Sensitivity of Plant and Soil Ecosystems of the Alps to Climate Change

84 Pages·2006·1.66 MB·English

5
SensitiovfiP tlya natn dS oiElc osystoefm s
theA lpst oC limaCthea nge
Jean-PTahuelu riFlrlaant�,Fo eilsb Peart,r iGceiias sler,
Jean-MiGcohbealMt a,r lyFsiee rAzn,d reFaiss chlin,
PhiliKpuppef eArn,d rSec hlusCsaetle,rV ie nlal uti,
Cui-FaZnhga oa,n dJa nnW illiams
5.1 CLIMATE AND VEGETATION
Climiasatm ea jdoert meirnaonftt h dei striobfeu ctoitsoeynmso sne arth
(e.Wgo.ow,da r1d9 8a7st) h,le a titzuodnianotafitl oh ne vegbeitoamteiso,n
sucahse vergrraeifenon r est, baonrtdeu anl,d a rsfba oerloetrzss ot n,a ­
biomferso tmh eeuq attoort hpeo lieulssl trCaltiemsfa.ut ned amentally
influtehndeci eissbt urtoifbo ont h sppleacanintevd se get(aet.Wigao.ln,t er
198149,8 W5a;l taenrBd r eckleE ll1ee9rn91gb19 ;81 69,8 8, I1nh9 i9g6h) .
moutnasi,cn limianutfeeln cneost eoncloys ysltaetmistd'ui disibtnuratli on
buatl stohe ailrt niatdlui dsitriIbnudteaielodtn,i. t zuodniantcaihlao rna cter­
iztehsve e getoafta ilholin gm ho unttahiafntos r smea prabtimeoe sc,a lled
oroiobmetsh,oa tc cwuirt htihzneo nobsiT.oh mueasno, r obicoomneso ifs ts
all vtehgee tzaottniiaoonfn r otmh beot totmot hteo opf t hmeo untain
ranTghege.r addueaclr oefma esaean it re mepratwuirtiechn r eaaslitnigt ude
(.05K5p e1r0 m0e teerlse'v aattmi iodndl laet itiusadbo eu1st,) 0 t0i0m es
steeipnEe urr oapnend or thAesrina 1(K ap beoru tk2 i0le0ot metrhsa)n the
latitduedcirnteaoalws aetr hdpe o l(eWsa l1t9e8r14 9,8 T5h)i.is sh a tlhfe
gradpireendti fcoNtroe rdAt mhe rbiycH ao pkilna'(wsM aAcrth1u9r7 2).
Hencvee,ga ettiboenl ts of aro5er00 ot bo1i ,o0mt0ei0sm n easr rotwhearn
thvee gaettizoonn oefzs o nobiaonmadem so,d ecshta nignme e aann nual
aitre mrpaetmuaryes ignifiacffaentcthtael l yt itduidsitnraiolbfp u ltainotns
anvde get(aet.iHgoa.nl, p1 i9n9 41a9,9 4Obz;e nadnaBd o r1e9l9 119,9 4).
Projecftogirlo onwbsaa rlm ibnytg h lea thtaeolrfft hnee xcte ntsuhroyu ld
modibfiyo mpersen'st re aensgA.c cortdoiH nagl pin (1994a, 1994b), high
moutnaisny stseumcsah st hEeu ropAelapansr lei kteolb yep artrilcyu la
vulnetroac blliemc ahtaena gnetd,h miasyh avsee vbeiroel oagniedcc aol­
nomciocn sequences.
Conseriidtnhgbe i oloagnisdco adcloo enoimmiopcr taonfbc oett hhf eo r­
esatnstd h e azlopnoiefnt heAe lp ss,e veqruaels tsihoonubslea d dr dessed
connciengrgl owbaarlm ipnotge'nsit mpiaaclt s:
Tow haetx tiecsnl ti miantfleu etnhceiesncgeo tseymss?
•
Inw hawta yasr e etchoetsseyeml ssi kteorl eys ptoocn ldi mcahntagee ?
•
Howm ayd iffecroemnpto noeftn htesss yes treemssp ond?
•
What bweti hlel orfca htaena gned ltihmeoi frt ess poofnts hee ssey se­co
•
temasn d tdhieffiecrroe mnpto nents?
Whawti blelt hmee chanoifrss empso nse?
•
How cwaebn e sats stehsiesm apctosfc limcahtaen ognef oerstasnd
•
alpeicnoets eymss?
Iti sa ne xecdeincgolmyp lteaxst koa sscelsism cahtaen gpeo'ntste ial
imapcotnse cosysMtoermes .f undraeemsaerinctsrha e lq ubierfeoadr r eel i­
abelvea luoaftt himeoa nj or ctarbneem n addsSe t.it lhdlee, fi nfienagt oufr es
thseu ballppiinenece-lo aicn(ei .ree.l,a tuinvmeoldyi fied,v ismduaiarflk­ ed
ferebnectew teheuenp per sfuboarleapsniatdnls ep mienaed omwask)ie at
uesfsuly stfeosmrt udycilnigmc ahtaincgp eot'esn tiimaapclot n m ounitna
ecosysTtheirmsas ia.sf eusr tqhueers tIiispo ton ss:i btlofie n mda rkientr hse
biospahnetdrh peee dsophetrome o nitthoeerv oluotfci loinm cahtaen ge's
imapcotne cosystems?
