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PROF. DR. FRITZ SCHWEINGRUBER Institute for Forest, Snow and Landscape Research WSL Zürcherstrasse 111 8903 Birmensdorf, Switzerland ANNETT BÖRNER PROF. DR. ERNST-DETLEF SCHULZE Max Planck Institute for Biogeochemistry PO Box 100164 07701 Jena, Germany Front cover illustrations (from left): Cross-section of a pollarded ash stem (Fraxinus excelsior). The light zones indicate small rings which are an expression of a pollarded crown. Valais, Switzerland. NKOOOA?PEKJKB=LAPNEłA@ BKOOEHAraucaria OPAI-APNEłA@#KNAOP NEVKJ= 20 Cross-section of dead wood from an Red Bearberry (Arctostaphylos uva-ursi). Most lumina of the cells =NAłHHA@SEPD>NKSJOQ>OP=J?AOSDE?DLNARAJPPDASKK@AJOPNQ?PQNABNKIIE?NK>EKHKCE?=H@ACN=@=PEKJĠI=CJEł?=PEKJk Cross-section of a birch stem (Betula pendula) with demarcation lines. They represent concentrations of mycelia of different fungus species.
Library of Congress Control Number: 2006927045
ISBN-10 3-540-32523-9 Springer Berlin Heidelberg New York ISBN-13 978-3-540-32523-9 Springer Berlin Heidelberg New York 1DEO SKNG EO OQ>FA?P PK ?KLUNECDP HH NECDPO =NA NAOANRA@ SDAPDAN PDA SDKHA KN L=NP KB PDA I=PANE=H EO ?KJ?ANJA@ OLA?Eł?=HHUPDANECDPOKBPN=JOH=PEKJ NALNEJPEJC NAQOAKBEHHQOPN=PEKJO NA?EP=PEKJ >NK=@?=OPEJC NALNK@Q?PEKJKJIE?NKłHIKNEJ=JUKPDANS=U =J@OPKN=CAEJ@=P=>=JGO!QLHE?=PEKJKBPDEOLQ>HE?=PEKJ or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permissions for use must always be obtained from Springer. Violations are liable for prosecution under the German Copyright Law. Springer is a part of Springer Science+Business Media springer.com © Springer-Verlag Berlin Heidelberg 2006 Printed in Germany The use of general descriptive names, registered names, trademarks, etc. in this publication does not imLHU ARAJEJPDA=>OAJ?AKB=OLA?Eł?OP=PAIAJP PD=POQ?DJ=IAO=NAATAILPBNKIPDANAHAR=JPLNKPA?PERA laws and regulations and therefore free for general use. "@EPKNġ!N!EAPAN VAO?DHEG %AE@AH>ANC $ANI=JU !AOGA@EPKNġ!N J@NA=0?DHEPV>ANCAN %AE@AH>ANC $ANI=JU KRAN@AOECJġ!AOECJ-NK@Q?PEKJ$I>% %AE@AH>ANC $ANI=JU =IAN=NA=@U>U JJAPP ÌNJAN 'AJ= $ANI=JU -NK@Q?PEKJġ)"1A5'AHKJAG 0?DIE@P3Ì?GHAN$>/ )AELVEC $ANI=JU Printed on acid-free paper
