udy (Fagus syl a ica, Pinus nig a, A hena he um ela ius, Alopecu us
p a ensis, Fes uca p a ensis, and Holcus lana us) we e iden i ied, whe e cu en clima e
condi ions a e close o hose p ojec ed o Bay eu h/Ge many o he 2071-2100 pe iod. Seed
ma e ial was ob ained om hese egions, whe e possible, as we assumed a highe
adap edness o hese p o enances/eco ypes o his speci ic clima ic en i onmen .
In a las s ep, 1-yea old plan s o he selec ed p o enances o he abo e men ioned species
we e exposed o a empe a u e ea men (wa ming and con ol) and o a p ecipi a ion
ea men (d ough and con ol). The clima e manipula ions we e ully c ossed esul ing in
ou ea men s (con ol, d ough , wa ming, wa ming & d ough ), which we e eplica ed h ee
imes each, esul ing in an expe imen al design wi h 12 expe imen al uni s (g eenhouses). All
plan s we e plan ed indi idually in po s.
3 Synopsis o he Thesis
19
Figu e 2a: Le : O e iew o he expe imen al si e wi h he 12 expe imen al uni s (g eenhouses).
Figu e 2b
:
Inside an expe imen al uni wi h wa ming ea men (wind-shel e s and UV-lamps) wi h
po ed F. syl a i a saplings
Figu e 3: Biomass ha es o g assland species in 2009
3 Synopsis o he Thesis
20
In 2009 p o enances o he ou g ass species and P. nig a we e es ed, in o de o iden i y
eco ypes ha a e op imally adap ed o he p ojec ed u u e clima e condi ions (Manusc ip s 2
and 4).
In yea 2010 p o enances o F. syl a ica we e exposed o he same ea men s (Nagy e al. in
p ep., no included in his hesis) A he same ime a simila expe imen was conduc ed in
Landau/Ge many wi h F. syl a ica, ye wi hou he wa ming ea men , bu wi h wo di e en
soil ypes, o asses he in e ac ion o p o enance, clima e and o he abio ic ac o s
(Manusc ip 3). Landau in Rhineland-Pala ina e lies be ween he Uppe Rhine Plain and he
Pala ina e o es and ea u es an annual mean empe a u e abou 2° abo e he annual a e age
o Bay eu h. A s udy in which he Bay eu h expe imen al wa ming is compa ed wi h he
“geog aphic” wa ming based on he wo es si es is in p epa a ion (no included in his
hesis).
Like men ioned in he in oduc ion, la e os e en s a e expec ed o be s ill pa o a Cen al
Eu opean clima e e en unde gene ally wa me condi ions in he u u e. The e o e, po en ial
candida e p o enances o eco ypes ha migh eplace o complemen local ones, ha e o
p o e ha hey a e able o cope wi h hese os e en s. P o enances o eco ypes o he lis ed
g ass species and o Fagus syl a ica we e exposed o a la e os ea men in o de o assess
whe he p o enances om less os -p one si es show a highe la e os sensi i i y.
Addi ionally, he impac o p eceding wa ming ea men s on la e os sensi i i y was es ed
(Manusc ip s 6, 7).
Figu e 4: Lea damage a e la e os ea men
o F. syl ica
Pinus nig a is a non-na i e coni e ous species ha is conside ed o be a po en ial candida e o
eplace clima e- h ea ened na i e coni e ous species like Picea abies. Ye he main pa s o i s
dis ibu ion ange a e loca ed in he Medi e anean egion wi h less se e e win e s compa ed
o Cen al Eu ope o Ge many. Cold ha diness o P. nig a p o enances was es ed in he lab
3 Synopsis o he Thesis
21
by he Rela i e Elec oly e Leakage me hod (REL) and he in luence o he p eceding clima e
ea men s on he cold ha diness was de e mined. Fu he mo e he os ole ance o P. nig a
was compa ed o he one o na i e coni e s (Manusc ip 5).
Figu e 5: Cold Ha diness measu emen s (REL) in he lab o
p o enances o P. nig a and o he local coni e species.
In he cou se o his s udy, no e idence was ound ha i will be likely o iden i y one single
“bes -adap ed” and so o say op imal p o enance o he u u e condi ions o a speci ic si e
such as Bay eu h. Clima e change is a mo ing a ge , bu addi ionally, p o enances exhibi
complex esponse pa e ns o combined and sequen ial changes. Ra he i seems o be
easonable o enhance gene ic di e si y and pheno ypic plas ici y o o es s ands, gene ally.
A model was de eloped in o de o es he hypo hesis ha a mixing o di e en p o enances
may educe he isk o a o al b eakdown o popula ions unde ex eme e en s o he isk o
high yield losses unde a ou able condi ions. The model was ed wi h inc emen da a o P.
nig a and F. syl a ica om he abo e men ioned expe imen s (Manusc ip 8).
3.2. Rema ks on Me hodological Challenges
Du ing he s udy many concep ual and me hodological challenges a ose. Fi s ly, when ying
o de e mine he a ge a eas, wi h clima e condi ions close o p ojec ed ones o Ge many and
om which o ob ain he seed ma e ial o he expe imen s, he p oblem was o se le on an
emission scena io on which p ojec ions should be based on. Finally he A1B scena io was
used, as his co esponds wi h he 2.0°C a ge and is “poli ically a ou ed”, e en i we we e
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5 Decla a ion o Own Con ibu ion
34
5. Decla a ion o Own Con ibu ion
Concep : Idea o he s udy and de elopmen o expe imen al design o de elopmen o
concep s o opinion a icle.
Da a acquisi ion: O ganiza ion and execu ion o da a acquisi ion measu emen s oge he wi h
he help o echnicians, s uden s and in e ns.
Da a analysis: S a is ical analysis o da a and illus a ion in ables and igu es.
W i ing: W i ing he manusc ip s, including li e a u e esea ch
Edi ing: P oo - eading and g amma edi ing, including commen s and inpu s om co-au ho s
and hei in eg a ion in he manusc ip and p epa a ion o esubmissions a e he manusc ip
was e iewed by he jou nals e e ees.
Manusc ip 1:
Res o a ion Ecology
19(4), 433-440 (2011)
Assis ed coloniza ion: a ques ion o ocal uni s and ecipien locali ies
Jue gen K eyling, To s en Bi ne , Anja Jaeschke, Anke Jen sch, Manuel Jonas S einbaue ,
Daniel Thiel, Ca l Beie kuhnlein
Concep : 10 %
Da a acquisi ion: ( e iew a icle wi hou da a)
Da a analysis: -
W i ing: 0 %
Edi ing: 10 %
5 Decla a ion o Own Con ibu ion
35
Manusk ip 2:
Fo es Ecology and Managemen 270, 200-208 (2012)
Uni o m d ough and wa ming esponses in Pinus nig a p o enances
despi e speci ic o e all pe o mances
Daniel Thiel, Lau a Nagy, Ca l Beie kuhnlein, Ge ha d Hube , Anke Jen sch, Monika
Konne , Jue gen K eyling
Concep : 50 %
Da a acquisi ion: 90 %
Da a analysis: 90 %
W i ing: 100 %
Edi ing: 50 %
Manusk ip 3:
Eu opean Jou nal o Fo es Resea ch (in p ess)
Di e en eac ions o cen al and ma ginal p o enances o Fagus syl a ica
o expe imen al d ough
Daniel Thiel, Lau a Nagy, Ca l Beie kuhnlein, Kolja Egen, Ge ha d Hube , Anke Jen sch,
Monika Konne , Jue gen K eyling, Cons anze Buhk
Concep : 25 %
Da a acquisi ion: 10 %
Da a analysis: 100 %
W i ing: 100 %
Edi ing: 25 %
5 Decla a ion o Own Con ibu ion
36
Manusk ip 4:
Jou nal o Ecology 99, 703-713 (2011)
SPECIAL FEATURE: ECOLOGICAL CONSEQUENCES OF CLIMATE EXTREMES
Eco ypes o Eu opean g ass species espond di e en ly o wa ming and
ex eme d ough
Ca l Beie kuhnlein, Daniel Thiel, Anke Jen sch, E elin Willne , Jue gen K eyling
Concep : 25 %
Da a acquisi ion: 100 %
Da a analysis: 75 %
W i ing: 25 %
Edi ing: 25 %
Manusc ip 5:
En i onmen al and Expe imen al Bo any 78, 99-108 (2012)
Cold ha diness o Pinus nig a A nold as in luenced by geog aphic o igin,
wa ming, and ex eme summe d ough
Jue gen K eyling, Guido L.B. Wiesenbe g, Daniel Thiel, Ch is ian Wohl a , Ge ha d Hube ,
Julia Wal e , Anke Jen sch, Monika Konne , Ca l Beie kuhnlein
Concep : 10%
Da a acquisi ion: 25 %
Da a analysis: 10%
W i ing: 10%
Edi ing: 10%
5 Decla a ion o Own Con ibu ion
37
Manusc ip 6:
Eu opean Jou nal o Fo es Resea ch 131(3), 717-725 (2012)
La e os sensi i i y o ju enile Fagus syl a ica L. di e s be ween sou he n
Ge many and Bulga ia and depends on p eceding ai empe a u e
Jü gen K eyling , Daniel Thiel , Lau a Nagy, Anke Jen sch, Ge ha d Hube , Monika Konne ,
Ca l Beie kuhnlein
Concep : 25%
Da a acquisi ion: 50 %
Da a analysis: 25 %
W i ing: 10 %
Edi ing: 25 %
Manusc ip 7:
Ecog aphy, 35(3), 268-275 (2012)
Eco ypic di e en ia ion and pas clima ic expe ience in luence he esponse
o la e sp ing os in ou common g ass species in Cen al Eu ope
Jue gen K eyling, Daniel Thiel, Ka in Simmnache , E elin Willne , Anke Jen sch, Ca l
Beie kuhnlein
Concep : 25 %
Da a acquisi ion: 50 %
Da a analysis: 25 %
W i ing: 10 %
Edi ing: 25 %
6 Manusc ip s
44
Table 2:
A gumen s agains assis ed coloniza ion
.
The cons Re e ences (no comp ehensi e)
Risk o ad e se e ec s on na i e species
composi ion and ecosys em unc ioning (as shown by
examples o in asi e species)
Chapin e al. 2007; Da idson & Simkanin 2008; Hoegh-
Guldbe g e al. 2008; Hun e 2007; McLachlan e al.
2007; Muelle & Hellmann 2008; Riccia di & Simbe lo
2009; Richa dson e al. 2009; an de Veken e al. 2008;
Willis e al. 2009
High isks o ecipien ecosys ems e en o sho
dis ance ansloca ions
Da idson & Simkanin 2008; Muelle & Hellmann 2008;
Riccia di & Simbe lo 2009
Single species app oach un imely in conse a ion Da idson & Simkanin 2008
Ex-si u conse a ion mo e e ec i e Da idson & Simkanin 2008
Impedimen o e o s o p ese e o es o e habi a s Hun e 2007; Riccia di & Simbe lo 2009; Richa dson e
al. 2009
P oblema ic iden i ica ion o ecipien locali ies wi h
impe ec knowledge on ecology and clima e change
del Cas illo e al. 2009; Hun e 2007; McLachlan e al.
2007
Technical easibili y gene ally ques ionable Pelini e al. 2009; Riccia di & Simbe lo 2009
Technical easibili y o a e and endange ed species
no p o ided
Hun e 2007; Muelle & Hellmann 2008
Lack o p edic i e me hods o isk assessmen Riccia di & Simbe lo 2009
Some cu en clima e condi ions a e wi hou u u e
de ini ion, assis ed coloniza ion o species om
hese clima es is no easible
Williams e al. 2007
Unknown cos s and accep abili y Richa dson e al. 2009
C ea ion o a biased lo a o auna Schwa z e al. 2009
Biological homogeniza ion an de Veken e al. 2008
An impo an esea ch ques ion wi h ega d o assis ed coloniza ion is whe he he a ge
species a e di ec ly limi ed by speci ic clima ic condi ions. A di ec dependence o species on
cu en mac o-clima e is ques ionable o many species (Pea son & Dawson 2003; Guisan &
Thuille 2005). E en cu en pa e ns o ec o he m dis ibu ions a e a om equilib ium wi h
cu en mac o-clima e (A aujo & Pea son 2005). The bio ic en i onmen wi h i s associa ed
mic o-clima es may be o highe ele ance o many a e species han mac o-clima ic
condi ions (Ellison e al. 2005; Ha sch e al. 2009; Pelini e al. 2009).
Ano he con ibu ion o unce ain y is he limi ed knowledge on e ec i e dispe sal a es
(Cla k e al. 2003; Higgins e al. 2003). Ra i y and s ochas ici y inhe en in long dis ance

6 Manusc ip s
45
dispe sal s ongly limi ou abili y o o ecas he sp ead o ocal uni s (Na han e al. 2008).
Finally, ecen indings indica e ha en i onmen al change, including clima e change, can
e en cause apid
pheno ypic change h ough bo h ecological and e olu iona y p ocesses
especially in small popula ions (Ozgul e al. 2009). Common species, on he o he hand,
gene ally possess high gene ic a iabili y which allow o adap a ion ia selec ion (Bischo e
al. 2010). Species ep oducing p ima ily h ough ege a i e means, howe e , may be a a
disad an age. Such kind o bio ic adap a ions a e neglec ed in scena ios o biodi e si y loss.
Focal uni s – wha should be mo ed?
Ra e and endange ed species a e challenged mos by clima e change and a e he e o e usually
discussed as ocal uni s o assis ed coloniza ion. We see, howe e , h ee easons why a e
and endange ed species a e no well sui ed o assis ed coloniza ion e o s:
(1) Fo many a e and endange ed species no adequa e ecipien locali y can be ound. Cu en
cen e s o a e species dis ibu ions a e loca ed in a eas wi h highly speci ic clima e
condi ions, which a e p edic ed o sh ink disp opo iona ely unde u u e clima e change
(Ohlemülle e al. 2008) o e en be los globally (Williams e al. 2007). No el condi ions may
u he be p oduced by in e ac ions among clima e, local s a ic en i onmen al condi ions (e.g.,
soils), and local species composi ions.
(2) The collec ion o su icien numbe s o indi iduals o es ablishmen needs is ha dly
possible wi hou ha ming local popula ions o endange ed species. Willis e al. (2009) used
wo bu e ly species and demons a ed he sho - e m (6 yea s) easibili y o assis ed
coloniza ion in a ield expe imen . They wo ked wi h common species and ansloca ed 500
and 600 indi iduals o he wo species espec i ely. Based on a me a-analysis o published
s udies, T aill e al. (2007) sugges a minimum o 1,650 – 100,000 indi iduals o iable
popula ion sizes o insec s in o de o gain a pe sis ence p obabili y o 99% wi hin a ime
ame o 40 gene a ions. Collec ing such numbe s would cause ex i pa ions o sou ce
popula ions wi hou he insu ance o success ul coloniza ion a he new locali y.
(3) Declining popula ions may ha e al eady passed h ough a gene ic bo leneck and assis ed
coloniza ion is doomed o ail in such cases. Remnan popula ions (E iksson 1996) ep esen
only a limi ed pa o he p e ious gene pool. An excess ex inc ion isk o na u ally small
popula ions o bu e lies, o ins ance, can be ela ed o inb eeding dep ession (Sacche i e al.
1998). The his o y o ein oduc ion p ojec s (Muelle & Hellmann 2008) implies ha he isk
o ailed ansloca ions is conside ably highe o a e species (54% ailing) han o common
species (only 14% ailings) (G i i h e al. 1989, Wol e al. 1996). Such ailings a e no only
6 Manusc ip s
46
de imen al o he ansloca ed indi iduals, bu also o he sou ce popula ions due o i s
deple ion o indi iduals.
Based on hese conce ns and aking also he decision ee p o ided by Hoegh-Guldbe g e al.
(2008) in o accoun , assis ed coloniza ion migh only be an op ion o a e y limi ed numbe
o endange ed species wi h su icien ly la ge gene pools and well-known clima ic and
ecological cons ain s ha can be me in new a ge locali ies. Those species mos h ea ened
by clima e change would no appea o be sui able in mos cases. The p oblem o a global loss
o a e species canno be add essed by his means. We conclude ha assis ed coloniza ion o
a e and endange ed species hemsel es is isky, ine ec i e and p obably haza dous o bo h
he ocal uni s as well as o he ecipien ecosys ems in many cases.
The c ea ion and conse a ion o clima e-sa e habi a s, i.e. habi a s ha can be expec ed o
wi hs and clima e change wi hou changes o hei o e all s uc u e and unc ionali y, may be
mo e p o i able in many cases. We al eady discussed ha a di ec dependence o species on
cu en mac o clima e appea s ques ionable o many species. Fine scale dis ibu ions can
u he be expec ed o depend s ongly on mic o-clima ic a ia ion wi hin a landscape.
Tempe a u e a ia ion due o exposi ion and ege a ion co e wi hin he same egion and
ele a ion is s onge han he p ojec ed inc eases in mean empe a u e un il he end o his
cen u y (Sche e and Ko ne 2010). Based on hese conside a ions, a e species migh be
conse ed wi hou he need o mo e hem by adap ing hei habi a s. Such clima e-sa e
habi a s depend mainly on he clima ic ole ance o he dominan plan s, which de e mine
s uc u e and mic o-clima e. Acco ding o Ellison e al. (2005), such species could be called
co e species, keys one species, s uc u al species, ecosys em enginee s o , as u he used
he e, ounda ion species, i.e. “single species ha de ine much o he s uc u e o a communi y
by c ea ing locally s able condi ions o o he species, and by modula ing and s abilizing
undamen al ecosys em p ocesses”.
Fo es y has a long adi ion in assis ed coloniza ion o ounda ion species (Zobel e al. 1987;
Chapin e al. 2007; McKenney e al. 2009). Al hough he main conce n in o es y is no he
p ese a ion o biodi e si y, o es ees p o ide speci ic en i onmen s ha se e as habi a s
o en i e communi ies o plan s, animals, and mic oo ganisms. Ensu ing he p esence o
hese species supplies clima e-sa e habi a s o a mul i ude o dependen species. Fo ins ance,
se e al housand species, such as plan s, insec s o ungi, depend on he Eu opean beech
(Fagus syl a ica) as a ounda ion species (Kölling e al. 2005). The loss o such ounda ion
species is expec ed o ha e cascading, ad e se e ec s on biodi e si y and ecosys em
unc ioning. Replacing na i e species by exo ic ones may sa egua d biomass p oduc ion in
6 Manusc ip s
47
o es y, bu would nega i ely a ec conse a ion alue. Fu he mo e, pollina o sys ems
p o ide ample examples o a e species pe o mance, some imes e en su i al, depending on
he p esence o common species (e.g. Gibson e al. 2006). Co al species (Ac opo idae) wi h
wide anges may se e as ano he example he e. In co als, low-la i ude popula ions exis
which show highe empe a u e ole ances han hose a highe la i udes (Be kelmans & an
Oppen 2006). The la e ha e al eady declined o died o due o inc eased he mal s ess.
In oducing lowe -la i ude, hea -adap ed eco ypes o hese deg aded si es may he e o e se e
as a use ul managemen s a egy (Be kelmans & an Oppen 2006). These examples and
se e al ela ed e iews (e.g. Simbe lo 1998; Booge e al. 2006) sugges ha ounda ion
species and hei ela ionships wi h biodi e si y a e common phenomena. The main ques ion
is he e o e how o e ain o es o e he ounda ion species in imes o change.
Mo ing ounda ion species ou o hei na u al ange (i.e., he po en ial cu en ange in he
absence o human in e e ence), can be expec ed o c ea e e en la ge ecological p oblems
(e.g., in asi eness) han mo ing a e species due o he gene ally highe compe i i e powe o
ounda ion species (Hun e 2007). Founda ion species a e commonly wide anging species
(Ellison e al. 2005) ha exhibi la ge gene ic a ia ion (Ham ick 2004). This is commonly
displayed in s ong local adap a ion, especially o he clima e (Joshi e al. 2001, McKay e al.
2005). Eco ypes can be ound wi hin he cu en dis ibu ion o he ounda ion species ha a e
adap ed o u u e clima e condi ions a loca ions u he no h o a highe al i udes (Figu e 1).
Mo ing such p e-adap ed gene ic esou ces o si es whe e he species is al eady p esen o
whe e i was p esen be o e human in e e ence in case o si es subjec ed o es o a ion e o s
would ensu e ecosys em in eg i y by conse ing he p esence o ounda ion species and by
p o iding clima e-sa e habi a s o a hos o dependan species. Wi hin-species ange
ansloca ion migh also minimize po en ial p oblems wi h in asi eness (bu see below).