A comprehreenvsiioevftwe h see intsivtioct lyi mcahtaen ogfet he
diverrasnoegf ee c otseymissn the Alipbsiye noen tdrh esegc iooopftne h is
chaptetrh.ri esFa oswroe fn o,c muasni loyn t hpeot entrieaslp oonfs es oil
ecosysatnedm astl hpezi onnet eoc hantgeemdep rataunrpde r iepcitation
regipmeredsi cftoterhd ce o ming cOetnhtieumrrp yo.ra tsapneotcf gt lso bal
chan(geen hancoefam mteonstp hCe0r2ic co ncenteruattriootpnih,oi nc,a
acidifiicnactriieonUVan -s,Be c, h nageisnl anuds ew)i blelm entioonnleyd
whetrhee iyn tesryancettr igciwasiltlchyl imfaatcitco rs.
Herweea naltyhzpeeot entrieaslp oonfps leao nftt sha el pzionnete oc li­
matceh anugsei,pn ugb lisheda nlsdiut tdeiirenpas rt uorgera tet swsos ites
int hAel posft hVea lawihse,sr eo ialnsvd e get(abtriyoonp vhayustlceasr,
planatrbseie) n ign vestaitsg earvtaeellde voefcl os mpleTxhietssuyted. i es
are retfoae str hr"eeE dcc loipnreeo cjtF.i"g u51r. ree ndtehroesr ngiaza­
tiolneavlae nlidsn teraccotnisoinidsnte hrEeec do cplorijneec t.
In cthhaipswt ee r,
1.b rierfleyv itehbwei odivoefbr rsyiotpyh ayntvdea sus lcaprl ainntt sh e
EuropAelapns
2.o utltihmneae i n ecaonlbdoi goilcoagli cpaerlv ialfiafncobgtro o trshso ils
and vegetthasetu iboanla pniiadnnl ep zionneoe fst hAel sp,em phzaisnig
thiem oprtaonfcc leai tmfiacc tors
3.p reensatn i ntegarsasstemesedn bta,s oende xampflreotsmh peas to,f
howa lpienceo sysmtaeyrm ess pdoinffde arleltnyotc ilimcahtaen agned
whatth ceo nuseenqcoefts h miasyb ef obri odiversity.
Exploroaftt hieposonnei t iscs o upwlietadhs u mmaorfky e ays ssmseenotfs
vegetaantdis ooenic lo smyssr'te espotnocs leai tmceh nageT.h ec hapter
conclbuypd reosp oasni nign tveigsraiabotonetu dht pe a radoifsg epmc ies'
indivriedsupaoln se.
226 Chapt5e r
Figu5r.e1 Domaiannsdo rgaantiizolneavleo lfts h bei osphienrr eel atiotnots hhfieie pl odfs
studmye,t hoodfsi nvestigaantdti iomne,s cRaellee.v ainnccer ewaistehs bsoptahts icaall ing-up
andt imPer.e ciisnicorne ianst ehsoe p posdiitree ction.
5.2T HE DISTRIBUOTNI OF BRYOPHYTESA ND VASCULAR
PLANTSI NT HE ALPS
5.2.1V egetatBieotlns F,o resatnsdA, l pinEec ossytemosft heA lps
Chagensi ne levarteisiounntl htfe o rmaotfmi oornoe rl ehsosm ogeneous
vegeta­
sectoifvo engse teaxttieognno d vie7nr0 m0e teirnes l evactailolne d
tiboenl ts
(gfiur5e. 2G)i.v tehnre geu ltaerm perdaetcurrweeia teshle e vation,
av geetatbieolcnto rrestpoao r nadgensi nm eaann nutaelme pratoufr e
arou3n8.Kd . A cocrdgit noS chr(o1t29e3r1) 0,m0 e teorfes l evasthioornt ­
entsh ger owgis neasbyon ni ndea yosna, v egera,an nu(asliilnsxy p rgai nnd
thrienae u tumn).
227 SensitoifPv liatanynt d S oiElc osystoeftm hsAe l ptso C limaCthea nge
Elevat[imo]n
Dayosf Internal
growign Alps
season
3,100
2,400
1,700
1,000
<300
Vegetation
belts
Figur5e. 2V egetatioinnt hbteee lmtpse rpaatroetf t hAel psw,i tihn dicaotfit ohnes leonfg th
thger owisnega s(odna wysi tmhe adna ialiytr e mpeurraoet fa tl ea5s°tC ).
sub­
Thhie ghebsetwl hte froer ceaspntot etniasltlgiyrl oliw st ermtehde
alvpeitgniebelot etna.
