Certain critical environmental factors influencing the growth-form of three Southern California Suffrutescents

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CERTAIN CRITICAL ENVIRONMENTAL FACTORS INFLUENCING THE GROWTH-FORM OF THREE SOUTHERN CALIFORNIA SUFFRUTESCENTS

A T h e s is P re s e n te d to t h e F a c u lt y o f th e D e p a rtm e n t o f B o ta n y The U n i v e r s i t y o f S o u t h e r n C a l i f o r n i a

In P a r t i a l F u lfillm e n t o f th e R e q u ire m e n ts f o r th e D egree M a ste r o f A rts

by R o b e rt F ra n k lin L e s lie June

1942

UMI Number: EP70561

All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion.

UMT D is se rta tio n P u b lish in g

UMI EP70561 Published by ProQuest LLC (2015). Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code

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T h is thesis, w ritten by ......................

under the direction of hl $ ~ F a c u lt y C o m m i t t e e , a n d a p p r o v e d by a l l its m e m b e r s , has be en pr es e n te d to a n d a c c e p te d b y the Co un c il on G ra du ate S t u d y an d R ese ar ch in p a r ti a l fulfill­ m en t of the req u ire m e n ts f o r the d e g re e of

Mas.£.ex....Q£...&r.£.s.

D ean

S ecreta ry D a te.

F a c u lty C o m m ittee

C h airm an

/

ACKNOWLEDGMENTS G r a t e f u l a c k n o w l e d g m e n t i s m ad e t o D r . H ow ard de F o r e s t , u n d e r w hose d i r e c t i o n t h i s h i s m an y h e l p f u l

w o rk w a s d o n e ,

fo r

s u g g e s tio n s , c a r e f u l g u id a n c e , and c lo s e

su p e rv isio n in th e p re p a ra tio n o f t h is

stu d y .

A c k n o w le d g ­

m e n t i s a l s o m ade t o D r . G. R. J o h n s t o n e a n d M rs . Tema G la re f o r t h e i r

v a lu a b le a id .

F u rth e r a p p re c ia tio n i s

ex p ressed f o r

c o n trib u tio n s

o f m a t e r i a l s , a d v i c e , an d e q u ip m e n t by D r. E lm e r N o b le o f S a n ta B arb ara S t a te

C o l l e g e a n d b y M r. M a u n s e l l Van

R e n e s s e la e r , D i r e c t o r o f th e S a n ta B a rb a ra B o ta n ic G arden.

TABLE OF CONTENTS CHAPTER I.

PACE THE PROBLEM AND DEFINITION OF T E R M S ...........................

1

T h e p r o b l e m .......................... . . ...................................................

1

S ta te m e n t o f th e p ro b lem

..........................

1

I m p o r t a n c e o f t h e p r o b l e m ..........................................

2

D e fin itio n s

o f t h e t e r m s ..............................

2

S u f f r u t e s o e n c e ......................................................... .....

II.

2

W o o d i n e s s .......................................................................

4

H erb aceo u sn ess

...................................................................

4

REVIEW OF THE L I T E R A T U R E .....................................................

5

T he n e e d f o r r e s e a r c h .........................................................

5

C o n tr ib u tio n s o f v a lu e to t h i s

5

R e la te d l i t e r a t u r e

stu d y

. . . .

..............................................................

13

...............................

13

.....................................................

14

L im ita tio n s o f p re v io u s s tu d ie s III.



MATERIALS AND PROCEDURE

D e te rm in a tio n o f s u i t a b l e

sp e c ie s

..........................

14

B a s e s f o r s e l e c t i n g s p e c i e s .........................

15

L o c a t i o n o f e x p e r i m e n t a l s t a t i o n s ...........................

16

L o c a tio n of th e s p e c ie s in

24

o th e ra re a s

. . .

T ypes o f S o u th e rn C a l i f o r n i a e n v iro n m e n ts e x a m i n e d ...................................................................................

26

E s t i m a t i o n o f c r i t i c a l e n v i r o n m e n t a l c o n d i t i o n s 30 C ritic a l fa c to rs

..............................................

E x p e r i m e n t a l m e t h o d ..............................................................

32 32

iv CHAPTER

PAGE THE ANATOMICAL S T U D Y ..................................................

36

T h e d e t e r m i n a t i o n o f w o o d i n e s s .....................................

36

P r e p a r a t io n o f p erm an en t s l i d e s

...............................

37

T h e a n a t o m i c a l s t u d y ..............................................................

39

I V .'

T he s p e c i e s a t t h e T he s p e c i e s i n V.

s t a t i o n s .......................................

39

o t h e r a r e a s ...........................................

40

RESULTS AND D IS C U S S I O N ..................................... C ritic a l fa c to rs a t th e s ta tio n s

................................

42 42

B e h a v io r o f th e s p e c i e s u n d e r e x p e rim e n ta l c o n d i t i o n s ..... ............................................................................. C ritic a l fa c to rs

................................

49

S . a p i a n a ...................................................................................

49

S.

l e u o o p t i y l l a ........................................................................

51

E.

o a l i f o r n i c a ........................................................................

$2

R e s u l t s o f t h e a n a t o m i c a l w o r k ....................................

53

SUMMARY AND C O N C L U S IO N S ..........................................................

86

S u m m a r y .............................................................................................

86

C o n c lu s io n s

.........................................................

87

BIBLIOGRAPHY............................................................................................................

88

V I.

in o th e r a re a s

48

. . . . .

LIST OF TABLES TABLE I.

PAGE C e r ta in E n v iro n m e n ta l C o n d itio n s a t th e S t a ­ tio n s

II.

..................... .

.............................. .............................. .....

P e rc e n ta g e s o f S o i l M o istu re a t O c to b er

III. IV .

. . . . . .

. . . . . . . . . . . . . .

Jj».

c a l i f o r n i c a ...................................................................

...............................

62

K e w -G ro w th C h a n g e s i n P l a n t s T a k e n f r o m t h e .

63

H e w -G ro w th C h a n g e s i n P l a n t s T a k e n f r o m t h e .

64

N e w -G ro w th C h a n g e s i n P l a n t s T a k e n f r o m t h e le u c o p h y lla S ta tio n . . . . . . . .

65

H e w -G ro w th C h a n g e s i n P l a n t s T a k e n f r o m t h e S u ffra te s c e n t S. le u c o p h y lla

X II.

61

N e w -G ro w th C h a n g e s i n P l a n t s T a k e n f r o m Woody

W oody S . X I.

60

M o r p h o l o g i c a l C o m p a r i s o n s o f t h e G r c M th - F o r m s

H e r b a c e o u s S . a p i a n a S t a t i o n .............................. .. X.

59

M o r p h o l o g i c a l C o m p a r i s o n s o f t h e G r o w th - F o r m s

S u f f r u te s c e n t S. a p ia n a S ta tio n IX .

58

.......................................................................

a p ia n a S ta tio n V III.

57

M o r p h o l o g i c a l C o m p a r i s o n s o f t h e G r o w th - F o r m s

of V II.

th e S t a ti o n s in

D i s t r i b u ti o n o f E x p e rim e n ta l P la n ts

o f S. le u c o p h y lla ¥1.

56

.........................

o f S. a p ia n a V.

.

S ta tio n

. . . .

66

H e w -G ro w th C h a n g e s i n P l a n t s T a k e n f r o m t h e H erb aceou s S.

le u c o p h y lla S ta tio n

. . . . . .

67

TABU X III#

PAGE Mfew-Growth C h a n g e s i n P l a n t s T a k e n f r o m t h e Woody 2.

c a lifo rn ic a

S ta tio n to th e S u ffru te sc e n t

E . c a l i f o r n i c a S t a t i o n a n d V i c e V e r s a ....................... X IV .

68

T y p e s o f E n v i r o n m e n t a n d T h e i r E f f e c t s U pon S u f f r u t e s c e n t G r o w t h - E o r m s ..............................................

69

LIST OF FIGURES FIGURE 1.

PAGE T h e Woody S a l v i a a p i a n a S t a t i o n G e n e r a l V i e w ..........................................................................................

2.

Tiie Woody S a l v i a a p i a n a S t a t i o n D e t a i l ...................................................................

3.

....................................

. .......................................................................

29

The S u f f r u t e s c e n t S a l v i a l e u c o p h y l l a S t a t i o n D e ta il .

11.

27

T h e S u f f r u t e s c e n t S a l v i a l e u c o p h y l l a St a t i o n G e n e r a l V i e w ........................................................................................

10.

27

T h e Woody S a l v i a l e u c o p h y l l a S t a t i o n D e t a i l .............................................................

9*

25

T h e Woody S a l v i a l e u c o p h y l l a S t a t i o n G e n e r a l V i e w ..............................................

8.

25

The H e rb a c e o u s S a l v i a a p ia n a S t a t i o n D e t a i l .........................................

7*

23

The H e rb a c e o u s S a l v i a a p ia n a S t a t i o n G e n e r a l V ie w • • • • ...................................................................

6.

23

The S u f f r u t e s c e n t S a l v i a a p i a n a S t a t i o n D e t a i l ..................... ,

5*

21

The S u f f r u t e s c e n t S a l v i a a p i a n a S t a t i o n G e n e r a l V iew

4.

21

.............................................................................................

29

T he H e r b a c e o u s S a l v i a l e u c o p h y l l a S t a t i o n G e n e r a l V i e w .......................................................................................

31

v iii FIGURE 12.

PAGE The H e r b a c e o u s S a l v i a l e u c o p h y l l a S t a t i o n D e t a i l ........................................

13.

T he Woody E n c e l i a c a l i f o r n i c a S t a t i o n

The S u f f r u t e s c e n t I n c e l i a

33 c a lifo rn ic a S ta tio n

G e n e r a l V i e w ............................................................. 16.

31

...............................................................................33

D e t a i l .............................. 1$.

.

T he Woody E n c e l i a c a l i f o r n i c a S t a t i o n G e n e r a l V iew

14.

.

34

The S u f f r u t e s c e n t E n c e l i a c a l i f o r n i c a S t a t i o n D e t a i l ............................................................................................................. 34

LIST OF PLATES PLATE I.

PAGE Woody F o rm o f S a l v i a a p i a n a T ra n sv e rse S e c tio n

II.

....................................................................

W oody F o rm o f S a l v i a a p i a n a L o n g itu d in a l S e c tio n

III.

. . . .

..............................................................

...................................................................

75

7&

..............................................................

77

S u f f r u t e s c e n t F o rm o f S a l v i a l e u c o p h y l l a 78

S u f f r u t e s c e n t F orm o f S a l v i a l e u c o p h y l l a L o n g itu d in a l S e c tio n

X I.

7A

W oody F o rm o f S a l v i a l e u c o p h y l l a

T r a n s v e r s e S e c t i o n ................................................................... X.

73

Woody F o rm o f S a l v i a l e u c o p h y l l a

L o n g itu d in a l S e c tio n IX .

72

H e r b a c e o u s F o rm o f S a l v i a a p i a n a

T ra n sv e rse S e c tio n V III.

..............................................................

...................................................................

L o n g itu d in a l S e c tio n V II.

...............................................

H e r b a c e o u s F o rm o f S a l v i a a p i a n a T ra n sv e rse S e c tio n

V I..

71

S u f f r u t e s c e n t F o rm o f S a l v i a a p i a n a L o n g itu d in a l S e c tio n

V.

..............................................................

S u f f r u t e s c e n t F o rm o f S a l v i a a p i a n a T ran sv e rse S e c tio n

IV .

70

..............................................................

79

H e r b a c e o u s F o rm o f S a l v i a l e u c o p h y l l a T ra n sv e rse S e c tio n

...................................................................

80

X

PLATS X II.

PAGE H e r b a c e o u s F o rm o f S a l v i a l e u c o p h y l l a L o n g i t u d i n a l S e c t i o n ............................................................

X IX I*

Woody F o rm o f E n c e l i a c a l i f o m i c a T ran sv erse S e c tio n

X IY .

...................................................................

Woody F o rm o f E n c e l i a

..............................................................

83

S u f f r u t e s c e n t F o rm o f E n c e l i a c a l i f o r n i c a T ra n sv e rse S e c tio n

X V I.

82

c a lifo rn ic a

L o n g itu d in a l S e c tio n XV.

81

...................................................................

84

S u ffru te s c e n t F o m o f E n c e lia c a lif o r n ic a L o n g itu d in a l S e c tio n

..............................................................

85

CHAPTER I THE PROBLEM AM) DEFINITIONS OF TERMS T h is and c a r r ie d

i n v e s t i g a t i o n w a s b e g u n d u r i n g t h e sum m er o f 1 9 4 0 on f o r a b o u t tw o y e a r s .

P re lim in a ry o b se rv a tio n

s u g g e s t e d s e v e r a l p ro b le m s w h ic h m ig h t w e l l be co m b in ed i n one s tu d y due t o t h e i r o b v io u s ly c l o s e f r u t e s c e n t p l a n t s w ere o b se rv e d t o C a lifo rn ia and, as th e w o rld .

c o n n e c tio n .

S u f­

b e common t h r o u g h o u t

in d ic a te d in th e

lite ra tu re ,

th ro u g h o u t

C e r ta in s p e c ie s w ere se e n t o be s u f f r u t e s c e n t i n

one l o c a l i t y and e i t h e r

c o m p le te ly h e rb a c e o u s o r c o m p le te ly

woody i n o t h e r r e g i o n s , d e p e n d in g u p o n m i c r o c l i m a t i c d i f f e r ­ ences. I.

THE PROBLEM

S ta te m e n t o f th e p ro b le m .

The c h i e f p u rp o se o f t h i s

i n v e s t i g a t i o n w as t o com pare t h e a n a t o m i c a l s t r u c t u r e o f th e ste m s i n c e r t a i n

s u f f r u t e s c e n t p l a n t s , an d t o c o r r e l a t e th e

phenom enon o f s u f f r u t e s e e n c e w i t h c e r t a i n o f th e

e n v ir o n m e n t w h ic h in d u c e i t .

c r itic a l fa c to rs

I t w as n e c e s s a r y t o

in ­

c lu d e a c o n c is e d e f i n i t i o n an d l i m i t a t i o n o f th e te rm s " s u f f r u t e s c e n t , " " h e r b a c e o u s , " a n d "w o o d y ," d r a w in g s i n b o t h tr a n s v e r s e an d l o n g i t u d i n a l s e c t i o n s o f th e g ro w th -fo rm s o f th e th r e e

sp e c ie s used,

and th e t r a n s p l a n t i n g o f sp e c im e n s

from one c r i t i c a l l o c a t i o n to a n o t h e r i n o r d e r to a s c e r t a i n

2 th e e n v iro n m e n ta l f a c t o r s

in s tr u m e n ta l in b rin g in g ab o u t th e

change i n th e g ro w th -fo rm . Im p o rta n c e o f th e p ro b le m . w o rk h a s d e a l t

A lth o u g h t h i s

e x p e rim e n ta l

o n ly w ith t h r e e n a ti v e C a li f o r n ia

s p e c ie s , th e

d a t a h e r e m i g h t a p p l y t o som e c o m m e r c i a l l y i m p o r t a n t p l a n t s w h ic h u n d e r c e r t a i n

c o n d itio n s a re s u f f r u te s c e n t in h a b it

( 3 3 ).