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(IPLZHSIH 7PUHJLHL :PS]LY-PY (IPLZWPUZHWV 7PUHJLHL :WHUPZO-PY (JHJPHHI`ZZPUPJH -HIHJLHL -SH[[VW(JHJPH (JHJPHJ`TIPZWPUH -HIHJLHL (JHJPHKLJ\YYLUZ -HIHJLHL )SHJR>H[[SL (JHJPHKYLWHUVSVIP\T -HIHJLHL -S\[L(JHJPH (JHJPHLY\ILZJLUZ -HIHJLHL )S\L;OVYU (JHJPHOV[^P[[PP -HIHJLHL (JHJPHTLSSPMLYH -HIHJLHL )SHJR[OVYU(JHJPH (JHJPHW`JUHU[OH -HIHJLHL .VSKLU>H[[SL (JHJPHYLÄJPLUZ -HIHJLHL 9LK<TIYLSSH(JHJPH (JLYJHTWLZ[YL (JLHYHJLHL *VTTVU4HWSL (JLYWZL\KVWSH[HU\Z (JLHYHJLHL :`JHTVYL (KHUZVUPHKPNP[H[H )VTIHJLHL (MYPJHU)HVIHI (KHUZVUPHNYLNVYPP )VTIHJLHL (\Z[YHSPHU)HVIHI (KLUVJHYW\Z]PSSVZ\Z -HIHJLHL *VKLZVKL*\TIYL (LVUP\THYIVYL\T *YHZZ\SHJLHL )SHJR;YLL(LVUP\T (LZJ\S\ZOPWWVJHZ[HU\T /PWWVJHZ[HUHJLHL /VYZL*OLZ[U\[ (NH[OPZH\Z[YHSPZ (YH\JHYPHJLHL 2H\YP (SU\ZNS\[PUVZH )L[\SHJLHL )SHJR(SKLY (SU\ZPUJHUH )L[\SHJLHL .YL`(SKLY (SU\Z]PYPKPZ )L[\SHJLHL .YLLU(SKLY (SVLKPJOV[VTH (SVHJLHL 2VRLYIVVT (TIVYLSSH[YPJOVWVKH (TIVYLSSHJLHL (TIVYLSSH (UHIHZPZHY[PJ\SH[H )YHZZPJHJLHL (UPZ\TZH[P]\T (WPHJLHL (UPZL (YH\JHYPHHUN\Z[PMVSPH (YH\JHYPHJLHL 7HYHUH7PUL (YH\JHYPHHYH\JHUH (YH\JHYPHJLHL 4VURL`W\aaSL;YLL (YJOHLVW[LYPZ (YJOHLVW[LYPKHSLZ (YJ[VZ[HWO`SVZ\]H\YZP ,YPJHJLHL 9LK)LHYILYY` (YNHUPHZWPUVZH :HWV[HJLHL (YNHU;YLL (YPZ[VSVJOPHJSLTH[P[PZ (YPZ[VSVJOPHJLH )PY[O^VY[ (ZWHYHN\Z[LU\PMVSP\T 3PSPHJLHL (ZWHYHN\Z (ZWSLUP\T]PYPKL (ZWSLUPHJLHL .YLLU:WSLLU^VY[ (Z[LYV_`SVUTHJRPLP (Z[LYV_`SHJLHL ([YPWSL_ZW *OLUVWVKPHJLHL :HS[I\ZO (aVYLSSHJVTWHJ[H (WPHJLHL @HYL[H )HURZPHIYV^UPP 7YV[LHJLHL )YV^U»Z)HURZPH )HURZPHZLYYH[H 7YV[LHJLHL :H^)HURZPH )LYILYPZZW )LYILYPKHJLHL )HYILYY` )LY[LYVHPUJHUH )YHZZPJHJLHL /VHY`(SPZVU )L[\SHUHUH )L[\SHJLHL +^HYM)PYJO )L[\SHWLUK\SH )L[\SHJLHL :PS]LY)PYJO )L[\SHW\ILZJLUZ )L[\SHJLHL +V^U`)PYJO )V\NHPU]PSSLHZWLJ[HIPSPZ 5`J[HNPUHJLHL )V\NHPU]PSSLH )YHJO`JOP[VUY\WLZ[YL :[LYJ\SPHJLHL 8\LLUZSHUK)V[[SL;YLL )Y\N\PLYHN`TUVYYOPaH *VTIYL[HJLHL )\YTH4HUNYV]L )Y`VUPHKPVPJH *\J\YIP[HJLHL >OP[L)Y`VU` )\_\ZZLTWLY]PYLUZ )\_HJLHL )V_>VVK *HSHTP[LZZW *HSHTP[OHJLHL *HSS\UH]\SNHYPZ ,YPJHJLHL /LH[OLY *HWWHYPZZW *HWWHYPKHJLHL *HWLY *HWZLSSHI\YZHWHZ[VYPZ )YHZZPJHJLHL :OLWOLYK»Z7\YZL *HYL_WLUK\SH *`WLYHJLHL 7LUK\SV\Z:LKNL *HYPJHWHWH`H *HYPJHJLHL 7HWH` *HYSPUHZW (Z[LYHJLHL *HYSPUL;OPZ[SL *HYULNPLHNPNHU[LH *HJ[HJLHL :HN\HYV *HYWPU\ZIL[\S\Z )L[\SHJLHL *VTTVU/VYUILHT *HZ[HULHZH[P]H -HNHJLHL :^LL[*OLZ[U\[ *LPIHWLU[HUKYH )VTIHJHJLHL 2HWVR;YLL *OHTHLKHWOULJHS`J\SH[H ,YPJHJLHL 3LH[OLYSLHM
