1975 naveh - evolutionary significance of fire in mediterranean region

7/29/2019 1975 Naveh - Evolutionary Significance of Fire in Mediterranean Region

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Vegetatio Vol. 29, 3: 199-208, 1975

T H E E V O L U T I O N A R Y S I G N I F I C A N C E O F FI R E I N T H E M E D I T E R R A N E A N R E G I O N *

Z . N a v e hFacu l ty o f Agr i cu l t u ra l E ng inee r ing , T ech n ion I s rae l Ins t i t u t e o f T ech no logy , Ha i fa , I s rae l**K e y w o r d s :Ad ap ta t i o n , E co log i ca l d ive r si t y , E vo lu t i ona ry st ra teg ie s , F i r e , Med i t e r r anean r eg ion , Rege ne ra t i on , Ro o t sp r ou t e r s

Introduction

D u e t o t h e d e s t ru c t iv e c o m b i n a t i o n o f o v e r g r a z in g a n dl and abus e wi th burn ing , wi ld l and f i re s have been r ega rdedi n t h e M e d i t e rr a n e a n R e g i o n a s a w h o l l y c o n d e m n a b l e ,m a n - m a d e e l em e n t . T h i s m i g h t b e t h e r e a s o n f o r th e f e w ,sys t em a t i c s t ud i e s o f f i r e e f f ec t s and neg l ec t o f t he i revo lu t i on a ry ro t e i n Med i t e r r an ean vege t a t i on . In p rev iouss tud ie s , f i r e ha s bee n m e n t ione d o n ly b r i e fl y a s a de s t ruc -t i ve f ac to r (Z oha ry , 1962) o r i n connec t i on wi th m an-induced r eg re ss ion s t ages , l e ad ing away f rom the "m aqu i -c l i m a x ' ( B r a u n - B l a n q u e t, 1 9 2 5 ; B h a r u c h a , 1 9 5 2 ; K u h n -ho tz -L orda t , 1939) . On ly i n a r e cen t , m os t com prehens ived e s c r ip t i o n o f M e d i t e r r a n e a n v e g e t a ti o n t y p e s , W a l t e r( t 968) ha s r eco gn ized f ir e a s one o f the m a jo r e co log i ca lf a c t o r s w h i c h s h a p e d t h e M e d i t e r r a n e a n l a n d s c a p e a n da f fec t ed it s p re sen t m osa i c - l i ke pa t t e rn o f r egene ra t i on a nddegrada t i on s t ages .

Shan t z (1947) r ecogn ized t ha t sh rub com m un i t i e s o fm ed i t e r r anean c l im a te s , no t on ly i n Ca l i fo rn i a bu t a l so i nS o u t h A f r i c a a n d S o u t h A u s t r a l i a a n d i n t h e M e d i t e r r a -n e a n R e g i o n p r o p e r , s h o u l d b e c o n s i d e r e d a ' f i r e t y p e 'o r ' f i re c l im ax ' becaus e o f t he i r l ong sub j ec t i on t o r e -cu r r i ng f i r e s . Naveh (1967) d rew a t t en t i on t o t he s t r i k ings im i l a r it y i n f ir e r e sponse o f i nd iv idua l p l an t s an d co m -m u n i t i e s o f m e d i t e r r a n e a n s h r u b e c o s y s te m s i n C a l i f o r n iaand I s rae l . T h i s i s m an i fe s t ed i n bo th coun t r i e s by r ap idregene ra t i on o f t he sam e ind iv idua l s and /o r b y a sh if t s in

* Nomenclature followsM. Zohary, N. Feinbrun & A, Eig.Analytical Flora o f Palestine, 2nd edit ion I965.Ham agdir. Jerusalem. i965, Sifriat Hapoalim. Tel Aviv.** Publication No. 209.

c o m p o s i t io n t h r o u g h v o l u n t e e r se ed s o f b u r n e d m o t h e rp l an t s . Such a p rocess o f ' au to suc ces s ion ' (Hanes , 1971) i sobscured , i n gene ra l, by a sho r t i n t e r lude o f he rbaceousp l an t do m in a t ion (M ul l e r e t al ., 1968) . Na veh (1967) al sopo in t ed t o t he com parab l e ro l e o f f i r e i n bo th m ed i t e r -r a n e a n e n v i r o n m e n t s i n t h e e v o l u t io n o f s h r u b e c o s y s t e m sa n d i n t h e i r m a i n t e n a n c e i n a r e j u v e n a t e d a n d v i g o r o u ss t a te , coup led wi th t he m obi l i z a t i on o f ti ed -up nu t r i en t sa n d t h e r e m o v a l o f h e a t- u n s t a b l e p h y t o t o x i c , a ll e lo p a t h ica g e n t s . A f t e r c o m p a r i n g m o n o t o n o u s , i m p e n e t r a b l e a n dc o m p l e t e l y p r o t e c t e d m a q u i w i t h b u r n e d a n d d i s t u r b e d ,b u t b i o l o g i c a ll y m u c h r i c h e r m a q u i a n d b a t h a ( a b i bl ic a lt e rm app l i ed by E ig (1927) i n I s r ae l t o hea the r - l i ke dwar f -s h r u b a n d h e r b a s s o c i a t i o n s ) , N a v e h ( t 9 7 1 ) c o n c l u d e dtha t p re sc r ibed burn ing shou ld becom e an i nd i spens ib t et o o l i n th e e c o l o g ic a l m a n a g e m e n t o f n a tu r e p a r k s a n drese rves i n t he m aqu i be l t fo r conse rv a t i on o f b io log i ca ldiversity.

L e Ho ue r ou (1973) r ecen t ly d i scuss ing t he e f f ec t o f fi r eo n M e d i t e rr a n e a n v e g e t a t io n a l s o c a m e t o t h e c o n c l u s i o nt h a t i t i s d o m i n a t e d b y p y r o p h y t e s a n d t h a t p e r i o d i c a l l yb u r n e d f o r e s t a n d s h r u b l a n d i s m o r e c o m p l e x i n c o m -p o s i t io n a n d s t r u ct u r e t h a n t h a t n o t b u r n e d a t a ll o r b u r n e dt o o f r e q u e n t ly . M o o n e y & D u n n ( 19 7 0) o u t l in e d t h e c o n -ve rgen t evo lu t i on a ry s t ra t eg i es o f m ed i t e r r anean c l im a te -type sc l e rophy l l sh rubs and i nc luded f i r e a s one o f t hem a jo r env i ronm enta l s t r e ss f ac to r s . T hey re fe r red , how-eve r , on ly t o exam p le s o f f ir e - s t im ula t ed g e rm in a t i on an dr e s p r o u t in g b e h a v i o u r o f C a l i fo r n i a p l a n ts . S u c h a d a p t i v em e c h a n i s m s f o r C a l i f o r n i a ' f i r e t y p e s ' h a v e a l s o b e e ndesc r ibed i n m ore de t a i l by Sweeney (1967) and Vog l(1967).

T he ob j ec t o f t h is pap e r i s t o d i scuss i n m ore de t a il t hee v o l u t i o n a r y s i g n i f i c a n c e o f f i r e b e h a v i o u r b y w o o d y a sw e ll a s h e r b a c e o u s p l a n ts f r o m t h e M e d i t e r ra n e a n R e g i o naga ins t t he backg roun d o f a l ong f i re h i s to ry .

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T h e f i re h is to ry o f M ed i terra n ean eco sy s tem sA t p re sen t , l i gh tn ing cause s wi ld f ir e s on ly o n 0 ,6 -4 % o ft h e a n n u a l l y b u r n e d a r e a s i n t h e M e d i t e r r a n e a n R e g i o n(Susme l , 1973) . In I s r ael , l i gh tn ing ou t o f c l ea r sk ie s m ayoccu r on seve ra l d ays i n Ap r i l - Ju ne and cause wi ld - fi re s .In anc i en t t imes , howeve r , even such a s i ng l e l i gh tn ingf ir e on a d ry day , i f no t pu t o u t immed ia t e ly , cou ld ca t cht h e u n d i s t u rb e d , d e n s e a n d h i g h l y in f l a m m a b l e w o o d y a n dhe rbaceous vege t a t i on and sp read r ap id ly ove r va s t a rea s .Such f i r e s on wi ld l and and pa s tu re s we re a l r eady men-t i oned i n t he B ib l e i n connec t i on wi th l i gh tn ing a s t he' f ir e o f G o d ' i n t h e B o o k o f J o b ( I /1 6 ) a n d i n c o n n e c t i o nw i t h ' t h e h e a t o f t h e s u m m e r d r o u g h t ' b y t h e p r o p h e tJoel (1/11).

The g rea t geographe r and eco log i s t , Saue r (1961 ,p . 262) c l a imed tha t 'moun ta inous a rea s i n a r i d and semi -a r id l ands do se t up d ry - sea son t u rbu l ence wi th l i gh tn ingd i scha rges i n to in f l amm able p l an t ma t t e r . Such f ir e s occurund e r cond i t i ons o f l ow ihumid ity and a re accom pan iedby l it tl e o r no r a in . . . . The cap tu re o f f i re m ay we l l havecome abou t a s a d i scove ry on food co l l e c t i ng , fo l l owinga f t e r a na tu ra l f ir e. A s t he sou rce o f such f i re s I sho u ldpre fe r vu t can i sm; I t a ly and t he ea s t e rn Medi t e r r anean ,t h e E a s t A f r i c a n R i f t V a l l ey ?

A c c o r d i n g t o R o s e n a n ( p er s. c o m m u n i c a t i o n ) s u c hl igh tn ings a s we l l as vu l can i c e rup t ion caused m any l a rge-sca le f i res in geologica l t imes, especia l ly s ince the LastIn t e rg l ac i a l de s i cca t i on , when the ma in Medi t e r r aneanc l ima te pa t t e rns and vege t a t i on t ypes became f i na l l ye s t ab l i shed (Bu tze r , 1972 ; Horowi t z , 1968) .

The ea r l ie s t a rcheo log i ca l ev idence o f t he p re sence o ff i re in t he maqu i be l t o f the M edi t e r r anea n Reg ion r eachesb a c k t o f i n d i n g s o f w o o d e n a s h a n d h e a r t h s i n t h e f in a lAch eu l l i an and Leva l lo i so -M ous t e r i an l eve ls o f the Me -s o l i t h i c p e r i o d a t t h e M o u n t C a r m e l c a v e s i n I s r a e l b yGar rod & Ba te (1937) and a t t he Kas t r i t sa c aves nea r l akeIoan ina i n G reece (Higgs e t a l. , 1970) .

W e h a v e n o d i r e ct e v i d en c e o f i n te n t io n a l b u r n i n g b ythese pa l eo l i t h i c hun t e r s and food ga the re r s . Bu t i t i sr ea sonab le t o a ssume tha t , a f t e r r e a l i z ing how na tu ra lf ir es open ed t he dense and un pene t rab l e fo re s t and m aqu i sth i cke t nea r t he i r hab i t a t s and c rea t ed b io log i ca l l y r i che reco tones an d success iona l s tages , t hey migh t have used f i rede l i be rat e ly t o f ac il i ta t e hun t ing and foo d ga the r ing and t oinc rea se ed ib l e food fo r man and beas t by t he l ush r e -gene ra t i ng t r e e s and sh rubs and i nvad ing g ra sse s , he rbs ,bu lb and t ube r p l an t s , t yp i ca l fo r t he se burns . Tha t f i r ebecame , t he reby , t he f i r s t fo rce fu l t oo l fo r e cosys t em200

m a n i p u l a t i o n b y t h e P l e i s t o c e n e h u n t e r - g a t h e r e r e c o n o -mies ha s been sugges t ed no t on ly by Saue r (1957), bu t a l soby eminen t an th ropo log i s t s and a rcheo log i s t s such a sSt ewar t (1957) , Oak ley (1961) and o the r s , c i t ed by Ko-m a r e k ( 1 9 6 7 ) w h o d i s c u s s e d t h i s p r o b l e m m o s t c o m -prehensive ly .