The a n a t o m i c a l s t u d y o f s u f f r u t e s c e n c e h a s b e e n l a r g e l y o v e r ­ lo o k e d , i n v e s t i g a t o r s h a v in g p u b lis h e d l i t t l e o n o m ic d e s c r i p t i o n s .

m ore t h a n t a x ­

The p r e s e n t i n v e s t i g a t i o n d e a l s w i t h

t h e p ro b le m o f s u f f r u t e s c e n c e fro m t h e s t a n d p o i n t o f an ato m y as

in flu e n c e d by c e r ta in c r i t i c a l The r e s u l t s

e n v iro n m e n ta l f a c t o r s .

c o r r o b o r a t e o t h e r r e s e a r c h w h i c h h a s sh o w n

t h a t c lim a tic f a c t o r s in flu e n c e a n a to m ic a l m o d ific a tio n s (6 ).

The f a c t t h a t l i t e r a t u r e

e x is te n t fu rth e r ju s tif ie s II.

on th e

(4 ),

s u b j e c t i s a lm o st n o n ­

th e i n v e s t i g a t i o n .

DEFINITIONS OF THE TEEMS

S u ffru te sc e n c e .

T he t e r m " s u f f r u t e s c e n t " a s u s e d h e r e

a p p l i e s to a n y p a r t l y h e r b a c e o u s , p a r t l y woody s h ru b o r h a l f ­ s h ru b w h ic h e x h i b i t s t h e h e rb a c e o u s h a b i t o f g ro w th i n t h e ste m s f o r a c o n s i d e r a b l e d i s t a n c e b e h in d th e e n tia tio n . due to

zone of d i f f e r ­

As w i l l b e sh ow n i n C h a p t e r Y , t i s s u e s l i g n i f y

c e r t a i n c r i t i c a l e n v iro n m e n ta l c o n d itio n s ,

o f a n n u a l g ro w th phenom ena. o f h a b i t s am ong t h e s e p l a n t s .

irre sp e c tiv e

T h ere a r e v a r i o u s c o m b in a tio n s C e rta in

sp e c ie s of P ap av eraceae»

3 L a b i a t a e , L e g u m in o s a e . an d c e r t a i n

s p e c ie s o f A n tirrh in u m , in

d e s e r t l o c a t i o n s o f e x t r e m e l y h o t and d r y sum m ers, p r o d u c e ste m s above th e g ro u n d w h ic h a r e e n t i r e l y h e rb a c e o u s an d w h ic h d i e down d u r i n g t h e c r i t i c a l . p e r i o d , ste m s re s u m in g a c t i v i t y

t h e i r woody u n d e rg ro u n d

d u rin g th e w in te r r a i n s

A n o th e r ty p e o f s u f f r u te s c e n c e

( 2 6 )*

in c lu d e s th o se

sp e c ie s

w i t h p e r e n n i a l u n d e rg ro u n d rh iz o m e s w h ic h p ro d u c e a n n u a l a e r i a l p a r ts d u rin g th e fa v o ra b le in th e R osaceae

(7 )*

seasons; th is

i s e v id e n c e d

C e r t a i n le g u m e s, s p e c i e s o f th e O le a c e a e

a n d R h a m n a ce ae , a s w e l l a s S t a n l e y a , S o la n u m , and L e p id iu m h a v e s u f f r u t e s c e n t s te m s , t h e h e r b a c e o u s p a r t s o f w h ic h d i e back a n n u a l l y to th e tio n (7 )

o f sp in e s th e lo n g ,

o l d e r p a r t o f t h e b ra n c h , a ssu m in g th e (9 ),

(2 6 ),

s tra ig h t

I n Rhom neya c o u l t e r i , H a r v .

s t e m s m ay be q u i t e

b a c e o u s w ith no e v id e n c e w h ile

(7 ).

of s e c o n d a r y g ro w th o r w o o d in e s s ,

w i l l be fo u n d h e rb a c e o u s abo v e th e

c o n d itio n s

g ro u n d a n d woody n e a r o r

I t has a ls o

c o m p le te ly woody o r t o d i e sto c k .

su c c u le n t and h e r ­

t h e sa m e s p e c i e s u n d e r s l i g h t l y m o r e x e r i c

b e lo w t h e s o i l s u r f a c e *

func­

been se en to

be a lm o st

b ack t o a p e r e n n i a l , woody r o o t ^

When I r i o g o n u m p a r v i f o l i u m , S m i t h , o c c u r s on s t e e p ,

d ry slo p e s o r

c liffs

t h e h e r b a c e o u s ste m s l i g n i f y d u r in g th e

sum m er m o n th s e v e r y y e a r .

U n d e r m ore m e s ic c o n d i t i o n s t h e

p l a n t a p p e a rs a lm o st c o m p le te ly h e rb a c e o u s

( 2 6 ).

A w id e v a r i e t y o f f u r t h e r c o m b in a tio n s o f h e r b a c e o u s ­ n e s s and w o o d in e ss e x i s t d e p e n d in g upon s p e c ie s and e n v ir o n ­ m e n t.

O th e r te rm s i n t e r p r e t e d to c lu d e (9 ),

" su ffru tic o se "

( 9 )* " s u f f r u t e x "

"h erb aceo u s s u b -s h ru b ” (9 ),

shru b "

( 26),

"n ear-w oody"

W o o d in e ss.

in d ic a te

(26),

The te r m

su ffru te sc e n c e

( 3 )> " o b s c u r e l y

" lig n e sc e n t" (10), "n ear-h erb aceo u s"

t i o n o f t h e x y lem e le m e n ts a n d u n i t s o f th e sc lere n ch y m a f i b e r s .

sh ru b b y ”

" h a lf(6 ).

"w o o d y " w a s i n t e r p r e t e d t o

t o p l a n t s w i t h ste m s e x h i b i t i n g m ore o r l e s s

in ­

refer

even l i g n i f i c a p h lo em , p i t h ,

and

T he g r o w t h - f o r m w a s d e c l a r e d t o b e w o o d y

o n ly w here l i g n i f i c a t i o n

b e h in d th e

b ra n c h o f th e p l a n t e x te n d e d to th e H erb aceo u sn ess.

The te rm

s p i c a l m e riste m o f e ac h zone o f d i f f e r e n t i a t i o n .

" h e rb a c e o u s " w as a p p l i e d t o

p l a n t s a f t e r t h e v a rio u s t e s t s f o r l i g n i n p ro d u ced n e a r ­ n e g a tiv e r e s u l t s .

T h is

was th e m ost v a r i a b l e ,

g ro w th -fo rm o f th e h o w ever, due to th e

c o n d i t i o n s tin d e r w h ic h t h e fo rm g row s to be no s h a rp l i n e

of d is tin c tio n

th e s u f f r u t e s c e n t form s a s and th e woody.

(1 9 ).

s p e c ie s u n d e r stu d y g reat v a rie ty of T here a p p e a re d

b e tw e e n t h e h e r b a c e o u s a n d

t h e r e w a s b e t w e e n t h e s e tw o f o r m s

T h e r e f o r e , t h e p l a n t s c o n s id e r e d t o be h e r ­

b a c e o u s w e re t h o s e w i t h s u c c u l e n t , nonw oody ste m s and r o o t s y s te m s w h ich w e re o l d e r t h a n one y e a r .

CHAPTER I I REVIEW OF THE LITERATURE Much h a s b e e n w r i t t e n u p o n a n a t o m i c a l s p e c i a l i z a t i o n s bro u g h t ab o u t by s p e c if ic

c lim a tic

i n s t a n c e s th e s p e c ie s r e f e r r e d g ro w th h a b i t ;

but s c ie n tific

fa c to rs,

and in

sev eral

to w ere s u f f r u t e s c e n t i n

lite ra tu re

d ire c tly p e rtin e n t

to th e p r e s e n t i n v e s t i g a t i o n i s a lm o st n o n e x is te n t. I.

THE IsIEED FOR RESEARCH

The c h i e f j u s t i f i c a t i o n th e

fact

fo r th is research e x is ts

in

t h a t no i n v e s t i g a t o r s have o f f e r e d an y e x p la n a tio n

o f su ffru te s c e n c e as a c o n d itio n f a c t o r s w h ic h w ere m e a s u r a b le .

in d u ce d by e n v iro n m e n ta l T here a re

no l i s t s

of su f­

f r u t e s c e n t p l a n t s a n d t h e i r h a b i t s , a n d a p p a r e n t l y fe w h a v e re c o g n iz e d th e

su ffru te sc e n t as a p la s tic

b u t d i s t i n c t g ro w th

fo rm . II.

CONTRIBUTIONS OF VALUE TO THIS STUDY

B rew er ( 6 ) i n 1900 w orked w i t h s u c c u l e n t s . s tra te d

t h a t p l a n t s g ro w n i n m o i s t a i r a s c o m p a r e d w i t h t h o s e

grow n i n tis s u e , crease

He d e m o n ­

n o rm al a i r

sh o w e d c h a n g e s i n t h e

fib ro v a sc u la r tis s u e ,

sto ra g e

a i r sp a c e s, d ry w e ig h t, d e­

in a sh c o n te n t o f le a v e s ,

in c re a s e

fo llo w in g :

in c re a se

in c h lo re n c h y m a ,

i n i s o d i a m e t r y of c h l o r o p h y l l c e l l s , w avy e p id e r m a l

6 w a lls , and in c re a s e i n sto m a ta . f a c t o r s , n o t th e

He b e l i e v e d t h a t t h e a i r

s o i l f a c t o r s , d e te rm in e d th e s e

f u r th e r b e lie v e d th a t th e

changes.

He

c h an g e s a n d th e m o d i f i c a t i o n s w ere

h e re d ita ry . E b erh ard t

( 1 7) in th e

caused in c re a s e in w e ig h t,

in

in

o f th e l e a f ,

i n t h e am ount o f d r y

th e am ount o f c h l o r o p h y l l, and i n r o o t d e v elo p m e n t;

w h ile d ry a i r tic le ,

th e s iz e

sam e y e a r s h o w e d t h a t h u m id a i r

caused an in c re a s e

in th e th ic k n e s s o f th e cu­

th e num ber o f s to m a ta ,

woody t i s s u e ,

and p a lis a d e

B o n n ie r (4 )

in t h e am ount o f s c le re n c h y m a ,

c e lls .

i n 1902 s t a t e d t h a t h e to o k p l a n t s from

F o n ta in e b le a u t o T o u lo n , F r a n c e .

W ith in s e v e r a l w eeks he

n o te d ch an g e s w h ic h u l t i m a t e l y m o d if ie d th e fo rm o f t h e l e a v e s , c a u s e d t h e x y lem c e l l s

to

lig n if y so o n er, and a lte r e d

e r a l a p p earan ce o f t h e

p la n ts.

C lem en ts ( 1 4 ) i n 1905 s t u d i e d t h e stru c tu re

to p h y s i c a l f a c t o r s .

th e gen­

r e l a t i o n o f th e l e a f

She h a d t h r e e

o b je c tiv e s:

(1 ) To c o r r e l a t e t h e h i s t o l o g y o f t h e l e a v e s w i t h t h e m ea su re d p h y s i c a l f a c t o r s of th e e n v iro n m e n t; (2 ) To d e te r m i n e th e k in d and am ount o f t h e m o d i f i c a ­ t i o n t a k i n g p l a c e i n t h e sam e s p e c i e s i n d i f f e r e n t h a b ita ts; ( 3 ) To t h r o w l i g h t u p o n t h e e n t s p e c ie s and g e n e ra .

p la s tic ity

of th e d i f f e r ­

U nder p h y s i c a l f a c t o r s she in c lu d e d c lim a te and s o i l a n d sh o w e d t h e m ad e t o

v a r io u s m o d if ic a tio n s w h ic h th e

each c o n d itio n .

Of a l l h e r s t a t i o n s ,

s p e c ie s have o n l y tw o h a d

d i r e c t v a lu e t o th e tio n ,

stu d y a t hand.

H e r " F o o t h i l l M esa” s t a ­

w ith i t s

2 t o 4 p e r c e n t a v a i l a b l e w a t e r , 28 t o 7 7 p e r o o c e n t h u m i d i t y , a n d 57 t o 89 F . t e m p e r a t u r e , m i g h t b e com­

p ared w ith th e o c e a n -fa c in g s t a ti o n s stu d y .

T he t h i c k e t

sta tio n

e m p lo y e d i n t h e p r e s e n t

o f C le m e n ts, w ith i t s

11 p e r

o

c e n t a v a i l a b l e w a t e r , 38 t o 8 0 p e r c e n t h u m i d i t y , a n d 53 t o o 85 F . t e m p e r a t u r e c o m p a r e s f a v o r a b l y w i t h t h e t w o c h a p a r r a l sta tio n s

o f th e p re s e n t stu d y .

th ic k e r d ip lo p h y ll,

e p id e rm is,

c h lo re n c h y m a i n th e f o r m e r . each c ase g o t th e a n tic ip a te d

The r e s u l t s w e re t h e c u tic le

sam e:

(so m e e x c e p t i o n s ) , a n d

She v a r i e d th e f a c t o r s an d i n r e s u l t w ith in th e

sp e c ie s,

con­

c lu d i n g t h a t t h e l i g h t was a g r e a t e r f a c t o r th a n d r y n e s s . T h is w as,

in p a r t ,

b o rn e o u t by th e p r e s e n t i n v e s t i g a t i o n .

The w o r k o f M a c D o u g a l a n d S p o e h r ( 2 9 ) w a s i n d i r e c t to

c o n tra st

t h a t o f C le m e n ts. C a n n o n ( 1 0 ) i n 1 9 1 4 e m p h a s i z e d t h e f a c t t h a t t h e m any

v a ria tio n s

in c lim a te in C a li f o r n ia have in d u c e d m o rp h o lo g i­

c a l s p e c ia liz a tio n s . a r e due t o s p e c i f i c l i m i t th e

He p o i n t e d lo c a l

d is trib u tio n

out th a t th e m ic ro c lim a te s

c o n d itio n s and t h a t

o f th e

sp e c ie s.

c o n d itio n s

T he p r e s e n t i n v e s t i ­

g a t i o n a t t e m p t s t o show t h a t s u f f r u t e s c e n t d istrib u tio n

th e se

s p e c i e s h a v e a w id e

and a r e c a p a b le th ro u g h a n a to m ic a l changes o f

a d a p t i n g t h e m s e l v e s to v a s t l y d i f f e r e n t c i r c u m s t a n c e s o f m o is ­ tu re ,

a ltitu d e ,

s o i l , e x p o s u r e , a n d p l a n t c o m m u n i ty .

H anson (2 3 ) i n 1917 r e f e r r e d t o

th e w o rk s o f B o n n ie r,

U b e r h a r d t , a n d C l e m e n t s , a l l o f whom h a d f o u n d i n t r a s p e c i f i c v a r i a t i o n s w h ic h h ad b e en i n t r o d u c e d b y e n v ir o n m e n ta l d i f f e r ­ ences.

H a n so n ’ s p u rp o s e was t o

in v e stig a te

th e e x a c t d i f f e r ­

en ces in th e s t r u c tu r e o f th e le a v e s an d to c o r r e l a t e t h i s w ith f a c t o r s o f th e e n v iro n m e n t, pow er o f th e

a ir,

such a s l i g h t ,

e v a p o ra tin g

t e m p e r a t u r e , h u m i d i t y , a n d w in d *

He sh o w e d

t h a t d r y w e i g h t s w e re g r e a t e r i n a r e a s o f m ore i n t e n s e f a c t o r s a n d i n c o m p a r i n g s t r u c t u r e s h e found t h a t t h e l e a v e s i n t h e sh a d e w ere t h i n n e r and t h a t th e d i f f e r e n c e w as c a u se d by th e in c re a se

in p a li s a d e and d e c re a s e in spongy t i s s u e .