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7HNL
30:;6-:7,*0,:
*OLUVWVKP\THSI\T *OLUVWVKPHJLHL >OP[L.VVZLMVV[ *OLUVWVKP\TZW *OLUVWVKPHJLHL .VVZLMVV[ *OVZLUPHHYI\[PMVSPH :HSPJHJLHL *PYZP\THY]LUZL (Z[LYHJLHL *YLLWPUN;OPZ[SL *SLTH[PZ]P[HSIH 9HU\UJ\SHJLHL ;YH]LSSLY»Z1V` *VJVZU\JPMLYH (YLJHJLHL *VJVU\[7HST *VTTPWOVYHZH_PJVSH )\YZLYHJLHL 9VJR*VYR^VVK *VVRZVUPHKL]VUPJH 9O`UPHJLHL *VYLTHHSIH ,TWL[YHJLHL *VY`S\ZH]LSSHUH )L[\SHJLHL /HaLSU\[ *YH[HLN\ZTVUVN`UH 9VZHJLHL 6ULZLLK/H^[OVYU *`H[OLHJHWLUZPZ *`H[OLHJLHL *`JHZHYTZ[YVUNPP *`JHKHJLH *`[PZ\ZZJVWHYP\Z -HIHJLHL *VTTVU)YVVT +HKV_`SVUZW *VYKHP[HJLHL +HWOULTLaLYL\T ;O`TLSHLHJLHL 4LaLYLVU +HWOULZ[YPH[H ;O`TLSHLHJLHL .HYSHUK-SV^LY +H]PLZPHTVSSPZ -HIHJLHL +PJRZVUPHHU[HYJ[PJH +PJRZVUPHJLHL ;HZTHUPHU;YLLMLYU +PNP[HSPZVIZJ\YH :JYVWO\SHYPHJLHL +\ZR`-V_NSV]L +YHJHLUHKYHJV +YHJLUHJLHL +YHNVU;YLL +YHJHLUHZLYY\SH[H +YHJLUHJLHL +YHNVU;YLL +Y`HZPU[LNYPMVSPH 9VZHJLHL (YJ[PJ4V\U[HPU(]LUZ +Y`TPZWPWLYP[H >PU[LYHJLHL +Y`VW[LYPZH\Z[YPHJH (ZWPKPHJLHL )YVHK)\JRSLY-LYU +Y`VW[LYPZÄSP_THZ (ZWPKPHJLHL 4HSL-LYU ,WOLKYHOLS]L[PJH ,WOLKYHJLHL 1VPU[7PUL ,WOLKYHTHQVY ,WOLKYHJLHL 1VPU[7PUL ,WOLKYHZW ,WOLKYHJLHL 1VPU[7PUL ,X\PZL[\TOPLTHSL ,X\PZL[HJLHL 9V\NO/VYZL[HPS ,X\PZL[\TTH_PT\T ,X\PZL[HJLHL .PHU[/VYZL[HPS ,YLTVWOPSHZW 4`VWVYHJLHL ,T\)\ZO ,\JHS`W[\ZKP]LYZPJVSVY 4`Y[HJLHL 2HYYP ,\JHS`W[\ZQHJRZVUPP 4`Y[HJLHL 9LK;PUNSL;YLL ,\JHS`W[\ZTHYNPUH[H 4`Y[HJLHL 1HYYHO ,\JHS`W[\ZWH\JPÅVYH 4`Y[HJLHL :UV^.\T ,\JHS`W[\ZZHSTVUVWOSVPH 4`Y[HJLHL :HSTVU.\T ,\JHS`W[\ZZVJPHSPZ 4`Y[HJLHL 4HSSLL,\JHS`W[ ,\JHS`W[\ZZW 4`Y[HJLHL ,\JHS`W[ ,\WOVYIPHJ`WHYPZZPHZ ,\WOVYIPHJLHL *`WYLZZ:W\YNL ,\WOVYIPHWLZJH[VYPH ,\WOVYIPHJLHL ,\WOVYIPH]PYVZH ,\WOVYIPHJLHL ,_JVLJHYPHHNHSSVJOH ,\WOVYIPHJLHL )SPUK@V\Y,`L4HUNYV]L -HN\ZZ`S]H[PJH -HNHJLHL ,\YVWLHU)LLJO -PJ\ZJHYPJH 4VYHJLHL *VTTVU-PN -PJ\ZVISPX\H 4VYHJLHL :[YHUNSLY-PN -PSPWLUK\SH\STHYPH 9VZHJLHL 4LHKV^Z^LL[ -YH_PU\ZL_JLSZPVY 6SLHJLHL *VTTVU(ZO .HSP\TTVSS\NV 9\IPHJLHL /LKNL)LK[YH^ .PURNVIPSVIH .PURNVHJLHL .PURNV .SLKP[ZPH[YPHJHU[OV -HIHJLHLZ /VUL`3VJ\Z[ .SVI\SHYPHJVYKPMVSPH .SVI\SHYPHJLHL /LHY[SLH]LK+HPZ` .UL[\TNULTVU .UL[HJLHL .