W i t h i n c r e a s i n g h u m a n p o p u l a t i o n s a n d a d v a n c e dh u n t i n g a n d f o o d c o l l e c t i n g e c o n o m i e s , s u c h a s t h e N a -t u f i an c u l t u re s o f th e C a r m e l C a v e s , d e sc r ib e d b y G a r r o d& Ba te (1937) , Ga r rod (1958) and Vi t a -F inc i & Higgs(1970) t he ex t en t and i n t ens i t y o f bu rn ing p rob ab ly i n -c r e a s e d s t e a d i l y a n d b e c a m e m o r e a n d m o r e l i n k e d w i t hgraz ing by wi ld and semi -domes t i c a t ed ungu la t e s , such a sfa l low deer , gaze l le , wi ld goat and ca t t le . The steadyinc rea se i n Med i t e r r anean ga r igue , s t eppe and rock dwe l -l i ng roden t s , r e l a t i ve t o more mes i c fo re s t and maqu idwe ll er s, f rom th i s pe r iod onw ards ha s been sh ow n inrecen t pa l eon to log i ca l f i nd ings i n t he se caves by Tche r now(1968) and i nd i ca t e t he g radu a l en l a rgem en t o f d r i e r,m ore expo sed and ro ck i e r hab i t a t s i n t he v i c in i t y . As t he set r ends we re no t a ccompan ied by ove r -a l l c l ima t i c de s i cca -t i on (van Ze i s t , 1969 ; Bu tze r , 1972 ; Ucko & Dimbleby ,1969) , they can be interpre ted as the f i rs t , f i re - induceds t a ge o f t h e a n t h r o p o g e n i c b i o f u n c t i o n w h i c h h a s l e dt h r o u g h v a r i o u s c y c le s o f d e g r a d a t i o n a n d a g g r a d a t i o n t ot h e p r e s en t M e d i t e r ra n e a n l a n d s c a p e ( N a v e h & D a n , 1 9 71 ).

F i r e ha s p l ayed a no l e ss a c t i ve ro le i n fu r the r phase s o fth i s an th ropogen ic b io func t i on i n t he pa s to ra l and ag r i -cu l t u ra l e cosys t em modi f i c a t i on and conve r s ions , i n fo re s tand l and c l ea r ing fo r cu l t i va t ion and appa ren t ly a l so i n t hed o m e s t i c a t i o n o f c e r ea l c r o p s .

As r ev i ewed ex t ens ive ly by W hyte (1961) and Ucko &Dimbleby (1969) , t he t r ans i t i on zones be tween t he semi -a r i d s t e p p i c I r a n o - T u r a n i a n a n d t h e s u b - h u m i d M e d i t e r -r a n e a n w o o d l a n d s a r e a m o n g s t t h e f i r s t k n o w n c e n t r e so f success fu l c e rea l and s tockbreed ing economies . Theser e g i o n s i n c lu d e t h e m o s t f i r e - w o n e M e d i t e r r a n e a n b i o m e s(Naveh , 1973a ), i n wh ich t he p ro gen i to r s o f ou r c e rea ls ,t he l a rge g ra ined wi ld ba r l ey = Hordeum spontaneum a n dt h e E m m e r w h e a t = Triticum dicoccoides, a r e a b u n d a n t( Z o h a r y , 1 9 6 9 ) . T h e a n n u a l g r a ss e s a r e a m o n g s t t h emos t p ro l i f i c f i r e fo l l owers , and pos t - f i r e co l l e c t i on o fseeds , a s a l so sugges t ed by Komarek (1967) cou ld havebeen on e o f t he f ir s t phase s i n the i r dom es t i c a t i on . The i rf i r e - sco rched seed d i spe r sa l un i t s wi th b ig pa rche d ke rne l scan easi ly be col lec ted af te r a wi ldfi re , an d ac cor ding toHa r l an (1967) the se p r imi t ive g lum ed ce rea ls needed t o bep a r c h e d b e f o r e t h e y c o u l d b e t h r e s h e d a n d w i n n o w e d .H a r t a n & Z o h a r y ( 19 6 6) s u g g e s te d t h a t a c t u a l d o m e s t ic a -


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t i o n a n d f a r m i n g m a y n o t h a v e t a k e n p l a c e w h e r e th e s ec e r ea l s w e r e m o s t a b u n d a n t a n d c o u l d b e c o l le c t e d f r o mn a t u r a l s t a n d s , b u t i n a d j a c e n t a r e a s . T h e s e i n c l u d e t h em a q u i b e l t , in w h i c h t h e s e m i - s e d e n t a ry N a t u f i a n c u l t u r e su s i n g f l i n t s ic k l e s a n d g r i n d i n g s t o n e s w e r e l o c a t e d , a s w e l la s t h e d r i e r J o r d a n V a l l e y w h e r e t h e f i r s t p r e - p o t t e r yn e o l i th i c s e t t le m e n t ( J e ri c h o ) a n d t h e J a r m u k i a n c u l t u r esh a v e b e e n f o u n d . T h e o r i g in s o f d o m e s t i c a t i o n a n d c u l -t i v a t i o n i n t h e N e a r E a s t a r e , t h e r e f o r e , s t i ll v e r y m u c h am a t t e r o f s p e c u l a t io n ( F l a n n e r y , 1 9 69 ). B u t K . K e n y o n -t h e d i s c o v e r e r o f J e r i c h o - c l a i m e d ( K e n y o n , 1 9 69 , p . 3 3 I ) :' I t w o u l d b e q u i t e i n k e e p i n g w i t h t h e t r e n d o f a r c h e o -l o g i c a l e v i d e n c e t h a t t h e f i r s t e x p e r i m e n t s i n a g r i c u l t u r ew e r e m a d e b y t h e N e o l i t h i c N a t u f i a n s o f , f o r i n s t a n c e ,t h e M o u n d C a r m e l c a v e s , a n d t h e m i g r a n t s t o t h e J o r -d a n V a l l e y b r o u g h t w i t h t h e m g r a i n t h a t h a d a l r e a d yp r o d u c e d m u t a t i o n s t h a t m a d e i t m o r e s u i t a b l e f o r a g r i -c u l t u r e .' A s m e n t i o n e d a b o v e , f r o m t h e s e c a v e s w e a l r e a d yh a v e t h e f i r s t e v i d e n c e o f u s e o f f i r e a n d i t s p o s s i b l e a f t e r -e f f e c t s o n e x p a n s i o n o f d r i e r a n d m o r e e x p o s e d s i t e s .O n e s h o u l d n o t d i s c o u n t , t h e re f o re , t h e i m p o r t a n c e o f fi rea s o n e o f t h e e n v i r o n m e n t a l a n d c u l t u r a l t ri g g e rs o f c e r e ald o m e s t i c a t i o n i n t h e f i r e - s w e p t d r i e r g r a s s l a n d s a n d i n t h ef i r e - i n d u c e d m a q u i - e d g e s .

T h r o u g h o u t t h e f o l l o w i n g p h a s e s o f i n t e n s i v e a g r i -c u l t u ra l a n d p a s t o r a l l a n d u s e a n d e s p e c ia l l y f r o m t h eB r o n z e a n d I r o n A g e u n t i l th e H e l l e n i s ti c a n d R o m a np e r i o d , t h e w i d e o c c u r r e n c e o f f i r e i s w e l l d o c u m e n t e d i nt h e B i b le , t h e T a l m u d , a n d H e l l e n is t ic a n d R o m a nl i te r a t u re . B u r n i n g w a s a c o m m o n p r a c t i c e fo r c l e a r i n g o ff o re s t, w o o d a n d s h r u b l a n d , f o r p a s t o r a l u se a n d c u l t iv a -t i o n a s w e l l a s f o r c l e a n i n g o f t h i s t l e s a n d s t u b b l e s i nf i e ld s , a n d o u t o f r e v e n g e a n d s h e e r c a re l e s s n e ss . A . D e r -m a n & Z . N a v e h ( u n p u b l . , c i t e d i n N a v e h , 1 9 7 3 b ) h a v el i s t e d 2 8 r e f e r e n c e s o f f i r e a n d v e g e t a t i o n f r o m t h e O l dT e s t a m e n t a n d L i a c o s (1 97 3) h a s c i t e d m a n y G r e e k a n dR o m a n s o u rc e s w h i c h s ho w h o w c o m m o n n a t u r al a n dm a n - m a d e f ir e s w e r e i n th o s e d a y s . A c c o r d i n g t o L e H o u e -r o u ( 19 7 3) t h e r e i s a l s o e v i d e n c e o f a n c i e n t b u r n i n g o ff o re s ts a n d m a q u i s f o r p a s t u r e i m p r o v e m e n t i n S o u t h e r nF r a n c e . T h i s p r a c t i c e w a s u s e d b y h e r d s m e n i n a l l M e d i t e r -r a n e a n c o u n t r i e s t h r o u g h o u t h i s t o r y ( S e m p l e , 1 9 3 1 ;S h a n t z , 1 9 47 ). V i v i d d e s c r i p t i o n s o f s u c h ' B r a n d k u l t u r e n 'a r e p r o v i d e d b y A n d e r l i n d ( 1 8 8 6 ) i n t h e l a s t c e n t u r y a n dm o r e r e c e n t l y b y R i c k l i ( 19 4 2) .

A t p r e s e n t t h e h u m a n a n d l i v es t o c p r e s s u re i n m a n yM e d i t e r r a n e a n c o u n t r i e s , e s p e c i a ll y i n t h e L e v a n t , h a sb e c o m e s o s e v e r e t h a t l i t t l e g r a s s fu e l a n d d e n s e w o o d yv e g e t a t i o n h a s b e e n l e f t f o r w i l d f i r e s . N e v e r t h e l e s s , L e

H o u e r o u ( 19 7 3) e s t i m a t e d t h a t e a c h y e a r c l o s e t o 2 0 0 0 0 0 h ao f M e d i t e r r a n e a n f o r e s t a n d s h r u b l a n d s a r e b u r n e d a n -n u a l l y ; a n d t h e c a u s e s o f t h e s e w i ld f i re s h a v e n o t c h a n g e dm u c h s i n c e b i b l i c a l t i m e s .

