Kobe University Repository : Kernel. Title. Author(s). Citation. Model analysis of the importance of reiteration for branch longevity in Pseudotsuga menziesii ...
Kobe
University Repository : Kernel
Title
Model analysis of the importance of reiteration for branch longevity in Pseudotsuga menziesii compared with Abies grandis
Author(s)
Ishii, Hiroaki / Kennedy, Maureen C. / Ford, E. David
Citation
Canadian Journal of Botany, 82(7): 892-909
Issue date
2004-07
Resource Type
Journal Article / 学術雑誌論文
Resource Version
publisher
URL
http://www.lib.kobe-u.ac.jp/handle_kernel/90000776
Create Date: 2012-01-06
897
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Can.｣.巨ot . Vol , 82, 2004
an dvi s u a l l yl e S e mbJ et hech a r a c t e r i s t i cma i na x i swe nz i e . u p i .Do t he s ec ons t i t ut e ha v ede s c r i be df orP.me SCUsa swe H? ( 2)Wha ta r et hea dv a nt a ge sofr e pe t i ( i onsofa r c hl t e C t u r a l s t r uc t ur et hr ou gh outabr a n c hi ngs ys t e m?T s hi ia ndFor d n( 2 00J )s t r es st ha tp r o] e pt i cr e i t e r a t i ona l l owsP me z i e L n ' it o ma i nt a i nf ol i a get h r ou ghouti t sc r own wh e n 】 i mi t a t i onsofc r ownw i dt hpr e v e ntmu c hf ol i a g epr oduc t i on a tt h et e r mi na le ndsorbr a nc hes( I s hi ia nd Wi l s on2001 ) .Ho we f ' f e ct i v ei spr ol e pt j cr el l e r a t l Oni n c ompe ns a t l ngf ors L L C hs i z el i mi t a t i ons ? To' l nS We rt hef l r S tque s t i on.t hede s c r l Pt l OnOrt h eSCU pr ov i de dbyl s hi ia nd For d( 20 01 )wa sus e dt oor ga ni 7 J C f ol i a gei r 7S i mul a t e dbl a nC he sofbot hA.gra1 7 di . va T l dP.me n z t ' e s i i .I nt hi spa pe r ,SCUsa r ec ons i de r e d or ga ni z a t i ona l u ni t si ft he i rr e i t e r a t i v epr o er p t i es( a c t i ng a saf l r S トor de r ma i na xi sa n dt he r e bybe i ngc or ) s i d er e dphys i o) ogl C a l 】 yl ndep e nde nt )C anb ea ppl i e dt oa l ls u c hgr oupi ngst hr oug hu山 t heb r a nc h,r e ga r dl e s sofl oca t i on.Toa ns we l 't hes e c ond andi sa nd que s t i on,weus et h ec ol nPa r i s onb e t we e nA.gr P .y ne nz i e s i i .Wes u r mi s et ha tA.gr a ndi si sa bl et omai nt a i n i t sl i v ec r ownt hr o ughac on c e nt r a t i onoft i g ht l ypa c k e df ol i a gel oc a t e ddi s t a l l yonb r a n c h e sa n dah l g he rbr a nc hde ns i t yt hr oughi t sc r o wn( nL l mb e r( ) どbr a nc he spe rme t r eh e i g ht gr owt h;H.l s hi ia n dD. C.Sha w,unpu b7 i s he dda t a ) ,whe l ' e a S P ･me nz i e L m ' iha samol . es pr a wl i n gb r a nc h i ngs t r u c t ur e,Wi t h F ol i a gema i n t a i n e dpr oxi ma l l ya l o ngt heb r a nc hi nga xi s( s e e l s hi ia n dWi l s on2001f orbr a nc hi ngde ns i t yofP.me nz i e s L ' i ) . Toe x pl or ebot hqu e s t i ons ,W ede v e l ope da nda s s es s e da c omput e rs l mL l l a t i onmode lofbr a nc hgr owt h.I nac omput e r s i mu一 a t i onwec a ne x pl or et h ee f f e c tofa d j us L l ngpa r al ne t e r S ofgr owt h,ma nl Pul a L i onst ha ta r edi f f i c ul to ri mpos s i bl ei n t her l el d.Wi t hc omput e rs i mul a t i onswcc a na l s oobs e r v et he d e v e l opmentoft h e br a nc hs t r uc t u r et hr o ugh t he e nt i r e c our s eofde v e l o pme nt ,r a t he rt h a ni nf e r r l ngde v e l opme nt f r om ac r os s s e c t i ona ls t udyo fbr a nchs t uc r t u r e sOfdi f f e r e nt a ge s( f ore xa mpl esofpl a ntr node l ss e eHonda1 971 ;Hon da a n d Fi s he r1 978;Fr a n c o1 98 6;Sut he r l a nd a l l d St i l l ma n 1 988;For de La l ,1 990;Sor r e ns onCot he r ne ta l .1 993) .We de v e 一 o pe das t o cha s t i cmodelt ha ta c c ommoda t e sr e i t e r a t i on, whe r er e i t e r a t i onoc cu r sone xi s t l ngb r a nc hi ngs t r u c t u r e s . Wedono ts i mul a t et her e i t e r a t i onofne wl e a de r sorne w br a n c h e spr oduc e dont hema i nt r ul l k. Th emode li sde s i gn e d t os i muL a t egr o wt hf or t hepot e nt i a ll i f e s pa noft l l ebr a nc h f orb ot hP .me nz i e . 如iandA.gr ar l dL . T . -
Met hods Dat ac ol l e c t i onandanal ys i s !me nz i e s ' iw. e r eobt a i n e df r om l s hi i( 2000)a nd Da t af orF u s e dt oge ne r a t er e gr es s i onr e l a t i ons hi psf o rbot hf ol l a g e ma s sa nda r e apr e di c t e db yf o l i a gea gea ndor de r .Ne wda t a we r ega t h e r e df orA gr andT ' S( Ke nne dy2 002)i nt hes a me ma nne ra sl s hi i( 2 000) ,Wi t ht hee x c e pt i ont ha tor l l yt hr e et oL a lbr a n c he swe l ･ ehal ･ ヽ e S t e da ndme a s ur e dbe c a us eofl i mi t a t i on st OS a mpl i ngi nt h eFor es tSe r vi c eol dgr owt hNa t u l て l l Res e a r c hAr e a :Onebr a nc hwasha r v e s t edf r om t h el owe r c r own,onef r om t heml ddl ec r own,a ndハn ef r om t heuppe r c r o wnoft hr e edl f f e r e ntt r e e s .The s eda t awe r eu s e dt opr oandi st o pr e di c t du c er e gr e s s i on r e l a t i ons hi ps f orA.gr f ol i a gema s sと I n da r ea .Addi t i ol l a lnon de s t r u ct i v eme a s ur e -
me nt sofa r c hi t e c t u r a lbr a nc hi ngc ha t ' a c t e r i s t i c s ,i n cl u di ng l a t e r albr a nc hi nga ngl ea ndt her a t i oofl eng t hsb e t we enp a r e nta ndl a t e r a】s hoot sf o rb ot hP.mL y nZ . i L , . T t ' T -a n dA.gl ･ an di s , we r emZ l dca HheWi ndRi ve rCan opyCr a n eRe s e i l r C hFL I Ci ] i t y( WRCCRF) . Thegr owt hofyounge pi c ol Ⅶi cs hoots t r u c t u r e swa sa ] S ome a s u r e d:t h el e ngt ha nda geofdL l u ght c r s hoot soft h ei ni t 】 a t e de pI C Or ml Ca r L dt hen umbc l 'orda ught e r s hoot sp ernodewa sr e c or de df orf i v ecl us t e r sone a c hl n C と 卜 s ur e dbr a nc h.Ma psoft hes enewl yr or l 川1 gC l us t e r sr cv e 乙 l 】 r s ,t h e n) n c r ca s l nggr owt h l i mi t e dg r owt hf ort h ef i r s t2ye a t h e r e a f t e rbot hi nt hebi f ur c a t i onort h ee pI C Or mi cと L Xi si l nd i nt heav e r a gel e ngt hofs hoot s( Fi g.2) ･ ConS t r u( : l i onl I rt hes i mul at i onmode l
T e r t l L S . ･S hoo tor de t ･andge ne t ･ aL i on Shool sa r eor de r e da c c or di ngt ot hebot i l ni c alol ` de nng s ys t em ( Bor c he r ta ndSl a °e】 981 ) ,whi c hl l S S umC Sf un c =01 1 山 r el a t i ons hi psa mong nes hoo t si nt h eb r と I nC hi ng s ys t ems ( Uyl i ngse ta l .J 975) .Shoot st h a tc oI T l edi r e c t l yf r om t h e t r unka r eor de r e done;l a l e l ' a ]s l l OOt Sa r ea S S l gne da nor de J ' oT l egl ea t e rt ha nt he i rpa r e nt ,a nds hoot st hatr e pr e s e nte x t e ns i onoft hepa r enta xi sa r et hes a meor de ri t st hepa r e nt ( Fi g･1 ) ISe quent i a lg r owt hi sgr o wt hwi t hi nt h eus ua lt = ni n . g oft het r e e( a f t e ras i ngl ew i nt eT ' ordor r ml n C y) ,a ndpl . Ot e P l l C mi L n tF ormor e gr owt h] ･ qgr o wt hf r om ab udt ha tha sbe e nL I ur t ha nonewi nt e r .Si mi l a r l y ,s e que l 1 i a lr e i t e l a t i oni st h edc di f f e r e nt i a t i onofal a t e r l L la ext p of un c t l Ona Si lf i t S トor dc l . ma i na x i swi t hi nt h eus uaLt i ml ngOl ､gr o wt h,a n dpr oL c pt l e r e i t e r a t