6 Manusc ip s
48
Figu e 1:
The p oposed assis ed eco ype coloniza ion o ounda ion species exempli ied o a gi en
si e a Bay eu h, Ge many ( iangle) whe e Eu opean beech (Fagus syl a ica) is he ounda ion species
o he main enance o (semi-) na u al o es s which con ain a high numbe o specialiced species
(Kölling e al. 2005). I s cu en dis ibu ion acco ding o EUFORGEN (2009) is shown in da k g ey.
Cu en clima e equi alen s o he a ge si e a he end o his cen u y ( egional clima e model:
REMO-B G (MPI-M Hambu g); SRES: A1B) based on mean annual empe a u e +/- 0.5°C, colde
mean win e empe a u e, and lowe summe p ecipi a ion a e shown in black (cu en clima e based
on wo ldclim; Hijmnans e al. 2005). Ou lined a e di e en geno ypes o he a ge species (based on
isozyme simila i y; Mag i e al. 2006) which imply ha p e-adap ed eco ypes o di e en gene ic
he i age would be a ailable.
Gene ic di e si y o a ge species inc eases es o a ion success (Bischo e al. 2010). I has
also been p oposed ha a wide selec ion o “mix u es o geno ypes om clima ically local
popula ions” migh bene i sho - e m es ablishmen and long- e m adap a ion po en ial
(McKay e al. 2005). We sugges mo ing one s ep u he by adding gene ic esou ces om
eco ypes o cu en clima ic condi ions compa able o he expec ed u u e clima e o he a ge
a ea (Figu e 1). Including his app oach in o gene al es o a ion concep s seems o be a
p omising scheme o he ecological adap a ion o landscapes o clima e change, which is an
ine i able challenge o any es o a ion e o nowadays. Based on all a gumen s made abo e
and on he associa ed unce ain ies, he assis ed coloniza ion o p e-adap ed eco ypes may
mo e o en be a success ul managemen s a egy han he assis ed coloniza ion o a e and
endange ed species (Figu e 2).
6 Manusc ip s
49
a e and endange ed species p e-adap ed eco ypes
o ounda ion species
1 Selec ion o a ge species
(= endange ed species)
1 Selec ion o si es
(= ecipien locali ies)
• isk o ex inc ion
• in asi e po en ial
• a ailabili y o es o a ion
• p esence o endange ed
species
2 Selec ion o ecipien locali y
• u u e clima e a ecipien
locali y simila o cu en
clima e a o igin
• adequa e bio ic condi ions
• a ailabili y o conse a ion
4 Sampling o seeds/ indi iduals
5 Es ablishmen a ecipien locali y
low high
ce ain y o success
3 Selec ion o p e-adap ed eco ypes
o he iden i ied ounda ion species
• cu en clima e a o igin
simila o u u e clima e a
ecipien locali y
2 Iden i ica ion o ounda ion species
• a ailabili y o seeds/
indi iduals
• es ablishmen
3 Sampling o seeds/ indi iduals
4 Es ablishmen a ecipien locali y
• a ailabili y o seeds/
indi iduals
• es ablishmen
• one o ew widely
dis ibu ed species which
de e mine s uc u e and
unc ioning o he sys em
Figu e 2: Concep ual compa ison o he basic s eps in assis ed coloniza ion o di e en ocal uni s.
The “p e-adap ed eco ypes” app oach s a s om a gi en loca ion which ei he is a ailable o
es o a ion o con ains an endange ed species and aims a he clima e-sa e adap a ion o his gi en
loca ion. The expec ed ce ain y o success o each s ep is based on subjec i e expe knowledge by
he au ho s.
This assis ed eco ype coloniza ion o ounda ion species does no come wi hou isks. Two
majo aspec s o gene ic pollu ion, i.e. he in oduc ion o non-na i e gene ic in o ma ion,
ha e o be conside ed acco ding o McKay e al. (2005). The i s conce n is ha he
in oduc ion o nonlocal geno ypes will c ea e a la ge gene ic load ha causes he es o a ion
o ail. To coun e his, we s ongly ad ise o es he sui abili y o a ge eco ypes
expe imen ally be o ehand and o use o igins om ma ching clima ic and en i onmen al
condi ions (Figu e 1). The usage o mo e han one o igin appea s o be a u he insu ance o
success. The second conce n is ha he nonlocal geno ypes will elimina e locally adap ed
geno ypes. He e, we a gue ha a high pe cen age o es o a ion e o s should be ca ied ou
a e he a ge species is al eady los om he si e and local adap a ion, consequen ly, is los
as well. I he a ge species is s ill p esen , any supe io i y o local adap a ion should also
esul in be e i ness and ( e-) selec ion o he bes adap ed geno ypes.

6 Manusc ip s
50
The consequences o an assis ed coloniza ion o p e-adap ed eco ypes wi hin he ange o he
a ge ounda ion species should be p edic able. No ad e se e ec s on he ecipien sys em
compa able o he expec ed majo isks o mo ing species ou side o hei na u al ange and
e en ou side o hei biogeog aphical con ex a e expec ed simply because he species is o
was al eady p esen , e en dominan , in he sys em be o e he ac ion (Figu e 2). Figu e 1
displays an example o he p oposed app oach. In acco dance wi h Hun e (2007), we s ongly
wa n agains he assis ed coloniza ion o ounda ion species ou side hei biogeog aphical
con ex s.
The e a e u he ca ea s o he idea o assis ed coloniza ion in gene al, as i migh no su ice
o selec ecipien locali ies based on he simila i y o hei expec ed u u e a e age clima e
condi ions (i.e. mean empe a u e and p ecipi a ion sum) wi h he clima e o he ocal uni s’
o igin. Ex eme wea he e en s migh pu much highe selec i e p essu e on su i al han
a e age clima e condi ions while ha dly being quan i iable a a su icien spa ial esolu ion
bo h in he pas and in u u e expec a ions (Jen sch & Beie kuhnlein 2008). Expe imen al
es ing o he ole ance o a ge geno ypes in he ace o expec ed ex eme e en s is,
he e o e, ecommended.
Fu he mo e, he p oblem o a ge ing ecipien locali ies wi h sui able clima e condi ions is a
ques ion o he app op ia e ime scale, as he ecipien locali ies hemsel es a e also subjec o
a changing clima e. McKenney e al. (2009) sugges ha he selec ion o ecipien locali ies in
o es y should depend on he sil icul u al o a ion leng hs o he ocal uni s, wi h clima e
condi ions simila enough o he clima e p e e ences o he ocal uni s o enable a good
su i al a e while also ensu ing good adap a ion owa ds he end o he o a ion. Compa able
sugges ions o he conse a ion o endange ed species a e missing. I is he e o e impo an
o explo e how a species’ dis ibu ions lag behind he shi ing clima ic condi ions, especially
in he an h opogenically modi ied landscape. Placing p e-adap ed eco ypes o species a he
leading edge o hei cu en dis ibu ion may be he mos e ec i e (and conse a i e)
in e en ion.
Conclusions
The deba e abou assis ed coloniza ion as a managemen ool o conse e biodi e si y in he
ace o clima e change is based la gely on wo opposing a gumen s. On he one hand, high
ex inc ion isks a e p ojec ed due o he ac ha ocal uni s migh no adap as enough o
clima e change h ough mig a ion o gene ic adap a ion. In asion biology, on he o he hand,
p o ides e idence ha he in en ional in oduc ion o species may ad e sely a ec ecipien
6 Manusc ip s
51
ecosys ems. A mo e impo an conside a ion may be ha assis ed coloniza ion will no be
easible o many endange ed axa due o hei limi ed a ailabili y and due o missing
ecipien locali ies. We conclude ha he assis ed coloniza ion o single endange ed species is
isky and no use ul in many cases.
The e o e, we p opose ha he s uc u e and mic o-clima e o habi a s con aining a e and
endange ed species could be he ocus o clima e change adap a ion (i.e. clima e-sa e
habi a s) ia he es ablishmen o p e-adap ed eco ypes o he ele an ounda ion species. We
sugges mo ing eco ypes o ounda ion species wi hin he species’ na u al ange in o de o
minimize possible ad e se e ec s. The assis ed coloniza ion o ounda ion species eco ypes
migh p o ide o a means o sus ainable clima e change adap a ion in es o a ion e o s.
The his o y o conse a ion biology in o ms us ha he es o a ion o conse a ion o sui able
habi a including i s dynamic p ocesses (e.g. dis u bance egimes) is he only sus ainable
op ion o manage endange ed species. Building clima e-sa e habi a s by he use o p e-
adap ed eco ypes o ounda ion species may he e o e be a be e in es men owa d he
conse a ion o biodi e si y han aiming a single endange ed species.
Examples om o es y show ha his app oach is applicable wi h ega d o socie al o
legisla i e dimensions. We see a s ong need o u u e esea ch on he ole o ounda ion
species, and on bio ic in e ac ions o ecosys em unc ioning in changing clima es. Ul ima ely,
he human dimension o assis ed coloniza ion, e.g. species selec ion, socie al accep ance,
legisla i e amewo ks and cos s equi e de ailed conside a ion e en i a consensus on
ecological ques ions can be achie ed.
Implica ions o p ac ice
• Assis ed coloniza ion o a e and endange ed species is isky o he a ge species and
he ecipien locali ies and canno be ecommended in gene al.
• Some endange ed species could be conse ed on si e by de eloping clima e-sa e
habi a s h ough he assis ed coloniza ion o p e-adap ed eco ypes o he ele an
ounda ion species (e.g. majo o es ees o co als). This app oach would conse e
s uc u e and mic o-clima e o he habi a and a oid mo ing species ou o hei
na u al ange.
• Rega dless o he conse a ion app oach, es o a ion e o s need o design clima e-
sa e communi ies. The in eg a ion o eco ypes adap ed o he expec ed u u e clima e
should become one pa o es o a ion concep s.
6 Manusc ip s
52
Acknowledgemen s
This wo k was pa ly unded by he "Ba a ian Clima e P og amme 2020" in he join esea ch
cen e “FORKAST” and by he Fede al Agency o Na u e Conse a ion (B N) in he p ojec
“Impac s o clima e change on auna, lo a and habi a s, as well as adap a ion s a egies”. The
au ho s hank wo anonymous e iewe s and he associa ed edi o o aluable commen s on
an ea lie e sion o his manusc ip .
6 Manusc ip s
53
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Ma e ial and Me hods
Expe imen al Si e
The p o enance- ial (EVENT 3) was es ablished in Ma ch 2009 in Bay eu h, Ge many, in
he Ecological-Bo anical Ga dens o he Uni e si y o Bay eu h. I is pa o he EVENT-
expe imen se ies (Jen sch 2010).
The mean annual empe a u e a he si e is 8.2°C wi h a mean annual p ecipi a ion o 724 mm,
which is dis ibu ed bimodally wi h wo majo peaks in June/July and Decembe /Janua y
(da a : Ge man Wea he Se ice).
Subspecies and P o enances
Seeds o six p o enances o Pinus nig a we e ob ained and b ough o ge mina ion a he
Ba a ian Ins i u e o Fo es Seeding and Plan ing (ASP) in Teisendo , Ge many in Ap il
2008. The p o enances s em om au och honal popula ions, excep he p o enance om
No he n I aly o which he s a us is unclea and he Ge man p o enance, which mos
p obably has an Aus ian o igin. Two p o enances (I aly-S, F ance) belong o he subspecies
la icio, he o he s (Ge many, Aus ia, I aly-N, Se bia) a e a ibu ed o he subspecies nig a.
Table. 1: Si e in o ma ion (DE Ge many, AT Aus ia, SRB See bia, IT I aly, FR F ance) o he
p o enances o Pinus nig a used in he EVENT 3-Expe imen (Hube , 2011). Tempe a u e (T),
p ecipi a ion (P) and p ecipi a ion o wa mes qua e (PJJA) ep esen long- e m mean annual alues
(sou ce: Wo ldClim).
Loca ion Ssp. Au och honous
La Lon Ele a ion
(m.a.s.l.) T (°C) P
(mm)
P
JJA(mm)
DE Zellingen nig a no 49°53'17"
09°43'16" 290 9.2 587 206
AT D eis e en nig a yes 47°46'00"
16°11'00" 369 8.4 712 256
SRB K emanske nig a yes 43°49'39"
19°35'22" 866 8.7 964 265
IT-N T ies ino
(Ca so) nig a unclea 45°42'00"
13°49'00" 372 11.4 1212 301
IT-S
Calab ia
(Sila
Cosenza)
la icio yes 39°18'08"
16°20'22" 1500 9 1300 105
FR Pon eils-e -
B esis
la icio
(Co se) yes 44°24'18"
03°58'39" 581 10.8 745 171
In Ap il 2009, he seedlings we e anspo ed o Bay eu h and indi idually plan ed in o 4-li e
po s in a sandy loam subs a e om a local o es op-soil dug-ou (pH 7.27, o al C 1.89%,
o al N 0.15%, plan -a ailable NH4
+
1.79 mg L
−1
, plan -a ailable NO3
−
22.50 mg L
−1
).

6 Manusc ip s
61
Indi iduals we e selec ed andomly o each p o enance o subspecies om all li ing plan s
a plan ing da e. Mean plan size a he s a o he expe imen was 12.2 cm (± 2.5 cm SD).
Expe imen al Design
The po ed indi iduals we e exposed o a empe a u e ea men (wa ming and con ol) and o
a p ecipi a ion ea men (ex eme summe d ough and con ol). The wo clima e ea men s
we e ully c ossed, esul ing in ou clima e manipula ions (con ol, d ough , wa ming,
wa ming and d ough ), which we e eplica ed h ee imes, esul ing in 12 expe imen al uni s
in o al. The p o enance ea men was nes ed wi hin each expe imen al uni , esul ing in a
spli -plo design. Each p o enance was u he eplica ed wi h se en plan s pe expe imen al
uni (nes ed eplica es), esul ing in 21 indi iduals pe ac o ial combina ion o he h ee-
ac o ial design and 672 plan s o e all.
Each expe imen al uni was co e ed by one la ge ain-ou shel e cons uc ed o a s eel ame
(GlasMe all Rieme GmbH, Rahden-Sielho s , Ge many) and co e ed wi h a anspa en
polye hylene shee (0.2 mm, SPR5; He mann Meye KG, Rellingen, Ge many). The edge o
he ain-ou shel e s was 80 cm all and pe mi ed he pene a ion o nea ly 90% o he
pho osyn he ically ac i e adia ion.
The con ol p ecipi a ion egime simula ed he local daily 30-yea a e age p ecipi a ion. The
applica ion was done wice a week wi h collec ed ain wa e .
In 2009 he d ough ea men consis ed o a 42-day pe iod wi hou i iga ion, which
ep esen s a local d ough e en wi h a s a is ical ecu ence p obabili y o 1000 yea s. The
ea men s a ed on May 27
h
2009. Soil mois u e was measu ed hou ly o e he whole
du a ion o he expe imen using h ee ECH20 EC-5 mois u e senso s (Decagon De ices,Inc.,
USA) pe ea men . The pe manen wil ing poin was de e mined ia soil ype using a
pedological soil su ey manual (Ad-hoc-AG Boden 2005). App oxima ely h ee weeks a e
he s a o he ea men soil mois u e d opped below he pe manen wil ing poin (pF = 4.2)
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62
Figu e 1: Ai empe a u e a plan heigh and soil mois u e o e he wo yea s o manipula ions. F om
Oc obe 2009 o Ap il 2010 no wa ming ea men ook place. Howe e , he plan s we e unde snow
co e , so hey we e shel e ed om he empe a u e minimums in win e 09/10. In he uppe g aphic
(ai empe a u e) he do ed line ep esen s 0°C, in he lowe one (soil mois u e) he do ed line
ep esen s app oxima ely he pe manen wil ing poin (pF = 4.2). The g ey a ea ep esen s he
wa ming ea men in he uppe g aphic and he d ough pe iod in he lowe .
In he e-wa e ing phase a e he d ough pe iod each indi idual ecei ed 240 ml and ano he
300 ml ou days a e he i s e-wa e ing in o de o p e en he soil mois u e om d opping
below he pe manen wil ing poin again. A e his, he po s we e wa e ed acco ding o he
con ol p ecipi a ion ea men .
In he second yea (2010), no d ough ea men was applied and all plan s ecei ed he same
wa e ing in o de o examine he long- e m e ec s o he p e ious yea (2009) expe imen al
d ough e en .
The wa ming ea men was applied om May 26 h un il Oc obe 25
h
in 2009 and om Ap il
1
s
o Decembe 7
h
in 2010. Wa ming was a ec ed passi ely by wind-shel e s, which educed
wind speed by 70 %, and by black loo -co e s in con as o whi e loo co e s. This
inc eased he a e age ai empe a u e in he wa ming ea men a plan heigh by 1.5 K on
a e age (Figu e 2) compa ed o con ol. In he second yea addi ional IR-lamps we e ins alled
(IR- adia ion wi h app oxima ely 30 W pe m²), which only esul ed in a wa ming by 1.6 K on
a e age (Figu e 2). Wi hin he con ol ea men he empe a u es we e 1.5K highe on
a e age han he ambien condi ions ou side o he expe imen al uni s. The mode a e
addi ional wa ming e ec due o IR-lamps mos p obably esul ed om di e en wea he
pa e ns in he second yea , as he wa ming e ec due o passi e means was mos p onounced
unde di ec sun ligh . The ou h ea men was a combina ion o d ough and wa ming. The
6 Manusc ip s
63
addi ional wa ming ele a ed he d ough ea men e ec by educing he soil mois u e by
abou 1.5% o o al soil mois u e on a e age compa ed o he d ough only ea men (Figu e
2).
Du ing he win e 2009/2010, plan s we e kep ou side he shel e s in-g ound in a sand bed
om Oc obe o Ap il and ecei ed na u al p ecipi a ion.
Response Pa ame e s
In 2009 he heigh o he plan s was measu ed a plan ing ime in Ap il and a e he g owing
pe iod in Oc obe 2009. In 2010 heigh was measu ed again a he end o Oc obe . Inc emen
was calcula ed by sub ac ing he ini ial alue om he alue a he end o he g owing pe iod.
Indi iduals ha died a e he i s yea we e coun ed wi h inc emen se o ze o o he second
yea . In o de no o unde es ima e heigh g ow h due o he ze o inc emen o he dead
indi iduals, he model was un again wi h he dead indi iduals le ou comple ely and yielded
simila esul s.
The numbe o la e al shoo s was eco ded in bo h yea s a he end o he ege a ion pe iod.
Mo ali y was quan i ied a he end o bo h yea s by coun ing he li ing indi iduals.
Fo he phenological analyses, he da e o he s a (needles clea ly isible in lea shea h) and
he end (needles as long as o longe han las yea ’s needles) o olia ion was eco ded o
each indi idual in 2010. Phenological da a o 2009 is missing because he olia ion p ocess
was al eady in p og ess when we ecei ed he plan s.
S a is ics
Analysis o Va iance (ANOVA) combined wi h linea mixed e ec models (package nlme)
we e applied o es o he main and in e ac i e e ec s o he h ee ac o s d ough , wa ming
and p o enance on heigh g ow h, numbe o shoo s and phenology. Da a was es ed o
homogenei y o a iance and no mali y o he esiduals p io o analysis and me he
p e equisi es o linea models. The expe imen al uni was included in he model as a andom
ac o , accoun ing o he spli -plo design (Pinhei o & Ba es 2004). Pos -hoc es s we e
ca ied ou wi h he Gene al Linea Hypo hesis unc ion glh (package mul comp) (B e z
2010).
The main and in e ac i e e ec s o he h ee ac o s comp oising o d ough , wa ming and
p o enance on mo ali y we e es ed by Analysis o Va iance (ANOVA) combined wi h a
Cox p opo ional haza ds eg ession model (package su i al) (The neau 2000). Fu he mo e,
6 Manusc ip s
64
o e i y he esul s o he Cox p opo ional haza ds eg ession model, gene alized linea
mixed models using a binomial dis ibu ion we e applied. The expe imen al uni was included
as a andom ac o again. Pos -hoc es s we e ca ied ou wi h he Gene al Linea Hypo hesis
unc ion glh . All s a is ical analyses we e conduc ed wi h he so wa e R.2.13.0 (R
De elopmen Co e Team, 2011)
Resul s
G ow h
Absolu e heigh g ow h di e ed signi ican ly be ween p o enances in bo h yea s o he
expe imen (ANOVA: 2009: F = 6.1, p = < 0.001; 2010: F = 3.9, p = 0.002; Fig. 2 a and b).
Especially he F ench p o enance exhibi ed signi ican ly less inc emen han he wo I alian
p o enances in 2009 and signi ican ly less han he Sou he n I alian p o enance in 2010.
No signi ican di e ences be ween he p o enances we e ound o ela i e g ow h (ANOVA:
2009: F = 2.1, p = 0.065; 2010: F = 0.9, p = 0.277; Fig. 2 c and d).