The a da pt a t i o n o f me d i t e r r a ne a n p l a n t s t o f ir eT y p i c a l M e d i t e r r a n e a n ' f i r e b i o c li m a t e s ' (N a v e h , t 9 7 3 b )a r e c h a r a c t e r i z e d b y l o n g , h o t a n d d r y s u m m e r s e a s o n sw i t h m a x i m u m a v e r a g e d a i ly t e m p e r a t u r e s a r o u n d 3 0 C ,a v e r a g e re l a t iv e a ir h u m i d i t e s o f 5 0 - 6 0 % a n d f r eq u e n th e a t w a v e s o f ' s h a r a v ' ( W i n s t a n l e y , 1 9 7 2 ) o r ' s i r o c c o '( w e s t e r n M e d i t e r r a n e a n ) i n t h e b e g i n n i n g a n d t h e e n do f t h e d r y s e a so n , w h e n t e m p e r a t u r e s r i s e a b o v e 4 0 Ca n d r e l a t i v e h u m i d i t i e s c a n d r o p b e l o w 3 0 % .

U n d e r t h e s e c i r c u m s t a n c e s t h e c h a n c e s f o r t h e h i g h l yi n f l a m m a b l e M e d i t e r r a n e a n w o o d y a n d h e r b a c e o u s v e g -e t a t i o n c a n o p i e s t o e s c a p e t h e h i g h t e m p e r a t u r e s o f w i l df i r e s a r e s l i m . A s h a s b e e n s h o w n a b o v e , t h i s v e g e t a t i o nh a s b e e n e x p o s e d t o r e c u r i n g f ir e s f o r s u c h a l o n g p e r i o d ,t h a t o n l y s p e c ie s a n d b i o t y p e s a d a p t a t e d b y p o s t - f ir er e g e n e r a t i o n f r o m u n d e r g r o u n d o r f i r e t o l e r a n t p a r t ss u r v i v e d . T h e s e a d a p t a t i o n s c a n b e u n d e r s t o o d i n ac y b e r n e t i c c o n t e x t a s h o m e o s t a t i c f e e d b a c k c o n t r o l r e -s p o n s e s t o c h a n g e s i n t h e p l a n t - e n v i r o n m e n t s y s t e m , i n -d u c e d b v t h e f i r e. I n t h e se , t h e i n f o r m a t i o n o f t h e c a t a -s t r ophic f i r e event and i t s eco logica l a f te r - e f f ec t s i s pa r t lyt r a n s f o r m e d o r r e - c o d e d , b y s u r v i v a l a n d i n c r e a s e d f r e -q u e n c y o f g e n e s c o n t r i b u t i n g t o f i r e t o l e r a n c e , w h i l s t g e n esc o n t r i b u t i n g t o v u l n e r a b l e n e s s t o f i r e a r e d e l e t e d . . A c -c o r d i n g t o t h e L a w o f R e q u i s i t e V a r i e t y ( R o s s - A s h b y ,1 9 56 ) t h o s e g e n o t y p e s i n w h i c h t h i s i n f o r m a t i o n f e e d b a c kw a s l a r g e e n o u g h t o p e r m i t e n d u r a n c e o r e v a s i o n o f f i reh a d t h e b e s t c h a n c e s o f s u rv i v a l.

P o s i t i v e f e e d b a c k r e s p o n s e s a r e t h o s e w h i c h h e l p i no v e r c o m i n g t h e f i r e h a z a r d a n d i t s a f t e r - e f f e c t s b y i n -c r e a s e d p h y s i o l o g i c a l a c t i v i ty . T h i s i s c h i e fl y e x p r e s se d b yv e g e t a t i v e , f i r e - s t i m u l a t e d r e s p r o u t i n g a n d - o r f i r e - s t i m -u l a t e d s e e d g e r m i n a t i o n a n d p o s t - f i r e f l o w e r a n d s e e dp r o d u c t i o n . E x a m p l e s o f t h es e f r o m r e p r e se n t a t iv e s o f th em a j o r e c o l o g i c a l p l a n t g r o u p s i n t h e M e d i t e r r a n e a nR e g i o n f r o m f i r e - p r o n e u p l a n d e c o s y s te m s a r e s h o w n i nT a b l e 1 .

A s n e g a t i v e f e e d b a c k r e s p o n s e s , o n t h e o t h e r h a n d , c a nb e r e g a r d e d a l l t h o s e d e f e n c e m e c h a n i s m s w h i c h e n a b l e t h ea v o i d a n c e o f t h e f i r e h a z a r d , e i t h e r b y d i r e c t f ir e t o l e r a n c eo f s e ed s o r p l a n t o r g a n s o r b y t h e i r r e d u c e d p h y s i o l o g i c a la c t i v i t y d u r i n g c r i t i c a l f i re p e r i o d s . O u r p r e s e n t k n o w l e d g e

201


4/10

T A B L E i . P O S I T I V E F E E D B A C K R E S P O N S E S O F P O S T - F I R E R E G E N E R A T I O N O F S O M EC 01,NON M E DI T E R R ANE AN PL ANT S

R e g e n e r a t i o nEcological groups a n d p l a n t s p e c i e s Veget. Re,rod,P h a n g r o p h y t e ~Quercus cocoifera, Q , calliprinos~ Q, ithaburensi$ +)iP~st acia lentiscus, P,terebinthus~ P.palaestina, P.at lanti ca +Phyl lire a media~ Olea europea, ~har~lus alaternus,

R . p a l e a s t i n a +C e r a t o n i a siliqua, Laurus nobilis, St yrax officinslis,

M y r t u s c o m m u ni s +T e t r a c l i n i s a r t i cu l a t a, J u n i p e r u s p h o e n i c i a + ) 2A r b u t u s unedo, A,andrachne, S p a r t i u m j u n c e u m + *C a l y co t o m e v i l l o s a ) * E r i c a a r b o re a ~ E . m u l t i f l o r a + +Pinna halepensis)*, P,brutia, P.pin ea +

C =aephyt~sS a c r o p o t e r i u m s p i n os a ) * , R o s m a r i n u s o f f i c i n a li s ,

H e l i a n t h e m u m t u b e r a r i a + ) 3 +T h y m u s c a p i t a t us , S a t u r e j a t h y m b ra ~ T e u c r i u m s p p .

Phlomis lynchnitis + +S a l v i a t r i l o b a, S a l v i a s p p , ~ m~%y o t h e r L a b i a t a e * +C i s t u s v i l I os u s ) *, C . s a l v i f o l i u s ) * + +C i s t us m o n s p e l i e n s i s , C i s t u s s p p . +

C l i m b e r sA r i s t o l o c h i a p a r v if o l i a, A . s p p . , A s p a r g u s ~ h y l l u s ,

Aspar agus acutefolius, A,spp. + ) 4Smil~ x aspera, Tamu s ccam~unis *Clematis, eirrbosa, Rubi a tenuifolia, R. spp. +Geo~h ztes , Hemicr~p%o~h~tes~ P e r e n n i a l h e r b sO p h r is , s p p . , O r c h i s s p p . , A s p h o d e l u s m i c r o a r p u s , A . s p p, +I r i s a p p . , A l l i u m s p p . ~ N a r c i s u s tazette,

C y c l a m e n p e r s i c u m a n d m a n y o t h e r g e o p h y t e s + +Eryllgium falcatum, Dory cnum suffructosum, Echinops spp. + +

C a r l i n a i nv o l u c ra t a , C e p h a l a r i a l e u c ~ t h a , O n o n i s a p p . + +g a n g u i s o r b i a m i n o r , G a li n a c o r r u d a e f o l i ~ ~ m a n y o t h e r s +

Perennial herbs

P e r e n n i a l g r a s s e sO r y z o p s i s m i l i a c e a ) % O , c a e r u l e s c e n s ) * , S t i p a b r o m o i d e s ) * + +B r a e h y p o d i u m r a m o su m , B . p i n na t u m , P e s t u c a a r u n d i n a c e a + +A r r h e n a t h e r u m elatus, Phalaris tuberosa, gromus syria cus * +Dactylis glomerata, }lordetgnbulbosum)*, Poa bulbosa, + +H y p a r r b e n i a h i r t a ) * , A n d r o p o g o n distachius)* + +

T h e r o p h z t e sA v e n a sterilis)*, A. spp., Stipa tortilis)*, gromus spp.A e g i l o p s s p p . a n d m a n y o t h e r a n n u al g r a s se sL o t u s p e r e g ri n u s )* , P s o r o l e a b i t u m i n o s a , T r i f o l i u m s p p .

M e d i c a g o p o l y m o r p h a) * M , s p p . a n d m a n y o t h e r l e g u m e sA n t h e m i s a p p. C r e p i s s p p . , S e n e c i o s p p . , S o n c h u s a p p . *S c a b i o s a s p p . +E r o d iu m s p p ., G e r a n i u m s p p . , C e p h a l a r i a j o p p i ca ) * +A n a g a l l i s c o e r a l a ) *, L a g o e c i a c o m m u ni s ~ m a n y o t h e r

a n n u a l p I ~ n t s +

) 1 = r e s p r o u t i n g f r o m u n d e r g r o u n d s t u m p s a n d r O O t c r o w n s a n d a d v e n t a t i v er o o t s ( s u c k e r s )

) 2 = r e s p r o u t i n g f r o m u n d e r g r o u n d s t u m p s a n d r o o t c ro w n s , n o s u c k e r s)g = r e s p r o u t i n g f r o m r o o t cromls)4 ~ r e s p r o u t i n g f r o m ro o t s a n d u n d e r g r o u n d b u l b s o r corms, tubers an d

o t h e r r e g e n e r a t i o n buds.)* p r o n o u n c e d p o s t - f i r e g e r m i n a t i o n o b s e r v e d i n f i e l d e x p e r i m e n t s in

Israel.

20 2

i s fa r to o l im i ted to g iv e a d e ta i l ed a c co u n t o f a ll th ese f i rea d a p ta t io n s in d i f f eren t p la n t s a n d th e i r g en et i ca l , p h y s io -l o g i c a l a n d m o r p h o l o g i c a l m o d e o f a c t i o n . F o r t h i sp u r p o s e m u c h m o r e e x t e n s i v e f i e l d a n d l a b o r a t o r y w o r ki s n ecessa ry th a n th e s tu d ies n o w ca rr i ed o u t in th i s reg io na n d s u m m a r i z e d i n t h e T a l l T i m b e r F i r e E c o l o g y C o n -feren ce a t T a l la h a ssee , F lo r id a in M a rch 1 9 7 3 , b y L eH o u e r o u , L i a c os , N a v e h a n d T r a b a u t .

L e H o u e r o u ( 1 9 7 3 ) , r e v i e w i n g t h e e f f e c t o f f i r e o n M e -d i t er r a n e a n v e g e t a t i o n t y p e s , d i s t i n g u i s h e d b e t w e e n a c t i v ea n d p a ss iv e p y r o p h y te s . T h i s c la s s i f i ca t io n i s c lo se ly re la tedt o d o m i n a n c e o f p o s i t i v e o r n e g a t i v e f ir e f e e d - b a c kr e s p o n s e s . T r a b a u d ( 1 9 7 3 ) s t u d y i n g t h e r e s p o n s e o f g a r i g u ep l a n t s i n S o u t h F r a n c e t o d i f f er e n t f r e q u e n c i e s a n d t i m i n g so f f i re , h a s g ra d ed th ese p la n t s a cco rd in g to p o s t - f i rereg en era t io n ca p a c i t i e s to f req u en t b u rn in g . In ev a lu a t in gt h e s e r e su l t s it s h o u l d b e k e p t i n m i n d t h a t o n l y d e n s e a n dw e l l d e v e l o p e d m a q u i a n d g a r i g u e s t a n d s , w h i c h h a v ereco v ered 5 y ea rs a n d m o re a f t er th e f i re , a re h ig h ly in -f l a m m a b l e . W e c a n , t h e r e f o r e , a s s u m e t h a t t h o s e f i r ef r e q u e n c i e s w h i c h h a d g r e a t e s t e v o l u t i o n a r y s i g n i f i c a n c ef o r t h e se w o o d y p l a n t s a r e p r o b a b l y n o t g r e at er t h a n o n c ei n 5 - 1 0 y e a r s .