i oni st her e l e as eF r om dor ma n c yolと I ne Pl e Or l l l i c s h oot ,whi cht he nl ･ e i t e r a t e st h e. q t l ･ u C t ur eOl ､t hemi t i ni l Xi . q ( Fi g.J ) ･Al le pi c or r ni cs hoot si l 】 ` Ca s s i gne dl oor de rl( ゝ u t ha tt he yr e i t e r a t et hes t uc r t u r eorL hebr a n c hmai l la xi s ) Ia n d t hema ina xi sora n yi l l dep e r ) de ntSCU i sa s 凱l me dt ob e or de rL Ge ne r a t i onsa r ca s s i gne dt odi s t i ngui s h( h egr owt ho( 1 e p1 c o r i cs m hoot sf r om r e gul a r) a t e r a 】s h oot s .Ther l r S = hoot s gr o wni nt hes i mul a t i ona r ea s s l gn e dl oge n e r a t i onon e∴l n d a l ls hoot spr odl l C e dt h r ot l ghs e que nt i a lgr owt ha r et h es i H T t C g e ne r a t i ona st hei rpa r e nt .Ane pl C O r mi cs pl . Outi sa s s i gn e d t oage ne r a L i oI 1Onegr e a t e rt ha nt ha tofi t spar e nt( Fi g･L. l s hi ia ndFol . d2001 ) . Mo de ls L r uc f E L r e Muc hoft hel a ngua gef o rt h et wodi men s i ona l1 1 1 OL l elr ' 01 1 l o †sFor da ndFor d( 1 990)a ndFor de ta l l(1 ' ) 9 0) I Th esin-u1 a t 1 0nmod ell ungOnaye a r l yt j mcs t e pと l ndp e r r or mSt h e f ol l o wl ngF u nc t i onse a c hye a r : ( 1 )Re gul a rs h ootg r owt h:Ne ws hoot sa r epr odu c e di l lt he t e r mi na len dofa c t i v en ode sa ndと I r ea S S 】 gne d1 ) Ⅰ ` C di c t e d . l e ngt hs ,f o一 i a gema s s e s ,之 I ndar eas ( 2)Epi c or ml ci ni t i a t i on:Th eJ T I Odc]t e 舶 wl l C l h erpL . e Vi ous l ys u ppr e s s e dbudsi l l ' edu eL ober el eと I S C df ) ･ om i nhi bi t i on. ( 3) SCU i nde pe ndc n c c :I fac oHe c t i onofs h ooL ss a t i s r i c L qE h c c r i t e r i ont obe c omean e wL yi nde ende p ntSCU,l h e t Hh之 l t c o】 L e c t i onofs h oot si sc ons i de r e dt of or n lan e w SCU di s t i nc tf r om t hepar e nt' l Xi s . Mo de lpar ame t e r s Th es i xmo dclpa I ･ a me t e r sa r eV al u e si r npor t L I ntL ' Orgr owt h t ha ta r eus e di nt hemode]b utwe t . enotde t enl l l n e dt hl Ough の2 0 ( ) 4NRCCi l n i J d i l
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r ea s s i gned t hr e e da ught e rs hoot s .Thi s gr e at e rt h an 3 a cha ngest hepr oba bi 一 i t yordr a wul gat hr e ea sf ol l ows :
r 2l
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wher et heme a noft hedi s t r i but i oni snol onge re xa ct l yt he九 pa r a me t e r .Fore xa mpl e,i ft heval uef orav e r a gebi f ur c a t i on i s3. 0,t heac t ua ]meannumberofda ught e rs h oot sw il lbe 2. 328.An ybi f ur ca t i onse xpl or e dmus tt he r e f or ebec ons i d e r e di nt hecont e xtoft hi sr educ e de xpe c t e dva l ue.The r ear e uni que P a r a met er sf ort hea ve r a gebi f ur c at i on ofor de r1 ( r ba) ,or de r2( r bb) ,or de r3( r bc)andne we pi c or i cs m huot s f ort hef i r s t2ye ar s( e pi r ba) ,al l ddef a ul tva l ue sf ort hepa r a me t e r st ha twe r ede t e r r ni nc dt hr ough i ni t i aLs i muJ at i ons ( Ta bJ e] ) . l ti st hege ne r alca s ef ort hes es pe cl eSt ha ta ver a gebi f ur c at i on dec r e as esw it hi ncr e as l ng ( ) r der .T fbi f ur ca t i onsof l q h oot sor de r e dgr ea t e rt ha n3a r es i mpL ygl Vent heor de r3 par ame t e rv a l ue,t he r ear eunr e a l i s t i cal l yhi h vat g u esoft ot a l s hoot sa tye a r9 0f orA gr andi ss i r nul a t i ons ,wi t hs i gni f i cant ･Topr e ventt her e qui r ementf orauni 9ue v ar i a bi l i t y( Fi g･3a) par amet e rf oral lpos s i bl es hootor der sgr ea t e rt ha n3as t m-
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Thi sr e duc t i ons ucce s s f ul l yr educ e dt hemnge壬 I ndv i l r i abi l i t yoft ot als i mul a t eds hoot sa tyear9 0f orA,gr l "7 ` I L L y ( Fi g･3b) . I ni t i als i mt l ht i onsa l s o demons t r a t e ds t r ongcumuht i v e ef f ec t sofepI C Or mi cgr owt hont heout comeofs i mul L t ( i ons . ForP.menzi eL u' L 'r na ny i ni t i a 一s i mt l ] at ed br i l nChcsdi d not s ur iveunt v i lt hemi ni mum a geobs e r v e dbyJ s hi iと I ndFor d ( 2001 ;Tabl e4) ,a ndf ort hes i mul at e d br anc hest ha tp er s i s t e dt heout col neSWe r ehi hl g y val ' 1 a bl e (Fl gゝ ･4( I ,4b). Somes i mul a t e dbr a nchesha dmor et hant hr e el i mest het ot al f ol j a t eds hoot st ha twer eobs e r vedbyI s hi iandFor d( 2OOl ) ･ Ourgoa lwast oobs er v el ongt er m b1 a nC hdevel opment ,と I nd i Ls e el l l Sne ces s a r yt Oenf or ceami ni mum i l mOuntOfgr owt h t opr e ventt hepr ema t ur edea t hofas i mul a t e dbr anc h,The modelwast he r e by modi f i e dt o al l ocと I t edet e r i ni m s t i c al l y t hr eedaught e rs hoot sea chyea rt ot het e l . mi nalnodeort he r e gul a rma i nz I Xi sf ort h ef i t ' s t50year sofs e quent i と l lgr owt h ( r a ndma H ､ =50).ThereaftertheregulとIrmとIillaxisbecol l l C L q . s ub j e i ; 上土 ol l l el l B J l dQ_lI HPQi s _ 5 9nPr OC eS S ･ r ni t i a ls l mul at i onsal s os howe dt hatr e i t er a t i onha ds i gni f i ca ntcumu一 a t i vee f f ec t sont T l enumberOrr Ol i a t e ds ho( ) t s( ) n ￠ )2( ) ( ) 4NR Cei l n i l d i l
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Fi g･4･Pl ( ) t soft ot alr ｡】 】 a l e ds h oot =) v c rl i mef L ｡rps e u d l J I ･ u gaM , , n z l e ' V l l ,Wh e r ee i l Chc l l r V et ･ e Pr e S e nt SaS i ngl er L l n.( a)The r ei sa r lunr c a 日s l i c i 1 日yhi ghn umbe rofs hoot si nt h eml l j E l lmodcl ,wl b outt l l l yr e dl l C t l ( ) Al l -e PI C ( ) r mi ci nl t i a t i onwi t hl l l C r e a S l ngge ne r a t i on,wl t h 50r unsa Hh cs a mepa l ､ i i n l e t e rV al ue s (/ , 日. bl l a t e ds hoot sI ors i xS e e dv a l ue sc hos e nf ordc t a i l edと t r l a l ys i s ･( C )Re gul a rs hoot sf ort h e mXs e e dva l l l e H how s l mi l a rgr owt hov e rt l mel( 4)Di ve r ge n c eofgr owt hこ I mOngt hes i xs e e dv a l u e si se vi de nti nt hec our s eofe pi e nr mk S hoot sov e rt i t t l e,i l di ca t l n gt ha tdi f f e r e nt i a ls t l CC C S SOfl ･ e i t e r a l i one a J ･ l yl ngr owt he xpl ai nt qt heva r i a bi 】 i t yobs e r v e di nbr a nc h de v e l 叩1 1 1 Cnla n dl h cH i gni nCa nteu1 1 1 u] at i v ee f f e c t sofr e i t er a t i on･( a)A d e l e r n l nl S t i cma l na x i sr e s ul t si npos i t i v ea ndva r i a bl egr owt h r t ) r501 u1 S. ffJA r e duc t l Ol li ngr o wt hwi t hi nc r e a s 】 nge pi c or mi cge n e r a t i on( e q.4)r e s t r l C【 st heuppe rr a ngeoft ot a ls hoot sf ( ) r5( ) r u ns . C J t . J
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Tl r ni ngo fEpI C Or ml CPr oduCtl0n e r at i onoff oura ndabov e.Thi sr e duct i onOnl yappHe st o Rh. H . ,t h a tl S ,t Ot hef l r S t2yea r sofgr owt hF oranew e pL c or i C,s m uc ht hati ft hei ni t i a t e ds hoots uc ces s f ul l yel ongat e s s ubs e quer l tgr owt hpr oce e dsr egar d] es sofgener a t i on.Thi s r e duc t i onr es t r i c t L St hepr oba bi l i t yt ha tanewepI cor i cs m hoot s ucc es s f ul l y de vel opsi nt oa nSCU f orhi ghergene r a t i ons a ndr e s t r i c t sba s e l i nemode lout c ome scl os e rt ot her i l nge obs er ved( Fl g･4f 二Ta b】 e4) .