Genrealtlhybi,les t l'osw leirmi intt h Eeu ropAelapinss
betw1ee,n2a 0n01d , 7m0 a0.. lsa.n,id t s ulpipmieisbrt e twe1e,n9a 0n0d
2,4m0 0( LanadnodAl etcs hima1n89n6 L;a nd1o9l29t;F avar1g99e5rb ).
Formatoifpo hnasn ero(pwhoyotpdelysa nttaslt lhear5n 0c entimeters­
treaensds h rucbhsa)r acttheivrsei gzeet baetliwtoh,ni ch subporpeoarlt s
(Paibcieeas
conifefroorueossfs t pse cieassN osruwcsahpy r uce (LK.a)r st.),
(Ldaerciixd ua (Picneaum sb r
larch MillAerro)lp,li an e L.a)n mdo untpaiinne
(Pimnuugso uncinata
supsp. (DCD.om)i nT)h.es ubaldpeicnied fuooruess ts,
patricubleaherwclooyd sg,r oiwnt hleo wpearro tft hviesg etbaelttoi no n
theex tebrnoarldo eftr h Ael spa nadr seu bjteoac m to rAet lacnltiimca te
(Pache, Micha1l99e6t)T,.h esa unbda lvApeiigmaneete ti boenl mte'asn
annutaelmr paetiusbr eet w0e.eCn5 a °n4d° Ca,n td hvee getgatrwiotohn
pereixotde onvde1sr01 t o1 8d0a y(sO ze1nd9a8T 5h;e ur1i9l)9l.1a t
alpvietngaeetb ieolnt
The begiantas r nodu1 ,9t0o02 ,4m0 0as ..aln.d
exteunpdt so 2,630,001m 0.Ft oom ratioofsn wsa radnsld o hwe atthhse,
228 Chapt5e r
abseonfpc hea neyrtoeapsd h,e creaabsudena niocnfae n nupalla (ntthse ro­
phytaensod)f b ulpblaar(n gtesyo tpehasn)td,h p ers eenocfce u shpiloann ts
characttheirsi (zebe.e Lgla.tn ,da onlAdt e cshima1n9n8L 6a;n d1o99l2t;
Ozen1d89a5 F;a var1g9e81r999 ,5) aA.l all ppilnaecn otm mnuitciaebnse
consi"dseprieeacdl i(ziel.die"m. i,tt ope adr tihcaubliastreas etce st;i5 o4.n. 3)
witghe omorphmoilcorgo,yra ,en ldsi ieetxfep ossutrreo dnegtleyr mining
thediirs triTbhuetfsicaeot nod.re srt meitnheme i crocalnitdmh arete el ated
lengotfsh n ocwo v(eTrh orn1w9a5i4t)e.
nibv,ea llt
Abov2e, 60,01-m03e0t eirfsos u tnhde wherteh ger owsien­ag
soinsl se st ha5n0d yasA.tt heeslee vastnioorwne sm,ao innfl sag tr ound
througthhsoeuu mtm Tehrse.ee nvironcmoenndtipatrlil eoucndtsehd ee vel­
opmeonfdt o spelda cnotm mtuinseiusca hss warVdass.c pullaaonrftt sh e
alpbienlfeot r lmo opsleac notm mnuitiinse csr aenerdso cfiksu sreisnt he
lowpearro tft hnei vbaellH tih.ge urp t,h epslea gnrtosow n lsyp oradically
inf avorracobklpyel ,a ces.
5.2.1.F1o restTsh feo reosftt hEseu ropAelapcnso nstaim taujeotcreo ­
sytseamn fdu lsfivelelri amlp orftuanncsttT. ih oernye gurlnuaotffte,h ereby
moderatthhieyn dgr olroeggiiacmnaepdl r eveenrtoisnoigfso tne selpo pes
(Brooakls1. 9 e9Tt1h )e.py r otheucmtas net tlemfernotamsv anlcahaensd
landsTlhiedpyeor sv.if dueep lu,l pwaonotddi ,m bfeohrru mauns aen di,n
conjunwcittaihlo pnmi enaed oawrtseh, de o mincaonmotpn eonfat n a ttrac­
tilvaen dsfcorarep cer eaatcitioitnviIaenals d .d ittoti hoenei cro normoilce s,
Alpifnoer easratems o ntghm eo snta turreamla ienciontsgey missnt hAel ps,
othtehraa nl peicnoes ysatnetdmh su,s ptlhaaeyniy m oprtraoniltneb io­
divercsointsye rOvnaa tl iaornsg.ce arul ned,te arnsdAilnpgif noer ecsltis­'
matsee tnisvimiatypy r oviimdpeo ritnasniitgn httthospe o etntrieaslp onses
ofm ountafionroeeucssot ts eymtssh rougthhwoeou rtl d.