A s c a n b e s e e n i n T a b l e 1 , m o s t s c l e r o p h y l l t r e e s a n dsh ru b s , a s w e l l a s c lim b ers , re ly so l e ly o n v e g e ta t iv e re -g e n e r a t i o n a n d a r e t h e r e f o r e ob l i g a t o r y r oo t r e sp r ou t er s .O n t h e o t h e r h a n d , a l l c h a m a e p h y t e s , a s w e l l a s h e r b a c e o u sp eren n ia l p la n t s a re f a cu l t a t i v e r oo t r e sp r ou t er s . T h e y m a yresp ro u t v eg e ta t iv e ly b u t a l so reg en era te b y f i re - s t im u la teds e e d g e rm i n a t io n a n d g r o w t h , a n d m o s t o f t h e m p r o d u c eseed s f r o m resp ro u t in g p la n t s a l rea d y in th e f i r s t y ea r a f tert h e f i r e a n d f r o m ( v o l u n t e e r i n g ' s e e d l i n g s ( f r o m s e e d sa lrea d y in th e so i l a t th e t im e o f f i re ) in th e seco n d y ea r(Naveh , 1960 , 1973a) .

M ed i terra n ea n co n i f ers , l ik e P inu s ha lepens i s a n d P . b r u -t i a d o n o t r e s p r o u t a t a l l , b u t r e l y s o l e ly o n p o s t - f i r e s e e dg e r m i n a t i o n f r o m c o n e s t h a t b u r s t o p e n f r o m t h e h e a t o fth e f i re . T h ey a re , th ere fo re , ob l i ga to r y seed regenera to r sa n d f i r e a p p a r e n t ly p r o v i d e s t h e o n l y o p p o r t u n i t y f o r t h e irn a t u r a l r e g e n e r a t i o n u n d e r a d e n s e m a q u i u n d e r s t o r y(Wa l ter , 1 9 6 8 ; N a v eh , 1 9 7 3 a ).F i r e r e g e n e r a t i o n b e h a v i o u r i s c l o s e l y l i n k e d w i t h r e -s p o n s e t o s e a s o n a n d w a t e r a v a i l a b il i ty . T h e o b l i g a t o r y r o o tr e s p r o u te r s a r e s c l e r o p h y l l o u s , m o s t l y e v e r g r e e n , d r ough t -endu r i n g a n d su m m er-a c t iv e t rees a n d sh ru b s .

B u t th e fa cu t ta t iv e ro o t resp ro u ters a re ty p ica l lydrough t -evade rs a d a p t i n g t o t h e d r y s u m m e r s b y m o r eres tr i ct ed p h y s io lo g ica l a c t iv i ty a n d esp ec ia l ly b y red u ct io no f th e i r t ra n sp ira t io n su r fa ce (O rsh a n , 1 9 7 2) . T h e y co rn -


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mence re sprout ing only after the f i rs t winter rains. Obl ig-atory resprouters , in contrast , s ince they rely on deepand wel l -branched root systems, can begin to resprout bythe midd le o f the summer immedia te ly a f te r the f i re andcont inue intensive growth at least through the f i rs t andsecond summer to rega in the i r fo rmer , dom inan t pos i t ionafter a few years ' t ime. As tong as their young suckers andshoots are soft and lush and their leaves are not yet l ignifiedand thorny, they are browsed heavi ly by herbivores . Thus,as s tated by Mooney & Dunn (1970), postf i re grazingpressure has acted as an addi t ional powerful select iveagent , favoring those species and biotyp es which very soondeveloped hard, thorny or dis tas teful leaves and twigs,but also those with highest vegetat ive regenerat ion ca-paci t ies to overcome defol iat ion s t resses or both f i re andgrazing. We may also assume that this re-coding of in-fo rmat ion f rom f i re and g raz ing has p readap ted them tofurther defol iat ion catast roph es from cut t ing and cop-picing.

As outs tanding examples of successful evolut ionarystrategies for maximizat ion of over-al l survival threesc lerophyt lous smal l t rees can be used- the Wes t Me-d i t e r ranean Kermes oak = Quercus coccifera and i ts EastMedi te r ranean v icar i ad = Q. calliprinos, and Pistacialentiscus. The f i rs t , dominat ing burned garigue on aneroded rendzina near Montpe l l i e r , Sou th France , p roduced374 g/m 2 above g round biomass in the f i rs t year after thefire and 874 g/m 2 in the sixth yea r (S pecht, 1969). T hesecond, in a mixed garigue on dar k rendzina in the Weste rnGal i lee, in Israel , regained i ts pre-burn height of 2 ,5meter in the second summ er and gained, together with otherobl igatory respro uters 60 ~o of the g roun d cover in thesecond summer an d 80 ~ in the thi rd summer after theburn (Naveh, 1960, 1973b). In the East Medi terraneanzone, Pistacia lentiscus i s the most tenaceous evergreenshrub, res is tant to drough t , f i re, grazing, and cut t ing, andremaining as the las t wood survivor in degraded maquisand wood lands (N aveh & Dan, 1973). By resprout ingvigorously, not only f i 'om extensive deep and lateral lybranched and rock penet ra t ing roo ts , bu t a l so by roo t ingfrom rapidly spreading, prostrate and very leafy twigs,i t very soon fo rms a dense and compact sh rub canopywith high soi l and water conserving features . (Figure 1).I ts smal t , sclerophyt lous leaves are highly resinous and verysoon become dis tasteful (Naveh, i960). In this way i tsp o t en t i a l s a r e max i mi zed -mo i s t u r e an d n u t r i en t ex -t rac t ion , energy and water t rapp ing and pho tosyn the t i cact ivi ty , carbohydrate reserve s torage, hydro-ecologicalefficiency, and post-def ol iat ion recuperat ive powers .

In most facul tat ive root respouters , posi t ive feedback offi rest imulated vegetat ive and reprodu ct ive regenerat ion iscoupled with morphological and physiological plast ici tyand agressiveness in invasion and establ ishment o f newly-opened, f i re-denuded but mineral-r ich habi tats . Thus f i rehas favoured no t on ly the t enaceous maqui dominan t s ,but also the evolut ion of opportun is t ic and poten t ial ly fast -g rowing subord ina te sp ec ies - dwarfshrubs , hemicryp to-phy tes , perenn ia l g rasses , geophytes , and therophytes -that remain for years as suppressed rel ics near t ree andshrub edges, rock outcrop s and shal low soi l patches, unt i ltheir next fire-induced upsurge (Naveh, 1960, 1973a).

Al l woody facu l t a t ive roo t resprou ters a re low un-pa la tab le thorny o r a romat ic chamaephytes , l ike Sarco-poterium spinosum, Cistus, Salvia and Erica spp . and manyothers . Under heavy postf i re grazing pressure they havebest changes of survival and can create monotonous and

Fig. 1. Fire-, drought- and grazing tolerant Mediterraneanplant community in the Western Galilee. Israel. In the back-ground Ceratonia siliqua an d Quer cus callipinos trees, in thecenter Pistacia lentiseus shrubs and in the front Sarcopoteriumspinosum dwarfshrnbs with tall flowerstalks of Hordeurn bul-bosum and flowering annual plants.203


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e v e n -a g e d b a t h a s t a n d s ( L e H o u e r o u , 1 9 7 3 ) . A m o n g s tt he mos t succesfu l he rbaceous f i r e fo l l owers and eco toneco lon i ze r s i n no t t oo heav i ly g razed maqu i s and ga r iguea r e x e r o m o r p h i c , s h a d e t o l e r a n t a n d m o s t l y c a lc i p hi lpe renn i a l bunchgra sse s . Thus B r a c h y p o d iu m r a m o s u m h a sshown the h ighes t ' pos t - f i r e r egene ra t i on i ndex ' (T rabaud ,1 9 7 3 ) a n d d o m i n a t e s f r e q u e n tl y b u r n e d , m o d e r a t e l yg r a z e d g a r i g u e i n S o u t h F r a n c e ( L e H o u e r o u , 1 9 7 3 ) .I t s Eas t Med i t e r r anean , e co log i ca l cou n te rpa r t i s O r y z o s p i sm i l i aeea , which exce l l ed i n vege t a t i ve and r ep roduc t iver e g e n e r a t i o n a n d f o r a g e p r o d u c t i o n a f t e r m a q u i a n d g a -r i gue burns i n I s r ae l (Naveh , 1960) . Inc rea sed ge rm ina t i onin t h is g ra ss , a f t e r hea t i ng t he seeds up t o 90 C , ha s beenshown by Me i r i (1959) . I t ha s a l so been wide ly used a s are seeded pa s tu re g ra ss - 'Smi lo ' - a f t e r chap a r ra l bu rn i nCa l i fo rn i a and ha s na tu ra l i z ed i t se l f we l l i n t he se con-d i t i ons a s a p ro l i f i c f i r e fo l l ower (Love & Jones , 1952) .These g ra sse s have adap t ed t hemse lves t o t he f l uc tua t i ngrad i a t i on , m oi s tu re and nu t r i en t r eg imes o f edge hab i t a t sby a r emarkab l e hydro - and pheno-eco log i ca l p l a s t i c i t y i ng r o w t h r h y t h m a n d h a b i t a n d i n t i m i n g o f r eg e n e r a t io nf r o m b a s a l b u d s , a s w e ll a s b y a u n i q u e s h r u b -l ik e ' w h o l eshoo t r egene ra t i on ' c apac i t y f rom in t e rca l a ry mer i s t ema n d a x i l a r y b u d s a l o n g t h e c h a r r e d o r d o r m a n t c a l m(Naveh, 1960) .