Rul e sf ore pi c or t l L i ci ni t i at i on T s hi i( 2000)r e c or de dL t l eL i ni ngofepi col mi ci ni t i a t i on, t ha ti s ,t he numb erofye a r sofdor r na nc yb ef or e a bud s pr out e di nt oanep] cor mi cs hoot ,∧na l ys I SOfhi sda t as hows t ha tt het i mi ngofe pi c or mi ci ni t i a t i on( Fi g.5)pe aksa r ound 5yea r saf t ert hebudi sf or me dandt he ndecl l neS .Thea s y7 1 1 met r i cs hapeoft hi sdi s t r l but i onr es embl e st hega mmapr oba bi L i t ydi s t r i but i on( Ca s eJ l aandt i e r ge r1 990) ,whi chi st h e s ul nOfe xpol l el t i aldi s t r i b ut i ol S .Thee xponent i a ldi s t r i but i oni SaWa i t J ngt i medi s t r i but i on,a ndt hi sc anr eas ona bl y bei nt e r pr e t e dast hewa it l ngt i met oepI COr i m ci ni t i at i on. Thi sdi s t l . i but i ona samodi f i c a t i onoft hee xpone nt i alma y r e f l ec tt ha ti li snoti ne vt f abl et ha tadonl l a ntbudL q Pr Out St O f or ma ne pI C Or i cs m hoot .Fort hes i mu) at i onsweas s umet ha t t hega mmadenS i L yl Sal la ppr opr i a t eS t a t i s t i call 1 1 0delf or t i ml ngOfc pI C Or n l i ci ni t i a t i oni nl hn e nz i e s i i ･s A s uch,t he a gea twhi cht ha ts hootma ys pr outa ncpI C Or mi ci sgener a t e df r om t hega mmadi s t r i but i on F ore a ch ne wl y gr own f i r s t 1 0r ders hoot( Pr es seta l 一1 992) ･Fore xa mpl e,i fapa r ent s hootdr a wst henumbe rf i ve,t he nt hes hooti st es t e df or e pJ c or mi ci ni t i a t i on du r i ng t he t i me s t ep a twhi c hi ti . 3 eCh os ent ha tqual i t a t i v eI ypr o5yea r sol d･P, d r a met el SW er
duce daga n l mi lPr obabi l i t ydl S t r i but i ont ha tn l と I t C hc dt heobs er ve d va l uesa l l d ha d as i mi l a rme i l n( α = 8,β= O. 7. Fi g.5) .Al t hought he s epar amet e l sdol l Ot1 こ l V ebi ol ogi cと I l i nt e r pr e t a t i ons ,t he ga mma di s t r i but i on der i ne d by t hem cl os el yr es e n l bl est heobs er ve ddi s t l ･ i but i on( Fi g･5) . Oncet hes hootr ea c he st hea get ha twと I mr a Wnfori tr r ol 1 t hegammadi s t r i but i on,andi ft hes hoots at l S r i e st hecl う t e r i onf ore pl C Or mi ci ni t i a t i onL I nderwhi cht hemodeli sI unni ng( s eebel ow) ,t hena nepI COr ml CS hootl SPr ( I ( 1 uc c d･l r not ,t h ebudi sa bor t e d.Epi c or mi csa r es i mul a t e dt o叩L Out onl yone xi s t l ngbr anc hi ngs t r uc t ur es .Th emca t y ur e L lr 山 co r e pI COr T T ) i ci l l i t i at i ol 1i l lA gr andi . t lWast ooi nf r e que ntl obc s i il m a r ) ya na l y7 , ed,a ndc pl COr mi csal ･ Cas s umedt o hi l VCi l ne gl i ibl g eef f e c tonA.gr undi ゞgr owt h.I tl mt S tbcnoL c dher e t ha tt heda t af ort het i ml ngOfe pI C Or mi ci ni t i L l t i onwer e紳 l l er e df ore pl C Oml i cs hoot st h; l Iha di t l r eヱ I dy s pr out e di l nd e xc 一 ude da ny nodest ha tdi d nots uc c cs げuL l ys pl outepl e.t hepr oba bi l i t yofs pr out i ngと I ti lPと H . l i cL卜 c or i cs m .Ther ef or 1 a ryea ri sl l kel yov er es t i ma t e d,whi chi scot 1 e n. p q i l t C dF ori 一 l t heal t e r n' l t i vcr ul esf orepI C Or mi ci ni t i a t i on. Thes er uL e swe r ede v el oped ul l dert heas s ul l 叩l I On t hi l l a pI Ca ldomi na nc eopcr a t c st oe s t a bl i s hi lC7 eu Imai ni l Xi -) l l br a nche si n P me nz i e s i F ' .bu ts omemech ani s l l l OE ■r el eas e l le p] r r onla Pi ca ldom i nanc ei sr e qui r e di nt hepl ` OC eS S( ) O e nI l bl cr ei L e r ; l t i on.T li s1 . C と t Wl i l hl cl hL l li l C or mi cs pr 01 1 1 ] ngt T a t er al l ygr owl ng axi spr oduci ng 乙 I uXi n mi l yC Ont J l i buL cl o ma i nt a i ni ngdor manc yofepI C01 ° i cbuds m ,whi cL ICi L nbcr ｡ L eas e dwl l e l lt l l eJ a t cr ala pexi sL os t .Thi si deai se xpl or e di n t h et h r c ca lt c r na t i v cr ul esf ol .e pi c or l l l i ci ni l i i l t i on( Ti t bl el ) : ( 1 )Si mpl ei ni t i と I t i or . 1 :Ul l dert l i sr ul et l l er ei l r enOl ' ur l h c川 ､ s t r i ct i onsone pl COr i cgr m owt handas hootpr oduc esi l n の2 ( ) 0 4NRCC乙 川I l l 1 i l
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c pI C Or mi cwheni t1 e乙 I Che si t sdes i gna t e dt i ml ng.Thi s i l mumeL qS OmCI l l C Ch･ ' Mi s I T l( ) t he rt hi l nl と I t e t ' als uppl ' eS S i ol l orL h c(lor mI l ntbud. ( 2)Onel ml C l l V eS i de'A pos s i bl et he oJ ･ yf ort heI , e l e a s eof donm ntbu dsi st her equl r en l enHha ta tl ea s toneoft he l L I L cr i 1 1･ L I X CSt hL l li ss ubt ende dr r OI nt hes amenodeast he s uppr cI S S Cdbudi H1 Ol ol l gergL ' OWl ng.l fbot ht l l el a t e【 一 a l s hool si l l ' CS L i 1 1と t C t i v catt hede s i gna t edt i me,t he nL hc I L L t Cl . 1 1 1乙 I XC Sこ I r Ca S S uI T I C dt obes uppl e s s l ngt hedo r ma nt bud.whi c hi . qt hcnと I bor t e d. ( _ n TwoL nと I Ct l VeS i de s :I l 一t hl SC as eb ot hoft h el a t e r と I Js ubt en dc di l X CSn l uS tnOl ongerbeg1 0Wl ngJ nOr de rf ort he budt o叩I ' OUti l ne pl C Ol ● mi c,gl VmgaS t r i ct e rf or m ofa pl Cと l ldomi ni t nC eOrt heL at e r aJs hoot sont hes upp r es s ed bu d,t h王 I ti ト ,i roncaxi si s. S t i l lgr owi ngt hedor l m ntbt l d r cl ml i n" uppr es s e d. Undcrと I L lt hL ､ C Cr L l l es ,c pI C Or mi css l r outOnl yonor de(1 I d XCHl ndこ l t t crt hefi r s t2yea r so1 -gr owt ht hee pl C Or i caxi m s i sgl V Cn t h es i H l l eと t V el a gebi f ur c i l t i on L I SOt he rI . i r s t or de r s hoot H .