Foreasnttdhs ae t mospfhoerarmnie n tertswyisntieenwdm h itcehme pra­
turper,ie pcitaatnwdii onnid,ns fl utehnsect er uocftf uorreea snftdos r,ie ns ts
turfne ebda ctkot hceml aitsiyces mtv ia chiananb lgeedssou ,r fraocueg h­
nesasn,ed v arpaontspi(re.a.W,tgo ioodnw a1r9d8R 7u;n niannNgde mani
1991O)n.a r egisocnaasllte u,dh iaevsseh owtnhp eot enotfit ahlef seee d­
bactkos arffeegcitco lniaml(a et.Lege .aa,nn Wda rri1l9o;8wP 9 ieeltak le.
1949).
In the ocfco lnitmceahxtaten f goer,ea sritems op rttaenrtr eesctorsiyasl­
tembse causec afintx ht,eh yr opuhgoht osyanntsdh teosaris esi ,g nificant
amouonfct a rbFoonir.n stoanna gc leo,sb caall e,e cfotoserymesssst to8 r0e
preceonftt h aeb ovoeu-ngadrn 4d0 p ecrenotft hbee loowu-ngndro fnols si
carbwohni,ac mho utnot6 s2 -7p8ec renta lcolaf r bsootrne idn ttehrer es­
tribaiploh searne�d 1 -7p7re cepnretc eonfat l cla rbionn wtohrel bdi'os­
mas(se .Dgi.x,ao nnTd u rn1e9r9W 1i;s nieewatsl 1k.9i ;9D 3ixeoatnl 1.94 9;
PerruchaonFuddi h slc1in99 5T)h.e resfinogirfiecc ahnnagtei snt heex toefn t
thweo rlfdo'rsec sotueslx da cecrhbaantigence lsi mbayrt eel ealsairngge
229 SensitoifPv liatannydt S oEiclo sysotfte hmAesl ptsoC limCahtaen ge
amounotfcs a rbionnt o the (aKtimanongsNd pe hielr1se99o 2nN; e inl so
1993;S miatnhSd h ug1a9r9tW3 a;t seotan l1 .96 9).
5. 2.1.A2l pinaen dN ival ZonIenas phytoggreaophsiecnastleh ,e
alpzionneoe ra, l pviengee tbaetliitost,n h a er ebae twtehetenr eea nldi ne
thsen ow Albionuoetn.f e o-ur(,t100h0 -1,s1p0e0co ifte hsAe)l spfl'o rias
fouinntd h ael pzionneie nc, o njunctiounp pwfeiortr zheo snttaeh n etd h e
nivzaoln Teh.e sspee cciaelsol,re odp hcyatbnee sc ,o nsidtehArele pds '
"primdairvye rtshiattty h"io sssp,ee cniaietvste ot hAel passo popsetdo
tho(smea iwneleydr su,d earnagdlr ,al sasnsdp ecwihesosen) a tudriaislbt ur­
tiohnab se ne wideelxyt entdherdo huugmha anc tiMvoitsoytr. o phytes
evolfvreodm s utrhreo uflnodroiaftn hgwe a rlmo wlainntd hlsea Tteer tiary
(PlioTcheendieir)s .t riibntu htaeil opnzi onneie sm oraef acotfom ri cro­
climgaetneesr baymt iecdr o(rteolcpiloeifm tahteaosnft) h gee necrlailm ate.
Thde iisbturtoifto hne osreo phiysat lescsoo nsitdoeb reie ned q uiulmi bri
witnha tucroanld ibteicoantushsa eel pbilenteu' ppse zro naetl, e aissct o,n ­
sidetrohe adve ex perimeinnciiemmdap laf crtoh mu man Iunts heri.es s tp,e c
thael painnndei vbaella trse ltaghrees ta trievulenlmyo dieficeodsm y isnt e
middEluer ope.
Thter leient ehc,eo ntbaecttw tehefeno raensAdtl pibneeli tsa,s tirking
bioclbioumnadtai(rcey . g., T1r79a9nW;qa urid1ll9le8i H3no;il tm1e89i9e;r
Tal1l9i9sS1 l;a tayneNdro b1le92 9)W.h ereed apahniodcr oagprhfiacc tors
abrupprtelvy engtr ottwhoh fte r etehste,r eeclaibnne ae s habropu nrdya
(aenc otownheec)rle;i mdaettee rtmhibenou ensd aart yr,a nsiztoin(oean na l
ecocloifhn eea)t hs, shtrurbeasen,smd ,e satduocnwatsone c duc rb etween
the floirmoeirstt it,m bleirna en,d ttrhleeie m oirtt r,le ien e. Atcoc ording
Tal(l1i9sa9 n1Sd)l atayneNdro bl(9e19 2t)h,te r eliein sed etermbiyn ed
"treest"h attnwa olm leetre( ris.m ei.c,r opphhaynteseresoe; saelcstoi ons
5..2154,.. 6)
Thissu balpineec-oacillpscii annletel h e"edk afmzponoert" h "ek rumm­
holzzo nIet.d"si striflbuucttiuioanrnt e essp toonl soecf aalc tsourcashst op­
ograapnhsdyi etxep osTuhrkeema .p fzonnoewr areexliyus ntdsne art ural
condiitnti hoAenl spb sea cusheu maanc tivhiatvleio ewse trheuedp per limit
off oraelsmstote verywthheubrsrea o,d entihnsegu balpineecc-olailnpei ne
(saeless eoc t54i..o6n) .