M acD ougal a n d S p o e h r (2 9 ) i n

1 9 1 8 a l s o m o v ed p l a n t s

fro m o n e a r e a to a n o t h e r w here c r i t i c a l in v o lv e d .

c lim a tic

f a c t o r s w ere

They a rg u e d t h a t w e l l- d e f in e d a n a to m ic a l a d j u s t ­

m en ts w ere n e c e s s a r y f o r s u r v i v a l in p l a c e s o f a r i d They f u r t h e r a t t e s t e d

th a t

c o n d itio n s

th e c h e m ic a l c o m p o s itio n o f t h e

p l a n t i s m o d if ie d by th e w a te r r e l a t i o n s d u r in g g ro w th an d t h a t m ore c e l l u l o s e a n d l e s s s t a r c h d ry a tm o sp h e re s.

is

d e v e lo p e d in p l a n t s in

A lth o u g h t h e i r w ork d e a l t p r i m a r i l y w i t h

s u c c u l e n t s , o n l y a few o f w h ic h d e v e lo p l i g n i f i e d

tis su e

a cam b iu m r i n g , t h e y d i d n o t d e a l w i t h t h e p o s s i b i l i t i e s

and of

su ffru te sc e n c e . R a m a le y ( 3 6 ) , (37) i n 1 9 1 9 s t u d i e d R o c k y M o u n t a i n p l a n t c o m m u n itie s i n C o lo ra d o and fo u n d i n c e r t a i n p l a n t s a h ig h ra te

o f a d a p t a b i l i t y to t h e r i g o r s o f t h e d i s t r i c t s .

l o c a t i o n s he was a b l e

In sandy

t o f i n d no h e r b a c e o u s p l a n t s b e tw e e n t h e

sh ru b s,

s u f f r u t e s c e n t s p e c ie s a lo n e b e in g a b le

th e e x tre m e s o f e x p o su re and s o i l s . g ra v e l e x h ib ite d p r a c tic a lly ta lu s

to w ith s ta n d

The a r e a s o f s a n d a n d

t h e sa m e s i t u a t i o n , w h i l e

on t h e

s lo p e s o f th e h i l l s A r te m is ia , s p . , a s u f f r u t e s c e n t in

som e l o c a l i t i e s

and h e rb a c e o u s i n

c o m p rise one h a l f t h e v e g e t a t i o n .

o th e rs

( 2 6 ) , was s e e n t o

On r o c k y r i d g e s a g e n e r a l

r e p e t i t i o n was o b s e r v e d . R a m a le y ^ w o rk , a lth o u g h c o n fin e d to o b s e r v a tio n o f q u a d ra ts and la c k in g in e x p e rim e n ta l in fo rm a tio n , ta n t. of th is

It

is p a r tia lly su b s ta n tia te d

is

im p o r­

in s e v e ra l c o n c lu sio n s

th e s is . I n a p a p e r on th e s u f f r u t e s c e n t h a b i t i n S o u th A f r i c a n

p la n ts,

B u rtt-D a v y (9 )

in 1922 a tte m p te d to ap p ro ach th e p ro b ­

lem o f s u f f r u t e s c e n c e fro m a n e v o l u t i o n a r y p o i n t o f v ie w , a rg u in g t h a t th e

"enough e v id e n c e h a s b een c i t e d to s u g g e s t t h a t

su ffru tic o se

h a b it i s an a d a p t a t i o n t o e n v iro n m e n t,

in g an in te r m e d ia te s ta g e

in th e

fo rm ­

e v o lu tio n o f an h erb aceo u s

fro m a n a r b o r e s c e n t t y p e . " He o b s e r v e d f r o m t h e l i t e r a t u r e

an d f r o m t h e f i e l d

s t u d i e s in S o u th A f r ic a th e v a r io u s g r a d a tio n s o f s u f f r u ­ te sc e n c e ,

b u t c o n c lu d e d t h a t

su ch o c c u rre n c e s m e re ly b rid g e d

t h e g ap b e tw e e n t r e e s a n d h e r b s .

In a su c c e e d in g p a ra g ra p h

he a s c r i b e s th e s u f f r u t i c e s t o a r i d i t y . th a t under c e r ta in c lim a tic under a rid ity ,

He f u r t h e r s t a t e d

c o n d itio n s in th e T ra n s v a a l,

s u f f r u t e s c e n t s p e c i e s w ere p ro d u c e d ,

i.e .

su g g e stin g

10 th a t th e s u ffru tic o s e

h a b it

" i s n o t m e re ly a tem p o rary ex­

p e d i e n t , b u t h a s b eco m e a f i x e d h a b i t w h i c h p e r s i s t s u n d e r m o re f a v o r a b l e c o n d i t i o n s . " sp e c ie s to

He e v e n t r a n s p l a n t e d c e r t a i n

"m ore f a v o r a b l e c o n d i t i o n s o f t e m p e r a t u r e a n d s o i l

m o is tu re " and g o t no change o f h a b it a f t e r " s e v e r a l" y e a r s . An o p p o s i t e r e s u l t h a s b e e n r e c o r d e d a t t h e S a n t a B a r b a r a B o ta n ic G a rd en , S a n ta B a rb a ra , C a l i f o r n i a , d u rin g th e co u rse o f t h i s

in v e s tig a tio n ;

i s t e n c e o f s p e c i e s w h ich a r e

and e ls e w h e re ,

fu rth e rm o re , th e ex­

s u f f r u t e s c e n t u n d e r some c o n d i ­

t i o n s a n d w oody u n d e r o t h e r s seem s to o f B u rtt-D a v y f a i l e d to ta k e i n t o

p o i n t o u t t h a t t h e w ork

acco u n t th e r e a l ly

c ritic a l

fa c to rs. In th e

sa m e a r t i c l e ,

th e a u th o r c i t e d th e e v id e n c e

fro m p a le o b o ta n y t h a t t h e e a r l i e s t a n g io s p e r m s w e re w oody; th a t h erb s occur in

t h e m ore r e c e n t p l a n t g r o u p s ;

th a t th e

h e rb a c e o u s ste m h a s b e e n r e d u c e d from th e a n c i e n t , s o l i d , r i n g o f se c o n d a ry g ro w th ; and t h a t p r o g r e s s i v e r e f r i g e r a t i o n h a s b e en th e m o st im p o rta n t f a c t o r i n He g o e s o n t o

sta te

th a t a r i d i t y h a s had th e re d u c in g e f f e c t

and t h a t th e a b s e n c e o f t r e e s i s due to m o is tu re .

c o ld ,

" p e rio d ic d ro u g h t" of th e re g io n

s i m il a r to t h a t o f S o u th e rn C a li f o r n ia .

W ilso n (5 6 ) re s u lts

in c e r t a i n S o u th A fric a n a r e a s

d e s i c c a t i n g w in d s an d t h e a b s e n c e o f s o i l

He e x p l a i n s t h e

(9 ) w h ic h i s

t h e d e v elo p m e n t o f h e r b s .

i n 1922 fo u n d t h a t s e c o n d a r y g ro w th o f t e n

in annual rin g s

i n t h e x y lem b u t n o t i n t h e p h lo em

11 due t o

th e

c r u s h in g a t t h e end o f th e g ro w th p e r i o d .

found t h a t i n H e lia n th u s sp p . th e and r a r e l y n ifie d .

s i e v e t u b e s , c o m p a n io n c e l l s ,

th e p aren ch y m a o f t h e p h lo em , n o r m a lly becam e l i g -

H is m eth o d w as s i m i l a r t o t h e

ent in v e s tig a tio n lig n in .

He

one u s e d i n

th e p r e s ­

i n t h a t he u se d th e p h lo r o g lu c in t e s t

fo r

T h e s t r i k i n g o b s e r v a t i o n w h i c h h e m ade w a s t h a t

lig n ific a tio n

does n o t o c c u r in

e v e r y i n d i v i d u a l o f t h e sam e

s p e c i e s , a l t h o u g h he s p e a k s o f t h e n o rm a l c h a r a c t e r o f t h e phenom enon.

He d i d n o t n o t e u n d e r w h a t c o n d i t i o n s t h e p h l o e m

e i t h e r becam e l i g n i f i e d o r r e m a in e d h e r b a c e o u s . B ew s ( 3 )

i n 1927 s u g g e s te d

w ith f u r t h e r d e c r e a s e i n b r a n c h in g t h e t r e e fo rm p a s s e s i n t o t h e sh ru b fo rm . A c o n ti n u a t i o n o f th e p r o c e s s p r o ­ d u ces d w a rf-s h ru b s , s u f f r u t i c e s , u n d e r-s h ru b s , and f in a lly h erb s. Even w i t h i n t h e l i m i t s o f a s i n g l e g en u s ( e . g . , E u p h o r b i a . H i b i s c u s . S o la n u m . e t c . ) t h e w h o le r a n g e o f f o r m s m ay b e f o u n d . The r e s u l t s o f t h e p r e s e n t is

stu d y in d ic a te

i n b o th d i r e c t i o n s fro m th e

t h a t th e m ovem ent

s u f f r u te s c e n t fo rm s.

U nder

m e sic c o n d i t i o n s a s p e c i e s w i l l o c c u r a s a n e a r l y c o m p le te h e r b , u n d e r s e v e r e d ro u g h t th e a s u ffru te s c e n t, w ill ren d er i t

same s p e c i e s w i l l b e f o u n d a s

and a c o n tin u o u s u p -d ra u g h t o f a i r c u r r e n t s

a lm o s t c o m p le te ly woody ( s e e C h a p te r V ).

a r e , o f c o u rs e , o th e r f a c t o r s c o n cern ed . th e o p in io n t h a t th e e v o lu tio n i s W h itfie ld

(5 3 ), (54)

Bews s e e m s t o b e o f

in one d ir e c tio n

in 1931 s t u d i e d

T here

th e

o n ly .

o sm o tic con­

c e n tr a tio n s o f S a lv ia a p ia n a . J e p . , S a lv ia le u c o p h y lla .

12 G reen e, e t a l .

in th e S a n ta B arb ara re g io n a t v a rio u s e le v a ­

t i o n s fro m 3100 f e e t ab o v e s e a ^ l e v e l t o s e a l e v e l . th e m ost x e r i c ist

c o n d itio n s in a l l

i n O c to b e r,

He f o u n d

c o m m u n itie s s t u d i e d t o ex ­

The p r e s e n t s t u d y c o n f ir m s t h i s

a l t h o u g h f o r m a t i o n o f new l e a v e s was s e e n t o

o b se rv a tio n ,

o c cu r d u rin g

e v e r y m o n th f r o m J u l y t o A p r i l a t e a c h s t a t i o n , S h a p io r and de F o r e s t t r a n s p i r a t i o n r a t e s in th e

(4 2 ) in 1932, i n

c h a p a rra l,

Jep* a " l a v i s h u s e r o f w a t e r . " w in d y d a y s h a d l i t t l e

c o m p a rin g

fo u n d S a lv ia a p i a n a ,

They a l s o

fou n d

e f f e c t upon t r a n s p i r a t i o n .

sh ow n i n C h a p t e r V, t h e

(42) t h a t As h a s b een

i n f l u e n c e o f t h e w in d a n d c o n ti n u o u s

u p -d ra u g h t w as se e n t o b e a c r i t i c a l f a c t o r , h o w ev er, i n th e g ro w th -fo rm o f S a lv ia a p ia n a . B au er (2 ) exp o su re

i n 1 9 3 6 e f f e c t i v e l y sh o w e d t h a t d i r e c t i o n a l

su p p o rte d o r i n h i b i t e d

g ro w th i n c h a p a r r a l s p e c i e s ,

d e c l a r i n g t h a t n o r t h e r n e x p o s u r e s w ere t h e m o st f a v o r e d w h ile s o u th e r n e x p o s u re s w ere l e a s t

so,

S u f f r u t e s c e n c e and w o o d in e ss i n

o w in g t o

s o i l m o is tu re .

t h e sam e s p e c i e s w e r e o b s e r v e d

to o c c u r in d e p e n d e n tly o f exp o su re or m o is tu re c o n d itio n s ; h o w ev er,

t h e am ount o f m o i s t u r e a lo n g w i t h e x p o s u r e seem ed to

be c l o s e l y c o n n e c te d w ith h e rb a c e o u s n e s s . W a tk in s

($ 1 )

i n 1 9 3 9 sh o w e d t h a t

c h a p a rra l sp e c ie s in

S o u th e rn C a lif o r n ia had ab u n d an t s to ra g e t i s s u e , t h e common s t o r a g e fo u n d t o be t r u e

su b s ta n c e i n

th e r a y c e l l s .

i n a l l w oody form s o f th e

s t a r c h b e in g T h e sam e w a s

sp e c ie s under

13

s c r u tin y d u rin g th e p r e s e n t s tu d y . f r u t e s c e n t fo rm s— w h e th e r fro m t h e b ru sh ,

T he h e r b a c e o u s a n d s u f ­ c h a p a rra l,

o r d e s e r t s c r u b r e g i o n s — show ed l i t t l e

c o a sta l sage­ o r no s t a r c h

s to r a g e i n p erm an en t t i s s u e s . III. ' D u rin g th e

r elated l it e r a t u r e

co u rse o f t h i s

stu d y i t

has been n e c e s s a ry

t o r e f e r t o num erous t e x t s , m a n u a ls , a n d p e r i o d i c a l s f o r te c h n ic a l in fo rm a tio n n o t r e l a te d of th e ir

to su ffru te sc e n c e .

n u m b e r a n d i r r e l e v a n c e n o a t t e m p t h a s b e e n m ad e t o

i n c l u d e a r e v ie w o f th e s e e v en w here a l t e r a t i o n s i n have been in tro d u c e d . fo u n d i n

B ecause

C re d it in

te c h n iq u e s

e a c h c a s e , h o w e v e r, w i l l be

th e b ib lio g r a p h y a s w e l l a s in th e t e x t o f th e t h e s i s . IV .

LIMITATIONS ON PREVIOUS STUDIES

As a l l l e a d i n g t a x o n o m i s t s l i s t

s p e c i e s w h ic h a r e

woody i n o n e l o c a l i t y a n d s u f f r u t e s c e n t o r h e r b a c e o u s i n a n ­ o th e r,

it

seem s s t r a n g e t h a t no s t u d e n t h a s p u b l is h e d a n i n ­

v e s tig a tio n

of th e

f a c t o r s w h ic h a r e r e s p o n s i b l e f o r

th e

p a r t i c u l a r g ro w th -fo rm in a g iv e n a r e a . The g r e a t e s t s i n g l e stu d ie s e x is ts

in th e

fact

l i m i t a t i o n n o te d in p re v io u s t h a t c o n c lu s io n s w ere e x t r a c t e d

fro m e x t e r n a l o b s e r v a t i o n s , - u n s u b s t a n t i a t e d by i n s t r u m e n t a l , c h e m ic a l, o r m ic ro sc o p ic e v id e n c e .

I n m an y c a s e s ,

in v e s ti­

g a t i o n s w e re c o n c lu d e d b e f o r e a d e q u a te r e s u l t s w ere o b t a i n e d . Many d i s a g r e e m e n t s i n d e t a i l s w e r e a l s o

n o te d .

CHAPTER I I I MATERIALS AND PROCEDURE D ue t o

th e la c k o f l i t e r a t u r e

a n d e x p e r i m e n t a l w ork

on s u f f r u t e s c e n c e , g r e a t c a r e w as e x e r c i s e d i n p r o c e d u r e , e q u ip m e n t, and p l a n t and s t a t i o n I.

se le c tio n s .