Rul e sI( J rde J i l u' l l gSCt iL ' nde pe nde nc e EdChyC と I rl hじ1 T l ｡L l els cヱII . Che ヽl ' urg r oupsoL 'r ol kI geL hi l L L l r Cr C C Ogni / . c da uc wl yi nde e nde p ntSCUs ,a ndea c hs h oot i nt h egr oupL St he ni l S S l gned乙 I nSCU numbel ･ .Twoc r i t er i a t odi ㌔ t i ngui s hi l l dep e nde ntSCUsc a nbedi s c er ne df r om t he l hc or yorls hl H1 ndトor d( 2( ) Ol ) : (] )( l ont i gL I l t yl l ul e:Al lSeU l l qC OnS i der edl ndep el l de ntof l hcpi l r enLuni ti ri lh a st l or meda… pe t i ol e l i k es e c t i on s epと I r L l t l ng ] l L SS hoot ss pa t i al l yr l ' Om t he pa r e ntSCU" ( I , l hi i2( ) ( ) ( ) ) .Un dct 't hi H∫ ul e,al と I t e1 ala xi sore pi col ● mi c i sitH S uI T t C di ndc pcndc ntoft hema i l laxi swhenf ol i i l gC S乙 I li t L qba s e,r c s ul いng] nl a ckorcont l gul n l ( ) r l 仙t yoc cuJ. l t yOHol i ut cds hoot swi t ht h ema i na xi s .Thes hoot son t h cm; l i n之 l XI SOH hene w SCU a l Cgl VenOT . de 1 -I ;t he r c1 1 1 乙 I i nl ngS hoot si l ] ' e1 C Ot ` dcr eda c c or di ng一 y.Thi sr u1 e r cl i c H㌔ ( ) l c】 yoT ll hcmol Phol ogl Ci t l l ydi nl n CLc ha r a c t e r i s 1 i corL hcobs c r vc dSCUs . ( 2)Epi c o] 一 mi cr ul c･OJ l l yepi c or mi cs hoot sf or mt h eba s i sof i lne w SCU,whi c l lJ Sと t S S ume dt obeI ndep e nden ti fi ti s okI cHhi L n1 ( )yc i 1 1 1 < ･Thi sl 〕 一 一 CS Cr V C㌧t hec ont l g1 1 1 t yC一 一 i l e r i ( ) n rot .epi cor mi cs hoot sdes cr i beda bov e,butl a t er al 血) oI L ll t r CnCVC rC OnS i dcl ● C dL O bei nde pendentSCUs , b cc i l u㍉ Ct l l C ydonotbi i ul . C 礼 t ei l Sf i l . S t Or de ra x es .Thi s gl V eSi lS L L . OngerPhys i ol ogl Ca lba s i sl ol 'i ndepe ndenc e･
Ab i e sgr a n dE ' S Pa r e n tor d e r ,p a r c nr l c n gt h,C r o wnp t ) S i t l O n Un i t l e ss P a r e L l tOr d e r .c r o wn Pa r e l l lo l d e r .c l l 0 Wn ponL i on p o L q i t i o n c ( ) o li l g C ,SCUr l ( S i l i on, m2 / g sh o o -a g e ,SOU I 0日=( n, Sh e r o wnpos i t i on c r o wnp o t q i t i on Sh o o ta g e ,c r o wn g / C m Sh oo ta g e ,c l ･ 州′ n po s j t i on p o t q i l i o n u1 1 Non e NOl l e r a d
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Mode lc uns t anl s Re gr es s i onsobt a i ne dL hr uughda L aと I ni 1 1 yHswer eus e di n t hemode 一t o det e mt i nef ol i a gea mount sf ol ･as hootofa gi ve na gea ndl oca t i onf orbot hs pe ci es( Ta bl e2) ･Whe nt he s t i L t l S t i ca lr el a t i ons hi ppr e di c t e dl es st hanz . e r of ol i a gema s s anda t ･ e之 1 ,0日 ft hes hootwa sol det .t ha nt hema xi l nul na ge obs e r ved (L O ye a r s ) ,t hes hootwa sc ons i der e d dc f o] i a t e d. Re gr e s d on e quと l t i onswer eul s oge ne r a t e df ora r c hi t e ct ur al pa r a me t ers ,i n cl udi ng br anc hi ng a l l gl e,a n dt he 1 at i o of l engt hsbet we enpal 一 ental l ddaught e rs hoot so土di f T er entol 一 一 de r( Ta bl e2) .Fol ･S i mpl i ci t y,as hoott ha ti st hee xt ens i onof t hepa r enti sassume dt oha vet hes a mel e ngt ha lt hepa r e nt ( i . e. ,cons t a ntgr ow【 hr a t ea 一 ong' l naxi sofpa r t i cul a ror de r ) . Se ns F ' t i v i y anal t ys i s Forea c hs eci p e sal oc a Js e ns i t i vi t yana l ys i s( Campol ongo e tal .2OOO)Wと I Sp er f or medf orva l uesofa ve r と I gebi f uL ' Ca t i ons i nc on j unct i onwi t hf a c t ors c r e enl ngOft hedi f f e r e ntr ul esf or i nde pe nde nc eofSC s( t wor ul es )i l nde pi c oL . mi ci l l i t i a t i on ( t hr eer ul es ) .l nt hi smodelt het wof a ct or s( SCU i nde pe nd e nc el l . 1 l esと I nde pi cor mi ci ni t l a t i onr l l e g )ha ves i xdl S t i nct c omb i na t i ons( Tabl e 3) ,a nd t he L oca ls el l S i t i vi t ya nal ys i s wa sI ◆ epe a【 edf ore a choft hes i xr ul ec ombi na t i ons( s e ebe l ow) .ForA.g/ . andi st l let hleedi f f e r entr ul e sf ore pi c or mi c i ni t i a t i on wer enote xpl or e d be ci l uS ee PI COr ic m swe r eas s ume dt omak eane gl i gi bl ec o一 t r i b ut i onl ogl . OWt h･ Fors ens i t i l ′ l t yanal ys i s80uni quePa r a me t e r i 7 ･ a t i onswe r e gene l a t e df ore之 I Chr ul ec ombi n壬 I t i on.Ea chc ont i nuouspa r . l me t e l .Wa sS a mpl e d201 】 mesf r om < ' Il 1 ni f or m di s r r l bl 山or l , wi t ht heot her ske pta tt hei rdef a ul lva一 ue s .ForP me nz i e s i i L hepu r 之 I meL e r swe r es a mp] e d on t her a nge( 0, 3) ,a ndf or A･gr a' l dL st hepa r amet er s . wer es t wT I Pl e dont he] ･ a nge( 0, 4) I The r ei l r eS t r ol l sS t OC ha s t l Cef f ec t si nt he! T I Odel( Fl g･ 4 f) , al l dL Ot r yt OC Ont r OH ort he mi nt hes ens L t i vL t yal l a l ys I Sa s l ngl emndon lS e e dwnく1 ] S e dI Orea chpn r ･ l me l ers e l lTwo r unsWi t ht hes ar r ) epa r a l ne t e rVal ueswoul dt he 1 ef ol ■ er e s ul t i ni dent i c a lbr a nc he s .A r a ndom s e e di nt egert ha tYi el deda c el l t r alv a l uei n乙 tPOpul と I t i oI IOf50modelout put swi t sChos enf orus et hr oughoutt hes e ns i t i vi t yana l ys i s( s ee d=21a nd
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Tab一 e3.Rul ec 川r l bi n a l i on su 六 e df o rs e J I S i t L Vl t ya na l y s e s . Rul ec o1 1 1 bi n a t i on
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SCUsd e v e l o p( ) nb ot hl a t e l a la l l de plC O川1 i ca x e s ;e pE C ( l ln l i c ss pr o u tOnl yi fon eI a t c r . l la xi si si n a c t i v e 2 SCUsd e v e l o ponb o t hl a t e r a la n de pI C O r ml Ca X e S lI l ol i mi t a t i onone pI C Or r ni ei 一 一 i t i a t J On 3 xe -r ci n a c l i v L ･ Sn7 sd e v e l o p｡r lb o t h1 ( l t e r a la n de pl C Or ica m x e s ;e pl C Or mi c ss p r o u to nl yb o t ho ft h el i l l e r a la 4 SCUsd e v e J t ) ps ol e l yone pI C nr r r t l Ci l Xe S ;n Ol l it m a l i o none pl C O 1 mi ch l l T l a l i on 5( d e f a ul lc ombi n a t i onJ SCu d e v e l ops ol e l yone pJ e Or mi ca x e s ;e p I C O r mI C S叩r Ou tOnl yi ton eh t e r a la xt si sl n a C l i v c 6 SCUsd ev e l o ps o l e l yone pl C C ･ r ica m x e s ;e PI C O r l l li c ss pr outonl yi rh ol holt h el a t e r a la x e Mr L I Jl t l i I L lT V e (二 U,s h o o tc l u ト I e ru 1 1 1 ( . No t e :S
n z l e ･ uia ndA.gr a ndi s .r e s ec p t i v el y) ･Ana l ys e s 1 7F orP･me wer er e pe a t e df orea c hs eci p es ,a ndt os a vec omput at i on L i met hemode lwa sr u･ -f c ) r90yea r sf orbot hs p e ci es( Y90) ･ Thi syea rwaschos ena sac ent r a lbr anc ha geobs er v edi n P.me nz F ' e . u ' L ' ,a l t houghnoA. gr a ndL ' sbr anc heswe r eobs er v e d a tt ha ta ge.Tot lf a ol i a t e ds hoot sa tY90wa spl ot t e daga l nS t t hes a mpl edbi f ur c E l t i onr ahOt OI nf ert hee f f ec tofi nc r ea s l ng bi f ur ca t i ona te a chbr a nchor de rons hootgr owt h. Anal ys i so fbr anc hL on 号e v i t y
] naddi t i ont os e ns i t i vl t ya na l ys i s ,mode lbeha vi orwa sobs er v e df orupt o400ye a r sa tt hede l ' { l ul tpa r ame t e rv E ll uesf or ea chs pe ci es .I ft hebr anc ht e r mi na t edbef or e400yea r . S ,t he s i mul a t i one nde d,a ndt h ecor r es pondi ngy ea rwと L Sr e c or de d a st het e r i na m la geoft hebr anc h.Thet ot a H. ol i at e ds hoot s ov ert l mCWa sObs e r ve df orbr a nc hest ha twer egene r a t e d f r om f i v es i mul a t i onsa tder au】 tpa r a me t erval uesf ore a c h s pe ci est oa s s es sva r i a bi l i t yi nt he l ongt er m pa t t e r n of gr owt h･I na ddi t l On,aP OPul a t i onof50br ancheswaspr oes ,a ndf r e quenc yhi s t ogr a msoft e r n l ト duc edf わrbot hs pe c i :me n z i e . u ' E t het ot a l na la geoft hebr anc hwe l ePl ot t e d.Fori SCUsov e rt i meandt henumberofs c qucnt i a landcpi cor mi c s hoot sp erSCU ove rt i mewe r epl ot t edf ( ) ras l ngl es i mul a t i on.Thede v el opmentofas l ngl eSCU i nPme n z i e s i L -was obs er ve dt hr oughf r e quenc yhi s t ogr am. qof -s hoot sofdi f f ere nta ge cl a s s esovert i me. For F !me n z i e s i it he e pJ C Or i c gene m r a t i on a t whi c h l ' educ e de pI C Or mi cgr owt h wa sa pp一 i ed i nt he s i mul a t 】 on ( genr e d ･e q･4)wase xpl or c dt hr ougha nal ys i ヲof50s i mul a t i onst ha twe r epr o duc e df ore a c hge n. e dr a ngl ngf r om onet o s e ven,Wi t ha l lot he rpar a me t e r sa tt he i rde f a ul tva l ue s .The t e r i n' m llye; 1 TOft hebr a nc handt hen l aXi mum numberof f ol i a t e ds hoot sobs er v edf ort hebr a nchwe r epl ot t edaga i ns t ) y ,V al ue sr a ngl ngf l . om ]t ol oo,i nde x edby ge n. i . A .Si r ni l と I r 1 0,ofr a ndn l i . i n( t heyea ra twhi c ht hema i n. a xi sbc ca mcs ubj ectt Ot hemndom Poi s s onpr oces s )wer el nVeS t l ga t e d. -
Resul t s Mo de )anal ys i s
Se ns i t i v i t yanal y s i s :Ps e l l dot s ugamenz i e s j ir ul e c o mbl l nat L ' o r l S1 -3 Al l owl ngSCUst ode vel oponbot he pI C Or mi ca n dl at e r al a xes( RCl3;Ta bl e3)hasapr of ounde f f e c tonmodelOUL put .Ev e na tt h ede f aul tbi f ur c a t i onst het ot aJf ol i a t e ds hoot s a tY90i si mpos s l bL yhi gh,Ca us t l l gamemor yer r orOnt he c onl pu t er .Thi sr ul ef orSCUi nde pe nde nc epl a c est hec ur nul i L L i v ecJ r e cL sorr c i t c r at i uT IOI la l ll at er a ls hoot s ,ef f e ct i ve l y i nc r ea s l ngt hebi f ur c a t i onofs hoot sofeve ri nc r eas l ngOr der .