5.2.2B ryopyhtes
5.2.2.C1h aracteriMsotisecsssl ,i verwaonrdht osr,n waorrceto sl lec­
tivkenloyw ansb ryophtyhtseee sc,ol nadr ggersotou fpl anpdl anctosm,­
prissionm2ge3 0,00s pecwioersli ddwTeh.e ayr peo iilkohypdlraintcth sa,t
ispa,ln twsh oswaet seatrt ucso mpledteepleoynn dt sh aotft haee reinavli ­
ronm(enWta l1t39e1r,1 98W5a;l taenrBd r ec1k9l58e,1 991S)u.c phl ants
toledrraotuebg yhr te ducpihnogt osyanntrdhe esspiisar nawdti itothnas nd­
incgo mplteitseds euhey drIantc ioonnt.tr oa stm otshsheeo sm,e odhryic
230 Chap5t er
vauslcaprl anthsa ve developeda dmtaoaprtpiahonondils no tgeirncaall
conducstyisnttgeh maestn abtlheet mom ainrtealiant ciovtneaslwnyta ter
contents ionftd heeepinervn idreonPntom ieknitl.po rhoyvdtirhdypee h sy­ s
iloogli caabtiosl uirtvyti ovt ehp,eo ionfdt e saitcicaoh ni,gv ha riaitni on
watceorn tIennd rtcy.o ndibtriyoonpsh,sy ttrgeolsrny e sfirssotatn hde at
(Long1t89o0nG) r.o twhi rse iscttrteopd e rioondlsy twhhmeeo nsi sws e t.
Somber yoypthecsap nh otohseysnaittzv ee rlyo iwn tewrnaatlce orn tents
(Pro1c9t28o)ar,ni mptoaranbti floicrto ym petpiatritoinc,u lcahralnyg ­with
inggr owctohn idoints.
Bryophlyitivenems i cnrvoieronwmheonsctoesn dimtaiydo inffsfe rro m
thoosfte h seur roundviengtgae tdieotne,r bmyi( nmead)c icrmloa atnetd,h us
thesyh ocwh aracstuesbrtirssapttceiiecfi cBirtyyo.p hcyactnoe lso bnairzee
soaiflt learsn ldiodreo sth deirusb rtanc(eSsl 1a9c)8kI.8 n o thseirtu ations,
thecyo nsttihteeud taepc hliicm( asxe e s54e..co3tn)ir oonci kncs r,e vices,
anidn s crfeeensas,n, sd t reVaamrsi.bo ruyso pchoymtmetu inaeirspe r esent
ass ynuisnpi laaecn otm mnuitdioemsi nbaytv eaudsl cpalra nts.
Inb ryoyptheass,ue axrle purcotdiioson f taem no riem pormteaanontfs
proagpattihoasnnxe uarle prod(uDcutriaionnnvdg a Tno or1e9n8 T7h)e.y
havaev arioefmt oyd eosfv egetpartoipvaeg Ianht airoesnnh.v irtosnm,e n
bryophmyatype erss iatsav t e getsattaigvee fpoerr iIlondod insog.e cious
tax(at hwoistesh e xoenss erpaatpel anstusc)sc,fes uselx uraelpr oduction
depeonndt shs ei multparneesooeufnbs co teh wsheixgceehso r,ga pmhayy
rest(rLiocntga tnodn Gr1e69e9naGe)e .n evtairci aibesis elpciithayil glihyn
moss(eWsy aSttntoe,b uranneOdrd ,r zsyski1ko 899W;ya t1t99 4a)si, ts h eir
phenotpylpaisct( iLcointgyt on 1B9l7i4as;nsH d,a B muisl1bt9yo7,n 8 ).
Unfortuenocaptheylsyi,o dlaotgai ucnaaarlve a ifloaarbl lpeib nrey o­
phytaelst,h oeuxgthe nescioyvpseohi logwiocrahkla bse ecna rroiueitdn
polraerg iBoecnasu.as rec (taincad n tarecntviicrn)ot nmsma eyt os ome
extebnect o mpartaob laecl opnidniet thieorsneess, um latyp sr ovsiodmee
insirgehltestv oat nhEteu ropAelpasIn.na ddittihseop ne,ci vineesst igated
aloscoc iuncr e ntEruarelo. p
Thpeor ductoifbv riytoyp hiylsto web,sut th eaibro vegprrooudnudc tion
mayb e sitmoi ltahvraa tus lcpaolrfna tisna rcatrie(caO se cahneSdlv e­inb
jorn1s79s8oI)nnC. a naBduas,bB yl,ia snsHd,a mil(t1o9na7 n8Rd)o binson,
Vitatn,dT imon(e18y99 h)a vset udbireydo phryetsepsot'no ms iec ro­
climaantedid ca pghriacd ieesnpiteasclw,la yt Berry.oy ptherso'il nee cosys­
tempsr'eo scsss eucahsn utrient ianntde rwcaaentprdete rit oenni tssit oinl l
poloyru ndersattlo eoadfs,ott r e mperate arnegdi (otLnroson pgitcoanl
1929)A.g aimno,rd ea taara ev aifloaprbo llreae rg i(oLnosn g1t9o81n49 ,8 8).