DETERMINATION OP SUITABLE SPECIES

S a lv ia a p i a n a , J e p . , h e r e a f t e r r e f e r r e d t o a s S. a p ia n a . is

d e s c r ib e d by J e p s o n (26) a s :

S h r u b b y , 3 t o 5 Cor 1 0 ) f t . h i g h , b r a n c h e s v i r g a t e ; le a v e s o b lo n g t o e l l i p t i c , 1 t o 4 i n . lo n g , s e r r u l a t e o r s u b e n t i r e , m o s t ly v e r y w h ite ab o v e a n d b e lo w , s h o r t p e t i o l e d ; p a n ic le v i r g a t e , 1 to 4 f t . lo n g ; c o r o l l a w h i t e , 6 t o 9 l i n e s l o n g , th e tu b e s h o r t (2 o r 3 l i n e s l o n g ) ; u p p e r l i p v e ry s h o r t o r a lm o st o b s o l e t e , t h e l o w e r l i p m uch d e v e l o p e d , t h e c o n s p i c u o u s p a l a t e c u r v e d upw ard s o a s t o c l o s e t h e t h r o a t ; m id d le l o b e f r i n g e d , th e l a t e r a l lo b e s s m a ll; s t y l e and sta m e n s lo n g e x s e r t e d — M esas a n d c a n y o n s , 1000 t o 2300 f t . : c o a s t a l S . C a l . , e . to w e s t s i d e o f C o lo ra d o D e s e r t (A u d i b e r t i a p o ly s ta c h y a , B e n th .) S a lv ia le u c o p h y lla , G reene. , h e r e a f t e r r e f e r r e d t o as S.

le u c o p h y lla , i s

d e s c rib e d by Jepson

(2 6 ) a s :

S te m s woody b e lo w , l e a f y , w h ite f a r i n o s e a b o v e , 3 t o 4 f t . h ig h ; le a v e s o b lo n g - la n c e o la te o r th e lo w e st o v a te , o b tu se , th e u p p e r w ith tr u n c a te b a se , f i n e l y ru g u lo s e a b o v e , w h ite to m e n tu lo s e b e n e a th , 1 to 3 l i n e s lo n g , s h o r t p e t i o l e d ; w h o rled h e a d s 3 to 5 , v e ry d e n se ; b r a c ts o v a l o r o b lo n g , d e n s e ly w h ite f a r i n o s e ; c a l y x s p l i t t i n g down i n f r o n t , a t l e n g t h e m a r g i n a t e b e h in d ; c o r o l l a l i g h t p u r p le , 5 to 7 l i n e s l o n g , th e t u b e h a r d l y e x s e r t e d ; s ta m e n s a n d _ s t y l e m uch e x s e r t e d ; c o n n e c t i v e a lm o s t c o n t i n u o u s w i t h th e f i l a m e n t . — M ts. a l o n g t h e c o a s t , 500 t o 1 5 0 0 f t . , f r o m S a n L o u i s O b i s p o Co. t o t h e S a n t a I n e z , S a n t a M o n i c a , a n d S a n G a b r i e l M o u n ta in s. M a y -Ju n e .(A u d i b e r t i a n i v e a , B e n th .)

15 E n o e lia c a l i f o r n i c a , N u t t . , h e r e a f t e r r e f e r r e d E.

c a lifo rn ic a , is

d e s c r ib e d by J e p s o n

to as

(2 6 ) a s :

S t e m s s l e n d e r , f o r m i n g a s p r e a d i n g c lu m p 2 t o 3 1 / 2 f t . h i g h , woody o n ly a t t h e b a s e ; l e a v e s o v a te t o l a n ­ c e o la te , a c u te , m o s tly e n t i r e , g re e n , m in u te ly sc a b ro u s g l a b r a t e , 1 1 /2 t o 2 1 /2 i n . lo n g , on s h o r t p e t i o l e s ; h ead s la r g e , s o l i t a r y , te rm in a tin g e lo n g a te d n e a r ly naked p e d u n c le s ; d is k p u r p le , 1 /2 to 3 /4 in . b ro a d ; in v o lu c r e s d e n s e ly v h ite v i ll o u s , e q u a lin g th e d i s k - c o r o l l a s ; ra y s 16 t o 2 5 , b r i g h t le m o n - y e llo w , 3 / 4 t o 1 1 / 4 i n . l o n g ; c o r o lla - lo b e s o f d is k e i t h e r g la b ro u s o r p u b e s c e n t.— D ry h i l l s i d e s and s lo p e s o f b a r r a n c a s , S a n ta B a rb a ra t o S an D ie g o a n d e . t o R i v e r s i d e . M a r .- M a y . II.

BASES FOR SELECTING- SPECIES

In th e s e le c tin g o f d e c id e d

s u ita b le p la n t sp e c ie s,

it

w as

t h a t t h o s e s h o w in g t h e m o st o b v io u s d i f f e r e n c e s

d iffe re n t

c o n d i t i o n s s h o u ld be u s e d .

under

Among t h e a v a i l a b l e

m a t e r i a l t h e t h r e e s p e c i e s c h o s e n sh o w ed a w i d e r a n g e a n d a d e c id e d ly u n sta b le

g ro w th -fo rm .

S.

c o n d itio n s in S o u th e rn C a lif o r n ia

a p i a n a g r o w s u n d e r m an y

(1 9 ),

l e u c o p h y l l a h a s a m ore l i m i t e d r a n g e ; v a rio u s a ltitu d e s

o f f r o m 30 t o

(2 6 ), w h ile S.

th e l a t t e r

o ccu rs a t

3100* f e e t above se a l e v e l

on a l l e x p o s u r e s a n d u n d e r v a r y i n g c o n d i t i o n s o f m o i s t u r e and c o m p e titio n

(1 9 ),

(2 6 ).

E.

c a l i f o r n i c a h a s a w id e r a n g e

of

o v e r f o u r h u nd red m ile s a lo n g th e c o a s t a t v a rio u s a l t i t u d e s , e x p o su re s, an d m o is tu re c o n d itio n s

(1 9 ),

(2 6 ).

T he v a r i e t y

o f c o n d i t i o n s u n d e r w h ic h t h e s e p l a n t s grow , a s l i s t e d T ab le I ,

p a g e 5 6 , was n o t

in

so g r e a t in th e S a n ta B a rb ara r e ­

g io n a s i t w as, f o r e x a m p le , i n t h e San D ieg o r e g i o n .

16

A n o th e r b a s i s

f o r th e

s e le c tio n o f th e above th re e

s p e c i e s w as t h e i r a v a i l a b i l i t y and a b u n d a n c e . sary to

w as n e c e s

t r a n s p l a n t m any sp e c im e n s an d o f t e n u s e e n t i r e

o f ste m s f o r s e c t i o n i n g d u rin g t h e m ic r o s c o p ic p a tio n o f a v a i l a b i l i t y le d to th re e .

As i t

F a c ility

sc h e d u le s o f m ic ro -

fig u re d in t h e i r

in u p ro o tin g and t r a n s p l a n ti n g a ls o had to III.

stu d y , a n t i c i ­

t h i s e a r ly c o n s id e ra tio n of th e

o f h a n d lin g d u rin g th e

te c h n ic a l p ro ced u res a ls o

le n g th s

se le c tio n .

E ase

be c o n s i d e r e d .

LOCATION OF EXPERIMENTAL STATIONS

The s t a t i o n

a t w h ic h S . a p i a n a o c c u r r e d i n t h e woody

g r o w t h - f o r m w a s l o c a t e d o n C am in o C i e l o , a m o u n t a i n r o a d w h ic h f o llo w s t h e

c r e s t- s u m m it o f t h e S a n ta I n e z M o u n ta in s

a b o u t se v e n m i l e s from S a n ta B a r b a r a , n o te d fro m F ig u r e 1 , page 21, t h e fan is

sta tio n

o f San M arco s Canyon w h ere a s t e a d y , c o n tin u o u s a lm o st d a i l y

d ark .

is

As m ay b e

i s on t h e a p i c a l o f f - v a lle y u p -d rau g h t

from d a y l i g h t u n t i l

T he s o i l a t t h i s s t a t i o n

M io c en e s a n d s t o n e th e

C a lifo rn ia .

sh o rtly a f te r

g e n e r a l l y decom posed

(1 1 ), w ith ab o u t 2 p e r c en t m o is tu re d u rin g

c r i t i c a l m o n th o f O c to b e r .

The e x p o s u r e i s

so u th e rn ,

and

t h e p l a n t s w e r e g e n e r a l l y n o t m o r e t h a n 30 t o 4 0 cm. i n h e i g h t and o f s c a t t e r e d o c c u rre n c e r a t h e r th a n i n

c lu s te re d

The a l t i t u d e h e r e a p p r o x im a te d 2800 f e e t .

H e re a fte r th is

s t a t i o n w i l l be r e f e r r e d to a s S ta tio n ."

I t ty p ifie s

th e

sta n d s.

"Woody S a l v i a a p i a n a

th e v a rio u s lo c a tio n s th ro u g h o u t

17 S o u th e r n C a l i f o r n i a w here S. a p ia n a w as s e e n to o c c u r a s a woody h a l f - s h r u b . T he s t a t i o n a t w h i c h t h e sam e s p e c i e s o c c u r r e d i n t h e s u f f r u t e s c e n t g r o w t h - f o r m was l o c a t e d R anch on a f o o t h i l l b o r d e r i n g th e

on t h e D a v i d G r a y

so u th w e st s id e o f th e S a n ta

I n e z V a l l e y , a b o u t t e n m il e s from t h e

v illa g e

o f S a n ta In e z .

A s m ay be n o t e d f r o m F i g u r e 3 , p a g e 2 3 , t h e p l a n t s w e r e d i s ­ t r i b u t e d a b u n d a n tly on open h i l l s i d e s a p p e a r in g w i t h O nerous a g r i f o l i a , v a rio u s a n n u a ls.

N o th in g i n

was n o t e d a t t h i s s t a t i o n , te c te d

o w in g t o

t h e form o f a c o n t i n u a l w in d

th e p l a n t s b e in g f a i r l y w e ll p ro ­

The s o i l w as h e a v y a d o b e c l a y

c o n s id e ra b le o rg a n ic a d m ix tu re ,

th is

at

th e

s t a t i o n was

p l a n t s r a n g e d i n h e i g h t f r o m 50 cm. t o

T he a l t i t u d e

h e r e a p p ro x im a te s 800 f e e t .

s t a t i o n w i l l be r e f e r r e d

a p ia n a S t a t i o n . M I t

(1 1 ) w ith a

th e m o is tu r e b e in g a b o u t 3 p e r

The g e n e r a l e x p o s u r e

s o u th e a s te rn , and th e a m e te r.

N ee. , C e an o th u s s p p . , and

th e i r r e g u l a r to p o g ra p h y and th e n e a rb y

p resen ce of tr e e s .

c e n t in O c to b e r.

an d i n w ooded c a n y o n s ,

to a s

t y p i f i e s th e

S. a p ia n a o c c u rs in th e

th e

H e re a fte r

" S u ffru te sc e n t S a lv ia

v a r io u s l o c a t i o n s w here

s u f f r u t e s c e n t g ro w th -fo rm .

T h e s t a t i o n a t vh i c h t h e

sam e s p e c i e s o c c u r r e d i n t h e

n e a r - h e r b a c e o u s g r o w th - f o r m was l o c a t e d a b o v e t h e S a n t a B a r b a r a B o t a n i c G a rd e n i n M is s io n C an y o n , a b o u t tw o m i l e s fro m S a n ta B a r b a r a ,

C a lifo rn ia .

As m ay b e n o t e d i n F i g u r e 5 ,

p ag e 2 5 , t h e p l a n t s w e re on an o p en h i l l s i d e p e t i t i o n w ith

o th e r s p e c ie s d u rin g th e

free

f r o m com­

c r i t i c a l p e rio d .

T hey

18 a p p e a re d a s p e r e n n i a l h e r b s , w i th v e r y s l i g h t l y woody u n d e r ­ g round p a r t s .

The s o i l i s a l i g h t

c l a y lo a m w i t h a n a d m i x t u r e

o f l e a f m o ld a n d s a n d .

M o istu re a t t h i s

to

sta tio n .

irrig a tio n

above th e

m a te s 500 f e e t . a s th e

H e re a fte r th is

s ta tio n v a rie d

T he a l t i t u d e

e x is t in

e n c o u n te re d a t a l l th e

S.

a p ia n a w as

t h e h e r b a c e o u s g r o w t h - f o r m , a n d w hen

s ta n d s w ere s m a ll ,

C o u n t y o n t h e n o r t h s l o p e o f M ount P i n o s , C o u n ty n e a r O c e a n s id e ,

h e re a p p ro x i­

s t a ti o n w i l l be r e f e r r e d to

"H erb aceo u s S a l v ia a p ia n a S t a t i o n . w

r a r e l y fo u n d to

o w in g

one i n V e n tu ra one i n S an D ie g o

an d a n o t h e r i n Los A n g e le s C o u n ty n e a r

t h e m o u th o f M in t C anyon.

Such i s o l a t e d

sp e c im e n s d id n o t

seem t o w a r r a n t t h e am ount o f t r a v e l i t w o u ld h a v e e n t a i l e d to

e s ta b lis h

c u ltiv a te d

s ta tio n s in

th o se re g io n s ,

s o t h e m ore o r l e s s

s t a n d s w ere em p lo y ed .

The s t a t i o n

a t w h ic h S.

woody g r o w th - f o r m w as l o c a t e d

le u c o p h y lla o c c u rre d in th e on t h e c o a s t s i d e o f a f o o t ­

h i l l o f t h e S a n ta I n e z M o u n ta in s b e tw e e n R a t t l e s n a k e

C anyon

a n d S y c a m o r e C a n y o n , a b o u t tw o m i l e s f r o m S a n t a B a r b a r a . m ay b e n o t e d f r o m F i g u r e 7 ,

page 2 7 , t h i s

sta tio n

a p i c a l e n d o f Sycam ore C anyon, u n d e r t h e i n f l u e n c e

As

i s a t th e of up­

d r a u g h t s fro m t h a t d e p r e s s i o n a s w e ll a s from R a t t l e s n a k e and M is s io n C anyons.

T he s o i l i s

com posed o f l i g h t

c la y w ith

som e l e a f m o l d , cow m a n u r e , a n d s m a l l p i e c e s o f b r o k e n s a n d ­ sto n e . to th e

I n O c t o b e r , a b o u t 3 p e r c e n t m o i s t u r e w a s f o u n d o w in g f a c t t h a t t h e p l a n t s g ro w i n

th e

im m e d ia te p r o x i m i t y

19 of la rg e

sa n d sto n e b o u ld e rs .

T he e x p o s u r e i s

g e n e ra lly w e st­

ern ,

a n d t h e p l a n t s w e r e f r o m 3 0 t o 6 0 cm. i n h e i g h t .

th is

sta tio n

a s w as t h e

th e

p l a n t s w ere c l u s t e r e d r a t h e r t h a n s c a t t e r e d ,

c a s e a t t h e Woody S a l v i a a p i a n a S t a t i o n .

t i t u d e h e r e a p p r o x i m a t e s 500 f e e t . w i l l be r e f e r r e d

At

to a s t h e

H e re a fte r t h i s

The a l ­ sta tio n

"Woody S a l v i a l e u c o p h y l l a S t a t i o n . "

T h e s t a t i o n a t v f o ic h t h i s

s p e c ie s o ccu rred in th e s u f-

f r u t e s c e n t g r o w th - f o r m w as l o c a t e d on t h e f l o o r o f t h e S a n ta I n e z V a l l e y on t h e San L u c a s R a n c h , a b o u t s i x m i l e s fro m t h e v i lla g e o f S a n ta In e z .

As m ay b e n o t e d f r o m F i g u r e 9 ,

page

2 9 , t h e s t a t i o n w a s on p r a c t i c a l l y l e v e l g r o u n d i n a s a v a n n a o f £^. a g r i f o l i a .