I n T a b o r o a k = Quercus i thaburensis , t h e m a i n t r e e o fs u b - m e d i t e r r a n e an o p e n w o o d l a n d s w i t h a d e n s e, s u m m e r -d ry he rbaceo us unde r s to rey , t he t h i ck , f i re - r e s is t an t ba rk i san e f f i c i en t de fence mechan i sm fo r f i r e su rv iva l , becausehe re t he f i r e t empe ra tu re s a re no t so h igh a s i n dense ,w o o d y m a q u i s t a n d s . A t t h e s a m e t im e , h o w e v e r , T a b o roak i s an e f f i c i en t ob l i ga to ry roo t - r e sp rou t e r . The annua lp l a n t s t h a t d o m i n a t e t h e h e r b a c e o u s c a n o p y o f t h e s ew o o d l a n d s , a s w e ll a s t h o s e o f o p e n M e d i t e rr a n e a n a n dsemi -a r id S t eppe g ra ss l ands which a re even more f i r e -p r o n e a n d m a y b u r n y e a r a f t er y e a r i f n o t g r a z e d t o o h e a v i l y(Naveh, 1973a) , a re most successful f i re -fol lowers. Na-tu ra l l y , i n t he se t he roph y te s r e spo nses t o f i re , a s we ll a s t om a n y o t h e r s t re s s f ac t o rs , a r e c e n t r e d a r o u n d r e p r o d u c t i v ea n d g r o w t h b e h a v i o u n T h u s e a r l y a n d p r o l i f i c s e e d p r o -duc t ion , e a r ly seed shedd ing , and d i s t r i bu t i on by e f fi ci en td i spe r sa l mechan i sms a re cha rac t e r i s t i c fo r mos t t he ro -p h y t e s ( Z o h a r y , 1 9 6 2) , S e e d d o r m a n c y a n d p o l y m o r p h i s m

- fou nd e spec i a ll y i n l egumes - a l so i nc rea se t he changesfo r a t l e a s t pa r t o f t he seed c rop t o e scape f i r e and env i ron-me nta l r igor . O f specia l va lue in th is respect i s t r y p a n o c a r p y

- t he capac i t y o f d i sseminu le s o f ann ua l a nd pe renn i a lg ra sse s and some o the r abundan t spec i e s l i ke E r o d i u m ,t o ' d r i l l ' t hemse lves i n to t he so i l wi th t he a id o f hyg roscop ic20 4

awns , c a l l ous t i p s , and o the r t o r s ion mechan i sms . Bu tthe re a re appa ren t ly a l so spec i f i c f i r e f e edback r e sponsesre l a t ed d i r ec t ly t o t he h igh f i r e tempe ra tu re s , such a s hea tt o l e rance . Th i s ha s been p roved fo r va r ious success fu lM e d i t e rr a n e a n g r a s s i n v a d e rs o f C a l if o r n i a, s u c h a sB r o m u s m o I l i s a n d B . r i g i dus , E I ym us capu t - m edusa , a n do the r s (Sam pson , 1944; M cKe l l e t al ., 1962 ) . In t heT a b o r - O a k w o o d l a n d i n I s r a e l , t h e d o m i n a n t A r e n aster i l i s no t on ly r e t a ined v i ab i l i t y a f t e r hea t i ng t he seedsf o r h a l f a n h o u r u p t o 1 25 C b u t e v e n g a v e i n c r e as e dge rm ina t i on a f t e r t em pe ra tu re s up t o 105 C . Af t e r aho t g ra ss f i re when su r face t empe ra tu re s r eached 350 Cfor severa l se cond s and 51 C in 1 cm soi l depth, A r e n aseeds wi th cha r red awn t i ps wh ich had a l r eady bur r i edthe i r c a ryops i s ve r t i c a l l y i n t he uppe r so i l su r face , ge r -m i n a t e d i n t h e f o l lo w i n g w i n te r . I n o u r r e c e n t b u r n i n gs t u d ie s ( A . N a v e h & Z . N a v e h , 1 9 7 3 u n p u b . ) , 2 0 ~ o o fm ore o r l e ss cha r red A rena s t e r i l i s seedl ings protec ted by adense g ra ss mulch l aye r emerged .

Stipa tort i l is, t h e d o m i n a n t a n n u a l g r a s s o f th e s e m i-a r i d a n d f i re - p r o n e s te p p e g r a s s la n d s o f t h e U p p e r J o r d a nVa l l ey , is one o f the mo s t r ap id seed bur i e r s because o f i t ssma l l seeds . In t he l a s t d ry w in t e r on ly t hose seeds whichw e r e c o v e r e d w i t h 1 - 2 c m s oi l em e r g e d b o t h i n t h e b u r n e da n d u n b u r n e d p l o t s ( Z . N a v e h , S . S h a l o m & Z . Z a i t h ,1973 unpub . ) . T hese bur i ed seeds cou ld no t on ly e scapethe h igh su r face f i r e t empe ra tu re s , bu t a l so ge rmina t eu n d e r m o r e f a v o u r a b l e m o i s t u r e a n d t e m p e r a t u r e r e g i m e sand p robab ly a l so e scape g raz ing and seed-co l l e c t i ngan ima l s . In t he se g ra ss l ands , pe renn i a l g ra sse s and geo-phy te s a l so demons t ra t ed mos t success fu l s t r a t eg i e s fo rmax imiza t i on o f ove r -a l l d rou gh t , f ir e, and g raz ing su rv iva lpo t en t i a l s . Thus H o r d e u m b u l b o s u m a n d P o a b u l b o s u mc o m b i n e a ll t h e a b o v e d e s c r i b ed r e p r o d u c t i v e a d a p t a t i o n sof t he ann ua l g ra sse s wi th vege t a t i ve pos t - f ir e r e sp rou t ingf r o m u n d e r g r o u n d b u l b s . G e o p h y t e s l i k e A s p h o d e l u srnicrocarpus , which a re comple t e ly unpa l a t ab l e , r e s i s teven t he g rea t e s t f i r e and g raz ing p re ssu re and domina t et h e m o s t d e g r a d e d s t a g e s o f th e s e g r a s sl a n d s a n d b a t h a s(W al t e r , 1968 ; Naveh & Dan , 1971) .

Of specia l in terest a re prol i f ic perennia l grass f i re -fo l l owers o f t rop i ca l o r i g in l i ke H y p p e r h e n i a h i r t a , a b u n -d a n t o n r o c k y a n d s u n n y s l o p e s in o p e n m a q u i a n d b a t h a ,a s we l l a s i n f r equen t ly bu rned med i t e r r anean and semi -a r id g ra ss l ands . Th e capac i t y o f t he se g ra sse s t o r egene ra t ef r o m t h e i n t e c a la r y m e r i st e m s o f c h a r r e d c u l m s a n d b a s a lbuds , even i n t he midd le o f t he sum me r i n a r i d hab i t a t s(Naveh , 1960, t 973a ) , m igh t be a r el ic o f t he i r rhy th m o fg rowth and pos t - f i r e r egene ra t i on i n t rop i ca l savannas .


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D isc uss ion and c onc lus ions

A l t h o u g h n a t u r a l f ir es a r e a t p r e s e n t o f m i n o r i m p o r t a n c ein t he Medi t e r r anean Reg ion , t h i s was no t t he ca se i nea r l i e r t im es when l i gh tn ing and vu l can i c e rup t ions cou ldhave caused sweep ing f i r e s ove r l a rge a rea s . As Axe l rod( t958) ha s shown in Ca l i fo rn i a , f i r e s a l so p l ayed an im -p o r t a n t r o l e t o g e t h e r w i th d r o u g h t i n p o s t - G l a c ia l e v o l u -t i o n o f M e d i t e r r a n e a n p l a n ts , c o m m u n i t ie s a n d e c o sy s te m s .Bu t i n add i t i on t hey appa ren t ly a l so se rved a s an env i ron-m en ta l s t im ulan t fo r cu l t u ra l evo lu t ion .

T he pa l eon to log i ca l r e co rds i n t h i s r eg ion a re s t i l l t oof r a g m e n t a r y t o p r o v i d e a b a s i s f o r a l l o u r a s s u m p t i o n s .Bu t t he pa lyno log i ca l ev idences o f t he abu nda nce o f f ir e-t o l e ran t sc l e rophy l l and f i r e -dependen t con i fe r t r e e s 60 -8 0 ,0 0 0 y e a r s a g o , a n d o f t h e p r e s e n ce o f w o o d a s h i n t h ec a v e s o f t h e P a l e s ti n i an N e a n d e r t h a l m a n 6 0 - 4 0 , 00 0 y e a r sa g o g i v e s t r o n g s u p p o r t t o t h e m . U p t o n o w o n l y i n n o r t h -w e s t E u r o p e , u n d e r m u c h m o r e f a v o u r a b l e c o n d i t i o n s f o rpa l eo -eco log i ca l r e sea rch , have cha rcoa l ho r i zons g ivenp r o o f o f l a rg e - sc a le a n d a p p a r e n t l y d e l ib e r at e b u r n i n g o fp ine fo re s t s by Uppe r Pa l eo l i t h i c hun t e r s (Bu tze r , 1972) .I n t h i s r e g i o n t o o , b e c a u s e o f t h e p r e o c c u p a t i o n w i thc l im a t i ca l l y de t e rm ined o rgan i c and cu l t u ra l evo lu t i on ,t oo l i t tl e a t t en t i on ha s been pa id t o t he im p ac t o f p r im i t ivem a n o n h i s h a b i t a t b y b u r n i n g a n d h u n t i n g . I n t h e M e -d i t e r ra n e a n r e g i o n t h e c o l d w i n t e rs a n d d r y s u m m e r s o f t h eU p p e r P l e i s to c e n e a n d e a r l y H o l o c e n e a p p a r e n t l y c r e a te dcond i t i ons w hich we re no l e ss f avourab l e fo r t he use o f f i refo r hea t i ng and c rea t i ng fo re s t t h i cke t open ings t han i nE u r o p e . H e r e a l s o t h e c o l le c t io n o f b ig , p a r c h e d g r a i nke rne l s a f t e r f i r e m ay have been a f i r s t s t ep t oward d ry -l a n d f a r m i n g a n d g r a i n d o m e s t i c a t i o n d u r i n g t h e l a t eMes01 i th i c pe r iod , m ore t han 10 ,000 yea r s ago . Fromea r ly h i s to r i c a l t im es onw ard , t h e wide occu rence o f f ir e onMedi t e r r anean wi ld l ands i s we l l docum ented i n t he B ib l eand l a t e r source s . Mo s t o f t he ea r l ie r c ause s , such a sb u r n i n g o f f o r e s t a n d s h r u b l a n d f o r p a s t u r e i m p r o v e m e n tappa ren t ly p reva i l ed t h roughou t h i s t o ry un t i l t he p re sen t .

W e m ay , t he re fo re , a ccep t t he ro l e o f fi r e a s a l ong- t e rmand im por t an t s t r e ss f ac to r and se l ec t i ve fo rce i n Me-d i t e r r anean ecosys t em s . F i r e ope ra t ed f i r s t a s an i ndepen-den t c l imat ica l and geolog ica l externa l s t ress, a nd la ter asp a r t o f t h e m u l t iv a r i a te a n t h r o p o g e n i c b i o f u n c t i o n , i nw h i c h m a n b e c a m e t h e d o m i n a n t c o n t r o l l i n g e c o s y s t e mf a c t o r (N a v e h & D a n , 1 9 71 ). A s M o o n e y & D u n n ( 19 7 0)have show n so c l ea r ly , f ir e ha s a l so been a se lec ti ve fo rce i no the r m e d i t e r r anean b iom es an d i s c lose ly i n t e rwoven wi tho t h e r e n v i r o n m e n t a l s t r e s s f a c t o r s t h a t b r o u g h t a b o u t

c o n v e r g e n c e i n f o r m a n d f u n c t i o n . A m o r e t h o r o u g h ,com para t i ve s t udy o f t he e f f ec t o f f ir e on s t ruc tu re andf u n c t i o n o f m e d i t e r r a n e a n -t y p e c l i m a t e p l a n t s a n d c o m -m uni t i e s wo u ld be de s i r ab l e i n o rde r t o de f ine m ore c lose lyth i s conve rgen ce i n r e spec t t o f i r e behav iour .