Fo bo h, absolu e and ela i e g ow h, no signi ican ea men e ec was de ec ed in he i s
yea (Fig. 2 a and c). Howe e , in he second yea a s ong e ec o he d ough o he
p e ious yea was ound (ANOVA: absolu e g ow h: F = 3.8, p = < 0.001; ela i e g ow h: F
= 15.1, p = 0.005). Plan s g own unde he con ol ea men inc eased in heigh by 6.2 cm o
51% on a e age, whe eas he plan s ha we e exposed o he d ough in 2009 only g ew 3.6
cm o 30% on a e age (Fig. 2 b and d).
6 Manusc ip s
65
Figu e 2: Response o Pinus nig a o wa ming, d ough and p o enance. a) shows he annual g ow h
in cm as a unc ion o ea men and p o enance in 2009 and b) in 2010. c) shows he annual g ow h in
pe cen o he ini ial heigh o 2009 and d) o 2010. e) Depic s he numbe o shoo s pe indi idual
as a esponse o ea men and p o enance in 2009 and ) in 2010 and g) illus a es he pe cen age o
su i ing indi iduals in 2009. In 2010 almos all indi iduals su i ed (>99%). Lowe case le e s
ep esen signi ican homogeneous g oups as e ealed by he pos -hoc es s. F- and p-s a is ics a e
p o ided o p o enance, wa ming, d ough and he in e ac ion be ween wa ming and d ough (w:d).
Signi ican p- alues (<0.05) indica ed by bold le e ing. No signi ican in e ac ions be ween
p o enance and wea he ea men s we e ound. The Cox p opo ional haza d model does no p o ide
F-sa is ics. P o enances a e a anged acco ding o hei geog aphic o igin om eas o wes .
Wa ming had no signi ican impac on heigh g ow h, nei he in 2009 (ANOVA: absolu e
g ow h: F = 0.0, p = 0.976; ela i e g ow h: F = 0.9, p = 0.380), no in 2010 (ANOVA:
absolu e g ow h: F = 0.3, p = 0.582; ela i e g ow h: F = 0.0, p = 0.986).
Sensi i i y o wa ming and o d ough did no di e among he p o enances (ANOVA:
in e ac ion be ween p o enance and d ough : 2009, absolu e g ow h: F = 0.9, p = 0.483;
ela i e g ow h: F = 2.1, p = 0.068; 2010, absolu e g ow h: F = 1.3, p = 0.262; ela i e
g ow h: F = 0.9, p = 0.478; in e ac ion be ween p o enance and wa ming: 2009, absolu e
g ow h: F = 0.8, p = 0.546; ela i e g ow h: F = 1.1, p = 0.379; 2010, absolu e g ow h: F =
0.7, p = 0.613; ela i e g ow h: F = 0.5, p = 0.769).
Numbe o Shoo s
The p o enances did no di e signi ican ly in e ms o he numbe o shoo s in any yea
(ANOVA: 2009: F = 2.1, p = 0.060; 2010: F = 1.9, p = 0.096). D ough , howe e , educed he

6 Manusc ip s
66
numbe o shoo s (ANOVA: 2009: F = 11.1, p = 0.01; 2010: F = 13.3, p = 0.007; Fig. 2 e and
). In 2009 he con ol plan s had 3.3 shoo s on a e age and he plan s unde going d ough
ea men 2.8 shoo s on a e age. In 2010 i was 4.5 (con ol) o 4.1 (d ough ) on a e age.
Numbe o shoo s was no signi ican ly a ec ed by wa ming in any yea (ANOVA: 2009: F =
3.1, p = 0.119; 2010: F = 4.2, p = 0.075). O e all, he p o enances showed no signi ican
a ia ion in hei esponse o he wea he manipula ions (ANOVA: 2009: in e ac ion be ween
p o enance and d ough : F = 1.0, p = 0.413; in e ac ion be ween p o enance and wa ming F =
0.8, p = 0.554, 2010: in e ac ion be ween p o enance and d ough : F = 0.5, p = 0.765;
in e ac ion be ween p o enance and wa ming F = 0.6, p = 0.730).
Needle Phenology
The onse o needle olia ion in 2010 did no di e signi ican ly among he p o enances (Fig.
3) (ANOVA: F = 1.1, p = 0.377).
The wa ming ea men led o a ea lie bud bu s . Needle olia ion s a ed 10.6 days ea lie
when he plan s we e exposed o wa ming compa ed wi h con ol indi iduals (ANOVA: F =
49.1, p = <0.001). Needle olia ion was comple ed 2.3 days ea lie on a e age o plan s unde
wa ming ea men , hough his was an insigni ican end a he 95% con idence le el
(ANOVA: F = 3.6, p = 0.079). The in luence o he d ough ea men on he phenological
de elopmen o lea es was no signi ican a all (ANOVA: F = 0.2, p = 0.655).
6 Manusc ip s
67
Figu e 3: Needle olia ion in 2010 o he di e en p o enances. Ci cles ep esen bud bu s and
iangle he comple ion o needle lushing. The di e en shades o g ey illus a e he ou ea men s.
The ho izon al ba s ep esen he s anda d e o . Wa ming sigini ican ly impac ed bud bu s (ANOVA:
F = 49.1, p = < 0.001).
Gene ally, he p o enances showed no signi ican a ia ion in hei esponse o he wea he
manipula ions in e ms o needle phenology (ANOVA: in e ac ion be ween p o enance and
wa ming: F = 1.3, p = 0.262; in e ac ion be ween p o enance and d ough : F = 0.6, p =
0.738).
Mo ali y
P o enances exhibi ed a signi ican di e ence in e ms o mo ali y in 2009 (ANOVA: p = <
0.001. The Ge man p o enance showed he lowes su i al a e (78.6%), while he Aus ian
p o enance showed he highes su i al a e (97.6%) (Fig. 2 g).
D ough signi ican ly inc eased mo ali y (ANOVA: p = < 0.001). Only 79% o he plan s
subjec ed o he d ough ea men su i ed, whe eas 99.6% o he plan s ha we e no
exposed o d ough (con ol and wa ming) we e ali e a e he i s season (Fig. 2 g).
Wa ming signi ican ly dec eased su i i al (ANOVA: p = 0.011). Howe e , he dec eased
su i al only occu ed in combina ion wi h he d ough ea men (- 13%), hough no
s a is ically signi ican in e ac ion be ween d ough and wa ming could be de ec ed wi h any
o he wo s a is ical me hods applied (ANOVA: in e ac ion wa ming and d ough : p = 0.137;
Fig. 5 a, Tab. 2).
The p o enances showed no signi ican a ia ion in hei esponse o he wea he
manipula ions (ANOVA: in e ac ion be ween p o enance and wa ming: p = 0.444; in e ac ion
be ween p o enance and d ough : p = 0.840).
In 2010 mo ali y was negligible (<1%) and no ends based on ea men o p o enance
e ec s became isible. The GLM model quali a i ely yielded he same esul s.
Discussion
Wi hin-species a ia ion in he absence o clima e pe u ba ions
Pinus nig a has a e y agmen ed dis ibu ion ange ac oss he (sub-) Medi e anean egion.
I is spli in o six subspecies, which a e again subdi ided in o a ie ies (Isaje 2004).
Especially in he wes e n pa o i s ange, P. nig a popula ions mos likely su i ed he las
glacial maximum in e ugia (A zal-Ra ii e al. 2007), which implies a long- e m sepa a ion o
6 Manusc ip s
68
popula ions. P. nig a shows a high gene ic dis ance be ween popula ions (Scal soyiannes e
al. 2009) and hus i can be expec ed ha pheno ypic di e ences also occu be ween
popula ions and p o enances. The di e en pe o mance o he six p o enances in e ms o
heigh g ow h in ou expe imen is he e o e no su p ising. The sou he n I alian p o enance
showed he highes o al heigh inc emen in bo h yea s, whe eas he pe o mance o he
F ench p o enance was lowes in 2009 and 2010. The signi ican dispa i y be ween he wo
p o enances is unexpec ed, because hey bo h belong o he same subspecies la icio (Hube
2011). This a ia ion in g ow h may indica e a dis inc in e -popula ion gene ic a ia ion e en
wi hin a subspecies.
In 2009, he No he n I alian p o enance pe o med a he same le el as he Sou he n I alian
p o enance wi h high g ow h a es ac oss all ea men s, despi e he ac ha i o igina es
om a si e wi h high annual p ecipi a ion and also high p ecipi a ion du ing he ege a ion
pe iod (Tab. 1), whe e a po en ial local adap a ion should lead o a highe d ough -
suscep ibili y han o si es wi h low p ecipi a ion. Howe e , So o e al. (2010) ound a
posi i e co ela ion be ween summe p ecipi a ion and wi hin-popula ion di e si y o Pinus
nig a, as egions wi h egula p ecipi a ion would suppo la ge and demog aphically mo e
s able popula ions, han egions wi h a highe summe d ough equency. This p obable high
gene ic di e si y may be he eason o he ela i ely s able g ow h and ela i ely high
su i al a e wi hin his p o enance.
Ano he su p ising pa e n is e ealed by he su i al a es. The Ge man p o enance exhibi s
a signi ican ly lowe su i al a e han he Aus ian p o enance. Howe e , he Ge man
p o enance was ounded in 1909/1910 wi h seeds om Aus ia (Hube 2011). Clima ic
condi ions and ele a ion a e also simila be ween he places o o igin o hese wo
p o enances. Ne e heless hey show dis inc pe o mances, which may poin again o he ac
ha popula ions s ongly a y gene ically, despi e a close spa ial ela ionship.
Budbu s was no in luenced by p o enance. Though he No he n I alian p o enance
comple ed needle olia ion signi ican ly la e han he o he p o enances, i seems ha
p o enance o gene ic di e ences do no in luence phenological beha io as much as he
empe a u e signal does. This is in acco dance wi h indings o o he ee species (Vi asse e
al. 2009) and indica es a s ong po en ial o adap a ion o changing clima ic condi ions.
Species Response o D ough and wa ming
A educ ion in summe p ecipi a ion by abou 20% is p ojec ed o Cen al Eu ope acco ding
o he A1B scena io (B G 2009), wi h an inc eased isk o d ough pe iods (Rowell 2009).
6 Manusc ip s
69
These al e ed clima ic condi ions will ha e a s ong impac on empe a e o es ecosys ems.
Especially indigenous coni e s will su e mos unde d y summe s (Kölling 2009, Lindne e
al. 2010). Pinus nig a has o be able o cope wi h he p ojec ed clima ic condi ions, in o de o
come in o conside a ion as a po en ial subs i u e o he endange ed locals.
D ough was he main sou ce o a ia ion in g ow h and su i al ac oss all p o enances in ou
expe imen . Ye , conside ing he ex emeness o he d ough ea men (42 days wi hou
p ecipi a ion and mo e han wo weeks soil mois u e below he pe manen wil ing poin ),
su i al and g ow h a es emained ema kably high. Su p isingly, he e was no d ough
e ec on heigh g ow h in he i s yea , when he ac ual d ough ea men ook place.
Howe e , in he second yea heigh inc emen s ongly dec eased o indi iduals exposed o
d ough in he p e ious yea , despi e no u he d ough ea men . Ac oss he p o enances he
saplings o P. nig a exhibi ed a lagged esponse o wa e sho age by main aining high g ow h
a es in he d ough yea and a heigh g ow h d op in he subsequen yea . Simila lagged
e ec s o d ough a e epo ed by o he s udies. Lebou geois (2000) ound signi ican
co ela ions be ween ea ly adial g ow h and empe a u e (nega i e) and p ecipi a ion
(posi i e) in Oc obe o he p e ious yea in a s udy on clima ic impac s on he g ow h o
Co sican Pine (Pinus nig a ssp. la icio a . Co sicana). Analogous esul s we e epo ed by
Ma in-Beni o e al. (2008). They show ha he ea ly wood g ow h o Pinus nig a in
Sou heas e n Spain was nega i ely impac ed by a p e ious yea d ough e en . Du ing wa e
s ess, newly ixed ca bon is a he used o osmo ic adjus men o imp o e d ough esis ance
o is alloca ed o oo g ow h in o de o acqui e mo e wa e . This physiological esponse may
educe ca bohyd a e ese es which a e necessa y o nex yea ’s g ow h (Lebou geois 2000,
Ma in-Beni o e al. 2008). Such lagged esponses ha e been con i med o o he coni e s
(And eu e al. 2007, de G andp e e al. 2011). Howe e , all hese ocused on adul ees, wi h
signi ican s o age capaci y o ca bohyd a es. A delayed g ow h esponse o seedlings, as
p esen ed in ou s udy is a no el inding. Compa ed o adul ees, seedlings ha e less s o age
capaci ies, due o a smalle sha e o woody issue. The e o e, a educed ca bohyd a e
p oduc ion o a ealloca ion o assimila es should lead o a mo e immedia e heigh g ow h
eac ion. Main aining a high g ow h a e despi e wa e s ess migh he e o e be ela ed o he
s ong inc ease in mo ali y o ju enile s ages du ing and a e wa e s ess. Fu he mo e he
d ough pe iod migh al e soil o ganic ma e decomposi ion and ni ogen mine aliza ion
p ocesses, wha can lead o ca y-o e e ec s in o he nex yea s ( an de Molen e al. 2011).
The su i al o he pine seedlings in 2009 depended signi ican ly on wa e condi ions.
Ne e heless he o e all su i al a e was ela i ely high (79%) in he d ough ea men s,
6 Manusc ip s
76
K eyling J., Wiesenbe g G., Daniel, T., Wohl ah C., Hube , G., Wal e , J. e al., 2012. Cold ha diness o Pinus
nig a A nold as in luenced by geog aphic o igin, ex eme summe d ough and g adual wa ming. En i on.
Exp. Bo ., 78, 99-108.
Kupa inen, A., Sa olainen, O., Schu , F.M., 2010. Inc eased mo ali y can p omo e e olu iona y adap a ion o
o es ees o clima e change. Fo . Ecol. Manage. 259, 1003-1008.
Lamy, J.-B., Bou ie , L., Bu le , R., Plomion, C., Cocha d, H., Delzon, S., 2011. Uni o m selec ion as a
p ima y o ce educing popula ion gene ic di e en ia ion o ca i a ion esis ance ac oss a species ange.
PLoS One 6, e23476.
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po en ial, pho osyn hesis, s oma al conduc ance and g ow h in wo black pine a ie ies. Ann. Fo es Sci. 55,
287-299.
Lebou geois, F., 2000. Clima ic signals in ea lywood, la ewood and o al ing wid h o Co sican pine om
wes e n F ance. Ann. Fo es Sci. 57, 155-164.
Lindne , M., Ma oschek, M., Ne he e , S., K eme , A., Ba ba i, A., Ga cia-Gonzalo, J. e al., 2010. Clima e
change impac s, adap i e capaci y, and ulne abili y o Eu opean o es ecosys ems. Fo . Ecol. Manage. 259,
698-709.
Lu, P., Man, R., 2011. Assessmen o assis ed mig a ion e ec s on sp ing bud lush in whi e sp uce (Picea
glauca [Moench] Voss) seedlings. Fo es . Ch on. 87, 391-397.
Lucic, A., Mladeno ic-D inic, S., S a e o ic, N., Isaje , V., La adino ic, V., Rakonjac, L. e al., 2010. Gene ic
di e si y o Aus ian Pine (Pinus Nig a ARNOLD) popula ions in Se bia e ealed by RAPD. A ch. o Biol.
Sci. 62, 329-336.
Mahe ali, H., Delucia, E.H., 2000. Xylem conduc i i y and ulne abili y o ca i a ion o ponde osa pine g owing
in con as ing clima es. T ee Phys. 20, 859-867.
Mahe ali, H., Williams, B.L., Paige, K.N., Delucia, E.H., 2002. Hyd aulic di e en ia ion o Ponde osa pine
popula ions along a clima e g adien is no associa ed wi h eco ypic di e gence. Func. Ecol. 16, 510-521.
Ma acchi, G., Si o enko, O., Bindi, M., 2005. Impac s o p esen and u u e clima e a iabili y on ag icul u e and
o es y in he empe a e egions: Eu ope. Clim. Change 70, 117-135.
Ma in-Beni o, D., Che ubini, P., del Rio, M., Canellas, I., 2008. G ow h esponse o clima e and d ough in
Pinus nig a A n. ees o di e en c own classes. T ees-S uc . Func . 22, 363-373.
Ma in-Beni o, D., del Rio, M., Canellas, I., 2010. Black pine (Pinus nig a A n.) g ow h di e gence along a
la i udinal g adien in Wes e n Medi e anean moun ains. Ann. Fo es Sci. 67, 401-401.
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Ma inez-Vilal a, J., Cocha d, H., Mencuccini, M., S e ck, F., He e o, A., Ko honen, J.F.J. e al., 2009.
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Ma a uga, M., Haase, D.L., Isaje , V., 2010. Dynamics o seed imbibi ion and ge mina ion o Aus ian pine
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San ac uz-Va ela, A., 2009. Al i udinal gene ic a ia ion in Pinus ha wegii Lindl. I: Heigh g ow h, shoo
phenology, and os damage in seedlings. Fo . Ecol. Manag. 257, 836-842.
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424.
Zobel, B.J., an Wyk, G., S ahl, P., 1987. G owing exo ic o es s, John Wiley & Sons, New Yo k.
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6.3. Manusc ip 3:
Di e en eac ions o cen al and ma ginal p o enances o Fagus syl a ica
o expe imen al d ough
Eu opean Jou nal o Fo es Resea ch (in p ess)
Daniel Thiel, Jue gen K eyling
a
, Sab ina Backhaus
b
, Ca l Beie kuhnlein
a
, Cons anze Buhk
c
,
Kolja Egen
c
, Ge ha d Hube
d
, Monika Konne
d
, Lau a Nagy
b
, Anke Jen sch
b
a
Biogeog aphy, BayCEER, Uni e si y o Bay eu h, Uni e si ä ss .30, 95447 Bay eu h,
Ge many;
b
Dis u bance Ecology, BayCEER, Uni e si y o Bay eu h, Uni e si ä ss .30, 95447
Bay eu h, Ge many;
c
Geoecology/Physical Geog aphy, Uni e si y o Koblenz/Landau, Fo s aße 7, 76829
Landau, Ge many;
d
Ba a ian O ice o Fo es Seeding and Plan ing, Fo s am spla z 1, 83317 Teisendo ,
Ge many
Keywo ds
Ex eme e en s, local adap a ion, pheno ypic plas ici y, plan -clima e in e ac ions, p o enance
ial, wi hin-species di e si y
Abs ac
Clima e ex emes a e expec ed o inc ease in equency and magni ude as a consequence o
global wa ming, h ea ening he unc ioning, se ices and goods o o es ecosys ems. Ac oss
Eu ope, he ecologically and economically impo an ee species Fagus syl a ica is expec ed
o su e pa icula ly unde such condi ions.
The egional in oduc ion o p o enances om d ie and wa me clima es is one op ion o
adap beech o es ecosys ems o hese ad e se e ec s o clima e change. Ma ginal
popula ions om he d ough -p one sou he n and no h-eas e n edges o he species’
6 Manusc ip s
79
dis ibu ion come in o ocus in sea ch o sui able candida es o Cen al Eu opean deciduous
o es s.
He e, we es h ee ma ginal p o enances (Spain, Bulga ia and Poland) and h ee p o enances
om he cen e o he dis ibu ion ange (Ge many) o hei esponse o d ough in wo
di e en soil ypes (sand, loam) in a ull- ac o ial common-ga den expe imen in Landau,
Ge many.
D ough impac ed all g ow h pa ame e s nega i ely (lea damage +22% (pe cen age poin s),
heigh -40% and diame e inc emen -41%) and he sandy subs a e exace ba ed his e ec .
Howe e , p o enances di e ed in hei esponse o d ough and soil ype. E idence o a local
adap a ion o summe d ough was de ec ed, especially in e ms o mo ali y a es. The
Bulga ian and Spanish p o enance showed a s able pe o mance unde d ough condi ions
(BG -27% in diame e inc emen ; ES -32%), compa ed o he Polish (-48%) o he mos
sensi i e Ge man p o enances (-57%), ye o Bulga ia on a low le el o o al inc emen . This
may indica e a ade-o be ween d ough - ole ance and g ow h.
The e o e, a sole ocus on d ough - esis an ma ginal p o enances seems o no be conduci e,
as hey migh be less adap ed o o he clima ic ac o s, e.g. os , as well. Howe e ,
in e mixed wi h local Cen al Eu opean p o enances hese may ac as unc ional insu ance in
u u e d ough -p one o es s ands.
In oduc ion
Alongside an inc ease in a e age empe a u e, Global Wa ming is expec ed o igge an
inc ease in magni ude and equency o clima ic ex emes such as p olonged d ough e en s
(IPCC 2012). This will especially a ec he unc ioning o ecosys ems wi h long-li ing
o ganisms, such as o es s, and may lead o a declining p o ision o o es goods and se ices
(Allen e al. 2010, Lindne e al. 2010). The eloci y o hese changes in ecosys em
unc ioning may exceed he na u al dispe sal a es o ees and he speed o e olu iona y
adap a ion p ocesses (Be and e al. 2011, Chen e al. 2011).