The m o i s t u r e was l e s s t h a n 4 p e r c e n t i n

O c to b e r, a lth o u g h th e S a n ta I n e z R iv e r flo w s w i t h in a h a l f m ile o f th e

sta tio n .

The s o i l i s

w ith ro ck s o f v a rio u s o r ig in . V a lle y a re

a s a n d y a l l u v i a l lo a m ( 1 1 )

A i r m o v e m e n ts i n t h e S a n t a I n e z

v a r i a b l e , t h e p r e v a i l i n g w in d s b e in g g e n t l e an d

from t h e w e st

(3 8 ).

The s t a t i o n ,

ly in g as i t

v a l l e y , w a s e x p o s e d on a l l s i d e s , a n d t h e a m e te r in h e ig h t. H e re a fte r th is

T he a l t i t u d e

does in

p l a n t s w ere a l l u n d e r

h e r e a p p r o x i m a t e s 20 0 f e e t .

s t a t i o n w i l l be r e f e r r e d to a s th e

c e n t S a lv ia L e u c o p h y lla S t a t i o n . " l o c a t i o n s w here S .

th e open

It

t y p i f i e s th e

" S u ffru te s­ v a rio u s

le u c o p h y lla o ccu rs as a s u f f r u te s c e n t

h a lf-sh ru b . The s t a t i o n a t w h ic h th e

sa m e s p e c i e s g r o w s a s a n e a r l y

h e r b a c e o u s p e r e n n i a l was l o c a t e d n e a r t h e H e r b a c e o u s S a l v i a

20 a p i a n a S t a t i o n b e lo w t h e S a n t a B a r b a r a B o t a n i c G a r d e n , c o n d i t i o n s o f w h ic h h av e been l i s t e d t h i s s t a t i o n w i l l be r e f e r r e d le u c o p h y lla S t a t i o n . " ist

S.

on p a g e 1 8 .

to a s th e

th e

H e re a fte r

f,H e r b a c e o u s S a l v i a

le u c o p h y lla w as o f t e n

fou n d t o

ex­

in th e h e rb a c e o u s g ro w th -fo rm b u t o n ly a s s i n g l e p l a n t s .

S e v e ra l w id e ly s e p a ra te d

s ta n d s w ere d is c o v e r e d a lo n g t r i b u ­

t a r i e s o f th e S a n ta M a ria R iv e r , b u t a g a i n t h e d i s t a n c e d id n o t se e m t o w a r r a n t e s t a b l i s h m e n t o f a s t a t i o n . page 3 1 , g iv e s a g e n e r a l id e a o f th e

F ig u re 11,

situ a tio n a t th is

sta ­

tio n . T he l o c a t i o n s a t w h i c h E .

c a lifo rn ic a

o ccu rred in -th e

n e a r-w o o d y g ro w th -fo rm was l o c a t e d i n a r e g i o n f i f t e e n m i l e s s o u t h o f S a n t a B a r b a r a , know n a s t h e R i n c o n d e l M a r . su m m it o f a c l i f f h ig h t i d e

o f th e

a b o u t tw o h u n d r e d y a r d s a i r ocean th e

re a c h e d a c lim a x .

lin e

At th e

from t h e

i n c r e a s in g w o o d in e ss o f th e p l a n t s

As may b e n o t e d i n F i g u r e 1 3 , p a g e 3 3 , t h e

s p e c i e s c o n s t i t u t e d th e d o m in a n t v e g e t a t i o n o f th e a r e a , w i t h sc a tte re d

isla n d s

exposed to th e

of o th e r g en era.

T he s t a t i o n

sta tio n

d u r in g th e c e n t.

is

s o u t h w e s t , a n d an a lm o s t c o n t i n u o u s o f f - s e a ,

u p -d ra t£ g h t p r e v a i l s d a i l y a g a i n s t th e p l a n t s . th is

its e lf

The s o i l a t

i s a v e ry f i n e l y b ro k en s h a l e , and th e m o is tu re

c r i t i c a l p e r i o d o f O c to b e r w as l e s s t h a n 3 p e r

A l l th e p l a n t s w ere l e s s

d e n se ly c lu s te r e d .

The a l t i t u d e

th a n one m e te r in h e ig h t and a p p r o x i m a t e s 300 f e e t .

a f t e r t h i s s t a t i o n w i l l be r e f e r r e d t o a s t h e

H ere

"Woody E n c e l i a

21

r L F i g .

1 .

Fig. 2.

T i l e

W o o d y

S a l v i a

a p i a n a

S t a t i o n .

The Woody Salvia apiana Station.

G e n e r a l

Detail

v i e w ,

22 c a lifo rn ic a

S ta tio n .”

It

t y p i f i e s th e

v a rio u s l o c a l i t i e s

fro m P o i n t C o n c e p tio n ( S a n ta B a rb a ra C o u n ty , E n s e n a d a , M ex ico , w h ere th e

C a lifo rn ia )

s p e c i e s was s e e n to

to

occur as a

n ear-w o o d y h a I f - s h r u b . T he s t a t i o n

a t w h ic h E.

c a lif o r n ic a o c c u rs in th e s u f­

f r u t e s c e n t g ro w th -fo rm was l o c a t e d

in a re g io n

s ix m ile s

n o r t h w e s t o f S a n t a B a r b a r a , kn ow n a s H e n r y ’ s B e a c h . tw e lv e f e e t o f th e t i o n o f th e

h ig h t i d e

s p e c i e s was i n

le v e ls

th e s u f f r u t e s c e n t c o n d i­

e v id e n c e ;

g e n e r a l l y th ro u g h o u t th e a r e a .

tio n

its e lf

se a w in d s.

th is

c o n d itio n p r e v a ils

A s m ay be n o t e d i n F i g u r e 15*

page 3 4 , th e s p e c i e s c o n s t i t u t e s th e w ith s c a tte r e d

d o m in a n t v e g e t a t i o n ,

i s l a n d s o f R h u s i n t e g r i f o l i a , B.&W.

i s exposed to th e T he s o i l i s

a t th is

le ss

o ff-

g e n e r a l l y decom posed l im e s t o n e and

s t a t i o n d u rin g

sank to s l i g h t l y

The s t a ­

n o rth w e st w ith v a r ia b le

s h a l e , w i t h p a tc h e s o f d ia to m a c e o u s e a r t h e x p o sed . tu re

W ith in

th e

th a n 2 p e r

c r i t i c a l p e rio d in c e n t a t 20 cm.

The m o is ­ O c to b er s

The p l a n t s

r a n g e d f r o m 50 cm. t o a m e t e r i n h e i g h t a n d w e r e c l o s e l y c lu s te re d .

The a l t i t u d e

w ith in th e b o u n d a rie s o f th e

a p p r o x i m a t e 2 0 t o 30 f e e t a b o v e s e a l e v e l . s t a t i o n w i l l be r e f e r r e d c a lifo rn ic a S ta tio n ."

It

to

as th e

ty p ifie s

sta tio n

H e re a fte r th is

" S u ffru te s c e n t E n c e lia th e v a r i o u s l o c a t i o n s from

P o i n t C o n c e p tio n t o E n s e n a d a , M ex ico , w h ere E .

c a lifo rn ic a

o ccu rs as a s u f f r u te s c e n t h a lf - s h r u b . D u rin g th e c o u rs e o f t h i s

in v e s tig a tio n E.

c a lifo rn ic a

F i g .

3 *

Fig* 4*

T h e

S u f j f r u t e s c e n f c

S a l v i a

a p i a n a

S t a t i o n .

The SuJffnatescent Salvia apiana Station.

G o n e x ’a l

Detail.

v i e w .

zu was n o t f o u n d i n t h e h e r b a c e o u s g r o w t h - f o r m . As t h e p r o b l e m became w e l l d e f i n e d ,

i t was o b v i o u s

t h a t the e s ta b lis h m e n t o f s t a t i o n s e x h i b itin g th e v a r io u s g r o w t h - f o r m s w o u ld be n e c e s s a r y .

The f i r s t r e q u i s i t e seemed

t o be t h e l o c a t i o n o f a s u i t a b l e s t a t i o n f o r ea c h o f t h e t h r e e s p e c i e s a t w h ic h t h e p l a n t s o c c u r r e d i n t h e woody grow th-form .

The same was t r u e f o r t h e s u f f r u t e s c e n t an d

herbaceous h a b i t s .

The t a s k was s u c c e s s f u l l y a c c o m p l i s h e d

e x c e p t f o r th e h e r b a c e o u s g r o w t h - f o r m ( i f i t e x i s t s a t a l l ) of S. c a l i f o r n i c a .

In o rd e r t o c o n s id e r a l l f a c t o r s of en­

v i r o n m e n t and d e t e r m i n e w h i c h c o m b i n a t i o n was c r i t i c a l , i t was f u r t h e r n e c e s s a r y t o s e l e c t , a s f a r a s p o s s i b l e , w i d e l y s e p a ra te d s t a t i o n s w ith d i f f e r e n t ex p o su re s, a v a i l a b l e m ois­ t u r e , w in d v e l o c i t i e s , a l t i t u d e s , an d s o i l c o n d i t i o n s .

Fur­

t h e r , d i f f i c u l t i e s were e n c o u n te r e d b e c a u se o f t h e f a c t t h a t many p l a n t s w e r e r e q u i r e d a t e a c h s t a t i o n f o r t r a n s p l a n t i n g and s e c t i o n i n g .

Upon t h e s e b a s e s t h e e i g h t s t a t i o n s w e r e

estab lish ed . IV.

LOCATION OF THE SPECIES IN OTHER AREAS

As a b a s i s o f c o m p a r i s o n , j o u r n e y s w ere made i n t o n e a r l y a l l i m p o r t a n t r e g i o n s o f S o u t h e r n C a l i f o r n i a w h ic h t h e t h r e e s p e c i e s w e r e known t o i n h a b i t .

D iffe re n c e s in h a b i t of

g r o w t h , s o i l s , m o i s t u r e , t e m p e r a t u r e , and o t h e r w e a t h e r c o n ­ d i t i o n s were checked.

C a r e f u l s c r u t i n y was e x e r c i s e d b e c a u s e

F i g *

5 *

Fig* 6*

T h e

H e r b a c e o u s

S a l v i a

a p i a n a

S t a t i o n *

The Herbaceous Salvia apiana Station*

G e n e r a l

Detail

v i e w *

o f th e p re s e n c e o f f i e l d

h y b rid s

c o n s i d e r a t i o n w as g iv e n t o c o m p a rab le w ith t h e

(1 9 )»

On t h e s e

jo u rn ey s

o t h e r s u f f r u t e s c e n t s w h ic h a r e

th re e of th is

e x p e rim e n t in

th e ir reac­

tio n s to e n v iro n m e n ta l c o n d itio n s . V.

TYPES OF SOUTHERN CALIFORNIA ENVIRONMENTS EXAMINED

The f o l l o w i n g w e re i n v e s t i g a t e d (3 3 ): L i t t o r a l a n d dune ( 4 9 ) , Sea c l i f f

(26),

(19)

C o a sta l sag eb ru sh C h ap arral ( 2 ) ,

(5 4 )

(19)

(15),

(45),

(46),

(51)

W o o d la n d a n d s a v a n n a ( 3 3 ) G ra ssla n d R ip a ria n S e m i-a rid

(33) (24) (19)

M o ja v e D e s e r t

(30),

(44)

M o ja v e D e s e r t m o u n t a i n r a n g e s M o ja v e D e s e r t s t r e a m b e d s M o ja v e D e s e r t v a l l e y s C o lo ra d o D e s e r t

(30),

(44),

(7)

C o lo ra d o D e s e r t m o u n ta in r a n g e s C o lo rad o D e s e r t stre a m

beds

C o lo ra d o D e s e r t v a l l e y s M o n ta n e v e g e t a t i o n b e l t s

(26),

( 3 8 ),

(20)

27

T h e

W o o d y

S e l v l a

T d e

W o o d y

S a l v i a

F i g . l e u s o p h : ; 1 1 a

7 S t a t i o n .

G e n e r a l

F i g .

B . S t a t i o n .

D e t a i l

v i e w .

28 The i n v e s t i g a t i o n o f t h e l i t t o r a l a n d dune r e g i o n s b eg an a t P o i n t C o n c e p tio n , S a n ta B a rb a ra C o u n ty , C a l i f o r n i a , a n d a t M oro B a y ,

C a lifo rn ia ,

a n d e n d ed a t E n s e n a d a , M e x ico .

S u f f r u t e s c e n t p l a n t s w ere l i s t e d ous e n v iro n m e n ta l f a c t o r s .

a n d n o t e s m ad e o n t h e v a r i ­

W h ere r o a d s m ade i t

h i l l s , m e s a s , v a l l e y s b e lo w t h e

ch ap arral a re a s,

o cean , and v e g e ta tio n of n o rth e rn f r o m t h e sam e p o i n t o f v i e w .

p o ss ib le

th e

fa c in g th e

s lo p e s w ere i n v e s ti g a t e d

O n ly t h e

c h a p a rra l o f th e

S a n ta B a r b a r a , S a n ta M o n ica, San G a b r i e l ,

S a n ta A na, a n d San

D ie g o r e g i o n s w as s t u d i e d t o a n y e x t e n t .

The w o o d la n d and

s a v a n n a a r e a s o f L o m p o c , S a n t a I n e z , H uenem e P l a i n ,

th e

C h a n n e l I s l a n d s , a n d s e v e r a l v a l l e y s o f t h e S a n D ie g o r e g i o n w ere v i s i t e d a n d n o te s

c o m p ile d .

S in c e

th e n a t u r a l g r a s s la n d s

and v a lle y s o f S o u th e rn C a lif o r n ia do n o t c o n ta in a c o n s p ic u ­ ous p a r t o f th e n a ti v e fra g m e n ta ry r e s u l t s p o s s ib le

flo ra

because o f a g r i c u lt u r a l p r a c tic e s ,

o n l y w ere o b t a i n e d

th e can y o n and r i p a r i a n

W h e re

c o n d i t i o n s w ere c h e c k e d b e ­

c a u s e o f t h e m ore m e s ic c o n d i t i o n s o f yond th e

in such a re a s .

scope o f t h i s p a p e r to l i s t

th e se a re a s . th e m a l l ,

It

is

be­

but such w e ll-

know n c a n y o n s a s S a n G a b r i e l , S a n t a A n a , S e s p e , S a n P a s c u a l , F o x en , G a v io ta , a n d T e c h a c h a p i w ere s t u d i e d .

T he s e m i a r i d

r e g i o n s o f A n t e lo p e T a l l e y , H e m et, M in t C an y o n , C a jo n P a s s , Ju lia n ,

Jacum ba, a n d th e

strip

D i e ^ o a n d E n s e n a d a k now n a s v isite d .

of co ast lin e

b e tw e e n S an

th e W est C o a s t F oggy D e s e r t , w ere

S u f f r u t e s c e n t p l a n t s w ere c o n s id e r e d in th e v i c i n i ­

t i e s o f J o s h u a N a t i o n a l M o n u m e n t, A n z a r e g i o n ,

B orego V a l l e y ,

T h e

S u f f r u t e s c e n t

S a l v i a

F i g . 9 . l e u c o p h y l l a

s t a t i o n .

Fig. 10. The Suffrutescent Salvia leucophylla Station.

G e n e r a l

Detail

v i e w .

30

L u c e r n e V a l l e y , V i c t o r v i l l e , a n d i n t h e S u p e r s t i t i o n Moun­ ta in s.