F r o m o u r s t u d y i n t h e M e d i t e r r a n e a n R e g i o n i t i s a p -pa ren t t ha t f i r e r e sponse i s c lose ly l i nked wi th t he eco -l o g ic a l s ta t u s o f t h e p l a n t w i t h in t h e c o m m u n i t y a n d t h ecoenoc l ine , i t s e co log i ca l r equ i rem en t s t b r ge rm ina t i on ,ece s i s and com pe t i t i on . As Ande rson (1956) ha s shown inCa l i fo rn i a , f i r e - i nduced eco tones and d i s tu rbed sh rubopen ings can p rov ide i dea l oppor tun i t i e s fo r fu r the rspec i a t i on , hybr id i sa t i on and geno type r ecom bina t i on .T h e y c r e a t e d n o t o n l y f a v o u r a b l e c o n d i t i o n s f o r l i g h t -d e m a n d i n g d w a r f s h r u b s a n d h e r b s b u t a l s o f a c i li ta t ed t h einvas ion o f m o re xe ri c e l em en t s f rom ad j acen t , d r i e rr eg ions , i nc lud ing weedy and ag re ss ive annua l g ra sse s ,am ong s t t he se t he p rogen i to r s o f ou r c e rea l g ra ins . I t is o fi n t e re s t t o no t e t ha t t h e m o s t success fu l i nvade r s o f Ca l i -f o r n i a fo o t h il ls f r o m t h e M e d i t e r r a n e a n R e g i o n a r e s o m eof t he se annu a l g ra ss f i r e - fo ll owers , such a s Bromus, Arenaa n d a l s o Erodium,w h i c h w e r e a l s o p r e - a d a p t e d t o p a s t o r a land ag r i cu l t u ra l d i s t u rbances , and we re i n t roduced t oC a l i f o r n ia b y t h e S p a n i a r d s ( N a v e h , I 9 6 7 ).

T h e e v o l u t i o n o f n a tu r a l p l a n t c o m m u n i t i e s h a s r e c e n t lyb e e n d i s cu s s ed b y W h i t t a k e r & W o o d w e t l (1 9 72 ). M a n ycen tu r i e s o f seve re h um an and l i ve s tock p re ssu re , r e -su l t i ng i n i nd i sc r im ina t e bu rn ing , Cu t t i ng and g raz ing onnon-a rab l e wi ld l ands and i n bushc l ea r ing , t e r r ac ing ,cu l t i va t i ng , and l a t e r -on abandon ing and neg l ec t i ng o fa r a b l e p o r t i o n s ( N a v e h & D a n , 1 9 7 1 ) , h a v e i n d u c e d s om any re t rogre ss ive s t ages f rom the p r i s t i ne c l im ax andhave d i s to r t ed t h i s evo lu t i ona ry p rocess so m uch , t ha t i t sr econs t ruc t i on ha s becom e ve ry d i f f i cu l t . T h i s ha s causedsom e quas i - s t ab l e vege t a t i on s t ages t o be confused wi thc l im ax com m uni t i e s , r a the r t han r ega rded a s vege t a t i ontypes t ha t p reva i l unde r spec i f i c com bina t i ons o f env i ron-m e n t a l c o n d i t io n s a n d p a r t i c u l a r r e g im e s o f h u m a n d i s-t u r b a n c e o r p r o t e c t i o n ( N a v e h , 1 9 7 1 ) . A n y a t t e m p t a tr e c o n s t r u c ti o n o f th e t r u e c l im a x c o m m u n i t i e s s h o u l dt ake i n to accoun t t he f ac t t ha t f i r e ha s been an i n t eg ra lpa r t o f t he i r evo lu t i ona r y p rocess , a ff ec t i ng t he i r com p os i -t i on , s t ruc tu re , p roduc t iv i t y , and nu t r i en t c i r cu l a t i on f romthe i r e a r l ie s t s t ages o f evo lu t i on .

W h i t t a k e r & W o o d w e l l ' s ( 1 9 7 2 ) s u g g e s t i o n t h a t a l -t e rna t i ve evo lu t i ona ry r e sponses t o t he e ff ec ts o f f ir e a sin f luenced by c l im a te , so il , t opo gra ph y and f i r e f requenc yhave been r e spons ib l e fo r t he c rea t i on o f g ra ss lands ,sh rub l ands , and wood lands i n sem i -a r id c l im a te s seem s

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a l s o t o b e r e l e v a n t f o r t h i s r e g i o n , I n t h e d r i e r r e g i o n s ,t h e r e s t r ic t e d w i n t e r r a i n f a l l a n d l o n g s u m m e r d r o u g h t ,c o u p l e d w i t h g r e a t f i re h a z a r d , h a s f a v o u r e d t h e d o m i n a t i o no f s h o r t -s e a s o n e d , d r o u g h t a n d f ir e a v o i d i n g t h e ro p h y t e sa s w e l l a s p e r e n n i a l g r a s s e s , h e m i c r y p t o p h y t e s a n d g e o -p h y t e s w i t h s i m i l ar g ro w t h r h y t h m s a n d e c o l o g i c a lr e q u i r e m e n t s . H i g h e r r a i n f a l l a n d b e t t e r s o i l - a n d - r o c km o i s t u r e r e g i m e s , l e s s f r e q u e n t b u t m o r e i n t e n s i v e f i r e s ,h a v e p r o b a b l y fa v o u r e d t h e d o m i n a n c e o f d r o u g h t t o l e r a n t,s c l er o p h y l l o b l i g a t o r y r e s p ro u t i n g p h a n e r o p h y t e s b u t h a v ea l s o o p e n e d n i c h e s f o r d r o u g h t a n d f i re a v o i d i n g f a c u l t a t i v er e s p r o u t i n g d w a r f s h r u b s a n d h e r b a c e o u s u n d e r s t o r e y .S o m e w h e r e a l o n g t h i s m o i s t u r e a n d f i r e e c o c l i n e , t h ed w a r fs h r u b - d o m i n a t e d b a t h a c o m m u n i t i e s c a n b e f o u n d .I t s h o u l d a l s o b e a s s u m e d t h a t i n t h e s e e c o s y s t e m s b e f o r et h e e f f e ct s o f m a n y , t h e g r a z i n g p r e s s u r e o f h e r b i v o r e s ,w i t h o u t b e i n g d e s t r u c t i v e , w a s s u f f i c i e n t ly se v e r e t o a c t a sa n a d d i t i o n a l s e l e c t i v e a n d e v o l u t i o n a r y f o r c e t h r o u g ha d a p t i v e f e e d b a c k r e s p o n s e s . A s s t a t e d b y W h i t t a k e r( t9 7 0 ) i t m a y h a v e i n d u c e d - a m o n g s t o t h e r e f f e c t s -t h e d e v e l o p m e n t o f s e c o n d a r y c h e m i c a l s u b s t a n c e s a sb i o c h e m i c a l d e f e n c e s o f p l a n t s a g a i n s t g r a z i n g . A t t h e s a m et i m e , s o m e o f t h e s e s e c o n d a r y c h e m i c a l s u b s t a n c e s , s u c ha s e t h e r i c o il s , m a y h a v e i n c r e a s e d t h e i n f l a m m a b i l i t yo f t h e s e w o o d y p l a n t s u n d e r s u i t a b l e m e t e o r o l o g i c a lc o n d i t i o n s . T h e y m a y a l s o h a v e c a u s e d t h e a c c u m u l a t i o no f a l l e l o p a t h i c , g e r m i n a t i o n - i n h i b i t i n g a g e n t s i n t h e l i t t e ra n d s o i l , a s i n d i c a t e d b y r e s e e d i n g t ri a l s in t h e l i t t e r a n d d u f fa f t e r r e m o v a l o f t h e d e n s e m a q u i a n d g a r i g u e c a n o p y( N a v e h , 1 9 60 ). T h e s e ' k o l i n e s ' a re a p p a r e n t l y d e s t r o y e d b yt h e h i g h f i re t e m p e r a t u r e s t h a t c o n v e r t t h e l i t t e r a n d u p p e rh u m u s l a y e r in t o a f a v o u r a b l e se e d b e d .

T h u s w e c a n r e a c h t h e c o n c l u s i o n t h a t f i r e h a s c o n t r i -b u t e d n o t o n l y t o b i o l o g i c a l d i v e r s it y o f g e n o t y p e s , b u t a l s ot o t h a t o f M e d i t e r r a n e a n c o m m u n i t i es , t h e ir c o m p o s i t i o n ,s t r u c t u r e , a n d n i c h e d i f f e r e n t a t i o n . T h i s i s a l s o t r u e f o rg r a z i n g . B o t h , t h e r e f o r e , c a n n o l o n g e r b e r e g a r d e d a se x t r i n s i c ' d i s t u r b a n c e ' f a c t o r s o f r e c e n t o r i g i n , b u t a r ei n t e g r al p a r t s o f M e d i t e r r a n e a n e c o s y s t e m s a n d t h e i re v o l u t i o n ; a n d t h e y s h o u l d b e s t u d i e d a s su c h .

SummaryF i r e h a s p l a y e d a d e c i s i v e r o t e i n P o s t - G l a c i a l b i o l o g i c a la n d c u l t u r a l e v o l u t i o n i n t h e M e d i t e r r a n e a n R e g i o n . i t se v o l u t i o n a r y i m p a c t o n p l a n t s h a s b e e n m a n i f e s t e d b yf e e d b a c k r e s p o n s e s , i n w h i c h t h e f i r e a n d i t s a f te r - e f fe c t ss e le c te d p l a n t s f o r p h y s i o l o g i c a l a n d o t h e r m e c h a n i s m s t h a t20 6

e n a b l e d i r e c t f i r e t o l e r a n c e o r p e r m i t a v o i d a n c e f o l l o w e db y v e g e t at i v e a n d r e p r o d u c t i v e r e g e n e r a ti o n .

T h e d o m i n a n t , s c l er o p h y l l, d r o u g h t - t o l e r a n t p h a n e r o -p h y t e s o f t h e m a q u i a r e o b l i g a t o r y r o o t s p r o u t e r s , w h e r ea st h e s u b o r d i n a t e , d r o u g h t - e v a d i n g c h a m a e p h y t e s , a s w e l la s h e r b a c e o u s p e r e n n ia l s , a re b o t h v e g e t a ti v e a n d r e p r o -d u c t i v e r e g e n e r a t o r s a n d a r e w e l l a d a p t e d t o n e w , f i r e -d e n u d e d h a b i t a t s . A n n u a l a n d p e r e n n i a l g r a s s fi r e - fo l l o w e r sa r e a l s o a b l e t o e s c a p e h i g h s u r f a c e f ir e t e m p e r a t u r e s w i t ht h e a i d o f t o r s i o n d e v i c e s o n s e e d s . E v o l u t i o n a r y s t r a t e g ie st o o v e r c o m e f i re a r e c lo s e l y i n t e r w o v e n w i t h t h o s e a g a i n s to t h e r e n v i r o n m e n t a l s t re s s e s s u c h a s d r o u g h t a n d g r a z i n g .T h e s e c o m b i n a t i o n s o f s tr e ss e s h a v e b r o u g h t a b o u t c o n -v e r g e n c e i n p l a n t f o r m a n d f u n c t i o n i n m e d i t e r r a n e a nc l i m a t e s .