Fagus syl a ica is a na u ally dominan and economically impo an , bu d ough -suscep ible
ee species in Cen al Eu ope ha dese es special a en ion in he ace o hese changing
condi ions (Fo elli e al. 2009). E en hough F. syl a ica is conside ed a high- isk species in
he ace o clima e change (Ohlemulle e al. 2006, Sche e e al. 2011), he a ea s ocked wi h
F. syl a ica cons an ly inc eases in Cen al Eu ope/Ge many due o o es con e sion om
coni e ous o mixed s ands. Acco ding o Polley e al. (2009) beech is he species in Ge many
6 Manusc ip s
80
wi h he highes inc ease in s ocked a ea be ween 2002 and 2007 (83.000ha). D ough e en s
mos p obably de e mine he sou he n edge o he dis ibu ion ange (Jump e al. 2006).
Especially a lowe ele a ions g ow h and dis ibu ion o F. syl a ica is expec ed o decline a
i s sou he n limi s (Ma yas e al. 2009, Hlasny e al. 2011,Jezik e al. 2011) and on xe ic si es
in Cen al Eu ope (Czucz e al. 2011). The obse ed ecen decline in beech o es
p oduc i i y in F ance could be linked o low wa e a ailabili y, especially in ea ly summe
(Lebou geois e al. 2005, Cha u e al. 2010). Fu he mo e Pio esan e al. (2008) showed ha
basal a ea inc emen o Eu opean beech s ands dec eased wi h dec easing wa e a ailabili y in
he Apennine Moun ains in I aly. The excep ional 2003 d ough pe iod o e Cen al Eu ope
gene a ed dis inc nega i e e ec s on he pe o mance o F. syl a ica (Czajkowski e al. 2005,
Leuzinge e al. 2005, Be sch e al. 2011). Mo eo e i seems ha F. syl a ica loses i s
compe i i e ad an age o less d ough -sensi i e species, e.g. Que cus pe aea, unde wa e
limi ed condi ions in Sou he n and Sou h-Eas e n pa o i s dis ibu ion ange (Bonn 2000,
Fo elli e al. 2001, F ied ichs e al. 2009, Cla k e al. 2011, Scha nwebe e al. 2011), whe eas
posi i e impac s o clima e change on compe i ion may p e ail on he no he n dis ibu ion
edge in Sweden (Bol e e al. 2010). D ough pe iods also inc ease pa hogen- and ungi-
suscep ibili y o F. syl a ica, as shown by Jung (2009) o he pos -2003 yea s in a Ba a ian
beech s and. The e o e, Rennenbe g e al. (2004) and Gessle e al. (2007) conside he ocus
on F. syl a ica in he ongoing o es con e sion as e y isky.
Howe e , F.syl a ica, wi h i s wide geog aphic dis ibu ion, co e s a b oad ange o clima ic
condi ions (see Kölling 2007). The in oduc ion o F. syl a ica p o enances om wa me ,
mo e d ough -p one egions, especially om he edges o i s dis ibu ion ange o Cen al
Eu ope migh he e o e be one po en ial adap a ion ool o dampen he ad e se e ec s o
u u e clima ic en i onmen s (Bol e e al. 2009, K eyling e al. 2011). The gene ic
composi ion o o es ees de e mines he pheno ypic plas ici y and wi h his hei adap i e
capaci y o en i onmen al s esso s such as d ough (Schabe g e al. 2008, Ma yas e al.
2009). In Cen al Eu ope, F. syl a ica exhibi s a high gene ic di e si y wi hin popula ions
(Konne 1995, Vo nam e al. 2004), whe eas on a con inen al scale he gene ic di e en ia ion
be ween popula ions becomes mo e dis inc (Comps e al. 1990, Mag i e al. 2006). These
di e ences in gene ic con igu a ion be ween popula ions, mos likely, display di e ences in
adap i e capaci y be ween p o enances. In nume ous p o enance- ials o e he las decade, a
con as ing pe o mance o p o enances om di e en geog aphic o igins has been
demons a ed (e.g. on Wuehlisch e al. 1995). Fu he mo e, i was shown ha di e en
p o enances exhibi speci ic esponses o abio ic s esso s such as la e os s (K eyling e al.
6 Manusc ip s
81
2012) o d ough (Sch aml und Rennenbe g 2000, Peuke e al. 2006). In a Eu opean –wide
p o enance- ial ne wo k, mac oclima ic adap a ion could be de ec ed: The pe o mance o
di e en p o enances was nega i ely co ela ed wi h clima ic dis ance (absolu e di e ences
in ce ain clima ic pa ame e s) be ween es -si e and o igin o he p o enance (Ma yas e al.
2009). Czajkowski and Bol e (2006) ound a signi ican ly highe anspi a ion and he e o e
highe wa e s ess unde d ough condi ions o Ge man and Wes -Polish p o enances han
o p o enances om he mo e con inen al Cen al-Polish eas e n dis ibu ion edge o F.
syl a ica. In he ex ao dina y d y yea 2003, NW-G eek beech popula ions only expe ienced
mild d ough s ess compa ed o he epo ed damages in beech o es s in Cen al Eu ope,
despi e compa able clima ic condi ions (Fo elli e al. 2009), he eby indica ing a local
adap a ion o G eek popula ions o wa e sho ages. The e o e, such ma ginal popula ions
om he d y edges o he cu en dis ibu ion ange o F. syl a ica may come in o ocus in
sea ch o d ough - esis an eco ypes (Rose e al. 2009, K eyling e al. 2011) The suscep ibili y
o clima ic s esso s such as d ough , howe e , s ongly depends on he in e ac ion wi h
addi ional bio ic o abio ic ac o s (Messaoud and Chen 2011). The impac o d ough and
wa e s ess on he pe o mance o F. syl a ica depends on ai ozone concen a ion (Pollas ini
e al. 2010), ca bon dioxide concen a ions (Penuelas e al. 2008), ligh a ailabili y
(Czajkowski e al. 2005, Lo e al. 2005) and soil chemis y (Wei ne e al. 2007). E idence
sugges s ha , in gene al, d ough educes nu ien a ailabili y and up ake in o es soils
(Rennenbe g e al. 2009, K euzwiese and Gessle 2010). The in e ac i e e ec s o soil
nu ien balance and wa e a ailabili y on ine oo s, mic oo ganisms and soil nu ien cycling
p ocesses a e poo ly unde s ood (K euzwiese and Gessle 2010). Howe e , especially o F.
syl a ica which g ows on a ious soil ypes h oughou i s dis ibu ion ange (Ellenbe g and
Leuschne 2010), he occu ence o d ough condi ions o wa e s ess is a complex in e ac i e
mechanism be ween clima ic and edaphic ac o s (Gae ne e al. 2008).
Conside ing he epo ed nega i e impac s o d ough on he pe o mance o F. syl a ica and
he in aspeci ic a iabili y in d ough esponse, i is impo an o know whe he ce ain
p o enances o F. syl a ica a e less suscep ible o hese nega i e e ec s and whe he si e
ac o s such as soil ype in luence speci ic esponses. This knowledge is c ucial o assess he
po en ial o selec i e ansplan ing o d ough - esis an p o enances as a ool o dampen
nega i e impac s o such clima e ex emes in he cou se o clima e change.
He e, seedlings o six p o enances o F. syl a ica, h ee om he cen e o i s dis ibu ion
ange end h ee om he ma gins (SE, SW, NE) we e exposed o d ough in wo di e en soil

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82
ypes (wi h con as ing nu ien a ailabili y and ield capaci y) in a ull ac o ial common
ga den expe imen . We hypo hesized, ha (1) p o enances di e in he ex en o lea damage,
heigh and diame e g ow h and (2a) d ough and (2b) sandy soil subs a e nega i ely
in luence hese pe o mance indica o s. We u he assumed ha (3) ma ginal popula ions a e
less d ough sensi i e, and we expec ed (4) ha soil condi ions and d ough in e ac ions
signi ican ly a ec plan pe o mance.
Ma e ial and Me hods
Expe imen al Si e
The p o enance- ial (EVENT 3-Landau) is a subp ojec o he EVENT-Expe imen se ies in
Bay eu h, Ge many (Beie kuhnlein e al. 2011). I was es ablished in 2010 in he icini y o
he Uni e si y o Koblenz-Landau, a he Julius Kühn-Ins i u (JKI), Fede al Resea ch Cen e
o Cul i a ed Plan s, Siebeldingen (49°13’03” N, 8°02’47” E, 202m a.s.l.). The mean annual
empe a u e a he si e is 10.2°C and he mean annual p ecipi a ion is 643mm, which is
dis ibu ed bimodally wi h wo majo peaks in Mai/June and No embe /Decembe (da a:
Ge man Wea he Se ice). Annual a e age empe a u e is expec ed o inc ease by 3.7K by he
end o he cen u y (2080s compa ed o 1950-2000), and p ecipi a ion is expec ed o dec ease
by abou 50mm pe yea , wi h a majo dec ease in summe (JJA) p ecipi a ion (- 60mm),
acco ding o he A1B scena io o he MPI-ECHAM5-model (da a: wo ldclim).
Plan Ma e ial
Seeds o six p o enances o F. syl a ica we e ob ained in au umn 2009 and b ough o
ge mina ion a he Ba a ian Ins i u e o Fo es Seeding and Plan ing (ASP) in Teisendo ,
Ge many in sp ing 2010. All six p o enances s em om au och honous popula ions (Fig. 1,
Tab. 1). The p o enances we e chosen o ep esen he cen e o he dis ibu ion ange (DE1,
DE2, DE3) and he sou h-eas e n (BG) and sou h-wes e n (ES) edges, whe e d ough limi s
he dis ibu ion o he species, and he no h-eas e n edge (PL), whe e besides summe
p ecipi a ion win e condi ions and la e os a e impo an limi ing ac o s (Tab. 1, Fig. 1). In
Janua y 2011, he seedlings we e anspo ed o Siebeldingen and in Ma ch 2011, hey we e
plan ed in 12-li e po s wi h wo di e en soil ypes. Indi iduals we e selec ed andomly o
each p o enance and ea men om all li ing plan s a plan ing da e. Mean plan heigh a he
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83
s a o he expe imen was 22.1 cm (± 6.6cm SD), mean plan diame e 2cm abo e he oo
colla 4.7mm (4.4 – 5.3mm) wi h a s anda d de ia ion o ± 0.8mm SD (0.6-0.8mm).
Figu e 1: Geog aphic o igins o p o enances used in he expe imen . G ey scales display he Summe Hea
Mois u e index based on wo ldclim da a (Hijmans e al. 2005). Dis ibu ion ange o F. syl a ica is ou lined
acco ding o EUFORGEN
Code Loca ion Coun y La i ude Longi ude ele a ion
(m a.s.l.)
SHMI
BG Ko el Bulga ia N 42°51'59" E 26°26'40" 600 61
ES Mon ejo de la
Sie a
Spain N 42°01'00" W 03°05'00" 1463 68
DE1 Hengs be g Ge many N 50°08'00" E 12°11'00" 569 47
DE2 Johannisk euz Ge many N 49°18'00" E 07°50'00" 570 42
DE3 Kemp en Ge many N 47°44'48" E 10°08'54" 803 26
PL M agowo Polen N 53°52'00"
E 21°20'00" 137 51
Table 1 Si e in o ma ion o he p o enances used in he expe imen . Summe Hea Mois u e index de i ed om
Wo ldClim (Hijmans e al. 2005).
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Expe imen al se -up
The po ed indi iduals we e exposed o a ully c ossed expe imen al design consis ing o a
p ecipi a ion ea men (d ough and con ol) and wo di e en soil ypes (sandy loam, loamy
sand). Each ea men combina ion (d ough wi h sandy loam, d ough wi h loamy sand,
con ol wi h sandy loam and con ol wi h loamy sand) was eplica ed wi h 9 indi iduals. Fo
wo p o enances (Spain, Poland) he numbe o eplica es had o be educed o 8 and 7
espec i ely, due o mice damage, esul ing in 204 plan s o e all. The expe imen was se up
comple ely andomized in a conc e e bed illed wi h sand. All po s we e pu on plas ic
sauce
Figu e 2: Soil mois u e o all ou ea men combina ions. The ligh g ey a ea shows he a i icial d ough
pe iod, he da k g ey a ea shows he e-wa e ing phase. The wo ho izon al lines depic he app oxima ed
pe manen wil ing poin s o he loamy (black) and he sandy subs a e (g ey).
On Ap il 13 h he expe imen was co e ed by a ain-ou shel e cons uc ed o a s eel ame
(GlasMe all Rieme GmbH, Rahden-Sielho s , Ge many) and co e ed wi h a anspa en
polye hylene shee (0.2mm, SPR 5; He mann Meye KG, Rellingen, Ge many). The lowe
edge o he ain-ou shel e was 80 cm abo e g ound. The shel e pe mi ed he pene a ion o
nea ly 90% o he pho osyn he ically ac i e adia ion. Addi ionally, as seedlings o F.
syl a ica a e sensi i e o di ec adia ion, a shading can as (Quad a 105ME, 105g, DM-Folien
GmbH, Reu lingen, Ge many) was a ached o he inside o he polye hylene shee , esul ing
in a pene a ion o abou 55 % o he pho osyn he ically ac i e adia ion. As he shade- ole an
species Eu opean beech has shown o yield highes g ow h a es when exposed o medium
ligh le els (e. g. Schall e al. 2012) mos likely no limi a ion by ligh a ailabili y occu ed in
ou expe imen . F om he plan ing da e un il he ain-ou shel e was se up, he plan s
ecei ed he ambien p ecipi a ion and we e addi ionally wa e ed wi h g oundwa e (Ma ch
6 Manusc ip s
85
14 h, 18 h, Ap il 1s , 4 h 8 h and 11 h). A e he ain-ou -shel e was se up, he plan s we e
wa e ed wi h g oundwa e on Ap il he 18 h, 28 h and 30 h, so ha a su icien wa e supply
and oo g ow h was gua an eed. F om May 2nd on all plan s we e exposed o he con ol
p ecipi a ion egime which simula ed he local daily 40-yea a e age p ecipi a ion. The
applica ion was done wice a week wi h g oundwa e .
The d ough manipula ion consis ed o a 36-day pe iod wi hou i iga ion. I s a ed on May
9 h end was comple ed when 20% o he plan s showed s ong d ough damage (76-100% o
he lea es damaged), on June 13 h. App oxima ely 12 days a e he s a o he ea men soil
mois u e d opped below he pe manen wil ing poin (pF = 4.2; Fig.2) o he loamy sand
subs a e and abou one week la e o he sandy loam subs a e. In he e-wa e ing phase a e
he d ough pe iod he plan s we e exposed o he con ol p ecipi a ion ea men again.
Addi ionally hey ecei ed he amoun o wa e which was gi en o he con ol plan s o e he
36-day d ough pe iod, e enly dis ibu ed o e one week, so ha a he end o he expe imen
bo h he indi iduals in he con ol as well as in he d ough ea men ecei ed he same
amoun o wa e .
The olume ic soil wa e con en was measu ed wi h dielec ic pe mi i i y senso s (EC-5
soil wa e senso , Decagon De ices, Inc., Pullman, USA). Al oge he 10 senso s wi h wo
da a logge s (ECH2O, Decagon De ices, Inc. Pullman, USA) we e ins alled in he po s in 5-
10 cm dep h, h ee o he wo soil ypes each in he d ough ea men and wo each in he
con ol ea men .
The soil ea men consis ed o wo di e en soil ypes. The i s one was a sandy loam
subs a e om a local o es op-soil dug-ou , om now on called loam o loamy subs a e.
The second soil ype was a loamy sand subs a e, consis ing o a mix u e be ween he loam
and a enaceous qua z sand om a local sand pi (50% each), om now on e e ed o as sand
o sandy subs a e (Tab.2).
Table 2 Nu ien con en and pH- alues o he wo soil ypes used in he expe imen
K
mg/kg
Mg
mg/kg
P
mg/kg
NO3
mg/kg
NH4
mg/kg
Co g.
%
pH-CAT
pH-wa e
N
%
Loamy Sand 54,8 121 11,1 14,1 2,02 0,56 6,3 8,5 <
Sandy Loam 118 267 48,3 31,0 3,98 1,92 4,7 7,6 0,14
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Figu e 5: Diame e inc emen o F. syl a ica in esponse o a) p o enance, b) d ough and c) soil ype. All
signi ican main and in e ac ion e ec s (d- ) o he h ee- ac o ial ANOVA a e displayed. P o enances a e
a anged wi h inc easing SHMI.
Mo ali y
Mo ali y occu ed solely wi hin he d ough ea men , and wi hin he d ough ea men
mainly in he sandy subs a e (d ough /loam = 7.8%, d ough /sand = 25.5%; ANOVA: p =

6 Manusc ip s
93
0.012). P o enance did no impac mo ali y signi ican ly (ANOVA: p = 0.097), e en hough
mo ali y co ela ed signi ican ly wi h summe d ough condi ions a he places o o igin o
he p o enances (see Resul s, Local adap a ion). Mo ali y ended o inc ease (non –
signi ican ly) wi h diame e inc emen educ ion (adjus ed ² = 0.39, p = 0.110) in he d ough
ea men compa ed o con ol.
Co ela ion o lea damage wi h inc emen
The a e age pe cen age o damaged lea es pe p o enance was s ongly co ela ed o a
educ ion in diame e inc emen (adjus ed ² = 0.99, p < 0.001). The educ ion in heigh
inc emen exhibi ed a non-signi ican end owa ds an inc easing educ ion wi h inc easing
lea damage (adjus ed ² = 0.42, p = 0.097).
Local adap a ion
The Summe Hea Mois u e index (SHM) a he geog aphic o igin o he p o enances did no
in luence he eac ion o d ough wi h ega d o heigh inc emen (adjus ed ² = 0.01, Fig. 6a).
Howe e , e en i non-signi ican , p o enances om o igins which a e mo e p one o summe
d ough e en s (highe SHM) ended o be mo e d ough ole an in e ms o diame e
inc emen and lea damage, as SMH a he geog aphic o igins explained 25% o he a iance
in diame e inc emen educ ion (Fig. 6b), 31% o he a iance in lea damage (Fig. 6c). A
signi ican co ela ion (p = 0.041) could be de ec ed be ween he SMH index and mo ali y
(Fig. 6d). He e he summe hea mois u e index a he geog aphic o igin explains 69% o he
a iance in mo ali y.
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Figu e 6: Co ela ion be ween he summe hea mois u e index a he o igin o he p o enances and he mean
educ ion in a) heigh and b) diame e inc emen , c) mean lea damage pe p o enance and d) mo ali y pe
p o enance.
Discussion
In a-speci ic a ia ion
The cu en dis ibu ion o F. syl a ica is a esul o mul iple glacial pe iods. F. syl a ica
su i ed he las glacial maximum in se e al e uge a eas in Eu ope (Mag i e al. 2006). Mos
p obably, e uge a eas in he Sou h-Wes e n and Dina ic Alps played an impo an ole in he
e-coloniza ion o Cen al- and Wes e n Eu ope, whe eas he Medi e anean popula ions seem
o ha e su i ed in se e al geog aphically dis inc e uges on he Ibe ian and I alian
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peninsulas and in he Balkan (Mag i e al. 2006, Mag i 2008, B us 2010). In pa icula he
Balkan (and I alian) popula ions a e gene ically dis inc om he o he Eu opean popula ions
(Mag i 2008). This pos -glacial mig a ion pa e n and gene ic di e en ia ion, mos p obably,
also esul ed in pheno ypic a ia ion be ween p o enances. Dis inc pe o mances o
Eu opean beech p o enances g own in common ga den expe imen s a e well-known and
o en demons a ed (e.g. on Wuehlisch e al. 1995, Nielsen and Jo gensen 2003, Giannini
and on Wuehlisch, 2009). The di e ences be ween p o enances in all o he examined
pa ame e s in his s udy a e consis en wi h hese p e ious indings and likely exp ess he
di e ences in gene ic make-up.
Especially he Bulga ian p o enance showed a dis inc pe o mance in all pa ame e s.
Bulga ian beech popula ions a e gene ically dis inc om Cen al Eu opean popula ions
(Mag i e al. 2006), which may explain he signi ican ly lowe inc emen . Su p isingly, one
Ge man p o enance (DE2) pe o med signi ican ly wo se in e ms o heigh inc emen han
he o he wo Ge man p o enances (DE1, DE3), despi e a mos likely simila pos -glacial
his o y and assumingly compa able gene ic make-up. Ye , in Cen al Eu ope gene ic di e si y
is highe wi hin popula ions han be ween popula ions (Konne 1995). Fu he mo e, he
sil icul u al his o y o pa icula beech s ands is o en ha d o be econs uc ed. In addi ion,
local soil condi ions and small-scale ain all-pa e ns may be also esponsible (see below).