0 0 cc u) -Onk l l cI L I li L Xe S Thi sa l l ows. S e que nt i a lr ei t er a t i on1 hi sl cvelor t hr oughoutt l l ebr anc l l .Gi ve nt he. S i mul a t i ons,t ･ s e que nt i alr ei t e r a t i oni ncombi na t i onwi t hpr ol cpt ) cr ei l e r こ l n z i e s i i . t i onca nnotoc c uri nP.me
Ps e L l do暮S l l gamenZ l ' e s i ir ul ec o mbL nat i l ) n4 Rul ec ombi n a t i ( ) ∩4( RC4;Ti l bl e3)I . Cqui r e sSCUドl ode ve l oponl yone pI C Or i caxe m s .a ndcpI C Ol mi ｡] ni t i i t l i oni Hl Ot s ub j e c tt oal l yJ eH t r i ct i ol l Sbe y( ) 】 1 dt l l C乙 t geL I LS Pl ' Outdct e r 一 mi ne d by t hegammadi s l r i bl l t 1 0n.Undcl .t hes er ul esl hc maxi mum oft ot als hoot sa tY90wi t hi ncr e a s i ngbi r ur cこ I t i on i sgr ea t ert ha nf orRC5a ndRC6' l C r OS S州 or dcr いt n dl S E l gaI n一 a r ges twl t hi nc r ca s l ng Or de r3 bi f ur c a t 1 0ni l l l dt he oL ゝ a ve r a ge bi f ur ca t i on ofn ewl y･i ni l i i l l e d cpI COl t nlC 山 一 ( Fi g･6) ･UnderRC4t her eと I r enO. r C S けi c t i onsonc pi c or L l l i c i ni t i a t i on,hcnc ct hepr oce s sofr e l t er i l t i oni senh乙 I nC Cd;t he e f f e c t sofi nc r e a s esi l 一t hepa r amet e l .V i L L uesi l l ' Ci l mPL i l ' i cd･ be caus et he ya r ei ne f f ec ti ngr e a t el . nul l l be r sorSCUs乙 I CrO " t hebr a nch. Unde rRC4,t he r ei sani ncr ea s ei nt ot alSCUswl l hb( ) t h Lm or der1a ndi ni t i と I t e depi c or mi cbi f ul . Ca t i ons ,butnor Ck l l i s hi pwi t ht h ebi f ur ca t i onsofor der s2i l nd3( Fi gl71 .The ma gni t udeoft hecf T cc t sofi nc l ' ea s l ngOr de rIi L ndne w epl Co I Tni cav e r agebi f ur c at i oni sgr e at ert hと I nf oI 'RC5.Ac l ' OS S a 】 lbi f ur c at i onst henumberofSCUsi sgr ea t ert hant hel l Umb erofSCUsf or me dundel .RC5こ l n dRCh. Ps t ! udot s ugame I I X i e s i ir ul ec o mb L ' nat L ' on5 UnderRul ec t ) mbl l l at i on5( RC5;Ti l bl e3) .SCUL qこ t r Cil.1s ume dt ode v el ops ol cl yonc pI C Or mi cs hoot s ,notonl と I t C r i l l ax es .a nde pl COr mi ci ni t i a t i oni sl i it m e dbyt h er c qul r emet I ofonei na c t i ve] i l t er a la xi s ,l nge nel 一 之 I l ,t ot L l 11 ■ o] i a l e ds ho( 山 s how 之 InOnl i ne( l ri nc r e a S ew il hi nc r ea s l ngPこ l r i l n l e t erVa l t I C . ∼ ( Fi g･6) .Thema や¶um t ot a lC ol l a t e ds hoot s乙 t tY' ) 0i sgl ■ C i l ト nl r CL L t i onor es Hbri nc r e as e sl 【 1Or de13bi r ur c a t i ol lt l l l dt hebl ne wl yI ni t i at e dcpI C Or i cs m hoot s ,wher e a . H hcm' l Xl r mum for i nc r e a s esi nbi f ur c a t i onsofor der s]and2a r emuCl lnT l こ l l l c r ( Fi g.6) .t tL q houl dbcnot e dt ha tt os a v ec omput a t i onl i l nei r t hes i mL J L a t e dbr anc hde vel opedmor et ht l n200, ( 一 oof 01 1 こ I t e d s hoot s ,t hes i mul a t i onwast e r mi na t ed.andt ha tl ot と I Lwと I Sl ･ C nli ) l C or de dast hee ndnumbe roff ol l aL eds hoot s .Anyp()l Fi g.6( ha ti sabove200000r ol i a t e ds hoot sa c t ui l l l yr c pr c I s ent st hef ol i a t eds hoot st heye a rt hes i mul i l l l Ont er mi ni L t C( L notne ce s s ar i 一 yY9O. Sl i ght l ydi f f e r entr e s ul t sa r eobt ai ne dwi t ht ol i I ISCUsL i l Y90.The r ei ss omei ncr e as ewi t hi ncl e乙 I S l ngOl . derLi l V t 汀と I ge bi f ur ca t i on( Fi g.7) ,butt her el SaS L l ght l ynega t i ve1 . el a t i onS hi pbe t weent ot alSCUsandi nc r e a s esi nol der2bi r ul ' ci l t i ( n ( Fi g.7) .The l . eE I Pp ea r St ObeL i 〔 t l et ol ュ ot l el l dwi t hi n cl ' L l i l " i nor der3a ver a gebi f ul ` c a t i or l ,al t houghr o暮 .hl ghc rhi r ul C と ト の2 ( ) 0 4NReCi l t l 乙 l t l i l
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Fi g･6･ド( ) l l a t e ds h ( , ｡ t Sa ty C l l rり( 1°fPs e u l I o t L 川. q u1 , Z e n Z i e L " Iwi t hi n c r e a s l l l ga V e J a g eb i r u r c a l J OnO r( a)r l r S t ,( b)s c C 州dl ,a l l d( I T )t hi r do r d c rs h o ot s , 州d( J)n e we p i c or mi cs ho o t sa lt hr e edi f f e r e n tr ul ec ombi n a l l ( ) nS .No t et h edi f f e r e n ts C a l cf ol ･f oHi J t C ds h ｡ ot =1 ft l l i r do r dc ri l n dn e we p I C ｡r r n i cs h oo t s II ng e l l C r a lf o l i a t e ds h o o t sl n C r C a S Cn O nl i n e a r l yヽ ∼ i t hi 1 C r e a S 】 n ga v e r a g ebi f u r c a t i onordi ne r en l l y 01 1 SO nr e i t e r a t i ol la l l di shi g h e s tf l ) 1l h J J dOl d e r ( ) r dc r c ds h ( ) ot 六 ･Th ema gl l i t uL l eort h ee f f c c li n c r e a s e swh e nt h e r ea r cl c we rl e S t r i c t 1 bi r u' c alm n ,I n e r e a H I I l g1 1 1l h ca v e r a g ebi f u l ･ C a t i ol 】Ofl l 亡 Wl yI ni t i a t e del ) l C O 川1i cs l l OO t S .