I n t h e s e r e g i o n s t h r e e t y p e s o f e n v i r o n m e n t w ere ob ­

s e rv e d : m ountain r a n g e s w ith d i f f e r e n t e x p o s u r e s , s tre a m b eds, t h e v i c i n i t y o f o a s e s , a n d what m i g h t be t e r m e d "o p e n d e s e r t . " On t h e s e j o u r n e y s t h e m o n ta n e b e l t s o f v e g e t a t i o n w e r e ob­ s e r v e d i n t h e f o l l o w i n g r a n g e s : San R a f a e l , San G a b r i e l , San B e r n a r d i n o , T e h a c h a p i , P a l o m a r , an d San J a c i n t o . V I.

ESTIMATION OF CRITICAL ENVIRONMENTAL CONDITIONS

No d e t a i l e d a n a l y s i s o f t h e

s o i l s a t the v ario u s s t a ­

t i o n s seemed n e c e s s a r y a s t h e y w ere n e a r l y a l l d i f f e r e n t i n c o m p o s i t i o n , o r i g i n , a n d t e x t u r e , r a n g i n g from l i g h t s a n d y a t t h e Woody S a l v i a a p i a n a S t a t i o n t o t h e h e a v y ad o b e o f t h e S u ffru te sc e n t S a lv ia apiana S ta tio n .

F iv e k ilo g ra m sam ples

o f s o i l w e r e t a k e n f r o m e a c h s t a t i o n a t a d e p t h o f 20 c m . , s e a l e d i m m e d i a t e l y , oven d r i e d , w e ig h e d a g a i n , and t h e p e r ­ c e n t a g e o f m o i s t u r e d e t e r z a i n e d on t h e d r y w e i g h t b a s i s : w e t w t . o f s o i l — d r y w t . x 100 = fo o f m o i s t u r e dry w t. of s o i l Wind v e l o c i t i e s a n d d i r e c t i o n s w ere c h e c k e d w i t h t h e U n i t e d S t a t e s F o r e s t S e r v i c e and t h e w e a t h e r r e c o r d i n g s t a ­ t i o n s ; p r e v a i l i n g d r a u g h t s w e r e o b s e r v e d , a n d t h e h i g h s and low s o f t e m p e r a t u r e o f t h e g e n e r a l r e g i o n s w ere n o t e d , a l ­ t h o u g h t h e d a t a was a d m i t t e d l y i n s u f f i c i e n t . The g e n e r a l t e m p e r a t u r e o f t h e e n t i r e r e g i o n o f t h e

T h e

H e r b a c e o u s

S a l v i a

P i g . 1 1 . l e u c o p h y l l a

S t a t i o n .

Fig. 12. The Herbaceous Salvia leucophylla Station.

G e n e r a l

Detail.

v i e w .

32 sta tio n s

i s ra ild .

T h e s t a t i o n s w e r e s o a r r a n g e d t h a t n o tw o

h a d e x a c t l y t h e sam e e x p o s u r e , f o r i t

w as n o te d t h a t d i r e c ­

t io n a l expo su re a p p a re n tly e x e rc is e d

some p a r t o f t h e i n f l u ­

ence upon th e p a r t i c u l a r At none o f th e

s t a t i o n s w ere t h e

cro w d ed o r e v e n d i r e c t l y g e n e r a e x c e p t a t t h e E. a g rifo lia t h e w in d t o

g ro w th -fo rm o f th e p l a n t s .

e ffe c te d

s p e c ie s un d er stu d y

by th e p r e s e n c e o f o t h e r

c a lifo rn ic a

sta tio n s.

No d o u b t t h e

a t th e S u f f r u te s c e n t S a lv ia a p ia n a S ta tio n so m e e x t e n t ,

e i t h e r s id e o f th e

b u t th e p la n ts f u l ly

s ta tio n a ls o

b roke

e x p o s e d on

ap p eared in th e s u f f r u te s c e n t

g ro w th -fo rm . V II.

CRITICAL FACTORS

The d e t e r m i n a t i o n o f t h e o f d e d u c tio n .

c r i t i c a l f a c t o r s w as a m a t t e r

The t r a n s p l a n t i n g o f s p e c im e n s fro m o n e a r e a

to a n o th e r and t h e i r su b seq u en t changes v e r i f i e d

by m ic ro s c o p ic

e x a m in a tio n c o n firm e d th e d e d u c tio n s . V III.

EXPERIMENTAL METHOD

On N o v e m b e r 15 a n d 1 6 ,

1941,

s i x p l a n t s o f t h e woody

fo rm o f S a l v i a a p ia n a w ere t r a n s p l a n t e d a p ia n a S t a t i o n to

th e

f r o m t h e Woody S a l v i a

S u ffru te s c e n t S a lv ia

a p ia n a

S ta tio n ;

s i x m o r e f r o m t h e W oody S a l v i a a p i a n a S t a t i o n w e r e t a k e n t o th e H erb aceo u s S a lv ia a p ia n a S ta tio n .

S ix p l a n t s w ere ta k e n

from t h e H e rb a c e o u s S a l v i a a p ia n a S t a t i o n

to t h e w oody s t a t i o n

33

T ile

W o o d y

E n c e l i a

P i g . c a l i f o r n i c a

1 3 . S t a t i o n .

G e n e r a l

T h e

W o o d y

S n c e l i a

P i g . c a l i f o r n i c a

1 4 . S t a t i o n .

D e t a i l .

v i e w .

T h e

S u f f r u t e s c e n t

E n c e l i a

F i g . 1 5 . c a l i f o r n i c a

S t a t i o n .

G e n e r a l

Ik Fig. 15. The Suffrutescent Encelia californica Station.

Detail

v i e w .

35 and s i x t o th e

su ffru te sc e n t s ta tio n .

S ix p l a n t s w ere ta k e n

f r o m t h e S u f f r u t e s c e n t S a l v i a a p i a n a S t a t i o n t o t h e W oody S a lv ia a p ia n a S t a ti o n and s ix t o th e H erbaceous S a lv ia a p ia n a S ta tio n .

T h i s m ade a t o t a l o f t w e l v e n ew p l a n t s a t e a c h

sta tio n .

T h e sam e p r o c e d u r e w a s f o l l o w e d a t e a c h s t a t i o n

S.

le u c o p h y lla and E.

tra n s p o rta tio n

c a lifo rn ic a .

In o rd er to f a c i l i t a t e

th e p l a n t s w e re c u t b a ck and c a r e f u l l y b a i l e d ,

p r e s e r v i n g a s m an y r o o t s a s p o s s i b l e . at

th e

slid e s

As a c o n t r o l ,

s t a t i o n s w ere u p ro o te d and r e p l a n t e d . fro m

n in e ty days

p la n ts

M ic ro sc o p ic

f r e s h m a t e r i a l w e re m ade a f t e r t h i r t y ,

s i x t y , and

(s e e C h a p te r IT f o r te c h n iq u e ) .

The t a b l e The r e s u l t s

of

on p a g e 58, i s

a reco rd o f

a re g iv e n i n C h a p te r T.

th e tra n s p la n tin g .

CHAPTER IV THE ANATOMICAL STUDY L a b o r a to r y w ork,

f o r th e

c o n firm a tio n o f w hat ap p eared

t o be m o r p h o lo g ic a l c h a n g e s due to th e

c e r t a i n e n v iro n m e n ta l

p e c u l i a r i t i e s , was b a s e d u p o n t e c h n i q u e s o u t l i n e d i n d e t a i l by Jo h a n se n in h i s P l a n t M ic ro te c h n iq u e

(27).

B ecause th e

s u f f r u t e s c e n t m a t e r i a l w as n e i t h e r woody n o r h e r b a c e o u s , i t w as o f t e n n e c e s s a r y t o m o d if y som ew hat t h e e x a c t i n g 'c h a r a c t e r in o r d e r t o T he c h i e f a i m o f t h i s

o b ta in th e

p a r t o f th e

c le a re st re s u lts .

i n v e s t i g a t i o n w as t o com­

p a re m o r p h o lo g ic a lly th e ste m t i s s u e s fo rm s w i t h in t h e

s c h e d u l e s o f m o re

of th e

th re e

g ro w th -

sp e c ie s.

I.

THE DETERMINATION OF .WOODINESS

The p h l o r o g l u c i n t e s t o f J o h a n s e n was u s e d w i t h b o t h f r e s h a n d p a r a f f i n

(27)

and W ilso n

(56)

se c tio n e d m a te ria l.

T he

f r e s h m a t e r i a l was s e c t i o n e d by m eans o f a h a n d m ic ro to m e a n d w ashed i n d i s t i l l e d

w a te r.

T h e s e c t i o n s w e r e p l a c e d on s l i d e s

in a dro p o f a s o lu tio n o f 0 .1 g r.

o f p h l o r o g l u c i n o l i n 10 c c .

o f 100 p e r c e n t e t h y l a l c o h o l and c o v e re d w ith a c o v e r s l i p . A f t e r a n h o u r o n e d r o p o f 25 p e r c e n t h y d r o c h l o r i c a c i d w a s a llo w e d t o d i f f u s e

in a t th e

edge o f th e

slid e .

The m a t e r i a l

w as e x a m in e d u n d e r t h e m i c r o s c o p e , a n d t h e a p p e a r a n c e o f a r e d - v io le t c o lo r a t th e c e l l w a lls

in d ic a te d th e p resen ce o f

37 lig n in .

The s p e e d o f t h e a p p e a r a n c e o f t h e r e a c t i o n w as i n

d ir e c t p ro p o rtio n to th e

f re s h n e s s o f th e

c h e m ic a ls.

The t e s t was c o n d u c te d w i t h e x a c t l y t h e

same p r o c e d u r e

f o r t h e p a r a f f i n s e c t i o n s a f t e r t h e y h a d b e e n b r o u g h t down th ro u g h th e a lc o h o ls

to w a te r , b u t. th e

s tre n g th o f th e

so lu ­

t i o n s w e r e c u t i n h a l f a n d 95 p e r c e n t a l c o h o l w a s s u b s t i t u t e d f o r th e a b s o lu te . A c o n tr o l t e s t w as in tr o d u c e d f o r b o th th e

f r e s h and

th e p a r a f f i n s e c tio n s a s su g g e ste d by Jo h an sen (op. p.

19 4 ).

To a 0 .0 0 1 p e r c e n t a q u e o u s s o l u t i o n o f m e th y l r e d ,

en o u g h p o ta s s iu m h y d r o x id e w as a d d e d t o r e n d e r i t A g a in t h e

y e llo w .

f r e s h m a t e r i a l w as c u t on a h an d m ic ro to m e , w ash ed

in d i s t i l l e d w a te r, above

c i t •,

so lu tio n .

and p la c e d on th e

P a ra ffin

slid e s

in a d ro p o f th e

s e c t i o n s w e r e b r o u g h t down t h r o u g h

t h e a l c o h o l s t o w a t e r , an d t h e t e s t w as c o n d u c te d i n e x a c t l y t h e sam e m a n n e r a s f o r t h e f r e s h m a t e r i a l . w a lls in

L ig n ifie d

c e ll

b o th c a s e s a c q u ir e d a b r i g h t r e d - v i o l e t c o lo r .

F u r­

t h e r d e t e r m i n a t i o n o f w o o d in e s s was b y a c t u a l i n s p e c t i o n o f th e

c o u n te r s t a i n e d p erm an en t s l i d e s . In te s tin g

( 0 .3 g r .

fo r sta rc h

p o ta ssiu m h y d ro x id e ,

sto ra g e , 1 .5 g r .

th e io d in e t e s t p o ta ssiu m

(27)

i o d id e , 100 c c .

w a t e r ) was u s e d . II.

PREPARATION OF'PERMANENT SLIDES

B e g i n n i n g 1 cm. b e h i n d t h e g r o w i n g t i p ,

se c tio n s

38 slig h tly 4 cm.

o v e r 1 cm. i n l e n g t h w e r e c u t f r o m l i v i n g

s e c t i o n s ' b e i n g d i s c a r d e d b e tw e e n t h e

u n t i l f i v e sp e c im e n s w ere o b ta in e d a lly

th is

p ro ced u re

v a rie d in th e

was o f t e n n e c e s s a r y t o

ste m s,

sa m p le s ta k e n ,

fro m e a c h s te m .

O c c a sio n ­

su ffru te sc e n t p la n ts, as

se c tio n e n tire

ste m s.

A ll m a te ria l

w as l a b e l e d by m eans o f num bered c a rd b o a rd t a g s a t t a c h e d th e

sp e c im e n s by lo n g c o tt o n s t r i n g s .

and e v a c u a te d o f w a te r in a s o l u t io n a lc o h o l (90 c c . ) , (5 c c .

g la c ia l a c e tic a c id

o f 40 p e r c e n t ) .

so lu tio n

th e y w ere p la c e d

it

to

T he t i s s u e s w e r e k i l l e d o f 70 p e r c e n t e t h y l (5 c c . ) ,

and fo rm a lin

A f te r th r e e h o u rs i h th e k i l l i n g in ru n n in g w a te r f o r

f i f t y h o u rs.

D e h y d r a tio n o f th e w oody a n d h e rb a c e o u s t i s s u e s w as by e t h y l a lc o h o l and t e r t i a r y

b u ty l a lc o h o l sc h e d u le s.

The s p e c im e n s

w e r e a l l o w e d t o r e m a i n f o r s e v e r a l d a y s i n 50 c c . a lc o h o l and x y l o l, th e

50 p e r c e n t o f e a c h , t h e

c o n c e n tra tio n of

l a t t e r b e in g g r a d u a lly in c r e a s e d u n t i l a l l

c o h o l w ere rem o v ed .

In filtra tio n

I t w as t r a n s f e r r e d

D e h y d ra tio n o f th e

in to

tra c e s of a l­

o f t h e p a ra w a x w as g r a d u a l ,

an d th e m a t e r i a l w as a llo w e d t o s t a y i n th e days.

o f a b so lu te

fresh

oven f o r seven

p a ra ffin ev ery o th e r day.

s u f f r u t e s c e n t m a t e r i a l w as a c c o m p lis h e d by

th e u se o f d io x a n and t e r t i a r y

b u ty l a lc o h o l.

in v o lv e d th e u s e o f a p a r a f f i n

o i l and a c o m m e rc ia lly p re p a re d

i n f i l t r a t i o n w ax.

A l l e m b e d d in g w as d o n e i n

In filtra tio n

ic e w a te r, and

t h e s e c t i o n s w e r e m i c r o t o m e d b e t w e e n 10 a n d 1 4 m i c r o n s .

They

s

w ere a f f i x e d

to t h e s l i d e s w i t h M a y e r 's a d h e s i v e a n d b r o u g h t

39 down t o 7 0 p e r c e n t a l c o h o l w h e r e t h e y w e r e s t a i n e d f o r f o r ty -e ig h t h o u rs in a 1 p e r cen t s o lu tio n m e t h y l c e l l o s o l v e a n d 30 p e r c e n t a l c o h o l .

of sa ffra n in

0 in

A f te r th e m ordant

and w a sh in g th e y w ere d i f f e r e n t i a t e d an d d e h y d ra te d w ith a 93 p e r c e n t e t h y l a l c o h o l a n d 0 . 3 tio n .

per cent p ic ric

G o p lin j a r .

c lo v e o i l ,

o f 95 p e r c e n t e t h y l a l c o h o l i n

s e c o n d s , a n d th e - s l i d e s w ere p lu n g e d i n t o

fo llo w e d by a l c o h o l an d x y l o l .

t h r e e w ashes in x y l o l, th e III.

a fte r

s p e c im e n s w e re m o u n te d i n b a ls a m .

sta tio n s.

p h lo em , a n d o t h e r s t r u c t u r e s

m ade.