E ec s o d ough and soil on ee pe o mance
D ough ad e sely impac ed all measu ed pa ame e signi ican ly: I inc eased he amoun o
lea damage and dec eased heigh and diame e inc emen . This is in compliance wi h
nume ous s udies epo ing on he d ough sensi i i y o F. syl a ica (Rennenbe g e al. 2004,
Lebou geois e al. 2005, Jump e al. 2006, Pio esan e al. 2008, F ied ichs e al. 2009, Be sch
e al. 2011, Hlasny e al. 2011, Jezik e al. 2011, Sche e e al. 2011).
As expec ed, he sandy subs a e wi h lowe nu ien a ailabili y and wa e s o age capaci y
esul ed in mo e se e e nega i e impac s on he pe o mance o plan s, compa ed wi h loamy
subs a e. The d ough impac on lea inju y a es and diame e inc emen was mo e se e e o
plan s g own in he sandy subs a e, which shows he impo ance o he in e ac ion be ween
p ecipi a ion a es and soil p ope ies (Gae ne e al. 2008). The soil mois u e in he sandy
subs a e d opped below he app oxima ed pe manen wil ing poin one week be o e his poin
was eached in he loamy subs a e. Plan s in he sandy subs a e, he e o e expe ienced ~24
days o wa e s ess, whe eas plan s g own in he loamy subs a e only ~17 days. The educed
6 Manusc ip s
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nu ien up ake due o lacking wa e abso p ion du ing he d ough migh u he mo e ha e
weake impac s in nu ien - iche soils han in poo e soils. Apa om nu ien up ake,
e idence sugges s ha also nu ien a ailabili y o he soil is educed due o d ough
condi ions (Rennenbe g e al. 2009, K euzwiese and Gessle 2010).
In his s udy we did no in es iga e biomass alloca ion o abo e- and belowg ound
componen s o he ee seedlings. Especially a a seedling age, he abili y o Eu opean beech
o plas ically eac o en i onmen al/d ough condi ions in e ms o ealloca ion o esou ces is
qui e high, as Schall e al. (2012) demons a ed. Eu opean beech seedlings inc eased he
in es men in o belowg ound biomass as a eac ion o d ough . Acco ding o he balanced-
g ow h hypo hesis (Hun 1975, Shipley and Meziane 2002) plan s ealloca e esou ces o he
o gan ha exploi s he limi ing esou ce, in ou case he oo sys em. Such po en ial
ealloca ion e ec o a d ough ea men migh lead o o e -es ima ion o g ow h educ ion,
as i migh mask a cons an biomass p oduc ion, when jus obse ing abo e g ound-g ow h.
When ees ge olde his mo phological plas ici y dec eases and physiological acclima iza ion
p ocesses becomes mo e impo an (Schall e al. 2012). The e o e a ans e o esul s om
seedlings o ma u e ees should be backed by mo e esea ch, e.g. in long- e m p o enance
ials.
P o enance-speci ic eac ions and local adap a ion o d ough and soil
D ough -sensi i i y in e ms o diame e inc emen a ied be ween he p o enances in ou
expe imen . The Bulga ian p o enance (BG) showed a weak pe o mance unde he con ol
ea men , ye he smalles educ ion o diame e inc emen due o he d ough ea men . The
inc emen educ ion be ween d ough and con ol in o he p o enances (DE1, DE3, PL)
amoun ed o 50%. Howe e , hei absolu e inc emen alues unde d ough condi ions we e
s ill highe han o he BG p o enance. This beha iou o p o enances co ela es s ongly
wi h lea inju y a es ( ² = 0.99). The in e ac i e e ec o p o enance and d ough ea men
was no signi ican in his case, ye his is due o he ac ha lea inju y almos exclusi ely
occu ed unde he d ough ea men . Gi en he ela i ely small numbe o p o enances in
ou ial, signi ican co ela ions be ween he d ough esponse o inc emen and lea damage
wi h he clima e a he o igin o he p o enances could no be es ablished. Ye , he eac ions
o bo h pa ame e s ended owa ds a co ela ion wi h he long- e m summe hea mois u e
index a he o igin o he p o enances. This end is suppo ed by p e ious indings by
Czajkowski and Bol e (2006) who demons a e a co ela ion be ween he d ie , con inen al
6 Manusc ip s
97
clima e a he o igin o a beech p o enance om cen al Poland wi h low wa e s ess
sensi i i y and he mois e clima e o p o enances om Ge many and Wes Poland wi h
highe d ough s ess sensi i i y. I is u he in compliance wi h he esul s o Nielsen and
Jo gensen (2003) who ound a signi ican in e ac ion be ween soil wa e con en and diame e
inc emen o 14 p o enances in a common ga den expe imen in Denma k. Howe e , hese
au ho s ound a g ea e adap abili y o sou he n p o enances o changing wa e condi ions. In
ou expe imen , he Bulga ian p o enance showed a low bu s able inc emen o e he
ea men s while mos no he n p o enances (excep DE2 Johannisk euz) pe o med a simila
low le els unde d ough condi ions bu showed a much mo e posi i e esponse o a ou able
condi ions han he Bulga ian one. Simila o he Bulga ian p o enance he Spanish one is no
e y esponsi e o d ough in e ms o lea damage unde d y condi ions and showed he
lowes mo ali y a es, howe e , in con as o he Bulga ian p o enance on a highe
inc emen le el and mo e no able educ ions in inc emen . Su p isingly he pe o mance o
he Ge man p o enance om Johannisk euz (DE2) esembles he mo e he sou he n
p o enances ( han he o he wo Ge man and he Polish p o enances, which a e
geog aphically close ) wi h low lea inju y a es, low heigh g ow h and lowe d ough
induced g ow h educ ions. Nielsen and Jo gensen (2003) show in hei s udy ha no he n
p o enances display a low bu s able g ow h le el and sou he n ones a mo e plas ic esponse.
The esul s o ou s udy and he esul s o Nielsen and Jo gensen migh be supe icially
in e p e ed as inconsis en . On close inspec ion, howe e , he me e geog aphic di ision o
p o enances in o No h and Sou h migh no e lec he ac ual clima ic and si e condi ions
popula ions we e exposed o in pas and p esen a hei places o o igin. The sou he n
p o enances used by Nielsen and Jo gensen (2003) do no s em om he e y edges o he
dis ibu ion ange, excep one om Moun E na, Sicily (IT) and he places o o igin o hese
sou he n p o enances ha e ela i ely high p ecipi a ion a es, whe eas he no he n
p o enances come om egions wi h ela i ely con inen al clima e wi h low p ecipi a ion
sums (e.g. eas e n Ge many and Cen al Poland). They migh be e en mo e exposed o d y
condi ions a hei o igins. The same migh be applicable o he concep o geog aphically
ma ginal and cen al popula ions, as his concep excludes mic o- and mesoclima ic e ec s,
and small-scale di e ences in soil and si e condi ions. E en in he cen e o he dis ibu ion
ange, e y un a ou able and ‘ma ginal’ si es can be ound, whe eas on he geog aphic
ma gins a ou able g ow h condi ions can occu on small scale, e.g. a he wind-wa d side o
small moun ain anges wi h o og aphic ain all-pa e ns.

6 Manusc ip s
98
In his s udy he summe hea mois u e index o he ES p o enance is e en highe han o
BG, s ill he d ough - ole ance in e m o s able g ow h is no as high as in BG, which may be
due o edaphic easons (Gae ne e al. 2008). This could mean ha he ac ual d ough
se e i y, de i ed om p ecipi a ion and soil cha ac e is ics, migh be lowe a he o igin o he
Spanish p o enance (ES). The beha iou o he Ge man p o enance (DE2) is also simila o
he ma ginal p o enance om BG, al hough i s ems om he cen e o he dis ibu ion ange
wi h clima ic condi ions compa able o he o he Ge man p o enances. DE2 s ems om he
Pala ina e Fo es , a Ge man moun ain ange wi h lowe T iassic sands one o ma ions. The
sandy soil he e migh c ea e much d ie condi ions han wha he o he wo Ge man
p o enances expe ience a hei poin o o igin. Fu he mo e in o ma ion on slope, exposi ion,
soil dep h and subs a e a he exac loca ion o seed sampling (i.e. o single mo he ees)
migh imp o e explana o y powe in u u e analyses.
Addi ionally, hese in ica e indings migh indica e a complex pa e n o wi hin and be ween
popula ion a ia ions, de eloped om he in e play o pos -glacial his o y, local clima ic
adap a ion, local g owing condi ions and human sil icul u al p ac ices. He be e e al. (2010)
and Wo emann e al. (2011) claim ha pheno ypic a iabili y in d ough ole ance, he e
measu ed in ca i a ion esis ance, is mainly caused by pheno ypic plas ici y and no by
geno ypic di e ences be ween popula ions. Ye , ou da a poin owa ds some deg ee o
inhe i ed local adap a ion o clima e condi ions a he o igins o he p o enances e en when
conside ing ou small selec ion o p o enances. Acco dingly, ma ginal beech popula ions
which ace mo e ad e se condi ions a e unde s onge gene ic selec ion (Wo emann e al.
2011) han popula ions on a ou able si es. In ou s udy, he Bulga ian p o enance (BG)
shows he mos dis inc p o enance e ec and p o enance-speci ic eac ion. I s ems om he
e y sou h-eas e n dis ibu ion edge o F. syl a ica, whe e summe d ough is mos p obably
he ange-limi ing ac o (Jump e al. 2006). This again emphasizes he impo ance o
ma ginal beech popula ions in he sea ch o d ough esis an eco ypes (Rose e al. 2009).
A ade-o be ween su i al unde d ough condi ions and high inc emen unde a ou able
condi ions has been shown o ou b oad-lea ed species in Kenya (Kondoh e al. 2006).
Acco dingly, F. syl a ica displays he lowes mo ali y unde shade, ye he lowes g ow h
a e unde ligh compa ed o o he deciduous ees (Pe i an e al. 2007). Likewise, a
con inen al and a Medi e anean Pinus syl es is p o enance did no di e unde d y
condi ions, whe eas unde we e condi ions he con inen al p o enance showed highe oo
biomass alloca ion and seedling ec ui men a es (Rich e e al. 2012). This con i ms
6 Manusc ip s
99
p e ious indings, ha high pheno ypic plas ici y appea s bene icial in highly a iable
en i onmen s.
The d ough manipula ion in ou expe imen can be conside ed as ela i ely mild. Only ew
indi iduals died due o he ea men . P o enance did no impac he mo ali y a e
signi ican ly in gene al. Howe e , he e is a clea and signi ican co ela ion be ween summe
d ough and mo ali y, wi h lowe mo ali y a es o he sou he n ma ginal p o enances due o
d ough ea men (BG = 11.1%, ES = 0%, all p o enances 15.3%), p o iding e idence o an
adap a ion o local clima e condi ions. Gi en he abo e men ioned po en ial ade-o
mechanisms his sligh end owa ds highe mo ali y wi h s onge diame e g ow h
educ ion, could poin owa ds highe su i al a es o less plas ically esponding
p o enances. A s onge gene ic selec ion in d ough -p one eco ypes (Wo emann e al. 2011)
could lead o lowe gene ic di e si y (Kawecki 2008) and hus low pheno ypic plas ici y
(Schabe g e al. 2008, Ma yas e al. 2009), hough Kawecki (2008) also epo s ha o some
ma ginal species no gene ic depaupe a ion o ecologically ele an ai s could be de ec ed.
The e o e ma ginal popula ions could po en ially secu e ecosys em pe sis ence unde e y
ex eme clima ic e en s, ye migh inc ease he isk o lowe g ow h a es unde a ou able
condi ions.
Nagy e al.(in p ep.) ound a simila pa e n in a s udy in which Bulga ian and Ge man Beech
p o enances we e exposed o d ough condi ions, wi h low g ow h, bu s able eac ion o
d ough in some p o enances and highe g ow h a es, bu s ong educ ion o o he
p o enances. Ye , he eac ion did no depend on he o igin o he p o enance.
Fu he mo e, he Bulga ian, Spanish and he Ge man p o enance om Johannisk euz (DE2)
showed ela i ely s able esponses in heigh and diame e inc emen in he sandy subs a e
compa ed o he loamy subs a e. This migh indica e ha hese p o enances a e no able o
exploi he imp o ed wa e and nu ien condi ion in he loam. I could also indica e a local
adap a ion o ce ain soil cha ac e is ics, which was e.g. ound o Pinus nig a (Va elides e al.
2001). S ill, such a po en ial ade-o be ween s ess- ole ance and high pe o mance unde
mo e a o able condi ions is ye o be suppo ed by mo e e idence in u u e p o enance ials.
A he no h-eas e n edge o he dis ibu ion ange he SHM is no as high as a he sou he n
edge. The Polish p o enance (PL) o igina es om he eco one be ween beech domina ed
o es and bo eal o es s. Beech domina es on mo aine loamy soil, while Sco s pine domina es
on d ie and sandie soils (Bol e e al. 2007). Especially a he no h-eas e n ma gin whe e
con inen al clima e condi ions a e becoming mo e p e ailing co ela ions wi h single mac o-
6 Manusc ip s
100
clima ic ac o s ail o explain dis ibu ion ma gins o F. syl a ica (Bol e e al. 2007). The e,
win e condi ions and os a e a leas as impo an as d ough . Gi ing he clea esponse o
d ough in ou expe imen i seems ha a s ong selec ion owa ds d ough - esis an eco ypes
does no ake place.
Like men ioned abo e, below-g ound biomass p oduc ion was no de e mined in his s udy,
ye he e migh be di e ences in plas ici y o biomass alloca ion be ween he p o enances
(Rich e e al. 2012), which dese e mo e a en ion in u u e s udies.
Conclusion and implica ions o esea ch and o es managemen
P o enances o F. syl a ica di e in inc emen and exhibi di e ences in hei sensi i i y o
d ough . This in aspeci ic a iabili y can p obably be used o adap o es ecosys ems o
u u e clima e condi ions. Ye , he impac o he in e play be ween clima ic and soil and si e
cha ac e is ics on he eme gence o d ough condi ions ha e o be accoun ed o as
p o enances eac speci ically o con as ing soil ypes. The e o e he concep o ma ginali y
should be ex ended om a geog aphical o a mo e si e- ela ed concep ; ye , especially
ma ginal popula ions om he d ough -p one sou he n edges o he dis ibu ion ange a e
po en ial a ge s in he sea ch o d ough - esis an eco ypes. Howe e , a po en ial ade-o
be ween high pheno ypic plas ici y wi h high pe o mance a es unde a ou able condi ions,
and s ess- ole ance and su i al unde d ough condi ions and i s impac on o es g ow h
equi es u u e esea ch. Long- e m p o enance ials wi h a g ea e numbe o cen al and
ma ginal (maybe also om ‘ma ginal’ si es in he cen e o he dis ibu ion ange)
p o enances, whe e he pe o mance o ees can be moni o ed beyond he seedling age, could
con ibu e in app oaching hese open ques ions. Di e ing om exis ing ials, a hypo hesis-
based selec ion o p o enances and he op ion o clima e manipula ions in la e on ogene ic
s ages should be applied. Fu he mo e, a single d ough manipula ion does no e lec a
change in long- e m mean alues and changes in equency o ex eme e en s and he e long-
e m impac s. The e o e, a ne wo k o se e al clima ically di e en ial si es, including
ma ginal si es is needed.
Fu he mo e, mo e mul i- ac o ial expe imen s could add ess he p oblem ha esis ance o
one abio ic ac o such as d ough does no necessa ily imply esis ance o o he clima ic o
abio ic ac o s such as os (K eyling e al. 2012) o o es pes s. Howe e , he es ablishmen
o mul i- ac o ial, mul i-si e and long- e m ials and expe imen s will ake longe han he e
is ime o s a adap i e ac ions in o es managemen wi h ega d o he eloci y o clima e
6 Manusc ip s
101
change. E idence sugges s ha he selec ion o a single bes d ough -adap ed p o enance o
ansplan a ions is no easonable, conside ing he limi ed knowledge o he long- e m e ec s.
None heless, ma ginal and d ough -adap ed eco ypes migh be in e mixed, oge he wi h local
p o enances and eco ypes adap ed o o he abio ic ac o s, in o de o secu e yield and
s abili y unde mo e ex eme clima ic condi ions in he u u e. Fo es managemen should
he e o e aim a inc easing he gene ic di e si y o o es s ands o secu e popula ions agains
b eakdown due o clima ic ex emes.
Acknowledgemen s
This s udy was unded by he "Ba a ian Clima e P og amme 2020" in he join esea ch
cen e “FORKAST” and he Ba a ian S a e Minis y o he En i onmen and Public Heal h
(ZKL01Ab 7_18456) oge he wi h he Uni e si y o Koblenz/Landau. We hank he Julius-
Kühn –Ins i u e in Siebeldingen o he p o ision o he si e and in as uc u e o he
expe imen . Fu he mo e we hank D . Do o a Dob owolska, Fo es Resea ch Ins i u e,
Poland, o p o iding he seed ma e ial o he Polish p o enance.
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108
2010). In ex ensi ely used hay meadows o Cen al Eu ope, he selec ed species o his
expe imen —A hena he um ela ius (L.) P. Beau . ex J. P esl & C. P esl, Fes uca p a ensis
Huds. s.l., Holcus lana us L, and Alopecu us p a ensis L.—play a majo ole. These species
we e pa o he Ge man BIODEPTH expe imen (Hec o e al. 1999), which is loca ed
nea by he EVENT-Expe imen .
Indi iduals o local and egional popula ions de elop a pool o pheno ypes ha can be
assumed o be bes adap ed o hei speci ic en i onmen i ime is su icien and he
en i onmen is s able. In ac , only an incomple e spec um o pheno ypes o a axon can be
ep esen ed in a local pool, and gene ic di e si y wi hin he popula ion is limi ed by dispe sal
his o y o il e s. Only a ew s udies ha e conside ed geno ypes o pheno ypic esponses in
g ass species expe imen ally (e.g. Fe che & Sha e 1990; Ryse & Aeschlimann 1999). In
ac , he ole o wi hin-species gene ic and unc ional di e si y o he esponse o a species o
clima e change has been widely neglec ed in ecen esea ch. Especially o widesp ead
species, spa ial and gene ic dis ances be ween popula ions ha e o be aken in o accoun .
The objec i e o his s udy was o explo e mechanisms c i ical o unde s anding he
p ese a ion and adap a ion o ecosys ems in he ace o clima e change. We wan ed o
iden i y whe he eco ypes ( ep esen ed in local popula ions) o ce ain key species om
di e en loca ions (p o enances) also di e in hei esponse o ex eme clima ic condi ions.
I his we e ound, species popula ions o eco ypes could be sough ha a e be e adap ed o
he expec ed clima e o he u u e han egional popula ions. He e, we ocus on plan ai s
ela ed o he key ecosys em unc ion p oduc i i y: biomass and nec o ic issue.
Ma e ial and Me hods
Expe imen al Si e
This common-ga den expe imen (EVENT 3) is pa o he EVENT-expe imen s (Jen sch,
K eyling & Beie kuhnlein 2007; Jen sch & Beie kuhnlein 2010) and was es ablished in
Ma ch 2009. I is loca ed in Bay eu h, Ge many, on he p ope y o he Ecological–Bo anical
Ga dens o he Uni e si y o Bay eu h in di ec p oximi y o EVENT 1 and EVENT 2
(49°55’19” N, 11°34’55” E). The long- e m mean annual empe a u e o he si e is 8.2 °C,
whe eas he long- e m mean annual p ecipi a ion is 724 mm. P ecipi a ion is dis ibu ed bi-
modally wi h a majo peak in June/July and a second peak in Decembe /Janua y (da a:
Ge man Wea he Se ice).

6 Manusc ip s
109
Unde he A1B scena io (IPCC 2007) egional clima e models (REMO, B G 2009) ha e
p ojec ed he ollowing clima ic condi ions o he si e. Annual a e age empe a u e (30-yea
mean) is expec ed o inc ease by 2.4 K by he middle o he cen u y (2041/2070 compa ed o
1971/2000) and by 3.5 K by he end o he cen u y (2071/2100 compa ed o 1971/2000).
P ecipi a ion is no expec ed o change conside ably acco ding o he annual mean alues (+ 2
mm yea
-1
2071/2100), bu summe s a e expec ed o become d ye (-17 mm) o e he same
pe iod.