\PQLYHWHY]PÅVYH 9\[HJLHL /HRLHZW 7YV[LHJLHL /HSV_`SVUWLYZPJ\T *OLUVWVKPHJLHL :HS[;YLL /LKLYHOLSP_ (YHSPHJLHL 0]` /LSSLIVY\Z]PYPKPZ 9HU\UJ\SHJLHL .YLLU/LSSLIVYL
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30:;6-:7,*0,:
3HI\YU\THUHN`YVPKLZ -HIHJLHL .VSKLU9HPU 3HYP_KLJPK\H 7PUHJLHL ,\YVWLHU3HYJO 3HYP_NTLSPUPP 7PUHJLHL +HO\YPHU3HYJO 3HYP_SHYPJPUH 7PUHJLHL ;HTHYHJR 3HYP_ZPIPYPJH 7PUHJLHL :PILYPHU3HYJO 3LIHJOPHWPUPMVYTPZ =VS[aPHSLZ 3LTHPYLVJLYL\Z^LILYP *HJ[HJLHL *HYKVU 3LWPKVKLUKYVUZW 3LWPKVKLUKYHJLHL 3L\JHU[OLT\T]\SNHYL (Z[LYHJLHL 4HYNOLYP[H 3PTVUP\TJHWYHYPLUZL 7S\TIHNPUHJLHL 3PU\T\ZP[H[PZZPT\T 3PUHJLHL 3PUZLLK 3P]PZ[VUPHZW (YLJHJLHL *HIIHNL7HST 3VILSPH[LSLRPP *HTWHU\SHJLHL .PHU[3VILSPH 3VPZLSL\YPHWYVJ\TILUZ ,YPJHJLHL 4V\U[HPU(aHSLH 3VUPJLYHW`YLUHPJH *HWYPMVSPHJLHL 7`YLULHU/VUL`Z\JRSL 3`JVWVKP\THSWPU\T 3`JVWVKPHJLHL (SWPUL*S\ITVZZ 3`[OY\TZHSPJHYPH 3`[OYHJLHL 7\YWSL3VVZLZ[YPML 4HJYVaHTPHJVTT\UPZ AHTPHJLHL 4HNUVSPHZ[LSSH[H 4HNUVSPHJLHL :[HY4HNUVSPH 4HNUVSPH]PYNPUPHU\T 4HNUVSPHJLHL 4HSJVSTPHHLN`W[PJH )YHZZPJHJLHL 4HS\ZZ`S]LZ[YPZ 9VZHJLHL (WWSL;YLL 4HS]HWHY]PÅVYH 4HS]HJLHL 3LHZ[4HSSV^ 4LSPHHaLKHYHJO 4LSPHJLHL *OPUHILYY` 4LU[OHSVUNPMVSPH 3HTPHJLHL /VYZL4PU[ 4LZWPS\ZNLYTHUPJH 9VZHJLHL 4LKSHY 4PJYVILYSPUPHZW -HIHJLHL ALIYH^VVK 4\SPU\TZWPUVZ\T (WPHJLHL 4`YPJHYPHNLYTHUPJH ;HTHYPJHJLHL .LYTHU;HTHYPZR 4`YTLJVKPHZW 9\IPHJLHL 5LVYHPTVUKPHHYLX\PWLUZP *HJ[HJLHL 5LWLU[OLZZW 5LWLU[OHJLHL 5VSHUHZW :VSHUHJLHL 5VSHUH 5V[OVMHN\ZW\TPSPV -HNHJLHL 5\`[ZPHÅVYPI\UKH 3VYHU[OHJLHL *OYPZ[THZ;YLL 6SLHL\YVWHLH 6SLHJLHL 6SP]L;YLL 6UVUPZZWPUVZH -HIHJLHL :WPU`9LZ[OHYYV^ 6W\U[PHZW *HJ[HJLHL 7YPJRS`7LHY 6ZT\UKHYLNHSPZ 6ZT\UKHJLHL 9V`HS-LYU 6Z[Y`HJHYWPUPMVSPH )L[\SHJLHL /VW/VYUILHT 7HJO`WVKP\TUHTHX\LUZPZ (WVJ`HUHJLHL 7HLVUPHZ\MMY\[PJVZH 7HLVUPHJLHL 4V\U[HPU7LVU` 7HYHKV_`SVUSL\[OHYKPP *VYKHP[HJLHL +HKV_`SVU
30:;6-:7,*0,:
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30:;6-:7,*0,:
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