Studoinen su trciyecnltii ntn hgAe l astkaain(g Oae cahneVdla nC leve
1896C;h apeitan l 1.89 7s)h owtehdat thb er yoplhayytfeenur c tiaosn sa
seelcitvfiel taecrq,u irinnugt rsiroeampneita dsnl ldyo stihnegsm l owly.
Byrophyttiesd seucepo omsmeusc mho rseol wltyh atnh aotfv ascular
plabnetcsa iuths aeas h igcho ncentorfla itginoincn o-mlpiokuetn hduss,
accumulaacdtiiifnygoi rngga mnaitct ientr h seo il (e.1g9.9, 6)H.o bbie
231 SensitoifPv liatannydt S oiElc osysotfe mAtslh pets oC limCahtaen ge
Herbivaonriomunasol rsm adlonl oygt r amzoes sbeesc aoufst eh eir low
digestibility.
5.2.2.P2a tternosf D isrtibutiBonr yophhyatveaesn a ncient ongm,
witthh eoilrd estd aftoitsnosg i utlphspe De erv onians.p eEcxhitaeavsne t
beefno uinnd cEeondepe oistIsnG. r eBarti taaniNdno rtAhm erifcoas,s il
alpsipneec iedsi swceoravetse irtfeeadfsr r omp rtehsede instt rirbauntgieo n
(Mil1l9e8rF3 o)tr.h E eu ropAelappnsa ,l ceooleogdiactaal stuhppepr oerst­
encoefb ryophiynpt oeslsta gcdieaplo saisit nps ae,tp rofiblueutsn ,f ortu­
natewliyt hnoouf tu rtihdeern titfihcaajntu is"otmon s seHso.we"v ewre,
maya ssutmhea tp rtehseae lnptbi rnyeo pvheygteet easttiaobnl iitssheeldf
int he early Holocene.
Int eromfss pec1i,e1sm5,o4 s saensd4 23l iverawrokern tosw fno r
Euroipnec,l utdhiAenz go r(eCso ralnedCy r nudwe1l99l1 G;r ol1l89e3 ).
Morteh atnw o-tohfitr hdEesu ropsepaenca irefeso uinndt hAel posf,
whicahb o4u0tp ercceanbnte c onsiodreorpehdby atsees,d Sowni stsh e
florfaow,rh i7c7hm6 o ssaensd l2i5v2e rswpoercti reesp oa(rrUtere emdti
Grimmia
al1.92 9)T.h hei ghest-mgorsiosntw hiAenl gpi sas n i ndeterminate
Hedws.p ecait4e ,s86 m 3etear..ssla .tt hteo opf t hDeu four(sMpointtzee
Grimremciuar va
Rosoan)t hSew i-sIstabloirad(neV rca ac1ri9 )1.1 Schwagr.
(G=.,s essitana
cf. DeN otE.M.;a iepreo,rn saclo mmcuanitiiotsnh h)ei ghest­
growiknngo wsnp ecait4e ,5s5,4m etear..ssla .tt hteo opf P unGtnai fetti
(MonRtoes)i anI taAloyssVt aal l(eVcyaa c1r91i 3).
Inc onttroat shtae b undaonfvc aecs ulpalra ntthse,ar reve e rfye w
Riccia breidleri
endebmriyco phiynt eAtslh pews i,t ohn ltyh lei verworts
Hrebesrettnnudesr i
Steapnhd. (NeLeisn)rd ehc.o gansi zesMduo cshss.p ecies
oncceo nsiednedreehmdai vce sinbceeed nie sictohovenera r neodtc hoenr­
tinoernr te dutcose ydon nymayf ttearx onroemviicMs ainoyn .a rlaprien e
specairemeso rfer equiena nrtc rteigci Ionnc so.n ttroat shtle o wnldas,
wherteh sei gnihfiucmaanintm pahcatsc onsequently tihmep overished
divertshiaetl pybi,rn yeo pvheygteet iasst tiqioulnil rt ieca hnw de ldle vel­
oped, imnsa uibnallyfp oiraneensh dte actohm mnuitaineidsn h umivde ge­
tattiyopnse usca hsf enssp,r ingssn,o wabnwedhd esr,e b rtyhoep lhayyteer
mays how conpsrioddeurcaatbnilddvei i vteyr sity.
Thger emaatj oorfib tryy ophhyatvaeel s a rdgitesr iburtainocgnie r,c um­
borfeoatrlh mea joorfia tlyps ipneec Tiherso.u gthhoAeul tpe sa,cs hep cies'
ecolorgeuiqicraelm veanrtlisie tsct olmep atrote hdo osfev ascpuallnatrs .