Selec ion o Species and Eco ypes
The species ha we e selec ed o his s udy a e common in Cen al Eu opean managed
g asslands. A hena he um ela ius is a allg ass ha con ibu es subs an ially o biomass
p oduc ion in meadows. I is a widesp ead and common species in Eu ope, wi h a high
abundance in pe manen empe a e g assland. I is ound on mois o mode a ely d y nu ien -
ich soils (Obe do e 2001). Due o i s impo ance as a o age plan , his pheno ypically
a iable g ass was cul i a ed and plan ed ou side o i s na u al ange. Fes uca p a ensis, wi h
i s wide dis ibu ion ac oss Eu ope, is a high-quali y o age plan ha is also sui able o
pas u es. I g ows p edominan ly on esh soils ich in nu ien s and humus and has been
p omo ed h ough cul i a ion (Obe do e 2001). Holcus lana us occu s in all Eu opean
coun ies, p edominan ly on we and boggy soils (Obe do e 2001), whe e i is o a ce ain
impo ance as a o age plan ; i is less common on pas u es compa ed o meadows.
Alopecu us p a ensis is a common g ass ound h oughou mos o Eu ope in mois and
nu ien - ich soils (al hough also less common on pas u es) ha p oduces abundan and high-
quali y o age (Obe do e 2001). Fo his eason, i is also plan ed and cul i a ed ou side o
i s na u al ange. As pe ennial clonal g asses, all ou species sha e compa able li e cycles and
s a egies o esou ce alloca ion.
6 Manusc ip s
110
Figu e 1: Selec ed eco ypes o (3a) A hena he um ela ius, (3b) Fes uca p a ensis, (3c) Holcus
lana us and (3d) Alopecu us p a ensis. The shaded a ea indica es he species dis ibu ion. Black a eas
ma k he egions wi h cu en clima ic condi ions simila o he p ojec ed u u e clima e o he Ge man
loca ion. The local eco ype is displayed as a ge (DE Ge many). The illed ci cle shows loca ion o
he eco ype om no he n e e ence popula ions (SE Sweden). The emp y ci cles indica e sou he n
eco ypes om egions ha a e clima ic analogues o u u e expec a ions o he local clima e (ES
Spain, IT I aly, HU Hunga y, BG Bulga ia).
Besides local eco ypes o hese ou species om Ge many (DE), we selec ed o he Eu opean
eco ypes o hese g asses on he basis ha he clima e o he egion o o igin was simila o
he local p ojec ions o ou si e in no he n Ba a ia in he u u e. Local p ojec ions o mean
annual empe a u e, mean win e empe a u e and annual p ecipi a ion o he pe iod 2071-
2100 we e aken om he egional clima e model REMO (B G 2009) based on he A1B
Scena io (IPCC 2007). Regions wi h cu en clima ic condi ions simila o he u u e
p ojec ions o he a ge a ea we e loca ed based on wo ldclim da a (Hijmans e al., 2005).
Seed ma e ial was ob ained om hese a ge egions (Fig. 1), abb e ia ed below as Spain
(ES), I aly (IT), Hunga y (HU) and Bulga ia (BG). Fu he mo e, in o de o es whe he
no he n popula ions di e , we added a egion om he no he n pa o he species’ anges
(in ou case Sweden, SE). Fo A. ela ius and F. p a ensis eco ypes we e a ailable om seed
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111
banks o all a ge egions. Fo H. lana us and A. p a ensis only ou eco ypes we e a ailable
(Table 1).
The Cul i a ion o he Plan s
The a ge eco ypes we e cul i a ed a he b anch o ice o he Leibniz Ins i u e o Plan
Gene ics and C op Plan Resea ch (IPK) in Poel, Ge many, om Feb ua y 2009 o Ap il
2009. The seedlings we e hen anspo ed o Bay eu h and indi idually plan ed in o 4-L
Loca ion
Accession
La i ude
Longi ude
Ele a ion
(m a.s.l.)
T
(°C)
P
(mm)
A . e l a i u s DE Blau elden,
Hohenlohe Ebene
GR 12752 49°17'46"N 09°58'25"E 460 8.3 732
SE Uppsala PI 235543 59°51'25"N 17°38'22"E 20 5.7 551
ES Mon alban PI 234465 40°50'07"N 00°47'55"E 900 11.0 450
IT Rizzolo, Emilia-
Romagna
GR 12733 44°55'12"N 09°44'46"E 110 12.9 739
HU Lókú RCAT064783 47°12'00"N 17°52'00"E 440 8.9 621
BG NA BGR: 2008-ARR-
EL-1
42°00'00"N 24°50'00"E NA NA NA
F. p a ensis DE Blau elden,
Hohenlohe Ebene
GR 12753 49°17'46"N 09°58'25"E 460 8.3 732
SE Må ens o pe ,
G ängesbe g
No dGen: 1191 60°00'00"N 15°00'00"E 350 4.3 738
ES San An on B 1592 42°37'08"N 00°09'47"W 1250 6.8 989
IT Lago Ne o, Fe ie e
Piacenza
B 1078 44°33'00"N 09°27'00"E 1600 8.5 981
HU Vé eskozma (Gán ) RCAT040707 47°27'00"N 18°28'00"E 270 10.2 571
BG Dolna Banya GR 6976/99 42°19'05"N 23°45'07"E 710 9.6 585
H . l a n a us DE Blau elden,
Hohenlohe Ebene
GR 12750 49°17'46"N 09°58'25"E 460 8.3 732
IT To ano, Emilia-
Romagna
GRA 312 44°53'33"N 09°41'20"E 160 12.6 758
HU Ho á he elend RCAT040972 46°10'00"N 17°55'00"E 200 10.9 675
BG Mihil si GR 6632/00 42°31'19"N 24°48'52"E 330 10.9 581
A. p a ensis DE Blau elden,
Hohenlohe Ebene
GR 12751 49°17'46"N 09°58'25"E 460 8.3 732
SE Må ens o pe ,
G ängesbe g
No dGen: 1183 60°00'00"N 15°00'00"E 350 4.3 738
HU Lókú RCAT064581 47°12'00"N 17°52'00"E 440 8.9 621
BG Iska Dam GR 6635/00 42°26'29"N 23°35'20"E 810 9.0 593
Table 2: Si e in o ma ion (DE Ge many, SE Sweden, ES Spain, IT I aly, HU Hunga y, BG Bulga ia) o
he accessions o eco ypes (p o enances) o A hena e um ela ius, Fes uca p a ensis, Holcus lana us and
Alopecu us p a ensis used in he EVENT 3 expe imen . Tempe a u e (T) and p ecipi a ion (P) ep esen
long- e m mean annual alues (sou ce: WORLDCLIM). Accession esponds o he code o he seed bank
a IPK Poel (codes beginning wi h GR om IPK seed bank, codes wi h PI om USDA-ARS-GRIN, codes
wi h No dGen, B and RCAT om Eu isco)
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112
plas ic po s illed wi h a local o es - opsoil. The soil subs a e was sandy sil (pH 7.27, o al C
1.89%, o al N 0.15%, plan -a ailable NH4
+
1.79 mg L
-1
, plan -a ailable NO3
-
22.50 mg L
-1
).
Fo he i s wo weeks a e plan ing, he seedlings we e wa e ed gene ously wi h ap wa e o
ensu e g ow h. All indi iduals we e hen cu o a heigh o 7 cm in o de o c ea e compa able
s a ing condi ions and exposed o ambien p ecipi a ion un il he s a o he expe imen al
ea men s on 25 May 2009. The expe imen ended in Sep embe 2009.
Expe imen al Design
The po ed indi iduals o he selec ed eco ypes we e plan ed in po s and exposed o
empe a u e ea men s (wa ming and con ol) and o p ecipi a ion ea men s (ex eme
summe d ough and con ol) in a spli -plo design. The wo clima e ea men s we e ully
c ossed, esul ing in ou clima e manipula ions (con ol, d ough , wa ming, wa ming &
d ough ), which we e eplica ed h ee imes, esul ing in 12 expe imen al uni s in o al. The
eco ype ea men was nes ed wi hin each expe imen al uni . Each eco ype was u he
eplica ed wi h se en plan s pe expe imen al uni (nes ed eplica es). The a ailable plan s
we e assigned andomly o he 12 expe imen al uni s o each species. Each expe imen al uni
was co e ed by a single ain-ou shel e cons uc ed o a s eel ame (GlasMe all Rieme
GmbH, Rahden- Sielho s , Ge many) and co e ed wi h a anspa en polye hylene shee (0.2
mm, SPR5, He mann Meye KG, Rellingen, Ge many). The edge o he ain-ou shel e s was
a a heigh o 80 cm and pe mi ed nea ly 90% pene a ion o pho osyn he ically ac i e
adia ion.
The con ol p ecipi a ion egime simula ed he local daily 30-yea a e age p ecipi a ion. The
applica ion was done wice a week wi h collec ed ain wa e .
The ex eme d ough ea men consis ed o a pe iod wi hou p ecipi a ion. The de ini ion o
du a ion is based he e on he species-speci ic esponse o o ganisms: A ou -s age key (0 o 3,
whe e 0 s ands o ‘comple ely undamaged’ and 3 s ands o ‘ o ally d ied ou and b i le’)
was de eloped o desc ibe he amoun o isible damage caused by he d ough . By he ime
wo- hi ds o he indi iduals o one species had eached s ages 2 o 3, o by he ime one hi d
o he indi iduals had eached s age 3, he d ough was s opped. The d ough ea men las ed
16 days o H. lana us, 18 days o A. p a ensis and F. p a ensis and 19 days o A. ela ius.
The d ough ea men esul ed in a d opping o soil mois u e below he pe manen wil ing
poin o he soil app oxima ely one week a e he s a o he ea men (Fig. 2).
6 Manusc ip s
113
Figu e 2: Ai empe a u e a plan heigh and soil mois u e (-2 o -7 cm) du ing he expe imen . Ai
empe a u e was measu ed in 10-min in e als a wo loca ions wi hin each expe imen al uni by
shel e ed he mis o es (B57863-S302-F40, EPCOS) connec ed o a da alogge (dl2, Del a). Soil
mois u e was measu ed hou ly by FD-senso s (Echo.EC-5/k, Decagon De ices, Pullmann (WA),
USA) a one andomly assigned po o each species wi hin each expe imen al uni (n = 12 pe
ea men ). Mean alues o e all species a e shown as no de ec able di e ence be ween species
occu ed. The ho izon al do ed line ep esen s he app oxima e pe manen wil ing poin (pF = 4.2).
The wa ming ea men was pe o med con inuously h oughou he whole expe imen . This
was done passi ely ia wind-shel e s and black loo -co e s, which inc eased he a e age
empe a u e by 1.5 K compa ed o he empe a u e con ol ea men and by 2.5 K compa ed
o he ambien empe a u e ou side o he expe imen al uni s (Fig. 2).
The ou h ea men was a combina ion o ex eme d ough and wa ming. The addi ional
wa ming inc eased he d ough ea men e ec by addi ionally educing he soil mois u e by
abou 1.5% on a e age (Fig. 2). In he e-wa e ing phase each indi idual in he d ough and
combined ea men s ecei ed 350 mL on h ee consecu i e days (1050 mL in o al), which
co esponds o 38 mm o p ecipi a ion. This e-wa e ing esul ed in a s eep inc ease o soil
mois u e (up o 28%). A e wa ds, he po s we e wa e ed acco ding o he con ol
p ecipi a ion ea men . Soil mois u e a e e-wa e ing emained highe in he d ough -
manipula ed po s han in he con ol po s o nea ly one mon h. Soil mois u e ell epea edly
below he pe manen wil ing poin o sho pe iods in Augus due o unusually high ambien
empe a u es.

6 Manusc ip s
114
Biomass
Based on local ag icul u al managemen ou ines o ex ensi e g asslands, biomass was
ha es ed wice o e he g owing season. The i s biomass ha es ook place en days a e
he d ough ea men ended, espec i ely, o each species, in o de o accoun o he
eco e y capaci y o he plan s. Each indi idual plan was cu a 3 cm abo e he soil and he
biomass was d ied o 48 h a 70 °C and weighed. Fo h ee ou o he se en nes ed eplica es
pe expe imen al uni , he biomass was di ided in o li ing and nec o ic ma e ial and hen
p ocessed as men ioned.
The second ha es was conduc ed 72 days a e he i s ha es on 3 Sep embe 2009 wi h
he same p ocedu e, excep ha he ha es ed ma e ial was no spli in o li ing and nec o ic
biomass.
S a is ics
Linea mixed-e ec models we e applied o each species sepa a ely o es o he main and
in e ac i e e ec s o he h ee ac o s: eco ype, empe a u e ea men and p ecipi a ion
ea men . The spli -plo design and he nes ed eplica es we e accoun ed o by he use o he
expe imen al uni iden i y as a andom ac o (Pinhei o & Ba es 2004). Da a we e log-
ans o med o imp o e he homogenei y o a iances and he no mali y o esiduals p io o
analysis, i necessa y (Fa away 2006). In case o signi ican eco ype o in e ac ion e ec s,
pos hoc compa isons (Tukey’s es ) we e un acco ding o Ho ho n, B e z & Wes all (2008).
All s a is ical analyses we e conduc ed wi h he so wa e R 2.11.1 and he addi ional packages
‘nlme’, ‘mul comp’ and ‘sciplo ’.
Resul s
Biomass
Biomass p oduc ion di e ed signi ican ly be ween he eco ypes o all ou species and bo h
ha es s (Table 2). No single eco ype pe o med supe io o he o he s when iewed o e all
ou species, and he local eco ype (DE) was signi ican ly ou pe o med by a sou he n eco ype
in only one ou o eigh cases (H. lana us a he i s ha es ; Fig. 3a). In e es ingly, a iabili y
in biomass p oduc ion wi hin species was compa able o a iabili y be ween species (Table
6 Manusc ip s
115
3). This pa e n was independen om d ough and wa ming manipula ions as i did no di e
when da a we e a e aged o e he whole da a se o analysed pe single ea men s. A he
i s ha es , A. ela ius showed highes a iabili y be ween eco ypes, which e en exceeded
a iabili y be ween species. The same was ue o H. lana us o he second ha es .
The d ough ea men esul ed in a signi ican educ ion o biomass p oduc ion o all ou
species in he i s ha es , while only F. p a ensis and H. lana us we e s ill nega i ely
impac ed by he d ough in he second ha es (Table 2). Wa ming did no al e biomass
p oduc ion signi ican ly excep o a sligh inc ease in biomass p oduc ion o F. p a ensis in
he i s ha es (Table 2).
In he i s ha es , eco ypes only di e ed signi ican ly in hei d ough ole ance o A.
ela ius (in e ac ion be ween eco ype and d ough : p = 0.005). He e, he Ge man, I alian,
Hunga ian and Bulga ian eco ypes showed he bes pe o mance wi h simila educ ions in
biomass p oduc ion due o d ough , while he Swedish eco ype exhibi ed highe d ough
sensi i i y and he Spanish eco ype gene ally p oduced less biomass (Fig. 3).
Di e ences in sensi i i y o d ough became mo e appa en o e ime, wi h A. ela ius, H.
lana us and A. p a ensis showing signi ican in e ac ion e ec s be ween eco ype and d ough
a he second ha es (Table 2). Fo A. ela ius, he eco ypes wi h a be e pe o mance om
he i s ha es ended o inc ease biomass p oduc ion in he d ough manipula ion compa ed
o he con ol (DE, IT, HU, BG), while he o he wo eco ypes (ES and SE) showed no simila
end o compensa o y g ow h (Fig. 3).
The I alian eco ype o H. lana us pe o med bes a he i s ha es (Fig. 3a) and— oge he
wi h he Ge man eco ype—also a he second ha es (Fig. 3b). No in e ac ion was ound
be ween eco ype and he clima e ea men s o he i s ha es , while he eac ion o d ough
di e ed be ween eco ypes a he second ha es (Table 2). He e, biomass p oduc ion by he
I alian and he Ge man eco ypes was no signi ican ly a ec ed when compa ing con ol and
d ough , while a educ ion in biomass due o he d ough ea men was e iden in he
Hunga ian and Bulga ian eco ypes (Fig. 3).
Alopecu us p a ensis exhibi ed he hypo hesized pa e n o be e pe o mance among he
sou he n eco ypes (HU & BG) and wo se pe o mance han he no he n eco ype (SE), wi h
he Ge man eco ype in e media e and no signi ican ly di e en om any o he h ee o he s
o he i s ha es (Fig. 3a). The Bulga ian eco ype, howe e , ell behind in e ms o biomass
compa ed o he Hunga ian and he Ge man one, while he Swedish eco ype no longe
di e ed signi ican ly om he o he s a he second ha es . I was only o he second ha es
6 Manusc ip s
116
ha a signi ican in e ac ion be ween eco ype and d ough was e iden (Table 2) wi h he
Ge man eco ype being mos p oduc i e in bo h he d ough and he con ol ea men s, while
he Bulga ian eco ype p oduced less biomass in he con ol ea men and he Swedish eco ype
p oduced less biomass in he d ough ea men .
Figu e 3:
E ec s o clima e
ea men s and eco ypes (DE
Ge many, SE Sweden, ES Spain,
IT I aly, HU Hunga y, BG
Bulga ia) on biomass p oduc ion
in g pe indi idual o he ou
g ass species. Lowe -case le e s
below he boxplo s show
homogeneous g oups acco ding o
pos -hoc compa isons in case o
signi ican main e ec s o he
ac o eco ype (see Table 2 o
ANOVA esul s). Lowe -case
le e s abo e he boxplo s indica e
homogeneous g oups acco ding o
pos -hoc compa isons in case o
signi ican in e ac ion e ec s
be ween eco ype and he speci ied
clima e ea men . The uppe and
lowe edges o he boxes ep esen
he 25%- and he 75%-quan ile,
he black line wi hin he boxes he
median (50%-quan ile) and he
uppe and lowe whiske s he
whole a iance in biomass
p oduc ion. 3a) Fi s ha es
(June). 3b) Second ha es (ea ly
Sep embe ).
6 Manusc ip s
117
Table 3:
ANOVA esul s o he applied mixed models. Analyses we e un o each species and esponse pa ame e sepa a ely. Displayed a e he deg ees o eedom
(d. .), F- and p-s a is ics. See Ma e ials and me hods o de ails on model speci ica ions.
A hena e um ela ius Fes uca p a ensis Holcus lana us Alopecu us p a ensis
num
d. .
Den
d. . F p
num
d. . Den d. .
F p
num
d. .
Den
d. . F p
num
d. .
Den
d. . F p
1s ha es eco ype 5
470
25.5
< 0.001 5
472
13.1
< 0.001 3
312
44.5
< 0.001 3
311
12.1
< 0.001
d ough 1
8
187.0
< 0.001 1
8
83.8
< 0.001 1
8
55.9
< 0.001 1
8
8.8
0.018
wa ming 1
8
0.6
0.466
1
8
7.9
0.023
1
8
0.1
0.709
1
8
0.0
0.981
eco ype×d ough 5
470
3.4
0.005
5
472
1.9
0.100
3
312
0.4
0.724
3
311
0.9
0.448
Eco ype×wa ming 5
470
1.6
0.169
5
472
0.7
0.635
3
312
1.9
0.125
3
311
1.2
0.318
d ough ×wa ming 1
8
1.0
0.342
1
8
1.4
0.266
1
8
0.4
0.570
1
8
0.1
0.739
eco ype×d ough ×wa ming 5
470
0.6
0.707
5
472
0.3
0.903
3
312
1.0
0.412
3
311
1.2
0.301
2nd ha es eco ype 5
470
8.3
< 0.001 5
472
15.3
< 0.001 3
312
20.6
< 0.001 3
311
5.9
< 0.001
d ough 1
8
0.9
0.366
1
8
7.4
0.027
1
8
20.0
0.002
1
8
0.0
0.953
wa ming 1
8
0.3
0.600
1
8
1.1
0.317
1
8
0.0
0.889
1
8
0.3
0.623
eco ype×d ough 5
470
2.3
0.046
5
472
1.3
0.258
3
312
3.6
0.014
3
311
4.4
0.005
eco ype×wa ming 5
470
0.9
0.508
5
472
2.9
0.015
3
312
1.2
0.313
3
311
0.6
0.632
d ough ×wa ming 1
8
0.1
0.793
1
8
0.9
0.375
1
8
0.5
0.488
1
8
0.5
0.501
eco ype×d ough ×wa ming 5
470
1.2
0.330
5
472
0.4
0.852
3
312
0.2
0.873
3
311
0.9
0.433
nec o ic
issue eco ype 5
183
13.5
< 0.001 5
183
4.1
0.002
3
120
11.8
< 0.001 3
120
0.1
0.969
d ough 1
8
172.1
< 0.001 1
8
40.4
< 0.001 1
8
160.4
< 0.001 1
8
74.7
< 0.001
wa ming 1
8
0.1
0.717
1
8
2.4
0.164
1
8
0.4
0.547
1
8
0.2
0.665
eco ype×d ough 5
183
2.1
0.064
5
183
12.8
< 0.001 3
120
1.1
0.358
3
120
4.5
0.005
eco ype×wa ming 5
183
1.8
0.117
5
183
0.9
0.481
3
120
0.5
0.708
3
120
0.3
0.841
d ough ×wa ming 1
8
1.8
0.216
1
8
0.8
0.386
1
8
0.6
0.457
1
8
0.7
0.423
eco ype×d ough ×wa ming 5
183
0.1
0.996
5
183
0.8
0.585
3
120
0.5
0.663
3
120
0.9
0.447
6 Manusc ip s
124
ime. This could hen delay he immig a ion and es ablishmen o po en ially be e -adap ed
pheno ypes o he same species. Ou esul s on selec ed eco ypes o key g ass species in
managed Eu opean g asslands imply ha he deba e o and agains ‘assis ed mig a ion’ has o
be ex ended o he ansloca ion o pheno ypes. Ob iously, we need mo e la ge-scale
p o enance and eco ype ials o he common species o pe manen g asslands.