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( Fi g.11 ) .Thi si ndi c a t est ha ti nt hes i mu] i l t ' L Ol 1 St l l Cav er a ge dynami csofSCU gr owt hovel ●t i l l l ea r eS t abl et hr oughout mos toft hel i f es pa noft hebr a nch;i ti sc ha nge sl nl hcpopul a t i onofSCUst ha tmos tl i kel ) ,a c c ountf orva l ' i a bl egr owt h i nt ot a ls hoot sont hebr anc hove rt i r nc･Dem0gJ ' aPhl CPI oh oft ot a le pI C Or mi ca ndr e gu】 a rs hoot sf oragl VCnS hoota ge gener a t e de ver y5yea r sf oras i ngl es i mul at e dSCU( Fi g･L 2) f oHow t hedi s t r i but i onsdes cr i be dbyl s hi il l ndFol d( 2 ( ) Ol , t hei rFi g.5)i nt hecl a s s i f i c a l i ol lOft hes t agesofSCU de v cト opme nt .Thes i mul a t e d SCU e xpl or e d wa sde cl a r cd i ndepende ntj nyea r3 5( a ta ge1 0)a nds ur vi vedt h r oughye a r6 0, gl Vl n gl taL i f es pa nof35yea r s ･ Thepot ent i a ll i r e s pa noft heSCU c a nbea s s es s edi nt he c ont e xtoft hePoi s s ondi s t r i but i onf ort henumb e rofda ught e rs hoot spe ra ct i v enode.Thede c r e as ei ne xpe ct e dva l ue f ort hemodi f i e dPoi s s ondi s t r i but i onus e di nt hes i mul a t i ons doe snotaf f e ctt hepr obi l bi l i t yofapa r t i cul a rnodedr a wl nga z , e r o( i . e. ,t hes t oc ha s t i cdeat hoft het e r mi na lbud) .I fone ca l las s umet ha tt henumbe rofda ught ers hoot spr oduc e dby t hemai naxi si si l l depende ntf ore a c hc ons e cut i veye r,t a hen t hea geaLWhi cht hema i na xi sofa nSCU i se xpc c t c dt o dr a wi t sf l r S tZ e r oC a nbemodel edbyage ome t r i cdi s t r i but i on( i . e. ,t henumberof" t r i a l s "t ot her l r S le v ent ) .1 fwe der met her a ndom var i a bl eX a st henumbeT 'ofyea I . Sunl H t her na j na xi sdr a wsaz er oa ndas s umet hepr obi l bi 】 i l yof 吃)2 ( ) ( ) 4NR CCと I n l l L t i t
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Can.J.Bo† .Vo1 .82,2004
dr a wl nga7 J er Oi sdet e r mi ned byt hePoi s s ondi l q t r i but i on, t hent h epr oba bi l i t yt ha tt hemal na xi sj sl os tor lye arXa nd i t sexp ec t e dva l uea r ea sf oJ l ows( Cas e Haa ndBe r ger1 990) :
Rb O !=e-
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Forexa mpl e,l for de rIave r a geb i f ur ca t i oni s2. 5,t heni t woul dbee xpe c t e dt ha tt hede v el opi ngOr de r1axi swoul d r a ndom一 ydr a w az e r oa tye a r1 2. 1 8.Foranor de rla v er a ge bi f ur c a t i onof3. 0thee xpe c t e dye a rwoul dbe20. 09.Onc e t h ema i r la xi si sl os HheSCU s t r uct ur equi c kl yde t e l ` i or a t es , wi t hr e ge ner a t i onf r om e pI C Or i csr m e s uL t l ngt nne w SCUs ( Fi g.1 2) ,Theol de s tSCU I s hi ia ndFor d( 2 001 )obs e r v ed wa s24,whi l et hea v e r a gea ge sofobs e r v e dSCUswas日. 27, 1 2. 43,a nd1I , 1 7f ort heupp er ,mi ddl e,a ndl owe rcr o wns ,I f . l nSOU i se xpe c t e dt ol os ei t smai na xi sa tye a r1 2. 1 8,L he J l i twoul ds t i l lha v ef ol i a gebeal l ng S hoot sup t o 22 yea r s ( wi t l l1 0-11ye ar sbe i ngt hemaxi mu mf ol i a gel ongevi t y) ･I t s e emst ha Hhi spr oba bi l i t ys t uct r u r ea de qua t el yma t c hest he t y . obs er v a t i onsofSCU l onge vl Di scussi on Ques t i ol l7:How doe se pi c or i cs m pr out i ngr e s ul ti n SCUs ? Cl i ne(J 997)des c r i be sf ours t a gesofa pi c aldom i na nce= l a t e r a lbu df or ma t i on,i mpos i t i onOfi nhi bi t i onofl a t e r a )bud gr owt h,r el ea s eofa pI C aldom i nanc e,a ndbr a nc hs hootde ve l opme nt .Thes es t a gesr e f l e ctwha toc c ur si nt hei ni t i a t i on ofepl COr mi cs hoot sa sde s c r i be dpr e ious v l y.] na ddi t i ont o hor monalcor ) t r ol( e. g. ,Cl i ne 1 991 ,1 994;BoHma r ke tal . 1 995;Ha oI J i eetaL ･1 996,Cl i ne1 997;Mi guelc tal .1 998: DeWi te ta 1 .2002) ,bot hnut r i ent sa ndl i h tha g vebe eni mpl i ca t e di nt her e l ea s eofa pi c l domi a nar L C e( Ma r t i n1 987･ ,Cl i ne 1 991 ) ･T ti spos s i bl et ha tbot ht he 一 os sort hes uhl e nde d l a t e r ala pe x( s e er l 】 l es2and3f ore pi c or mi ci ni t i a t i on)and e s pondi ngf a vor ab一 el i ghtandnut r i entC Ondi t i onsf or t hec orr gr owt ha l ongt hemai na xi sa r enec es s a r yf ort her el eas eof s up pr es s e de pI C Or mi cbuds .Thi swoul dena bl et hebr a nc ht o c ont i nuec xpl oi t l nga r e a sC Onduc i vet ogr owt hal ongI t sa xi s wh e r et he l i mi t s off ol i a ge mor t a l i t y ha ve l e f ta nonpho【 os ynt he t i ct wi gba r e Ps e u dol s u gnme nz L ' e ▲ ∼ i iha sal ower br a nch dc ns l t yt ha nA.gr andL ' S( numbe rofbr a nc he spe r met r eofs t em:da t af orPme nz i e . i . i ic anbef oundi nI s hi ia nd Wi l s ol l( 2001 ) ;f orA.gr andl S ,H.I s hi ia ndD. C.Sha w,unpubl i s hedda t a ) ,a ndi thasbe enobs e r ve dt ha tA.gr andi shas ahi ghe ra ve r a gebi f ur ca t i ont handoe sPme nz t l e . mjonl a t e r al a xe s( r s hi ietal .2003) .Wes ugges tt ha tt hemor edi f f us e c r own ofP me nz t ' P . T i tpr ovi desgr e a t e rl i ghtt r a 7 1 L S mi s s i on t hr ought ot heJ owe rc r ownandt hei nnercor eolt hecr owL l t ha nA.gl . andi s ,whi c hpr ovi de st hem ic l ･ O enVi r onme ntne ces s a r yf orr ei t er a t i ont ooc cu r .Whe nt hes uppr es s e dbudi s r e 】 ea s c d,i tgr owsa si fs l l r r Oundi ngt i s s uedoe snoti nnue nc e i t ,t ha ti s ,i ta c t sasa ni ndepe nde ntma i nbr a ncha xュ s . Ther ul esf orSCU i r l dep endenc el J S edi nt hemode lr e f l e ct l wo l a ye r sofwha ti sme a ntbya ni nde e ndentcl p us t erof f oL i a ge･Thr oughouHhi sana l ys i si ndepe ndenc ei sdef i ne dbv -
a pl c a ldomi nan c e.t n a一 i mi t e ds ens ei l nd underc cl ' l i I i n ci r c ums t a nces ,br a nche sc anbea s s ul ne dt oF unct i onL l uL ol l Omous l ywl t hr ega r dt oc a r bon( Spr ugele ta1 .1 99LSpr uge1 2002) ,butwema kenos uc has s umpt i onsaboutSCUs . I nt hef i r s tr ul eofi nde pende ncet ha twa sexpl or e d,SCUs L q , a r ei dent i f i e dt hr ough mor phol ogl C aldi s L mct u es s .t hi L ti a nyaxi st hati snol ongerc onl l e C t e dt ot hemai na xi svi と Ll i v c r ol i a geca nbec ons i der eda nSCU.Ani mpl i ci tL I S S umPt l On oft hi sr ul ei st ha tmor phol ogl Caldi s t i nc t nes si ss uf r I Ci entt o ca us er e 】 e as eoft hel a t e r ala xi sf r om domi ni l nC COfi t . ia s s oc i a t e dma inaxi s( s e que nt i a lr ei t e r a t i on) ,al l owi ngf o1 gl ◆ ea t e r bi f ur ca t i onoft hel a t e r a laxi s( r uL elf orSCU i 1 dcp en dcn c e) ･ l fs uchmor phol ogl CaI l ydi s t i nctl at er a lax est hr ough ouHhe C . .r ei t er a t e dt obeL i