F in a lly ,

THE ANATOMICAL STUDY

The s p e c ie s a t th e

stu d ie d

th e n ex t

C o u n te r s ta in in g w ith f a s t g re e n w as a llo w e d t o

p ro ceed fo r te n

in th e

The n a tu r e

o f t h e x y lem ,

ste m s o f e a c h s p e c i e s w as

i n m in u te d e t a i l and th e d ra w in g s i n C h a p te r V w ere S p e c im e n s c h o s e n f o r s e c t i o n i n g w e re random s a m p lin g s

and n o t s p e c i a l o r u n u su a l b ra n c h e s . ous s e r i e s w ere m ade, th e

g r o w t h - f o rm . se c tio n

U ach d ra w in g ,

on one s l i d e .

F ro m e a c h s t a t i o n numer*

d ra w in g s b e in g ta k e n d i r e c t l y fro m

s l i d e s w h ic h a p p e a re d t o

be m o st r e p r e s e n t a t i v e o f t h e h o w e v e r , w a s m ade o f a s i n g l e

P a r t i c u l a r a t t e n t i o n w as g iv e n th o s e

t i s s u e s w h i c h a r o s e f r o m a c a m b iu m m e r i s t e m , t h e p i t h w as c a r e f u l l y te s ts

so lu ­

T h e a c i d w a s s t o p p e d b y a d d i n g f i v e d r o p s o f ammonium

h y d r o x i d e t o 75 c c .

th e

a c id

ro le

o f th e

s t u d i e d , a s i t w as s e e n t o v a r y , a n d m any

w ere a p p l i e d to d e te r m in e t h e

am ount and n a t u r e o f

40 sc le re n c h y m a f i b e r s . o f th e p l a n t s a t th e

No d e t a i l e d sta tio n s

s tu d y o f th e

g r o s s a n ato m y

seem ed n e c e s s a r y a l t h o u g h t h e *

g en eral c h a ra c te ris tic s G reat care

o f th e

s e v e r a l sy ste m s w e re a s s e s s e d .

i n t e c h n i q u e a n d a n a l y s i s w as em p lo y ed

w herever th e tr a n s p la n te d w e r e m ad e i n e x a c t l y t h e

s p e c i e s w ere c o n c e rn e d .

S e c tio n s

sam e m a n n e r f o r t h e e x p e r i m e n t a l

p l a n t s a s f o r t h o s e w h ic h o c c u r r e d o r i g i n a l l y a t t h e s t a t i o n s . S a m p l e s w e r e t a k e n a n d s e c t i o n s m ad e a t t h e

tim e o f t r a n s ­

p l a n t i n g a n d a t i n t e r v a l s o f t h i r t y d a y s f o r one h u n d re d tw e n ty days.

C ro ss c o m p a riso n s w i t h th e

h a b i t a t w e r e m ade i n e a c h c a s e . th e

se c tio n s

from t h e

o rig in a l

W h e re d i f f e r e n c e s w e r e n o t e d

d e t a i l s w e re i n c o r p o r a t e d i n T a b l e s IV , V, a n d V I, p a g e s

59 t o

62. The s p e c i e s i n o t h e r a r e a s .

F o rm a lin -a c e to -a lc o h o l (12),

( 2 7 ) w as a lw a y s t a k e n a l o n g w hen j o u r n e y s i n t o w ere m ade, and

th e

sa m p le s p e c im e n s w e re k i l l e d

T h is w as b e c a u se a v e ry im p o r ta n t p a r t o f t h i s

o th e r v ic in itie s in th e f i e l d . s tu d y w as th e

c o m p a riso n o f m ic r o c l i m a t e s i n th e S a n ta B a r b a r a r e g i o n w ith th o s e o f o th e r r e g io n s w h ere s p e c i f i c fe c te d th e p la n ts

in e s s e n t ia ll y

c r itic a l d e ta ils a f­

t h e sam e w a y .

S e c t i o n s w ere

com pared w i t h t h o s e o b t a i n e d fro m t h e e x p e r i m e n t a l s t a t i o n s . H e re a g a i n m a t e r i a l was t a k e n fro m w e l l - e s t a b l i s h e d s t a n d s r a t h e r th a n fro m u n u s u a l p l a n t s . T a b l e X IV , p a g e 6 9 ,

show s th e

g en eral d iffe re n c e s

41 n o te d i n th e a n a to m ic a l s t u d i e s o f t h e p l a n t s found i n th e s e v e ra l re g io n s.

S p e c i f i c r e s u l t s o f th e

b e f o u n d i n C h a p t e r V.

in v e s tig a tio n w ill

CHAPTER Y RESULTS AND DISCUSSION A lth o u g h n o i n s t r u m e n t a l s t u d y w as m ade a t a n y s t a t i o n d u rin g th e

in v e s tig a tio n

o th e r

m o i s t u r e , t h e r e w ere c e r t a i n

th a n th e d e te rm in a tio n

of so il

o b v io u sly c r i t i c a l f a c t o r s .

At

e a c h s t a t i o n w h e re a s p e c i e s o c c u r r e d i n t h e woody g r o w th fo rm a m ore o r l e s s p re se n t.

c o n tin u o u s u p -d ra u g h t a i r

The F o r e s t S e r v i c e o f th e U n i t e d S t a t e s D e p a rtm e n t

o f A g r i c u l t u r e , Los P a d r e s D i v i s i o n , tio n

c u r r e n t w as

e stim a te d th a t th e

in v e l o c i t y w as fro m 2 t o 7 m .p .h .

d e p en d in g upon th e a l t i t u d e ,

d e c liv ity ,

to

v a ria ­

50 t o 6 0 m . p . h . ,

se aso n , and lo c a tio n .

V e l o c i t i e s , a s w e l l a s e v a p o r a t i n g pow er o f t h e a i r w ere n o t checked in s tr u m e n ta lly due to o b s e rv a tio n o f v a s t d if f e r e n c e s in e x tre m e ly d i f f e r e n t c lim a tic C a lifo rn ia . sam e.

r e g i o n s i n S o u t h e r n a n d L o w er

T he e f f e c t s u p o n t h e

C o n tin u o u s u p - d r a u g h ts

duced l i g n i f i c a t i o n

of c e rta in

p l a n t s w ere v i r t u a l l y th e

in e v ery case o b serv ed had in ­ e le m e n ts i n s p e c i e s w h ic h u n d e r

o t h e r c o n d it i o n s w ere e i t h e r s u f f r u t e s c e n t o r h e rb a c e o u s . I.

CRITICAL FACTORS AT THE STATIONS

T h e Woody S a l v i a a p i a n a S t a t i o n , o f th e

lo c a te d n e a r th e

crest

o u t e r S a n ta I n e z Range n e a r S a n ta B a r b a r a a t 2 ,8 0 0

f e e t e le v a tio n , w as exposed to u p -d ra u g h t due to

th e

so u th w ith a c o n tin u o u s

th e w i d t h o f S a n M arcos Canyon a n d t h e

43 g e n e r a l d e c l i v i t y , w h ic h e x te n d e d fro m t h e f iv e m ile s out in to tio n

is

th e G o le ta V a lle y .

se d im e n ta ry in o r i g i n

sta tio n

fo r about

The s o i l a t t h i s

sta ­

(1 1 ) w ith a m o is tu r e o f a b o u t

2 p e r c e n t d u rin g th e

c r i t i c a l m o n th o f O c t o b e r .

w e re a l l m ore o r l e s s

t h e sam e h e i g h t , r a n g i n g fro m 30 t o

4 0 cm.

As s e e n i n T a b l e I ,

p a g e 5 6 , an d from F i g u r e 1 ,

page 21, th e d e c l i v i t y a t t h i s A l l p l a n t s w ere in th e

s t a t i o n w as g e n e r a l l y s t e e p .

op en , a n d t h e r e w as a lm o s t no c o m p e ti­

t io n w ith o th e r p la n ts d u rin g th e The f a c t o r s a t t h i s est

The p l a n t s

l a t e w in te r and s p r in g .

s t a t i o n w h ic h e x e r c i s e d

i n f l u e n c e upon th e p l a n t s w ere e s tim a te d t o

th e

stro n g ­

be w in d , e x ­

p o su re , and s o i l m o is tu re . The S u f f r u t e s c e n t S a l v i a a p i a n a S t a t i o n , l o c a t e d on t h e D av id G ra y R anch i n t h e

fo o th ills

o f th e n o r th e r n s lo p e o f th e

S a n ta I n e z V a l l e y , a b o u t t e n m i l e s from t h e v i l l a g e o f S a n t a In ez,

a t 800 f e e t e l e v a t i o n , w as n o t e x p o sed t o an y p a r t i c u l a r

a i r c u rre n ts. and th e c lim a te s ta tio n

is

W in d s a r e v a r i a b l e i s g e n e r a l l y m ild

so u th e a s te rn .

f a i r l y w e ll d ra in e d . C e a n o th u s,

It

in th e S a n ta In e z V a lle y , (1 1 ).

The s o i l i s

The e x p o s u r e o f t h e

th e Z aca c la y ty p e

su p p o rts a v e g e ta tio n of

s p p * , a n d m an y a n n u a l s .

§•

a p ia n a .

a g rifo lia ,

T he m o i s t u r e d u r i n g t h e

c r i t i c a l m o n th o f O c t o b e r w a s a b o u t 3 p e r c e n t , p l a n t s d ra w in g fro m i t w e re th e

(1 1 ),

b u t th e

o n ly

s c a t t e r e d o a k s, C e a n o th u s, and

C o m p e titio n f o r l i g h t an d m o is tu r e was e n t i r e l y

am ong i n d i v i d u a l s o f S . a p i a n a .

A l l p l a n t s w ere i n t h e

open

uu and th e d e c l i v i t y

o f th e

on t h e s o u t h s i d e

o f th e v a l l e y , t h e h i l l w h ic h th e s t a t i o n

o c c u p ie s h as a fte rn o o n The e s t i m a t e d

slo p e

is

g e n tle .

A lth o u g h s i t u a t e d

shade.

f a c t o r s o f im p o rta n c e a t t h i s

sta tio n

a re e x p o su re and s o i l m o is tu re . The H e r b a c e o u s S a l v i a a p i a n a S t a t i o n ,

lo c a te d

ju st

a b o v e t h e S a n t a B a r b a r a B o t a n i c G a r d e n o n T u n n e l H oad n e a r S a n ta B a rb a ra a t an a l t i t u d e o f a b o u t 500 f e e t a b o v e s e a l e v e l , h a s a n e x p o s u r e to t h e w e s t . w in d a n d w ith o u t

B e in g p r o t e c t e d fro m a l l

c o m p e titio n w ith o t h e r p l a n t s f o r l i g h t and

m o is tu re , S. a p ia n a o c c u rs h e re a s a p e r e n n ia l h e rb . x y lem t i s s u e s ev er,

i m m e d ia te l y b e lo w t h e

sh o w t r a c e s o f l i g n i n .

c e n t, and th e

su rface

o f th e

M o istu re v a r ie d

s o i l w as a l i g h t sa n d y lo am .

The

s o i l , how­

fro m 4 to

8 per

T he s lo p e o f th e

h i l l s i d e w as g e n t l e . T he f a c t o r s a t t h i s s t a t i o n w h i c h i n d u c e d t h e h e r b a ­ c e o u s g ro w th -fo rm w ere e s t im a t e d t o be s o i l m o is tu r e an d p r o ­ te c tio n

fro m c o m p e tin g p l a n t s an d w in d . T h e W oody S a l v i a

h ills id e

above th e

l o c a t e d on t h e

c o n f l u e n c e o f R a t t l e s n a k e an d S ycam ore

C a n y o n s a b o u t tw o m i l e s o f a b o u t 500 f e e t ,

le u c o p h y lla S ta tio n ,

is

fro m S a n ta B a r b a r a , a t a n e l e v a t i o n

e x p o s e d to t h e w e s t and t h e r e

c o n t i n u o u s u p - d r a u g h t f r o m t h e s e tw o c a n y o n s ; never. o b s e rv e d a t t h i s

sta tio n .

The s o i l i s

is a

s t i l l a i r was com posed o f l i g h t

c l a y w i t h an a d m ix tu re o f o r g a n ic m a t t e r and s m a ll s t o n e s ;

th e

45 s o i l m o is tu re

i n O c to b e r w as fo u n d t o be a b o u t 3 p e r c e n t .

The p l a n t s w ere u n e v e n i n s t a t u r e ,

th o s e g ro w in g n e a r e r th e

la rg e

sa n d sto n e b o u ld e rs b e in g t a l l e r

tu re ,

p r o t e c t i o n and w a rm th .

v a rie d g re a tly ,

o w in g t o g r e a t e r m o i s ­

W in d v e l o c i t y a t t h i s

b u t a c o n tin u o u s d ra u g h t e x is te d .

s ta tio n I t w as n o t

deem ed n e c e s s a r y t o c h e c k v e l o c i t i e s a s s u f f r u t e s c e n t p l a n t s th r o u g h o u t S o u th e rn C a l i f o r n i a w ere o b s e rv e d to e x h i b i t l i g n ific a tio n under a l l

c o n d itio n s o f c o n tin u o u s o f f - v a l l e y and

o ff- s e a u p -d ra u g h ts, re g a rd le s s of t h e i r r a te A ll p la n ts a t t h i s

o f m o v em en t.

s t a t i o n w ere i n th e open a n d c o m p e titio n

w i t h o t h e r p l a n t s w a s l i m i t e d t o w i n t e r a n d sum m er a n n u a l s , a lth o u g h th e p l a n t s w ere o c c a s i o n a l l y s u b je c te d of g ra z in g g o a ts .

The s l o p e i s

T he f a c t o r s a t t h i s

to th e

rav ag es

g e n tle .

sta tio n ,

s o i l m o is tu re , ex p o su re,

and c o n tin u o u s u p - d r a u g h t, w ere e s tim a te d

to be r e s p o n s ib le

f o r t h e w oody g r o w th - f o r m . The S u f f r u t e s c e n t S a l v i a l e u c o p h y l l a S t a t i o n , a b o u t s i x m ile s fro m th e

v illa g e

o f t h e S a n t a I n e z V a l l e y on t h e tio n

o f a b o u t 200 f e e t ,

a lly

fro m t h e w e s t.

lo c a te d

o f S a n ta In e z on t h e

flo o r

S a n L u c a s JRanch a t a n e l e v a ­

i s exposed to

T he s o i l h e r e i s

v a r ia b le w in d s,

g en er­

know n a s M e t z v e r y f i n e

s a n d y lo au ^ 111) a n d h a d a m o i s t u r e c o n t e n t o f l e s s t h a n 4 p e r c e n t in O c to b e r.

T he s t a t i o n i s

a l m o s t on l e v e l g r o u n d i n a

s a v a n n a o f Q. a g r i f o l i a , w i t h w h i c h t h e f o r m o is tu re and s u n lig h t.

sage h a s to

c o m p e te

46 The f a c t o r s h e r e w h ic h i n f l u e n c e th e th e p la n t a re c o m p e titio n ,

g ro w th -fo rm o f

e x p o su re, and s o i l m o is tu re .