Acknowledgemen s
We hank Reinhold S ahlmann o ca og aphic wo k, and all membe s o he EVENT
expe imen s a he Uni e si y o Bay eu h o suppo . D . Hugh A.L. Hen y made help ul
commen s o he manusc ip . We a e g a e ul o Sa ah Gwillym, who suppo ed us in
language issues. The esea ch was unded wi hin he FORKAST p ojec by he Ba a ian S a e
Minis y o Sciences, Resea ch and he A s.

6 Manusc ip s
125
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6.5. Manusk ip 5:
Cold ha diness o Pinus nig a A nold as in luenced by geog aphic o igin,
wa ming, and ex eme summe d ough
En i onmen al and Expe imen al Bo any 78, 99-108 (2012)
Jue gen K eyling
1
, Guido L.B. Wiesenbe g
2
, Daniel Thiel
1
, Ch is ian Wohl a
1
, Ge ha d
Hube
3
, Julia Wal e
4
, Anke Jen sch
4
, Monika Konne
3
, Ca l Beie kuhnlein
1
1
Biogeog aphy, Uni e si y o Bay eu h, D-95440 Bay eu h, Ge many
² Ag oecosys em Resea ch, Uni e si y o Bay eu h, D-95440 Bay eu h, Ge many
3
Ba a ian Ins i u e o Fo es Seeding and Plan ing (ASP), D-83317 Teisendo , Ge many
4
Dis u bance Ecology, Uni e si y o Bay eu h, D-95440 Bay eu h, Ge many
Keywo ds
os ha diness, black pine, eco ype, cold ole ance, global wa ming, win e ecology
Abs ac
Adap a ion o he ad e se e ec s o clima e change is being in es iga ed mo e and mo e
h ough he in oduc ion o species om wa me and d ie clima es, such as he (sub-)
medi e anean Pinus nig a o d y si es in empe a e Cen al Eu ope. Win e su i al,
howe e , may pose a se ious h ea o his s a egy as cold ex emes, which na u ally
de e mine he polewa d ange limi s o o es ees, a e no expec ed o ollow he gene al
wa ming end in he nea u u e.
He e, ju eniles o P. nig a om eigh p o enances h oughou Eu ope we e exposed o
di e en clima e change scena ios ( ac o ial combina ions o 42 days o d ough and wa ming
by 1.6°C) in a common ga den expe imen in Bay eu h, Ge many. Cold ha diness (LT50) was
de e mined by he Rela i e Elec oly e Leakage me hod (REL) in wo consecu i e win e s.
Cold ha diness o oliage di e ed by 10°C be ween he p o enances s udied and a local
adap a ion o minimum empe a u e was ound. Cold ha diness was u he a ec ed by
ex eme summe d ough , inc easing cold ha diness by 3.9°C on a e age in he subsequen
win e , and by summe wa ming, inc easing cold ha diness by 3.4°C. Yea - ound wa ming
had no signi ican e ec on cold ha diness. Cold ha diness was ela ed o he con en o
soluble ca bohyd a es and o he composi ion o a y acids and alkanes in he needles.
6 Manusc ip s
129
Ju eniles o P. nig a exhibi ed a compa able cold ha diness as ju eniles o species na i e o
Cen al Eu ope (P. syl es is, Picea abies, Fagus syl a ica and Que cus pe aea) unde he
same clima ic condi ions. Cold ha diness o he ine oo s o P. nig a a e aged -16.5°C
compa ed o -23.8°C on a e age o needles.
Ou esul s imply ha he cold ha diness o he oliage is adap i e o bo h long- e m g owing
condi ions a he seed o igin (gene ic he i age) and sho - e m al e a ions o hese condi ions
(indi idual plas ici y), while i s hin s sugges ha cold ha diness o he oo s is high and
migh no be adap i e. Fo P. nig a, below- and abo e-g ound cold ha diness o selec ed
p o enances in mid-win e appea sui able o cul i a ion in empe a e egions.
In oduc ion
Species espond o clima e change by polewa d ange shi s (Pa mesan and Yohe, 2003). The
speed o wa ming, howe e , is expec ed o exceed na u al mig a ion a es in many cases
(Thomas e al., 2004). In o es y in pa icula , human-assis ed ange shi s a e p oposed o
coun e long gene a ion cycles and modes dispe sal abili ies o o es ees (Schabe g e al.,
2008b; McKenney e al., 2009). Ye , he impo ance o win e condi ions is o en o e looked,
especially in he ecology o empe a e egions (K eyling, 2010). Absolu e minimum
empe a u es ha e s ong implica ions o species dis ibu ions by o en de e mining hei
polewa d ange limi s (Sakai and Weise , 1973; Repo e al., 2008). A single cold ex eme can
o se any dis ibu ional adap a ions o he gene al wa ming end (Jalili e al., 2010) and in
spi e o he mean wa ming and hei dec eased equency o occu ence, bo h he in ensi y and
he du a ion o such cold ex emes may e en inc ease egionally wi hin his cen u y due o
a mosphe ic ci cula ion changes and in e nal a mosphe ic a iabili y which coun e ac he
wa ming end om g eenhouse o cing (Va us e al., 2006; Kod a e al., 2011).
Pheno ypic plas ici y and he adap i e po en ial o o es ees a e de e mined by hei high
gene ic di e si y, allowing o es ees o de elop local adap a ions o en i onmen al s esso s
(Hosius e al., 2006; Schabe g e al., 2008b). The cold ha diness o Pinus de oniana, o
ins ance, inc eases wi h inc easing os isk along an al i udinal g adien (Saenz-Rome o and
Tapia-Oli a es, 2008). Simila ly, changes o he cold ha diness o Fagus syl a ica indica e
local adap a ion o he p e ailing minimum win e empe a u es (Visnjic and Doh enbusch,
2004) and o la e sp ing os isk (K eyling e al., 2011b) ac oss Eu ope, and he os
ole ance o Tsuga he e ophylla is adap ed o os isk along la i udinal and al i udinal
g adien s in No h Ame ica (Kuse and Ching, 1980). P o enance ials demons a e a
di e en ial pe o mance be ween he p o enances o di e en geog aphic o igins o Pinus

6 Manusc ip s
130
nig a (Va elides e al., 2001), which is he a ge species o his s udy. P. nig a was selec ed
because i is discussed in o es y as a ge species o ansloca ions o Cen al Eu ope
(Kölling, 2007; Hube 2011) and because o i s high gene ic di e si y (Nkongolo e al., 2002;
Jagielska e al., 2007). Based on i s agmen ed submedi e anean ange, one could assume
ha i lacks adap a ion o win e os , a leas in some p o enances. P o enance ials sugges
ha os damage occu s a ound -20°C and in pa icula hose p o enances om Co sica do
no su i e -25°C (summa ized in Hube , 2011).
The cold ha diness o e e g een ee species luc ua es o e he cou se o he yea . Du ing
acclima ion in au umn, he plan o gans become inc easingly ole an o he damaging e ec s
o issue eezing, pa icula ly p o ec ing cellula memb anes which a e a p ime place o
eezing inju y (Big as e al., 2001). E en hough he gene ic con ols o he p o ec i e
p ocesses in coni e s a e complex and no ye su icien ly unde s ood (Holliday e al., 2008),
da a has been summa ized on he chemical componen s ha a e in ol ed (Thomashow, 1999).
Du ing acclima ion, lipid composi ion in he plasma memb ane shi s owa ds mo e
unsa u a ed lipids (Bakh e al., 2006) in addi ion o accumula ion o soluble ca bohyd a es,
hyd ophilic polypep ides, an ioxidan s and chape ones in he memb anes (Thomashow, 1999).
Inc eased concen a ions o all hese chemical componen s se e he gene al pu pose o
p e en ing in a-cellula ice c ys alliza ion (Big as e al., 2001).
Plan s g own unde gene ally wa me condi ions, howe e , may lose hei unc ional
adap a ions o os (Eccel e al., 2009). Plan s can u he cope wi h di e en en i onmen al
s esso s by simila esponses a he cellula and molecula le el when hese s esso s igge
simila signal chains. D ough and os , o ins ance, lead o simila physiological esponses
in a coni e ous o es ee - aiming o p e en cellula dehyd a ion (Blodne e al., 2005).
Mo e equen d ough e en s may he e o e make up o diminished acclima ion due o
wa ming.
The (sub-) medi e anean dis ibu ion o ou a ge species P. nig a is e lec ed in high
d ough ole ance (Isaje e al., 2004) ela i e o empe a e species such as Pinus syl es is o
Fagus syl a ica. The e o e, ansloca ion o P. nig a is discussed as one adap a ion s a egy
agains he ad e se e ec s o clima e change a d y si es in Cen al Eu ope (Kölling, 2007).
The minimum empe a u e in win e , howe e , is one o he mos impo an ac o s se ing he
no he n bounda ies o he na u al anges o o es ee species (Sakai and Weise , 1973;
Koe ne and Paulsen, 2004). The cold ha diness o one single p o enance o P. nig a was
lowes among eigh Pinus species (wi h P. nig a showing he sou he nmos na i e ange o
he es ed species) in a common ga den expe imen in T ondheim, No way (S imbeck e al.,
6 Manusc ip s
131
2007). As ee species a e gene ally well adap ed o he minimum empe a u es o hei
en i onmen (Sakai and Weise , 1973), he ange o os ole ance o P. nig a ac oss
p o enances needs o be examined in de ail be o e ansloca ions o o he clima es a e
unde aken. This holds pa icula ly ue because clima e modelling implies ha cold ex emes
will emain s able in hei magni ude h oughou his cen u y in spi e o clima e wa ming
(Va us e al., 2006; Kod a e al., 2011). Wi h ega d o he li e span o ees, he expec ed
dec ease in equency o cold spells (e.g. Va us e al., 2006; Kod a e al., 2011) is clea ly
less impo an han magni ude and du a ion o indi idual cold spells, as e en wi h dec eased
equency he likelihood o expe iencing a leas one cold spell is s ill close o 100%.
Fu he mo e, o es s g ow slowly and managemen ac ion aiming a s able and p oduc i e
o es s in u u e need o be s a ed now. Ta ge species o ansloca ions need he e o e no
only be adap ed o u u e condi ions, bu also su i e cu en condi ions wi h p e ailing
occu ences o cold ex emes.
Plan o gans di e in hei cold ha diness. Gene ally, oo s a e he leas os ole an
(Mancuso, 2000; Big as e al., 2001). The on-going decline o Chamaecypa is noo ka ensis in
he Paci ic No hwes o No h Ame ica, o ins ance, has been linked o oo os damage
due o clima e change-induced educ ions in he insula ing snow co e (Schabe g e al.,
2008a). A simila educ ion in snow co e is also p ojec ed o Cen al Eu ope (K eyling and
Hen y, 2011). In addi ion o shoo cold ha diness, oo eezing ole ance should he e o e be
in es iga ed.
He e, eigh p o enances o P. nig a om au och honous o igins and om sou he n Ge many
we e es ed o hei cold ha diness in a common ga den expe imen in sou he n Ge many.
We hypo hesized ha (1) cold ha diness di e s be ween p o enances, wi h p o enances om
colde o igins displaying supe io cold ha diness, and ha (2) cold ha diness is a ec ed by
clima ic expe iences o he indi iduals wi h d ough inc easing cold ha diness and wa ming
dec easing cold ha diness. We u he expec ed ha (3) di e ences in cold ha diness be ween
p o enances a e physiologically- ela ed o he con en o soluble ca bohyd a es and lipid
composi ion o he needles, and ha (4) he (sub-) medi e anean species P. nig a is less os -
ole an han ee species na i e o Cen al Eu ope, while (5) cold ha diness o he ine oo s
o P. nig a is high compa ed o cold ha diness o i s oliage as i na u ally occu s in egions
wi hou con inuous snow co e .
6 Manusc ip s
132
Ma e ial and Me hods
Ju eniles o P. nig a om eigh p o enances h oughou Eu ope we e exposed o di e en
clima e change scena ios (wa ming and ex eme d ough ) in a common ga den expe imen .
Cold ha diness was de e mined by he Rela i e Elec oly e Leakage me hod (REL) in wo
consecu i e win e s. The expe imen was es ablished in Bay eu h, Ge many (49°55’19” N,
11°34’55” E) in Ma ch 2009. The long- e m mean annual empe a u e o he si e is 8.2°C,
whe eas long- e m mean annual p ecipi a ion is 724 mm.
Expe imen al Design
Eigh p o enances o P. nig a (Figu e 1; Table 1) we e ob ained as seeds and cul i a ed a he
Ba a ian Ins i u e o Fo es Seeding and Plan ing (ASP) in Teisendo , Ge many om Ap il
2008 o Ap il 2009. These p o enances a e pa o an in e na ional long- e m p o enance ial
which s a ed in 2009 (Hube , 2011). The p o enances s em om au och honous popula ions
o P. nig a excep o he p o enance om Zellingen, Ge many, which was in oduced om
Aus ian sou ces in 1909. Subspecies iden i ies o he p o enances a e assigned
geog aphically and mo phologically (Table 1), as gene ic analyses a e no ye a ailable
(Hube , 2011). The C oa ian p o enance s ems om a loca ion e y close o one o he ew
au och honous s ands o P. nig a subspecies dalma ica and i s assignmen o he subspecies
nig a is somewha ques ionable. The seedlings we e anspo ed o Bay eu h and indi idually
plan ed in o 4-li e plas ic po s illed wi h sandy sil (pH 7.3, o al C 1.9%, o al N 0.15%,
plan a ailable NO
3-
-N 22.5 mg l
-1
; plan a ailable NH
4+
-N 1.8 mg l
-1
). Selec ion o he plan s
occu ed andomly o each p o enance om all hose plan s ali e a he plan ing da e. The
mean plan size a he s a o he
expe imen was 12.2 cm ± 2.5 cm SD.
Figu e 1: O igins o he a ge
p o enances (open ci cles) wi hin he
dis ibu ion o P. nig a (black lines and
do s o agmen ed popula ions Isaje e
al., 2004). X indica es he expe imen al
si e. Abb e ia ions o p o enances a e
speci ied in Table 1. G ey scales display
he mean minimum empe a u e o he
pe iod 1950 o 2000 wi h a 5’ spa ial
esolu ion (Hijmans e al., 2005).
6 Manusc ip s
133
Table 1
:
O igins o a ge p o enances used in he expe imen wi h co esponding clima ic
in o ma ion. Skie: Iden i ica ion numbe in an in e na ional p o enance ial (Hube , 2011). MAT:
Mean Annual Tempe a u e; MinT: Mean Minimum Tempe a u e; MAP: Mean Annual P ecipi a ion;
P ecip. Seasonali y: Coe icien o a ia ion in mean mon hly p ecipi a ion sum. All clima e da a o
he pe iod 1950 o 2000 om wo ldclim (Hijmans e al., 2005).
P o enance Coun y Subspecies Skie No h Eas Al i ude
(m)
MAT
(°C)
MinT
(°C)
MAP
(mm)
P ecip.
Seasonali y
DE Ge many nig a 01 49°53'17" 09°43'16" 290 9.2 -3.1 587 18
AU Aus ia nig a 07 47°46'00" 16°11'00" 369 8.4 -4.9 712 33
YU Se bia nig a 12 43°49'39" 19°35'22" 866 8.7 -5.6 964 17
HR C oa ia nig a/
dalma ica
14 43°26'00" 17°13'00" 256 13.2 1.2 1108 33
IT.N I aly nig a 17 45°42'00" 13°49'00" 372 11.4 -1.2 1212 17
IT.S I aly la icio 19 39°18'08" 16°20'22" 1500 9.0 2.2 1300 48
FR1 F ance nig a 23 44°09'10" 05°52'30" 549 10.7 -2.9 789 16
FR2 F ance la icio 24 44°24'18" 03°58'39" 581 10.8 -0.9 745 19
The po ed indi iduals we e exposed o he ully c ossed h ee old ac o ial combina ion o
(1) a d ough manipula ion (d ough and con ol) and (2) a con inuous wa ming manipula ion
(wa ming and e e ence) and (3) he p o enance ea men (eigh p o enances). The wo
clima e ea men s we e c ossed esul ing in ou clima e manipula ions (con ol, d ough ,
wa ming, wa ming & d ough ), ha we e eplica ed h ee imes, esul ing in 12 expe imen al
uni s in o al. The p o enance ea men was nes ed wi hin each expe imen al uni . Each
p o enance was u he eplica ed wi h se en plan s pe expe imen al uni (nes ed eplica es),
esul ing in 21 plan s pe ac o ial combina ion o he h ee- ac o ial design and 672 plan s
o e all. Each expe imen al uni was co e ed by a single ain-ou shel e (11 m by 7 m, 3.8 m
high) cons uc ed o a s eel ame (GlasMe all Rieme GmbH) and co e ed wi h a anspa en
polye hylene shee (0.2 mm, SPR5, He mann Meye GmbH) enabling an almos 90%
pene a ion o pho osyn he ically-ac i e adia ion. The edge o he ain-ou shel e s was a a
heigh o 80 cm.
The con ol i iga ion egime simula ed he local daily 30-yea a e age p ecipi a ion. The
applica ion was ca ied ou wice a week wi h collec ed ain wa e . The d ough ea men
consis ed o 42 days wi hou p ecipi a ion, which ep esen s he local s a is ical 1000-yea
ecu ence d ough e en . D ough du a ion was no a p io i se be o e he manipula ions. We
moni o ed plan pe o mance du ing he ea men and would ha e s opped he ea men
6 Manusc ip s
140
Clima ic Expe iences Al e Cold Ha diness
Cold ha diness was a ec ed by he clima ic expe iences o he indi iduals. The ex eme
summe d ough inc eased cold ha diness by 3.9°C on a e age in he i s win e and he e
was a non-signi ican end in he same di ec ion in he second win e a e he d ough
manipula ion (Figu e 3). Unexpec edly, he summe wa ming om he i s yea esul ed in
inc eased cold ha diness o 3.4°C on a e age while he yea - ound wa ming o he second
yea esul ed in no signi ican e ec , al hough he end ollowed he same di ec ion as in he
i s yea . In e es ingly, he d ough and he wa ming e ec in he i s yea we e no addi i e
(ANOVA, in e ac ion be ween d ough and wa ming: F = 16.0; p = 0.004), esul ing in lowe
cold ha diness in he un ea ed a ian and compa able, high cold ha diness in he o he h ee
clima e manipula ions (Figu e 3).
Physiological Reasons o Va ying Cold Ha diness
The amoun o soluble ca bohyd a es in he needles inc eased by 25.9% in a p o enance
exhibi ing high cold ha diness compa ed o a p o enance showing low cold ha diness (Table
2; ANOVA: F = 15.3; p = 0.004). The d ough manipula ion had no signi ican e ec on he
ca bohyd a e concen a ion (F = 0.0; p = 0.889).
The composi ion o in e nal a y acids (ACL) as main componen s o cell memb anes did no
di e signi ican ly be ween a p o enance exhibi ing high cold ha diness and a p o enance
showing low cold ha diness (F = 3.7; p = 0.092). Likewise, no e ec o he d ough
manipula ion was ound (F = 1.7; p = 0.222). Fo he epicu icula wax lipids, he p o enance
wi h he high cold ha diness was cha ac e ized by a sligh ly highe ACL (3.9 %; F = 5.2; p =
0.051) han ha wi h he low cold ha diness. The d ough ea men led o a gene al dec ease
in ACL (F = 22.0; p = 0.002), which was s onge o he plan s wi h a high (5.5 %) a he
han a low cold ha diness (1.5 %; in e ac ion be ween p o enance and d ough manipula ion:
F = 7.7; p = 0.024).
The g ea es di e ences among lipids we e obse ed o he CPI alues o he epicu icula
wax alkanes be ween he wo p o enances. CPI alues we e 22.7 % lowe o he p o enance
wi h high cold ha diness compa ed o ha o he p o enance wi h low cold ha diness (F =
45.8; p < 0.001). D ough led o an inc ease o he CPI alue by 11.5 % in he p o enance
wi h high cold ha diness, while no e ec o he d ough manipula ion was obse ed in he
p o enance wi h low cold ha diness, esul ing in no signi ican e ec s o he d ough
manipula ion (F = 2.3; p = 0.166) and he in e ac ion be ween p o enance and d ough (F =

6 Manusc ip s
141
3.4; p = 0.104). Hence, he di e ence be ween bo h p o enances dec eased a e d ough , bu
CPI alues we e s ill 15.1 % highe in plan s wi h low cold ha diness.