F i r e h a s th u s b e e n i m p o r t a n t i n t h e e v o l u t io n o f m o r ex e r ic g r a s s la n d s a n d w o o d l a n d s a n d m o r e m e s i c m a q u ia n d s h r u b l a n d c o m m u n i t i e s . C o n t r a r y t o t h e p r e s e n tv i e w o f f i r e a s s i m p l y d e s tr u c t i v e , b o t h f i re a n d g r a z i n gh a v e f a v o r e d g e n e t i c a l a s w e l l a s e c o l o g i c a l d i v e r si t y .T h e y s h o u l d b e s t u d i e d a s i n te g r a l c o m p o n e n t s o f M e -d i t e r r a n e a n e c o s y s t e m s a n d t h e i r e v o l u t i o n .

R~sum~*I1 e s t a d m i s q u e l e f e u a j o u 6 u n r 61 e d 6 c i si f d a n s l ' 6 v o l u t i o nb i o l o g i q u e e t c u l t u r a l e d e l ' 6 p o q u e p o s t g l a c i a i r e d a n s l ar 6 g i o n m 6 d i t e r r a n 6 e n n e . S o n i m p a c t s u r l ' 6 v o l u t i o n d e sp l a n t e s s ' e s t m a n i f e s t 6 p a r d e s r 6 p o n s e s d e ' r 6 t r o a c t i o n '( f eedbac k) . D a r t s ce l te s - c i, le s e f f e t s de l '616ment f eu e t deses r 6per cuss ions on t e f f ec tu6 une s61ec t ion des p lan tesp o u r c e r t a i n e s a c t i v i t6 e s p h y s i o l o g i q u e s e t a u t r e s m 6 c a n i s -r u e s , c e q u i e n t r a i n e u n e t o t 6 r a n c e d i r e c t e a u f e n o u u n ep r o t e c t i o n e t u n e r 6 g 6 n 6 r a t i o n p a r v o i e v ~ g 6 t a t i v e e t r e -p r o d u c t r i c e .

L e s p h a n 6 r o p h y t e s d o m i n a n t s d u m a q u i s , s c l b ro p h y l te s ,t o l 6 r a n t s g l a s 6 c h e r e s s e , p r 6 s e n t e n t o b l i g a t o i r e m e n t d e sr e j e t s d e s o u c h e , a l o r s q u e l e s c h a m 6 p h y t e s d o m i n 6 sf u y a n t l a s 6 c h e r e s s e a u s s i b i e n q u e l e s p l a n t e s h e r b a c 6 e sv i v a c e s s e m u l t i p l i e n t ~ l a l o i s p a r v o i e v 6 g 6 t a t i v e e tr e p r o d u c t r i c e , e t s o n t b i e n a d a p t 6 s a u x n o u v e l l e s h a b i t a t sd 6 n u d 6 s p a r l e s f e ux . L e s h e r b e s a n n u e l l e s e t v i v a c e s q u is u i v e n t l e p a s s a g e d e s f e u x s o n t a u s s i c a p a b l e s d ' 6 c h a p p e raux temp~ r a tm ' es 61ev~es des f eu x gr~ tce /~ des m6c anism esd e t o r s i o n , o u d e s g r a i n e s . L e s t e n d a n c e s 6 v o l u t i v e s p o u rs u r m o n t e r l ' e f f e t d u f e u s o n t 6 t r o i t e m e n t l i 6 e s ~ c e l l e s* Translated by Dr . G. Bouxin, Nam ur .


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d ' a u t r e s ' c o n t r a i n t e s ' (s t re s se s ) d e l ' e n v i r o n n e m e n t c o m m el a s t c h e r e s s e e t l a p r e s s i o n d u p g t tu r a g e. C e r t a i n e s c o m -b i n a i s o n s d e s c o n t r a i t e s o n t e n t r a i n 6 u n e c o n v e r g e n c ed e f o r m e e t d e f o n c t i o n .

L e f e u a d o n c j o u 6 u n r t l e i m p o r t a n t d a r t s l ' t v o l u t i o nd e s p e l o u s e s p l u s x t r i q u e s , d e s f o r ~ ts , d e s m a q u i s p l u sm t s i q u e s e t d e s c o r n m u n a u t ~ s d ' a r b u s t e s. C o n t r a i r e m e n ta u p o i n t d e r u e a c t u e l s u r l e r t l e d e s t r u c t e u r d u b r f i l a g e,l e fe u e t le p f it u r a g e o n t f a v o r i s 6 l a d i v e r si t 6 g t n t t i q u e a u s s ib i e n q u ' t c o l o g i q u e . I ls d e v r a i e n t ~ tr e 6 tu d i t s c o m m e d e sc o m p o s a n t e s i n t 6 g r a n t e s d e s 6 c o s y s t t m e s d e M t d i t e r -r a n t e e t d e l e u r 6 v o l u t i o n .

BibliographyA n d e r l i n g , A . 1 8 8 6. A c k e r b a u u n d T i e r z u c h t i n S y r i e n i n s -

b e s o n d e r e P a l a e s t i n a . Z e i t s c h r i f t d . P a l / i s t i n a V e r e i n s 9 : 6 2 - 6 3 .A n d e r s o n , E . 1 9 56 . M a n a s a m a k e r o f n e w p l a n t s a n d n e w p l a n tc o m m u n i t i e s . I n : M a n ' s r o t e in c h a n g i n g t h e f a c e o f t h e ea r t h ,e d . W . L . T h o m a s , p p . 7 6 3 - 7 7 7 , U n i v e r s i t y o f C h i c a g o P r e s s .

A x e l r o d , D . I . 1 9 5 8 . E v o l u t i o n o f t h e M a d r o - T e r t i a r y g e o f l o r a .B o t . R e v . 2 4 : 4 3 3 - 5 0 9 .B h a r u c h a , F . R . 1 9 5 2 . E t u d e e c o l o g i q u e e t p h y t o s i c o l o g i q u e d el ' a s s o c i a t i o n B r a c h y p o d i u m r a m o s u m e t P h l o m i s l y c h n i t i sd e s G a r i g u e s L a n g u e d o c ie n n e . C o m m . S . I . G . M . A . M o n t p e l -l i e r , 18 : t -53 .

B r a u n - B l a n q u e t , J. 1 9 25 . D i e B r a c h y p o d i u m r a m o s u m - P h l o m i sl y n c h i ti s A s s o z i a ti o n d e r R o t e r d e b o e d e n S u e d a f r a n k re i c h s .I n F e s t s c h r . C . S c h r o e d e r , V e r o e f f . G e o b o t . I n s t . R u e b e l ,H e f t 3 .

B u t z e r, E . W . 1 97 2. E n v i r o n m e n t a n d a r c h e o lo g y . M e t h u e n ,L o n d o n . 7 0 3 p p .E i g , A . 1 9 27 . O n t h e v e g e t a t i o n o f P a l e s t i n e . A g r . E x p . S t a . B u l l .,

I n s t . A g r . & N a t . H i s t . , T e l A v i v , 7 : 1 - 8 8 .F l a n n e r y , K . V . 1 9 69 . O r i g i n s a n d e c o l o g i c a l e ff e c ts o f e a r l y

d o m e s t i c a t i o n i n I r a n a n d t h e N e a r E a s t . I n : T h e d o m e s t i c a -t i o n a n d e x p l o i t a t i o n o f p l a n t s a n d a n i m a l s . E d . P . J . U c k o &G . W . D i m b l e y , p p . 7 3 - 1 0 0 , A l d i n e P u b l . C o m . , C h i c a g o .

G a r r o d , D . A . C . 1 9 5 8. T h e N a t u f i a n c u l t u r e . P r o c . B r i t. A c a d .T h e l i f e a n d e c o n o m y o f a m e s o l i t h i c p e o p l e i n t h e N e a rE a s t . 4 3 : 2 t l - 2 2 7 .

G a r r o d , D . A . C . & D . D . A . B a t e . 1 9 37 . T h e s t o n e a g e o f M t .C a r m e l . I . C l a r e d o n P r e s s , O x f o r d . 2 4 0 p p .

H a n e s , T . T . 1 9 7 I . S u c c e s s i o n a f t e r f i r e i n t h e c h a p a r r a l o fs o u t h e r n C a l i f o r n i a . E c o l . M o n o g r . 4 1 : 2 7 - 5 2 .H a r l a n , J . R . 1 9 67 . A w i l d w h e a t h a r v e s t i n T u r k e y . A r c h e o l o g y20 : 197-201 .

H a r l a n , J . R . & D . Z o h a r y . 1 9 66 . D i s t r i b u t i o n o f w i l d w h e a t sa n d b a r l e y . S c i en c e 1 5 3: 1 0 7 5 - i 0 8 0 .

Higgs , E . S . , C . Vi ta -F inz i , D , R . H ar r i s & A. E . Fagg . 1967 . Th ec l im a t e , e n v i r o n m e n t a n d i n d u s t r ie s o f S t o n e A g e G r e e c e .P a r t I I I , P r e s h i st . So c ., V o l . X X X I I I : 1 - 2 9 .

H o r o w i t z , A . 1 9 6 8 . U p p e r P l e i s t o c e n e - H o l o c e n e c l i m a t e a n dv e g e t a t i o n o f th e N o r t h e r n J o r d a n V a l l ey , P h . D . t h es i s,H e b r e w U n i v . , J e r u s a l e m ( H e b r e w ) .

K e n y o n , K . t 9 6 9 . A r c he o lo g y ' i n t h e H o l y L a n d . E r n s t B e n nL i m t . L o n d o n .

K o m a r e k , E . V . 1 9 67 . F i r e a n d t h e e c o l o g y o f m a n . P r o c . T a l lT i m b e r s F i r e E c o l o g y C o n f . , T a l l a h a ss e e ~ F l o r i d m 6 : 1 4 3- 17 0 .

K u h n o l t z - L o r d a t , G . 1 9 39 . L a t e r r e i n c e n d i e e . E d i t . M a i sC a r r e e , N i m e s .L e H o u e r o u , H . N . 1 9 73 . F i r e a n d v e g e t a t i o n i n t h e M e d i t e r -r a n e a n B a s i n . P r o c . T a l l T i m b e r s F i r e E c o l o g y C o n f e r e n c e ,T a t l a h a s s e e , F l o r i d a , 1 3 ( i n p r e s s ) .L i a c o s , L . G . 1 9 73 . P r e s e n t s tu d i e s o f h i s t o r y o f b u r n i n g i n G r e e -c e . P r o c , T a l l T i m b e r F i r e E c o l o g y C o n f e r e n c e , T a t l a h a s s e e ,F lo r ida , 13 ( in p ress ) .L o v e , R . M . & B . J. J o n e s . 1 9 52 . I m p r o v i n g C a l i f o r n i a b r u s hr a n g e s . U n i v . C a l . C i r c . 3 7 1 .3 5 p p .

M c K e l l , C . M . , A . M . W i l s o n & E . L . K a y . 1 9 62 . E ff e c t iv eb u r n i n g o f r a n g e l a n d s i n f e s t e d w i t h m e d u s a h e a d . W e e d s10 : 125-131 .M e i r i , A . 1 9 5 9 . T h e e f f e c t o f b u r n i n g o n t h e g e r m i n a t i o n o fp a s t u r e p l a n t s . M . S c . T h e s i s , H e b r e w U n i v . , R e h o v o t( H e b r e w ) .