The roefsb urlytose corleosrgeciaahcr taehl e reafpoprlei claabrlggeee o t­o
graphairceaals .
Int hAel psso,m gee neorfba r yophyhtiegsh dlaiyrv ee rasnidafi lesdo
possgersespa hte notpylpiaicscit Etcyoo.gl icmaold ificatthiaroten flse ct
unfavogrraobwlteh comnadyi theiartom anpxso neovmailcu attihoeny, yet
may ubsee ffouerln vironmmoennittaolFr oiernx ga.m pFlrei,s( v1o89l8l)
Racomhiettreiruoms tichum
harse cernetvliyts hceeod m polfe x (HedBwrd.i.),
consitdoed raetoden e spiencc einetEsru arlof poetr,h Neo rthHeermni -
232 Chap5t er
spheArtle e.afi svtwe e ll-detlaixmai,wt ieetdaahp c raht iceucloalrao rgey ,
reporftoterhd Ae l sp.A simiploragrr ahmab see ns eutp f otrh gee nus
GrimHemdiww,ah o stea xoniontm hyAe l spw aps oloyru ndrestuonotdi l
recenAtskl nyo.w leodnhg aeb irteaqtu itrshe amisemp nroviehtda ,bs e en
suggetshtasetod m sep ecmiiegsbh etu seadsi ndicsapteoc(riM easia enrd
Gesils1e9r9 .5 )
5.2.3V asculaPrl ants
5.2.3.C1h aracteriOsrtiocpsh y(tseessee c t5i2.o.n1a .r2se)p ecifically
adapttoce odlc do nditthiroonpushg yhs ioalnomdgo ircpaohlgo ilcchaalr ac­
teri(set.iAgct.sk, iB no,t maann,d L ambe1r9s9F 6r;i eannddW oodward
1990K;o me1r89 91,9 9119,29 ,1 99149,9 K5o;m er Dainedm1 e9r8 7;
Korneetar l 1.9 ;8K 9omearn Lda rc1h9e;8rL 8 arc1h9e8r10 9,9 T4h)e.s e
spieechsa vdee velsoepveedmr eaclhi asnmosfr esnicset atoT hfreoys t.
asmsiileaffitcei eanttt elmyp ernaeta0ur°r Cde,es s ptihlteoe w pearr ptrieasl­
suroefC 02e,x hiabh iitgm hi thoocndrreisapli (rdaatrriekos np irraattei on)
inc ompartiops loannf rtostm h leo wla(nsdeasel ssoe ct5i.o4n.a 2n.d1 ),
acmcuulaltaera gmeo uonfts sou lblseu gaTrhsea.yl hsaov aeh ignhi trogen
cotnenatn da bundsaennctdo acroym poundesx.ap mlFteoh,re p yo ssae ss
higlhe voefal n tioxsiudcaahnsa t sscb,oi arcc iads, aadnta apttioot nh e
inteinrsrea dtihaamttai yog ne neroaxitdea stusibtvaen (cWeisal ndLdiu tz
1996P)l.a inntt sh ael pbienlaetl seox himboripth oloagniadcn aatlo mical
adaptattoli ootwne sm peraanthdue rreesd fietaatrsuyur caehsts h ilceka ves,
Carex
smlaslzi ea,n sdl ogwr owTthhte.i lolfte hriesm poratlapnsiten deg e
cuar vul
Allf.oe,rx amplmea,yg roown t hoer doef0r . 5m-i1l leirpm eert
yea(rG rabMhaehrrar,n,R d e is1i9g7lH8 o)w.e vtehroi,cs c iunr s conjunc­
tiwoint dha ytciamneo tpeym pertahtmauatry re esa 2c0h- 3b0e°cCa oufs e
strornagd iaAtlipoipnnl.ea nitnshe'r selnogtwrl oyw itrsh e lattoa es dm all
leaarfep ael re baifo maal sism,in tuemdbo efdr i vidiantgt hgecr eolwlisn g
pointasnl,do nwi gthetm perianthuiitrbiemnsig't odvtiiiscai notdnh erefore
ceglrlo wBtahs.oe nd tehnev ironcmoenndtiaatltti hoaenl sp inpel anbte lt,
morpho,la ongdpy lnatl ongeovriotpyh,ay rtceeo sn sid"esrtertdeo sls­
erataocrcso"r tdoit nh"gec opmetivtei-sttroerlsaes-n rtudesrchaelm"e
Grim(9e17 9d)e veloHpoewvdee.rc ,l oosbeesrr vatiinto hnhesi gh-alpine
anndi vbaelli tnsd itchaoatrte o phdyont oectso mplfiettt ehcleoy n coefp t
stsrt eoslerbaetcoartushs,eaey l sshoo cwo mpetaintrdiu vdeet rraasliu tcsh,
ass uppioncrgot nsriadbheleer bibvyso mrayml la ls( Die1m9e9r6 ).