Table 2: Compa ison o ca bohyd a e con en and a e age chain leng h (ACL) o a y acids o cu en
yea needles be ween wo p o enances exhibi ing low (IT.S) and high (FR1) cold ha diness. Samples
aken in he i s win e o he expe imen , mean ± s anda d de ia ion p o ided, n = 3.
Cold ha diness o sou ce Con ol D ough
Cold ha diness (LT50 in °C) high -25.8 ± 2.3 -36.7 ± 1.0
low -17.5 ± 1.6 -26.8 ± 4.5
Soluble ca bohyd a es (‰TM) high 118.4 ± 9.4 111.9 ± 9.5
low 89.0 ± 15.9 93.7 ± 3.0
ACL o epicu icula wax a y acids high 18.8 ± 0.2 17.7 ± 0.2
low 18.1 ± 0.0 17.8 ± 0.1
ACL o in e nal a y acids high 17.8 ± 0.5 17.6 ± 0.1
low 17.5 ± 0.1 17.1 ± 0.3
ACL o epicu icula wax alkanes high 27.5 ± 0.1 27.6 ± 0.0
low 27.5 ± 0.1 27.5 ± 0.1
CPI o epicu icula wax alkanes high 9.3 ± 0.2 10.4 ± 0.2
low 12.0 ± 0.6 11.9 ± 0.4
Cold Ha diness among Species
Cold ha diness o he (sub-) medi e anean P. nig a eached simila le els o he cold
ha diness o ee species na i e o Cen al Eu ope in he icini y o he expe imen al si e
(Figu e 5, ANOVA o all ju enile lowland samples: F = 2.0; p = 0.163). Needles o adul
coni e s, howe e , showed supe io cold ha diness compa ed o ju enile ees (F = 2.6; p =
0.046; Picea abies and Pinus syl es is) and ju eniles om highland si es exhibi ed highe
cold ha diness compa ed o lowland si es (F = 7.4; p = 0.026; Picea abies and Pinus
syl es is).
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142
Figu e 5: Compa ison o he cold ha diness (LT50) o P. nig a wi h common o es ee species in he
icini y o he expe imen al si e (lowland, 350 m asl) and, o he ju enile s age o he o he wo
coni e s, om a highland si e (760 m asl). ju .: ju eniles (2-4 yea s old); ad.: adul s (>30 yea s old).
Que . pe .: Que cus pe aea; Fag. syl .: Fagus syl a ica. n = 3 mixed samples o 7 indi iduals each
pe ba (mean and SE). Fo P.nig a: p o enance DE (Zellingen, Ge many) in he con ol ea men .
Cold Ha diness o Roo s
Cold ha diness o ine oo s o P. nig a a e aged -16.5°C. The wo es ed p o enances (IT.S
and FR1) did no di e signi ican ly (F = 1.6; p = 0.239) in he cold ha diness o hei oo s in
he second win e o he expe imen (-15.4 ± 1.9°C and -17.5 ± 1.1°C espec i ely (±1SE), n
= 12). The d ough (F = 0.1; p = 0.805) and wa ming (F = 0.1; p = 0.754 ) ea men s also
esul ed in no signi ican e ec on LT50 o he ine oo s.
Discussion
Local Adap a ion in Cold Ha diness
Cold ha diness di e ed by abou 10°C be ween he s udied p o enances o P. nig a. Local
adap a ion o minimum empe a u e egimes was indica ed as p o enances om colde
o igins eached supe io cold ha diness. These esul s co espond well wi h indings om
o he o es ees such as P. de oniana (Saenz-Rome o and Tapia-Oli a es, 2008), Fagus
syl a ica (Visnjic and Doh enbusch, 2004), Tsuga he e ophylla (Kuse and Ching, 1980),
Fagus c ena a and Be ula e manii (Ganse e al., 1999), all showing local adap a ion o
win e cold ex emes. Ou da a indica es u he ha minimum empe a u e does no only
de e mine he no he n ange limi s o species (Sakai and Weise , 1973), bu ha wi hin
species a iabili y in cold ha diness also needs o be aken in o accoun . The p o enance om
C oa ia (HR), howe e , did no i well in o he o e all pa e n. We assume ha his
p o enance is ei he no au och honous, i.e. o igina ing om a wa me win e clima e, o
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143
belongs o he subspecies dalma ica, which is desc ibed o e y es ic ed a eas along he
C oa ian coas . Gene ic cha ac e iza ion o he species and subspecies will shed ligh on his
ques ion.
P. nig a is known o i s high gene ic di e si y (Jagielska e al., 2007) which su passes ha o
o he pines (Nkongolo e al., 2002). Al hough no consensus on i s axonomy has been eached
(Hube , 2011), six main subspecies a e ecognized wi h P. nig a ssp nig a being he mos
abundan in Eu ope (Isaje e al., 2004). P o enances u he mo e di e in g ow h and
ecological pe o mance, exp essed in local adap a ions o soil and mean annual empe a u e
and p ecipi a ion in p o enance ials (Va elides e al., 2001). The s ongly con as ing cold
ha diness in ou expe imen sugges s ha minimum empe a u e is ano he gene ically
selec i e pa ame e , no only o os sensi i e subspecies such as P. nig a ssp la icio
(Va elides e al., 2001), bu also o P. nig a ssp nig a, which is usually conside ed o be he
mos os ole an among he subspecies (Isaje e al., 2004, Hube , 2011).
Clima ic Expe iences Al e Cold Ha diness
Cold ha diness was a ec ed by clima ic expe iences o he indi iduals wi h d ough
inc easing cold ha diness by 3.9°C on a e age in he subsequen win e and no signi ican
ca y-o e e ec o he second win e . This inding can be explained by d ough and os
igge ing simila esponses a he cellula and molecula le el o p e en cellula dehyd a ion
(Blodne e al., 2005). Wi hou expe iencing d ough hemsel es, he newly o med needles in
he second yea o he expe imen lacked signi ican addi ional cold ha diness in he d ough
manipula ion. Mo e equen d ough e en s accompanying clima e change may he e o e
inc ease cold ha diness in single (d y) yea s, bu no gene ally.
I has been sugges ed ha ees g own unde gene ally wa me condi ions may lose hei
unc ional adap a ions o os (Eccel e al., 2009). Su p isingly, ou esul s con adic his
expec a ion wi h inc eased cold ha diness by 3.4°C on a e age in he wa ming ea men a e
he i s season. The wa ming, howe e , was s opped in Oc obe and acclima ion o he
o me ly wa med indi iduals e iden ly su passed he con ol plan s when subjec ed o he
same empe a u e om Oc obe on. Responsi eness o cu en yea clima es a e also epo ed
o deciduous o es ees (Repo e al., 2008). Ye , he yea - ound wa ming in he second yea
o he expe imen esul ed in no signi ican di e ence be ween he ea men s. Clea ly, u he
expe imen s on in e ac ing clima ic d i e s a e u gen ly needed, as he esponse o such
in e ac ions migh di e conside ably om single ac o expe imen s (Shaw e al., 2002;
K eyling e al., 2011b).
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144
He e, we ocused on he ealized maximum os ha diness, e.g. he ha diness di ec ly a e
he coldes days o win e . Much bigge di e ences han obse ed be ween he p o enances
(10°C) o be ween he clima e manipula ions (up o 3.9°C) occu wi hin each needle o e he
cou se o he yea (mo e han 60°C in a single p o enance o P. nig a; Su inen e al., 1992).
Cues which d i e his s ong seasonali y in ol e pho ope iod and minimum empe a u e
expe ience. Thei ela i e impo ance, howe e , is s ill un esol ed, di e s be ween species
(Kozlowski and Palla dy, 2002; Holliday e al., 2008) and may e en di e be ween
p o enances in he same species. Ou esul s add o his discussion by showing ha bo h
gene ic he i age (di e ences be ween he p o enances) and p eceding clima ic expe ience
(he e mainly summe d ough ) can a ec he absolu e os ha diness. Po en ial di e ences in
he empo al pa e n o os ha diness be ween p o enances a e o high ecological ele ance,
especially wi h ega d o ea ly o la e os e en s and phenological di e ences wi hin species
(Visnjic and Doh enbusch, 2004; K eyling e al. 2011b). These poin s call o mo e de ailed
in es iga ions on in a-speci ic di e ences in seasonali y o os ha diness.
Physiological Reasons o Di e en Cold Ha diness
Di e en ial cold ha diness be ween p o enances was ela ed o con en s o soluble
ca bohyd a es and a y acids in he needles. Con en o soluble ca bohyd a es is also epo ed
o be closely ela ed o local adap a ions in cold ha diness o di e en Que cus species (Mo in
e al., 2007). The lipid con en s o P. nig a ha e been epo ed p e iously o ma u e ees and
needles collec ed du ing la e summe (Ma ei e al., 2004). In con as o hese ma u e ees,
whe e n-C
29
and n-C
31
alkanes con ibu e 2.2 % and 37.2 %, espec i ely, o o al alkanes, he
ju eniles o di e en p o enances in ou s udy we e all domina ed by n-C
29
alkane (36.3 ± 2.8
%) and lowe con en s o n-C
31
alkane (9.6 ± 1.0 %). This di e ence is p obably due o
di e en needle and plan age when compa ed o he li e a u e esul s, whe eas di e ences
be ween p o enances a e no likely as hey did no di e in hei ela i e con ibu ion o n-C
29
and n-C
31
alkanes in ou s udy. In gene al, he hyd ophobici y o he waxes is imp o ed unde
wa e and cold s ess o p o ec plan s agains wa e loss by an inc eased u no e o wax
componen s owa ds hyd ophobic alipha ic compounds which is no necessa ily ela ed o
shi s in he o al amoun o waxes (Shephe d and G i i hs, 2006). Fo ees, such
in es iga ions a e s ill sca ce and limi ed o selec ed ee species (e.g. o di e en Picea
species: Cape and Pe cy, 1993; o Pinus palus is: P io e al., 1997). Ou obse a ions o
small changes in he lipid composi ion (ACL alues) con i m mino in luences o cold and
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wa e s ess on lipid biosyn hesis, as desc ibed elsewhe e (Cape and Pe cy, 1993; Shephe d
and G i i hs, 2006). The low CPI alues o epicu icula wax alkanes o he plan s wi h high
cold ha diness, howe e , indica e a s ong biosyn hesis a e associa ed by a p oduc ion o
byp oduc s and deg ada ion p oduc s such as e en alkanes. This inc eased p oduc ion o wax
componen s indica es he ole o alkanes o imp o e he cold ha diness (P io e al., 1997).
The d ough manipula ion led o a educ ion in he o ma ion a es o alkanes (highe CPI) o
he p o enance wi h highe cold ha diness simila as obse ed o sesame plan s (Kim e al.,
2007). Hence, biosyn hesis o epicu icula wax alkanes is in luenced by wa e s ess and
appea s o be ela ed o cold ha diness in P. nig a. F eezing ole ance in plan s is
accompanied by lipid emodeling a he ou e memb ane (Moelle ing e al. 2010), ano he
aspec i ing well o ou da a and indica ing ha he e ec o changes in he lipid composi ion
migh be mo e impo an o cold ha dening han p e iously assumed. I should be no ed,
hough, ha bo h he obse ed di e ences in cold ha diness and he di e ences in
composi ion and concen a ions o cell memb ane compounds could be d i en by o he
ac o s such as wa e s ess o e summe in he p o enance wi h supe io os ha diness.
Seasonali y o p ecipi a ion and mean annual p ecipi a ion we e h ee imes lowe a he
o igin o his p o enance (Table 1). Fu he causal and unc ional analyses o os ha diness
and ha dening a e clea ly equi ed (Holliday e al., 2008).
Cold Ha diness among Species
Ju eniles o he (sub-) medi e anean species P. nig a exhibi ed compa able cold ha diness as
ju eniles o species na i e o Cen al Eu ope in he icini y o he expe imen , i.e. unde he
same clima ic condi ions. Unde colde condi ions in No way i has been shown ha P. nig a
is mo e sensi i e o eezing inju y han bo eal coni e s (S imbeck e al., 2007). I s abili y o
adjus o p e ailing clima ic condi ions he e o e appea s limi ed in compa ison o bo eal
species such as P. syl es is o Picea abies. Ye , unde he same clima ic condi ions, hese
species did no di e om P. nig a in ou s udy, implying ha ealized os ha diness and
po en ial os ha diness need o be discussed sepa a ely. I should be emphasized he e ha
wi hin-species a ia ion in cold ha diness, i.e. di e ences among p o enances o P. nig a and
di e ences be ween lowland and highland si es o ju enile and adul indi iduals o he o he
species clea ly exceeded among-species a ia ion a he ju enile s age. Gene ally, a ia ion
among species a he same si e and unde he same clima ic condi ions appea s less impo an

6 Manusc ip s
146
han commonly assumed. Wi hin-species a ia ion and indi idual pe o mance migh be mo e
ele an o o es ecology (Cla k, 2010).
We used ju enile ees in hei second o hi d yea in his expe imen . Ou esul s conce ning
P. syl es is and Picea abies con i m p e ious indings ha seedlings a e mo e sensi i e
agains os e en s han olde ees (Bol e e al., 2007). Howe e , he ju enile s age is o high
impo ance o he na u al egene a ion o o es s ands. Mo eo e , he high selec i e p essu e
o single ex eme e en s such as os o d ough can educe he gene ic di e si y o u u e
s ands (Hosius e al., 2006).
Cold Ha diness o Roo s
Cold ha diness o ine oo s o P. nig a a e aged -16.5°C o e wo p o enances, which is a
high alue compa ed o he cold ha diness o i s oliage (-23.8°C on a e age o he con ol
ea men ). This migh be an adap a ion o he species’ na u al habi a whe e soil os e en s
occu mo e o less egula ly as no snow co e insula es he soil agains ai empe a u e
luc ua ions (K eyling, 2010). Chamaecypa is noo ka ensis se es as an example o a o es
ee om empe a e ain- o es s wi h low oo cold ha diness ( oo s do no su i e
empe a u es below -5°C) in esponse o deep snow co e in i s na u al habi a (Schabe g e
al., 2008a). Win e clima e change, howe e , is expec ed o lead o educed snow co e and,
in consequence o he educed insula ion, o colde soils despi e he gene al ai wa ming end
(G o man e al., 2001). Fo Cen al Eu ope, a educ ion in snow co e is al eady aking
place, while minimum empe a u e o he soil may no dec ease (K eyling and Hen y, 2011).
In e es ingly, no esponse in cold ha diness o ine oo s occu ed o he di e en clima e
ea men s in ou expe imen . In addi ion, we in es iga ed oo cold ha diness o wo
p o enances wi h s ongly con as ing shoo cold ha diness and did no ind signi ican
di e ences in he oo s. This suppo s Schabe g e al. (2008a) who conclude ha no
acclima ion occu s in cold ha diness o ine oo s. Mo e de ailed in es iga ions conce ning
his aspec a e clea ly needed, especially wi h espec o he ques ion i cold ha diness o oo s
lacks adap i e po en ial o changing clima e condi ions. Fo P. nig a ou esul s imply ha no
selec i e p essu e is expec ed as oo cold ole ance is gene ally high.
Assis ed Coloniza ion
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147
P. nig a, based on i s ecology and na u al dis ibu ion (Isaje e al., 2004), is well adap ed o
wa me and d ie condi ions expec ed o pa s o Cen al Eu ope unde clima e change
(Kölling, 2007, Hube , 2011). He e, we show ha cold ha diness, a leas o some
p o enances, is also no limi a ion o he use o his species in Cen al Eu ope e en i cold
ex emes emain cons an h oughou his cen u y (Va us e al., 2006; Kod a e al., 2011).
Assis ed coloniza ion o ansplan a ions a e widely applied in o es y and may se e as
adap a ion s a egy agains ad e se e ec s o clima e change on ecosys em unc ioning
(McKenney e al., 2009, Schabe g e al., 2008b). Nume ous examples o ailed
ansplan a ions (Zobel e al., 1987), howe e , wa n agains ushed ac ion. The assis ed
coloniza ion o p e-adap ed eco ypes o key species wi hin hei cu en ange is sugges ed o
con ibu e o unc ional in eg i y o o es s ands wi hou he need o in oduce exo ic species
wi h unknown isks (K eyling e al., 2011a). Ye , na u ally domina ing ee species may lack
p e-adap ed eco ypes a hei wa m and d y ange limi s. He e, congene ic species om
adjacen clima es a e p e e able o e o he species. P. syl es is and P. nig a may se e as an
example, wi h he la e po en ially eplacing he o me a wa mes and d ies si es o i s
ange while main aining ecosys em unc ioning. Ou esul s sugges ha cold ha diness is
signi ican ly ela ed o clima ic condi ions a he o igin o he p o enances, implying ha he
selec ion o os - ole an p o enances could be based on he cu en clima ic condi ions
wi hin he species anges. Howe e , ou inding ha clima ic expe iences wi hin he li e o
single plan s al e cold ha diness indica es ha p o enance ials unde con ol condi ions may
be misleading unde changing clima ic mean and ex eme condi ions. The mul i ude o
possible clima ic a iables o be selec ed o and unce ain ies conce ning u u e clima es
imply ha he sea ch o bes -adap ed p o enances should no be he only s a egy. In
addi ion, managemen ac ions which p omo e gene ic di e si y (e.g. suppo ing na u al
egene a ion and addi ion o gene ically di e se ma e ial) a e c ucial as gene ic di e si y
enables o ganisms o con inue adap ing and e ol ing o new condi ions wi hin one o se e al
gene a ion cycles (Hosius e al., 2006; Schabe g e al., 2008b). Fu he mo e, he ole o
he bi o es and diseases unde changing clima e equi es de ailed in es iga ions. Fo ins ance,
a needle bligh known as he “ ed band disease” (Do his oma sep ospo a) is epo ed o
inc ease in impo ance o e ecen yea s in P. nig a (Isaje e al., 2004), a de elopmen ha
may be ela ed o clima e change (Wa e al., 2011).
Ul ima ely, ee species esponses should be ega ded in he con ex o popula ions unde
compe i i e p essu e. The ad an age o common ga den expe imen s is ha hey can de ec
he spec um o possible species-speci ic esponses. Ne e heless, he e is a need o es he
6 Manusc ip s
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ob ained esul s in communi ies whe e he compe i i e balance migh ampli y o bu e
esponses.
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149
Conclusions
Cold ha diness o Pinus nig a oliage is highly a iable be ween p o enances and shows
signs o local adap a ion o p e ailing minimum empe a u es a he o igin. Bo h se e e
d ough e en s and summe wa ming can inc ease cold ha diness, indica ing ha he
in e ac ion o di e en clima e pa ame e s leads o unexpec ed esul s and ha win e su i al
can be al e ed by clima ic e en s du ing he g owing season. Physiologically, cold ha diness
is ela ed o soluble ca bohyd a e con en and lipid composi ion. In e es ingly, a ia ion o
cold ha diness o he needles wi hin he (sub-) medi e anean species P. nig a was highe han
be ween his species and o he species common o he empe a e zone o Cen al Eu ope.
Taken oge he , ou esul s imply ha he cold ha diness o he oliage o P. nig a is adap i e
o long- e m g owing condi ions a he o igin (gene ic he i age) and o sho - e m al e a ions
o hese condi ions (indi idual plas ici y), while i s hin s sugges ha cold ha diness o he
oo s is high and p obably no unde selec i e p essu e cu en ly. Ou da a om mid-win e
sugges s ha below- and abo e-g ound cold ha diness o selec ed p o enances appea o be
well adap ed o cul i a ion in empe a e egions as an adap a ion s a egy agains he ad e se
e ec s o clima e change in d y habi a s. Howe e , wi h espec o la e sp ing and ea ly
au umn os e en s, he empo al pa e n o os ha diness wi h po en ial in a-speci ic
di e ences should be in es iga ed in mo e de ail. Be o e ansloca ions a e ecommended,
u he in es iga ions a e equi ed, e.g. explo ing he ole o bio ic in e ac ions unde
changing clima ic condi ions. Gene ally, wi hin-species di e si y should be conse ed a he
species le el and imp o ed in an h opogenically ounded s ands in o de o allow o adap ion
o clima e change.
Acknowledgemen s
This s udy was unded by he Obe ankens i ung (OF a_02631) in coope a ion wi h he
"Ba a ian Clima e P og amme 2020" in he join esea ch cen e “FORKAST” and he
Ba a ian S a e Minis y o he En i onmen and Public Heal h (ZKL01Ab 7_18456). We
hank Ch is ian Schemm, Elke König, S e an König, Ch is ine Pilsl and nume ous s uden
wo ke s and in e ns o hei ou s anding help du ing he ield wo k.