ket h e br a nc har et hengi v e nor de rI( i . mai naxi s ) ,s i mul a t e dgr owt he xpl odest oi mpos s i b一 yhi gh number soff ol i a t e ds hoot . q ｣nA.gr andt S ,hi gher之 I Ve1 L l gel a l e r a 】bl f ur c at i onr es ul t si nl a t e r aLE I X e St ha tvi s ual l yr e s embl e SCUs ,bt t tt he s es hou7 dnotbec ons i de r edJ ･ ei t e l ･ a t i onsoft h e n l a i naxi s.I nt hedef i ni t i onoft hef i r 山r ul el t HSCU I ndepe nde nc et hemor phol ogl Ca lc har a ct e r i s t i cspr e c e det heこ I S S Oci at e dpl l yS 1 01 0gl Ca lf unc t i onoft hegr owi ngaXI S , I nt hes ec ondr ul ef orSCU i ndepen de nc et h ephys i oL og卜 catf uI I C t i oL lpr e c edest hea r c hi t e ct ur alc ha t . a C t C r i s t i c s ;i ti s on一 yone pl COr mi cst ha tha vebee nobs e r v e dt of unct i onas f i r s Lor de rs hoot st ha tSCUsar ea Howcdl odc v el oL l( i ･ e･ ,t he ma i na xi sf unct i onpr e c edest hecl a s s i f i ca t i oni nt oSCU詫 ) .T n F 'me nz i e s i l ,SCUst ha tde vel ope dF r o) T le PI C ( ) nlli c 叩】 Out L ng r e s ul t e di ns i mul a t edbr anchest ha twer er ca s onabl er cL a L i v c L ot heobs er vedbl ' a nc hes( I 7 i g.4 f ;Ta bl e4) .Theke yt ot hi s di s t i nct i onbct wcenA.gr a ndi sa ndf hne nZ t ' C , Y L ' t 'i . qt hel ' e C Ogni t i onoft hedey el opme nt alor l gl nOft hes hootgr oupl ng･l t i sa l s opos s i bl ef oras e quent i a ll a t er ali l Xi st of unct i ona si l f i r s t or de ra xi s ,S uc hasaf t e rat r aumat ot heJ T l ai nt c l l ni nal bud;i nt hi sc as et hes e quent i a la xi sma ypr odu c eと m SCU. Thei s s uei showt odet e r mi l l eWl l e t he rt hene w axi si sphys i ol ogl C ll a yi l l de pende ntf r om t hef i r s t or de rmai ni l Xi s ･ Thet r endsoft ot a lf ol i a L e dL l hoot sa l l dSCUswi t hi ncl ･ C a S 1 I l gav e r a gebi f ur ca t i ons e el li nt hes cns i t i vl t yanat yS I S°i l n bee xpl a i ne dbyt her el a t i ons hi pbet we ent her ul esf orepL I c or mi ci ni t i a t i ona nd SCU i n de pe nden c e.Underi t l ll ' ul c s , t he r ei sam i ni mum numbe l ･Ofs hoot st ha tan e pl C Oml i c c ompl e xr m] s tha ver oheca l l edane w SCU (1 0s hoot s) ,and epI C Or mi cscanonl yf or m onor der)ax e s .TheL ef oL . C,I nc r ea s esi nor de r1bi f ur ca t i onf a c i l i t a t et hepr oc es sorSCU f or ュ 1 a t i onbyi ncr e a s l ngt henumbe rofor der1node sa vai レ a bl ef orc pi c or T ni ci ni t i a t i on( Fi gs .6a,7a) .Hi gh e) .ol ■ der7 a nd2bi f ur ca t i onsal s ol nCr e aS et hel i kel i hoodt h' l t; In e wl y f or mi ngSCU pr oduc ess l 】 f f i c1 cnts hoot st or ea chl h cmi ni mum numberne c es s a r yf orSCU i nde pendel l C e( Fi gs .6,7) I Howe ver ,i fepI COr mi ci ni L i al i oE He qul r eSOr l COl ･bot hoft he s ubt er l de dL at e r a Jaxe st obet e r mi na t e d( RC5,RC6) ,t h cl l hi ghe ror de r2 bi f ur c at i on woul d de cr eas eth epr obi t bHi t y t ha ta ne pI COr i Ci m ni t i a t es.Duet ot he s ecom pe l l ngt l or c es unde rRC5t he r ei sas l i ght l yne gat i ver el at i ons hl l lhe t wc cn or de r2bi f u r c a t i onand T het ol aJnumbe rOr SCUso nt he br a nc h( Fi g.7b) .Thi si sf ur t he rampl i f i e di fbo t h. i ubL el l deL I s hoot smus tnol onge rbegr owL ng,aSi l lRC6. Wi t hr e s e ctt p o que s t i on 1 ,weconcl ude t ha tep7 COr mi c s pr out l ngr e s ul t si nSCUst hr oug h i lCOl l l bi nat i onot l phys L O1 ogl Ca lc ont l OI sa nd pl antar chi t e ct ur e.Epi cor mi cs pr ou【 L ng a l l owst heba s i ca r chl t CCt ur almode lofP mcn< : l ( , . u' I 'l obeI C ￠ )2り( ) 4NR CCi H l . I ( l i l
907
Ke nn edyo tal . i t er l L t Cdr cl aL i vel yi ndep endent l ya l ongt heh or l Z Ont aLma i nbl 一 之 I nCha xi s ,e ve nと L r L c rt hebr an chha sr ea che di t sma xi mum C･ ,cT Ownexpa ns i onhasce a s e d) . l engt h( i ･ Th cr el t er a t i onofSCUsder l n e df orP.me nz i e si iocc ur son l . L IS mal l ers ca l et ha nt h er ei t e r a t i onde s cL . i be dbyHa Heeta ( 1 978)i nt l ･ OPi c 乙 I ll r eeト .Ha 1 1 6e taL( 1 978)obs e l . V e dve r t i ca l r ci t c r a t i onsoft hea 】 ･ chi t ect ur こ t lmode loft hee nt i r et r e e,r e s ul t i ngT n" m川1t l ' e eS ' 't hr oughouHheC r own.Al t houghHa1 1 6 c tal .( 1 978 )des c r l bei 1L i ml t e dnumbe rofba s I Ca l ･ C hi t e c t ur a ] J 一 一 Odel si nt 1 Opi c･ 1 1t r e e s ,t h e ya l s on ot egr e atdi v cr s l t yamong 0del .Of t ent hi sdi v e r s l t yl S t r c cst ha ts ha r ea nal ･ chi t e ct ul ` と 1 11 1 1 duet ot hewi l yl nWhi cht hemodeli sr ei t er a t e ddur i ngc r owl l de vel opme nt .] n7 二/ ) / ( ' c at a,EdeZ s t ei na nd For d( 2002) oh. s el . VC ds maHe rs cと 1 1 ev er t i c alr ei t er a t i onsal onghor i zont a 1 i p . u 日heobs e r ve dr ei 1 bmnc hi nga x es .J nt heca s eofP.menz c mL i un oc c ur shor i z onL と l l l y; 1 1 0ng t he br a nc hi ng a xi sa nd r c pe之It St heA) owt hofbr a nche s ,noHhcver t i calgr owt hoft he mi l i ns t e] T l ･Thr oughL ei t er a t i onat r e ee xhi bi t soppor t ul l i s t i c i l r C hi t ec t ur et hi l tal l owsf ordi v er s l t yl nt hewa yt hecr own e xpl oi t sと I V乙 l i l i l b】 er e t q ot L l ･ Ce S . Que s l i on2:Whatar et headvant age sofr ei t er at i on? l Li scl c と 1 1 't hi l lr ei t er a t i onC Ont l . i but ess l gni f i ca nt 】 yt ot he gr owt horp .mc 1 7 Z : L ' eL l i T I J e S uJ t mgl nUbL Y er V e dbr a nc hesupt o 1 55yei u . iOl d( 1 s hl ia ndFor d20O) )a nds i muL a t e dbr anc hes 93ye a r s( Fi gs .90, t hi t Lc ont i l i ne df ol i と I t eds hoot sF orupt o3 i ) り.l nc onl L ' a S t ,t heol des tobs e r v e dbr anc hofA.gr andi s wa sonl y( 1 3ye i l 】 ● Sol d,E nds i mul a t edbr a nc hesl i ve dupt o l ● S .Wi t hnocont r i but i onorr ei t e r a t i ont ogr owt h( Ta 22( ,yei l ( ) .Se ns i t i vi t ya nal ys i ss howst ha ts ma l l bl e4,Fi gs .9b､9( i nc l . e i I S eSi na v er a gebi f ur c a t i onofhi gheror del -S hoot spr opa ga t et hr ough outt hebr anc ha ndr es ul ti ndr a s t i ci ncr ea s e si n t ot a lr ol i a t e ds hoot L q( Fi gs ･4ca nd 8b)t ha tar eden . s el yl oca t c〔 ldl S l i l Hyont heb) 一 ･ d nCh.Fur t her mor e,as mal lI nc r e a s e i nt hea bi l i t yoft hebr ancht or e l T e r ; l t e( e i t h erpr ol e pt i c a l l y orons e quent i a la x es )a l s ohasapr of oundef f ectont het ot a l ob) . numbe rofl l ol i at e ds hoot soE lt hebr a nch( Fi gs.4,l oo.l EvenaI-e L Nr i ct edcと t pa Ci t yf orr e i t er a t i onhasas t r ongcumu) a l i v eer r c clont hea bi l i t yoft hebr anc ht os ur vi vet hr ough r c geneT i l t i onofi t sf ol i