M o o n e y , H . A . & E . L . D u n n . 1 97 0. C o n v e r g e n t e v o lu t i o n o fm e d i t e r r a n e a n - c l i m a t e e v e r g r e e n sc l e r o p h y l l s h r u b s . E v o -l u t i o n , 2 4 : 2 9 2 - 3 0 3 .M u l l e r , C . H . , R . B . H a n a w e l t & J . K . M c P h e r s o n . 1 9 68 .A l l e l o p a t h i c c o n t r o l o f h e r b g r o w t h i n t h e f i r e c y c l e o f C a -l i f o r n i a c h a p a r r a l . B u l l . T o r r e y b o t . C l u b 9 5 : 2 2 5 - 2 3 1 .

N a v e h , Z . 1 9 6 0 . A g r o - e c o l o g i c a l a s p e c t s o f b r u s h r a n g e i m -p r o v e m e n t i n t h e m a q u i b e l t o f I sr a el . P h . D . t h es i s, H e b r e wU n i v . , J e r u s a l e m ( H e b r e w ) .

N a v e h , Z . 1 9 6 7. M e d i t e r r a n e a n e c o s y s te m s a n d v e g e t a t i o n t y p e si n C a l i f o r n i a a n d I s r a e l . E c o l o g y 4 8 : 4 4 5 - 4 5 9 .

N a v e h , Z . 1 9 7 1 . T h e c o n s e r v a t i o n o f e c o l o g i c a l d i v e r s i t y o fM e d i t e r r a n e a n e c o s y s t e m s t h r o u g h e c o l o g i c a l m a n a g e m e n t .I n : T h e s c i en t if i c m a n a g e m e n t o f a n i m a l a n d p l a n t c o m m u -n i t i e s fo r conse r va t ion . P roc . Syrup . Br i t . Eco l . Soc ., No rwic h1970 , ed . E . Duf fey & A. S . Wat t , 11 : 605-622 , B lackwel l ,L o n d o n .

N a v e h , Z . 1 9 73 . ( a ) T h e e c o l o g y o f f i r e i n I s ra e l . P r o c . T a l lT i m b e r F i r e E c o l o g y C o n f . , T a l l a h a s s e e , F l o r i d a , 1 3 ( i n p r e s s ).

N a v e h , Z . 1 9 73 . ( b ) E f fe c t s o f f i r e i n t h e M e d i t e r r a n e a n R e g i o n .I n : F i r e a n d E c o s y s t e m s , A c a d e m i c P r e s s ( in p r e ss ) N e w Y o r k .

N a v e h , Z . & J . D a n . 1 97 3. T h e h u m a n d e g r a d a t i o n o f M e -d i t e r r a n e a n l a n d s c a p e s i n I s r a e l . M e d i t e r r a n e a n T y p e E c o -s y s t e m s . E d s . F . d i C a s t r i a n d H . A . M o o n e y ~ E c o l o g i c a lS t u d i e s 7 : 3 7 3 - 3 9 0 . S p r i n g e r , H e i d e l b e r g .

O a k l e y , K . P . 1 9 6 1. O n m a n ' s u s e o f f i re , w i t h c o m m e n t s o nt o o l m a k i n g a n d h u n t i n g . V i k i n g F u n d P u b l . A n t h r o p o l . 3 1 :176-193 .

O r s h a n , G . 1 9 72 . M o r p h o l o g i c a l a n d p h y s i o l o g i c a l p l a s t i c it y inr e l a t i o n t o d r o u g h t . I n : W i l d l a n d s h r u b s - t h e i r B i o l o g y u n dU t i l i z a t i o n , e d . C . M . M c K e l l , J . P . B l a is de I1 & J . R . G o o d i n :2 4 5 - 2 5 4 , U S D A F o r e s t S e r v . G e n e r a l T e c h n i c a l R e p o r tI N T - I . O g d a n , U t a h .

R i c k l i , M . 1 9 42 . D a s P f l a n z e n k l e i d d e r M i t t e l m e e r l a e n d e r .S u r k a m p .

R o s s - A s h b y , W . R . 1 9 56 . A n i n t r o d u c t i o n t o c y b e r n e ti c s .C h a p m a n & H a l l , L o n d o n .

S a m p s o n , A . W . 1 94 4. P l a n t s u c c e s s io n o n b u r n e d c h a p a r r a ll a n d i n n o r t h e r n C a l i f o r n i a . C a l i f . A g r i c . E x p . S t a . B u l l .685: 1-144.

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S a u e r , C . O . 1 9 56 . T h e a g e n c y o f m a n o n e a r t h . I n : M a n ' s R o l ei n C h a n g i n g t h e F a c e o f th e E a r t h , e d . W . L . T h o m a s , p p .4 9 -6 9 . U n i v . C h i c a g o P r e s s.S a u e r , C . O . 1 9 61 . S e d e n t a r y a n d m o b i l e b e n t s i n e a r ly s o c i et i es .I n : S o c i a l l if e o f e a r ly m a n , e d . S . L . W a s h b u r n , V i k i n gF u n d P u b l . i n A n t h r o p o l o g y 3 1 : 25 6 - 2 66 .

S e m p l e , E . C . 1 9 31 . T h e g e o g r a p h y o f t h e M e d i t e r r a n e a nr e g io n . H e n r y H o l t & C o . N e w Y o r k .S h a n t z , H . L . 1 9 4 7. T h e u s e o f f ir e a s a t o o l i n th e m a n a g e m e n to f t h e r a n g e s i n C a l i f o r n i a . C a l . S t a t e B o a r d o f F o r e s t r y .

S p e c h t , R . L . 1 9 69 . A c o m p a r i s o n o f s c l e r o p h y l l o u s v e g e t a t i o nc h a r a c t e r i s t i c s o f M e d i t e r r a n e a n t y p e c l i m a t e s i n F r a n c e ,C a l i f o r n ia a n d S o u t h e r n A u s t r a li a .I I . D r y m a t t e r , e n e r g y a n d n u t r i e n t a c c u m u l a t i o n . A u s t . J.Bo t . 17 : 293-308 .

S t e w a r t , O . C . 1 9 5 6 . F i r e a s t h e f i r s t g r e a t f o r c e e m p l o y e d b ym a n . I n : M a n ' s r o l e in c h a n g i n g t h e f a c e o f t h e e a r t h , e d .W . I . T h o m a s , p p , 1 1 5 - 1 3 3 , U n i v . o f C h i c a g o .S u s m e l . L . 1 9 73 . S w i l u p p i e p r o b l e m i a t t u a l i d e l c o n t r o l l o d e g l ii n c e n d i n e l l a f o r e s ta m e d i t e r r a n e a n . F A O , R o m ( C i te d b y L eH o u e r o u , 1 9 7 3 ) .

S w e e n e y . J . R . 1 9 67 . E c o l o g y o f s o m e f i r e t y p e s i n n o r t h e r nC a l i f o r n i a . P r o c . T a l l T i m b e r F i r e E c o l . C o n f . , H o b e r g , 1 9 67 ,7 : 111-126 .T c h e r n o v , E . 1 9 68 . S u c c e s s io n o f r o d e n t f a u n a s d u r i n g t h eU p p e r P l e i s t o c e n e o f I s r a e l. P a u l P a r e y , B e r l i n .T r a b a u d , L . 1 9 73 . E x p e r i m e n t a l s t u d y o n t h e e f fe c t s o f p r e s c ri b e db u r n i n g o n a Q u e r c u s c o c c i f e r a L . g a r r i g u e ; e a r l y re s u l ts .P r o c . T a l l T i m b e r s F i r e E c o l o g y C o n f e r e n c e , T a l l a h a s s e e ,F lo r ida , 13 ( in p ress ) .U c k o , P . J . & G . W . D i m b l e b y . 1 9 6 9 . I n t r o d u c t i o n : C o n t e x ta n d d e v e l o p m e n t o f s t u d i e s o f d o m e s t i c a t i o n . I n : T h ed o m e s t i c a t i o n a n d e x p l o i t a t i o n o f p l a n t s a n d a n i m a l s :X V I I - X X I . A l d i n e P u b l . C o m p . , C h i c a g o.

V i t a - F i n c i , C . & S . H i g g s . 1 9 7 0 . P r e h i s t o r i c e c o n o m y i n t h eM o u n t C a r m e l a r e a o f P a l e st i ne . S i t e C a t c h m e n t a n a l ys i s.P roc . P reh i s t . Soc . 36 : 1 -37 .

V o g l , J. R . 1 9 67 . F i r e a d a p t a t i o n s o f s o m e S o u t h e r n C a l i f o r n i ap l a n t s . P r o c . T a l l T i m b e r s F i r e E c o l o g y C o n f . H o b e r g ,C a l i f o r n i a 7 : 7 9 - 1 0 9 .W a l t e r , H . 1 9 68 . D i e V e g e t a t i o n d e r E r d e . B d . 2 D i e g e m / i s s i g te nu n d a r k t i s c h e n Z o n e n . G . F i s c h e r , J e n a . 1 0 0 1 p p .W h i t t a k e r , R . H . I 9 7 0 . T h e b i o c h e m i c a l e c o l o g y o f h i g h e rp l a n t s . I n : C h e m i c a l E c o l o g y , e d . E . S o n d h e i m e r & J . B.S i m e o n e , p p . 4 3 - 7 0 . A c a d e m i c P r e s s , N e w Y o r k .

W h i t t a k e r , R . H . & G . M . W o o d w e l l . 1 9 72 . E v o l u t i o n o f n a -t u r a l c o m m u n i t i e s . I n : E c o s y s t e m S t r u c t u r e a n d F u n c t i o n ,e d . J . A . W i e n s . A n n . B i o l. C o l l o g . , O r e g o n S t a t e U n i v . , 3 1 :137-159 .

W h y t e , R . O . 1 9 6 I . T h e e v o l u t i o n o f l a n d u s e in S o u t h W e s t e r nA s i a - A h i s t o r y o f t a n d u s e i n a r i d r e g io n s . E d i t . D . S . S t a m p ,U N E S C O P a r i s : 5 7 - 1 1 8 .

W i s t a n l e y , D . 1 9 72 . S h a r a v . W e a t h e r 2 7 : 1 4 6 - 1 6 5 .Z e i s t , v a n W . 1 9 69 . R e f l e c t i o n s o n p r e h i s t o r i c e n v i r o n m e n t s i n

t h e N e a r E a s t . I n : T h e D o m e s t i c a t i o n a n d E x p l o i t a t i o n o fP l a n t s a n d A n i m a l s , p p. 3 5 - 4 6 . E d . P . J . U k o & G . W . D i m p l e -b y , A l d i n e P u b t . C o . , C h i c a g o .Z o h a r y , D . 1 9 6 9 . T h e p r o g e n i t o r s o f w h e a t a n d b a r l e y i n r e -l a t i o n t o d o m e s t i c a t i o n a n d a g r i c u l t u r a l d i s p e r s a l i n t h e O l dW o r l d . I n : T h e D o m e s t i c a t io n a n d E x p l o i t a t i o n o f P l a n t s a n dA n i m a l s , e d . P . J . U k o & G . W . D i m p l e b y , p p . 47 ~ 56 . A l d i n eP u N . C o m p . C h i c a g o .Z o h a r y , M . 1 9 62 . P l a n t l if e i n P a l e s t in e . R o n a l d P r e s s , N e wY o r k . V I + 2 62 p p .

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