amm
5.2.3.P2at ternosfD istributTihoeEn ureoapAnl sps uppaoh ritg h
levoefpl l abnito divienar r seiltayts imvaaelrlleyw a i,ta hb o4u,t5 v0a0s ­
culpalras npte cmioertseh, aa tn h iorftd he en tEiurreon pfl eoar(aw eosftt h e
UralFsva)a.r g(e91r7 2i)d ent1i,fi0eo4dr9 o phtyatxiiantc h Ael sp( about
23p ercoeftn htAe l sp'e ntfliorreTa h)eh. i ghgersotw ivnagsl capurl ant
·
233 SensitoifPv liatanynt Sd o iElc osysotfte hmAesl ptsoC limCahtaen ge
(Sfarxbaiigfla ora
obserivne dt hieas s A xalipfrsa ge Alla.t4) 4 5m0e tear.ss .L
att hDeo minV al(aAinsc Ih98i5s)i.
AccordtioPn agw low(sIk9i73 079) t,a xian,t hnea rrsoewn saer,e
endemtiotc h Ael sp.I na ddittihoenar,re5 e 5 s ubendteamx(iapc l ants
havitnhgem iarid nit sribiutnti hAoeln p bsue tx tenadlitsnooog t her moun­
tairna ensgA)m.o ntgh een demaincdss u bendemmoirtcehs a,hn a lafr e
oropsh.By atseoendP awlowfisgkruie'Fssa, v ar(gIe9r7m 2e)n tio3n3eId
(3p2e rnctee)n demaimcosnh giI s, 0s4p9e coifoe rso phHyotweesv.te hre,
staotfue sn demtiacxr ae mauinncsl beeacra oufst eh lea cokfa precise
assmeesnostft hAel spf'ol raasw elalst axondoimviecr sgI.en anb coreader,
convsaetrisveeno snelI,y5 p ercoeftn htpel asnpte coifte hsAe l pmsa yb e
endem(iFcasv apresgroenrca olmu mnication).
Orophydtiesst'r ipbtautterireonsnsu ltf rmoatmih inenl tye groaftt wioo n
dominfaancttc olrisma:an tmdea cerloirGelfa.c idautriitonhngPe l teoicsene
playaek de yr oliens hanpgti he presentp adtitsoetrfsrn piseb cuiteiso n
accortdoii nntsgri bicin oglioccahla ract(edsripiesrtcsiaicposan cc iotmeypt,i ­
tivaeb ilaidtayp,t aatnidpo enr,s isStpeacnicieens)d'.i avliidrsuetiscp onse
toc limvaitciics smiatkuedsde isffii cttu oul stt eh ediitrsr ibuttodi eofinn e
phygteoorgaphriecgailio ntn hsEe u ropAelpasna, n ndo a greeemxeinstt s
betwaeuetnh eovrefsno t rh mea jdoirv is(iTohnesue rtai llI.l9 a9tH4 o)w.­
evesro,mg ee nepraaetlrt nesx i(sOtz eIn9da8I 59,8 I89,9 F4a;v arIg9e8,r9
I99I59a,9; 5T bheurIi9l9l5a)t.
Foumra idni stripbtautteoirfoon nrs o phcyatbneed s i stinguished:
I.M ansyp ieecasrd eit sribtuhtreodtu hgwehh orlaen ge
2.M ansyp ecoicecosun rli ynt hwee snto erer atserpna rotft hAel posn,
eitshiedoref a l ifnreo Bmo dentsoLe aeg doiC omoo,ra l ifner oLma ke
GenetvoLa a gMoa ggiore
3.M ansyp ieecesx hiabn iotr th-dsiuosnujctthii nto hne diirs triabsu tai on
resouftl htae b seonflc iem esitnto hnceee ntpraarolftt hAel ps
4.S evesrpaelci inecsl,ub dohitt nhgo dsies trtihbruottuehgdeh ernatnigree
oft hAel passw elalse atseronrw eesrtdni sbturtiisohnosw, dwiijdsuen c­
tioornf ragmednittsreidb sum,ta iiondnluyte o g laceivaeln ts.
Ina ddittoti ohnef soeup ra ttseervnessr,pa elcp ioesssc ehsrso mosraocmeisc
(dipploloyipdl,to hiasdth) o aww e-setadsits tri(bputhtoyigoernao pghical
pesudovicFaarviaaIrng9ce6er2; ) .
Glaceixapla nhsaifdoo nuo hrte erff ecotnts h deit sribuotfoi rooynpt hes,
ina ddittoti hosoneo ng enedriaslt ripbautttieornn s:
I.t hea ncesotfot rhseA rcTteotr-iaflroyr wae rael msotc omlpetely
extirpated
2.m anoyr ophytleism iwtteorer edef uignti hnae or theassuottehrnse,tna e,r
ands uothwteesrmna rgoifnts h Aelp sw,h erteh eayr neo wd itsributed;
theefrotrheme,a in coefen ntdeermsai rstemh M ea irtiAmlep tsh,se uo htern
Alpbsew teeLna gdoiC omaon Lda goG adrida ant,dh leo ewa stAelrnp s
234 Chapt5e r