a ge( Fi g.4) . l thasb ccnc ) bs e l ● v e dt ha tgr owt hdec l i neswi t hi nc r ea s l ng 96 3;Fi s he l .a nd ヱ I gCandc ompl e xi t y( Moor bya ndWa r ei ng 1 977こZl r T l me r mと I nl 978;Cl a r k1 983:Bor c he r tand Honda 1 11 98 4,Bond2OOO) .wi t hma nye xpl a n, d t l OnSf orhow Hondi gr e at ercol 1 l ) 】 c xi t yc と l l lbcdel r i me nt aL .Wi l s on( 2000)C 一 ai ms と I PI Ca lc ont r olr c gul a t e st h eamountore一 onga t i ona nddi a met eI 'gr OWt hi nt r c csL l ndbr anc hes ,p os s i bl yt hr oughar e duc t i oni nt het r . l n叩Or tOrW之 I t erL l ndnut r i e nt st ol a t e r a ls hoot s . L n( 1 り78)f oL ) ndと Idi s t l nethydr a u一 i cc ons t r i ct i onat Zl mmeT 1 1 1 1 br i l n Chi ngj un ct i ons ,whi ) cBol . C her tandHondaH984)obs e. ･ v e dl h z t LJ l uX( I I Otde f i n e d by a ut hor s )dec r e a s edw it h i n cr c i l S ] ng Or del . ,a nd 1 . e duct i onsi l 一bi f uJ ' Ca t i on ofhi gher br L I nC hi ngol ◆ dc l ･ qWa sduet ot hcc xpor L ent l a lde c r e a s ei nt he l .(1 999)pl ` OpOS e r l ux oH と I t el . i l lbr a nc he s .Hubbと I r de ta i nc r e i l S J ngnt l r nbc r sofbr a nchj unc t i onsa l ongc on duct anc e pi l t hwとIyS とlH ( ) neOrl h cpos s i bl cと l l t e l ' na t i vee xpl ana t i onsf or di f r e r e ncc si nhydr i l ul i cc onduct a nc eと I ndl e afga se xcha nge -
be t we . enl al lと nd s hor Hr e es ･Ni i ne me t s( 2 002)c onc l ude d t ha ts l / . ehi l Sと lgr C a l e re f f e ctonde c r e a s l ngS t Oma l alc onduc 一 lf !( t bL ' e sa ndPi nuss ) ･ l v e s f r i s .Toe f f ec t t l nC Ct hi L nOn乙 I g ei t i v el y. i i l l l ul aL e br u l l Che s of lp me nz i e s i i ,we f ound i t
ne c es s a r yt oi ncl udeapa r ame t ert ha tr e s t r i ct e dc omp] ex】 t y c a us e dhyr e i t el ` a t i on( ge nT ｡ J)a ndt c ･r e duc ebi f ur c at i onw it h i ncr e as i ngge ner a t i ont Fi g.4,e q,4) .rhl SaL s oe f f ' e c t i vel yr e duc e dt hei nf ] uenc coft hes t r ongs t ochas t i ceff e c t sobs e r ve d l ,al t hought hes i mul a t i onss t i l lexhi bi t e de xt eni nt hemode s i ves t ochas t i cl t y.Thel i mi t at i on,l l OWeVer ,Wa si mpos e dL o obs el ● vede s i r e dr a nge sofmode lout p一 l t Sa nddonotyetha ye aba s i si nf i el dobs e r v a t i ons . Toa ns werques t i on2,wec oncl udet ha tr ei t e r a t i onoft he SCU i n P me J Z Z l e L utPr of oundl yl nC f ･ e･ l S e = hel i f e s pa nof bl anC l l e Sl ' el at i v et oA.gr undi s .a ndr ei t e r a t i oni ss hownt o aha veas l gni f i cantc un l ul a t i vee f f e ctul lgr owt h.Theut i l i z t i onofe xi s t l ngme r i s t e mst or e ge ner a t ef oL i こ I geOnes t abl i s hedbr a nc ha xesa ndi nt heabs enc eofaddi t i ona lbr a nc h l engt hi s a powe l lu)met hod ofcr own mai nt e na nce i n P me nz i L , s L ' l . l ti spos s i bl et ha tal i mi t edf or m oft r a uma t l C Seq l l ent j < l l r ei t e r at i ons houl dbes i mul a t e dont hes ebr a nchesa sar e a s ona bl ea ) t e r na t i vet of or c l ngt l l emai l la xi sl ope r s i s tf ol 't he f i r s t50ye a r sofbr nc a hgr owt h.Wi t ht hel os soft het e ml i na l bud,S e quent i all a t e r aLs hoot swoul dber e l e as e df r om a pI C a l domi na nc ea nda s s umet her ol eoft hemai na xi swi t hi nt he us ua lt i mL ngOfgr owt h,a shasbee nobs er vedi nt h ev e r t i ca l 9R 6) i gr owt hofbr a nche si nv ar i ouss pe ci e s( a. g. ,Ha r di ng 1 Tr auma t i cr ei t er a t i o( 1Woul dbeexpe ct edt oi nc r ea s et hel i f e s pan ofbot hs pcc i es ,al t hough l i it m a t i onsonc ompl e xi t y LI na ddi t i on, ma yne e dt obei mpos e di nt hi sca s ea swcl S omeba s a lr ei t e r a t i onha sa l s obee nobs e r ve di nF !me nz i e s i i ( I s hi ie tal .2002) ,whe l ' eS t l ppr eS S e dbudsi ni t i a t eonol de l ■ br a ncha xes( upt o60ye a r sol d,about23yea r sona v el ､ a ge) . The i nc or por at i on ofe pI COr ic s m pr out l ng On Ol ders uppr e s s e dbudsma yal l ow f oュ .1 ' ei t e l a t i ol lt Ooc curWi t houtr es t r i ct i on,wher el owergene r a t i onsofSCUsa r ecol l t i nua l l y f or mi nga ta l ls t a gesofbr a nc hde ve l opment .I nt hi sc a s e r ei t e r a t l Onpr OVl de sapot ent i al l yl i mi t l es ss oul C eOfne wf b) i a t e dt i s s ue( La nner2002)ont hebr E l nCh. Tl l eS t l l dyort heSCU Toa i df ur t he ri r l Ve S t l ga t i onwepr opos et hecl as s i f i c ･ l t i on ofSCUsi nt oL hei 1 -de vel opment alor i gi ns :pr ol ept i c( vi aepi c ot ･ mi cs pr out i ng)ors e que nt i al( onl a t eJ ･ a ls equel l t i u It l X eS . i n r es pons et ot hel os soft hea pi calme r i s t em) .Th epl ℃per t l eS oft he s eol ･ ga ni z at i ona luni t sca nbec ompa r e dt ode t e mi ne wh et hert heSCU i st r ul yaf unda ment a luni tt ht ･ Oughoutt he lmt ･ nz t ' e ･ " l iorwhe t heri t sdef i ni t i on br a nc hi ngs t r uc t ur eofl i sr es t r i ct e dt ogr owt hvi aepI COr mi cs pr ou t l ng.I na ddj t j on, t l l el oca le n vi r ol l me nt alc undi t i onsoft hebr a nchi nt hepr oxi ml t y Ofne wl y de ve 】 oped SCUsca n bei n yes t l ga t e da l l d c ompa r e dwi t ha r e aswi t houtSCUst odel i nea t et heca us eof ] y ,t he e pI C Or mi cs pr out l ng a nd SCU deve l opme nt ･Fi na J gr owt hofne w一 ys pr out ede pi cor mi cscl as s i f i e dbygener at i onc a nbeqL L a nt i f i e dt ode t e r i newhe m t he rgr owt hde cl l n es wi t hi ncl . ea S l nggene r a t i onori fge ner a t i on i nf l uenc e st he nr i e s l ' t -br L l nC h. s uc c es sofe pl COr mi cs pr out 1 1 1 gOnaP me Acknowl edgement s Th eda t af o rt t l i sr es e a r chweT ･ eg' l t heT ･ e dヱ l tt heWl ndRI V er Ca nopy cr a ne Res ea r c h Fa ci l i t y,a c ooper a t i v es ci e nt i f i c vent ur ea l l l Ongt heUni v e r s l L yofWa s hi ngt on,USDA For es t . Se r vi c ePa ci f i cNo) . t hwes tRe s ea r chSt a t i on,a ndFor es tSet ◎2 OO4NR CCヱ I J l i l d ( i
Can.J.Bot .〉ol .82,2004
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vi ceGi f f or dPi nchotNat i onaLFor es t .Fi nanci als uppor twas pr ovi dedt hr oughagr antf r c ) mt heAndr ew W .Mel I onFoundat i on.Theaut hor sw is ht ot hankt woanonynl OuSr evi ewer s f ort hei rhel pf l l lc( ) mment s,andDougl asSpr ugelandJudi t h Zeh f orus ef ulcomment sand s uppor tt hr oug h outt hi sr es ear chpr o j ect .W ewoul dal s oL i ket ot hankAnnet t eHami ] l ,DaveShaW,andMar kCr ei ght onf ol °t hei ras s i s t ancei n 1 ol t hes t udyand manyS t udentandpar Hi mewor ker sf ort hei r as s i s t ancei nt hef i el dandl abor at or y.
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