T he H e r b a c e o u s S a l v i a l e u c o p h y l l a S t a t i o n w a s l o c a t e d n e a r th e H erbaceous S a lv ia a p ia n a S t a ti o n a s th e o fte n

o c c u r r e d g ro w in g t o g e t h e r .

h ow ever, th e

tw o s p e c i e s

In o r d e r t o a v o id h y b r id s ,

s t a t i o n s w ere a h a l f m ile a p a r t , w h ic h gave t h e

H e r b a c e o u s S a l v i a a p i a n a S t a t i o n a b o u t o n e h u n d r e d f e e t m o re a ltitu d e . p o su re i s tio n

The s l o p e a t t h e s o u th e a s te rly .

in th e v i c in ity

th e r e g i o n and th e

sta tio n i s

g e n tle

H ere a g a in th e

and th e e x ­

p ra c tic e of ir r ig a ­

c a u se d th e s o i l m o is tu r e t o be h ig h f o r

season.

d r a u g h t was i n s u r e d by th e

P r o t e c t i o n a g a i n s t w in d an d l o c a l to p o g ra p h y and th e n e a rb y

p re s e n c e o f E u c a ly p tu s , s p p . , a lth o u g h th e l a t t e r c o m p e titio n f o r e i t h e r s o i l m o is tu re or s u n lig h t.

o f f e r e d no S.

l e u c o p h y l l a o c c u r r e d a l s o a s an h e r b a c e o u s p e r e n n i a l on p r e c i p ­ i t o u s s l o p e s p r o v i d e d t h e r e w e re am p le m o i s t u r e a n d p r o t e c t i o n fro m d r a u g h t.

O th e rw ise th e

s p e c i e s w as a lw a y s s e e n i n e i t h e r

t h e s u f f r u t e s c e n t o r t h e woody g r o w th - f o r m . The f a c t o r s a t t h i s s t a t i o n w h ic h e x e r c i s e d th e s t r o n g ­ e s t i n f l u e n c e u p o n t h e p l a n t s w ere e s t i m a t e d t o b e am p le s o i l . m o i s t u r e , p r o t e c t i o n fro m w in d a n d c o m p e titio n f o r l i g h t , m o is tu re , and s o i l space w ith o th e r s p e c ie s . T h e Woody E n c e l i a c a l i f o r n i c a S t a t i o n ,

lo c a te d f if te e n

m i l e s s o u t h o f S a n t a B a r b a r a on t h e K in c o n d e l M af c l i f f s a b o u t tw o h u n d re d y a r d s from t h e o c e a n a t an e l e v a t i o n

of

47 a b o u t 300 f e e t*

The s t a t i o n i s

d ra u g h ts d u rin g th e day. c liffs

S o il m o is tu re v a rie d a lo n g th e s e

b u t was n e v e r fo u n d t o

g e n e ra l ex p o su re i s

exposed to o f f - s e a up­

be lo w e r th a n 1 p e r c e n t .

s o u t h w e s t e r l y and th e

so il is

The

of sh a le

o rig in .

A f te r n o o n w in d s w e re e s t im a t e d t o r e a c h a v e l o c i t y

o f 18 t o

25 m . p . h .

n o t so g r e a t . p e n d ic u la r, v e g e ta tio n

h e re , a lth o u g h th e c o n tin u o u s d ra u g h t i s

The d e c l i v i t y

b u t w here th e

in t h i s re g io n i s

a lm o st p e r ­

s lo p e p e r m i ts th e g ro w th o f l i t t o r a l

im p e n e tra b le m a ts a r e fo rm ed .

S o il m o is tu re , u p -d ra u g h t, e x p o su re , and c o m p e titio n w ith o th e r p la n ts c a tio n

in th is

se e m e d t o b e t h e

c h ie f reaso n s

fo r

re g io n .

The S u f f r u t e s c e n t E n c e l i a o a l i f o r n i c a S t a t i o n c a te d on th e s t e e p h i l l s i d e m ile s fro m S a n ta B a rb a ra . f e e t above sea l e v e l , and

p o su re i s n o rth w e s te rly , th a n 2 p e r c e n t.

a t t h e Vfoody E n c e l i a

is

lo ­

b e h in d H e n r y ’ s B each a b o u t s i x T he a l t i t u d e

is

tw e n ty t o

th irty

th e a r e a i s n o t e x p o se d to a n y

c o n tin u o u s d r a u g h t , a l t h o u g h th e w in d s a r e

w as l e s s

lig n ifi-

and th e

v a ria b le .

s o i l m o is tu re

The e x ­

in O c to b e r

C o m p e titio n h e r e was k e e n e r t h a n

c a lifo rn ic a S ta tio n

due to th e m ild e r

d e c liv ity . An e s t i m a t i o n o f t h e f a c t o r s o f s t r o n g e s t i n f l u e n c e in c lu d e d e x p o su re, c o m p e titio n w ith o th e r p la n ts , m o is tu re •

and s o i l

48 II.

BEHAVIOR OF THE SPECIES UNDER EXPERIMENTAL CONDITIONS

To t h e W oody S a l v i a a p i a n a S t a t i o n o t h e r tw o f o r m s w e r e t r a n s p l a n t e d . tra n s p la n tin g th e

s i x p l a n t s o f the

W ith in n i n e t y d ay s a f t e r

t h e s u f f r u t e s c e n t f o r m h a d g ro w n h u t l i t t l e

ste m s w e re w oody f o r t h e i r e n t i r e

le n g th .

and

S te m s o f t h e

h e r b a c e o u s f o r m d i e d b a c k a l m o s t t o t h e g r o u n d a n d t h e new g ro w th was h e a v i e r b u t s h o r t e r a n d e n t i r e l y w oody. To t h e S u f f r u t e s c e n t S a l v i a a p i a n a S t a t i o n o f t h e o t h e r tw o f o r m s w e re t r a n s p l a n t e d . a f te r tra n s p la n tin g ,

six

p la n ts

W ith in n in e ty days

t h e w o o d y f o r m h a d g r o w n a b o u t 20 c m . ,

h a l f o f w h i c h new g r o w t h w a s w o o d y .

S te m s o f t h e h e r b a c e o u s

f o r m d i e d b a c k a b o u t h a l f w ay a n d t h e n ew g r o w t h f r o m t h a t p o i n t was p a r t l y w oody. To t h e H e r b a c e o u s S a l v i a a p i a n a S t a t i o n s i x p l a n t s t h e woody fo rm a n d s i x p la n te d .

of th e

of

s u f f r u t e s c e n t fo rm w e re t r a n s ­

W ith in n i n e t y d a y s a f t e r t r a n s p l a n t i n g ,

t h e woody

f o r m h a d g r o w n a b o u t 3 0 cm. , o n e t h i r d o f w h i c h n ew g r o w t h was woody.

W i t h i n t h e sa m e l e n g t h o f t i m e t h e s t e m s o f t h e

h e rb a c e o u s fo rm d ie d b ack c o m p le te ly t o th e

g ro u n d , and th e

new g r o w t h w a s n e a r l y a l l w o o d y , p r o b a b l y d u e t o t h e v e r y d i f ­ fe re n t c o n d itio n s. V i r t u a l l y t h e sa m e r e s u l t s w e r e r e c o r d e d f o r t h e g ro w th -fo rm s o f S a l v i a l e u c o p h y l l a , a n d th e

th re e

tw o g r o w th - f o r m s

o f E n o e l i a c a l i f o r n i c a w e r e r e v e r s i b l e w h en m o v e d f r o m o n e

49 c r i t i c a l a re a to a n o th e r. III.

CRITICAL FACTORS IN OTHER ARIAS

A l t h o u g h m an y o t h e r s p e c i e s p r e s e n t e d t h e fo rm s e ls e w h e r e , th e S.

a p ia n a . S.

th re e

g ro w th -

p r e s e n t i n v e s t i g a t i o n d e a ls o n ly w ith

le u c o p h y lla , and E.

c a lif o r n ic a , th e

la st

of

w h i c h w a s s t u d i e d i n b u t tw o f o r m s . §•

a p ia n a .

S. a p ia n a w as n o t o b s e rv e d in l i t t o r a l ,

dune, o r sea c l i f f l o c a l i t i e s ^ c h a ra c te ris tic w here i t

but i t

com ponent o f t h e

i s an im p o rta n t and

c o a s t a l sa g e f o r m a t i o n ,” (19)

o c c u rs m ost f r e q u e n tly a s a s u f f r u t e s c e n t .

I n M issio n

C anyon, S a n ta B a r b a r a , th e p l a n t o f t e n gro w s a s an h e rb a c e o u s p e re n n ia l, a s i t

does in canyons fa c in g th e

O c e a n s id e i n S an B ie g o C o u n ty . ra l,

S . a p ia n a was c o n s i d e r e d

ocean n e a r

As a c o m p o n e n t o f t h e c h a p a r ­ in th e

re g io n s n e a r S a n ta

B a r b a r a , S a n ta M o n ic a, S an G a b r i e l , S a n ta A na, and S an B ie g o . In th e se fiv e

s e c t i o n s , w here c o n d itio n s o f c o n tin u o u s u p ­

d rau g h t e x is t, le ss

th e

sev eral te s ts

c o m p le te w o o d in e ss o f th e s te m s .

so u th e rn ,

show ed m ore o r

When t h e e x p o s u r e i s

e a s t e r n , o r w e s t e r n , an d w h ere th e

posed to o r a r e i n s e r io u s

c o m p e titio n w ith

sp e c ie s is a s u f f r u te s c e n t. is

fo r lig n in

p la n ts are

ex­

o th e r p la n ts ,

th e

When p r o t e c t e d a n d w h e r e t h e r e

am p le s o i l m o i s t u r e , S . a p i a n a o c c u r r e d a s a n h e r b a c e o u s

p e re n n ia l,

slig itly

w oody a t

th e

w o o d la n d a n d s a v a n n a r e g i o n s t h e fru te sc e n t.

In p ro te c te d

base o r u n d erg ro u n d .

In th e

s p e c ie s a r e a lw a y s a s u f ­

v a lle y s i t

o fte n

o c c u r r e d a s m ore

50 h e rb a c e o u s th a n s u f f r u t e s c e n t , w h ile on g r a s s y p l a i n s i t su ffru te s c e n t.

¥ J h e re t h e

yon v e g e ta tio n i t .w a s

s p e c i e s grew i n

a lw a y s woody b e lo w .

r i p a r i a n and c an ­

h e rb a c e o u s u n le s s th e

o t h e r p e r e n n i a l s w as g r e a t ,

was

c o m p e titio n w ith

in w h ic h c a s e t h e p l a n t s w ere

U nder sh a d e and on n o r t h e r n e x p o s u re s

w i t h a m p le s o i l m o i s t u r e , S . a p i a n a w as h e r b a c e o u s . .

T h i s was

o b s e rv e d in u n m is ta k a b le fo rm i n S p sp e , G a v io ta , and T e h a c h a p i C anyons.

In th e

s e m ia rid r e g io n s o f A n te lo p e V a lle y ,

H em et, M in t C an y o n , C a jo n P a s s , J u l i a n , J a c u m b a , a n d a l o n g t h e strip

o f W est C o a s t F oggy D e s e r t i n L ow er C a l i f o r n i a t h e

c i e s w as s u f f r u t e s c e n t i n s i t u a t i o n

of little

spe­

w in d b u t i n

o t h e r s i t u a t i o n s w i t h w in d t h e t i s s u e s w ere w e l l l i g n i f i e d th e

e n t i r e l e n g t h o f th e

ste m s.

On t h e

ta lu s

M o ja v e D e s e r t S . a p i a n a w a s a s u f f r u t e s c e n t . v e g e ta tio n i t

o c c u rre d in th e

in v e s tig a tio n

it

?fh ere i t

s lo p e s o f th e In rip a ria n

sam e g r o w t h - f o r m .

was n e v e r s e e n a s a n h e rb

D u rin g t h i s

on th e d e s e r t .

a p p e a re d on t h e d e s e r t s i d e o f t h e San B e r n a r d in o ,

T e h a c h a p i, S an J a c i n t o , a n d B lac k M o u n ta in s , S . a p ia n a w as a l w a y s c o m p l e t e l y w o o d y , o w in g t o w i n d s a n d l a c k o f m o i s t u r e . O d d ly en o u g h m o st o f t h e s e l o c a t i o n s h a v e n o r t h e r n t o n o r t h ­ e a s te rn ex p o su res.

On t h e C o l o r a d o D e s e r t S .

a p ia n a o c c u rs

a s a s u f f r u te s c e n t a lo n g th e n o rth e rn s id e s o f t a l u s

slo p e s

i n t h e r e g i o n o f J o s h u a N a t i o n a l M o n u m e n t, t h e A n z a d i s t r i c t , and th ro u g h o u t th e S u p e r s t i t i o n M o u n ta in s. c r e s t s a n d w in d y r i d g e s lig n ifie d

tis s u e s .

It

it

A lo n g m o u n ta in

w as t o t a l l y w o o d y i n a l l t h e u s u a l l y

l i k e w i s e d i d n o t o c c u r on t h e C o lo ra d o

51 D e se rt a s an h e rb .

I n t h e m o n ta n e b e l t s

o f t h e B an R a f a e l ,

S an G a b r i e l , San B e r n a r d in o , T e h a c h a p i, P a lo m a r and San J a c i n t o M o u n ta in s S . a p i a n a w as fo u n d i n a l l t h r e e fo rm s.

I f exposed to

a woody p l a n t ;

if

c o n tin u o u s u p -d ra u g h t,

ap p eared a s

p r o t e c t e d a n d w i t h am p le s o i l m o i s t u r e ,

was s e e n i n t h e h e r b a c e o u s fo rm ; p la n ts,

it

g ro w th -

if

in c o m p e titio n w ith o th e r

o r on ro c k y w e s te r n , s o u th e r n ,

i t w as a s u f f r u t e s c e n t .

it

or e a ste rn ex p o su res,

The m o n ta n e b e l t s o f t h e r e g i o n s

m e n tio n e d w e re i n a d e q u a t e l y s t u d i e d , a n d t h e m any t r a n s i t i o n a l f o m n s s u g g e s t e d t o o m an y m i c r o c l i m a t i c r e s p o n s e s t o

be in ­

c lu d e d i n th e p r e s e n t i n v e s t i g a t i o n .

S.

le u c o p h y lla .

S. le u c o p h y ll a w as n o t se e n i n th e

d e s e r t r e g i o n s o r th e m o n ta n e b e l t s o f v e g e t a t i o n . com ponent o f th e

As a

l i t t o r a l a n d san d dune a s s o c i a t i o n s i t

c u rred a s is o la te d p la n ts ,

ra re ly

in la rg e

oc­

s t a n d s , f r o m M oro

B ay, S a n L u is O b isp o C o u n ty , t o C a p i s t r a n o B e a c h , O ra n g e C o u n ty .

In th is

c o m m u n ity i t w a s a l w a y s a s u f f r u t e s c e n t ,

s l i g h t l y woody w h ere i t

o c c u r r e d on s e a c l i f f s ,

and th is

con­

d i t i o n w as p a r t i c u l a r l y

o b v io u s a t P o i n t C o n c e p tio n , H uenem e,

S a n ta C ruz I s l a n d , S a n ta B osa I s l a n d , an d P a c i f i c P a l i s a d e s . S.

l e u c o p h y l l a was fo u n d i n a l l t h r e e g ro w th -fo rm s i n

c o a s ta l sage b ru sh a s s o c ia tio n s . w e ste rn , tu re

so u th e rn , and e a s te r n

A lo n g t h e

th e

c o a s t and on a l l

e x p o s u r e s , w h e re am p le m o is ­

e x i s t e d w i t h o u t w in d and c o m p e t i t i o n ,

t h e y w ere h e r b a ­

c eo u s; u n d e r c o n d itio n s o f c o n tin u o u s d ra u g h t,

even w ith

52 a m p le m o i s t u r e , t h e y w e r e w oody.

The sa m e c o n d i t i o n s w e r e i n ­

d i c e s of th e s e g ro w th p h a s e s i n a l l

o t h e r a r e a s w here th e

s p e c i e s was f o u n d , e x c e p t i n t h e s e m i a r i d r e g i o n s o f B ouquet C anyon, San F r a n c i s q u i t o ( 1 9 ) , w h ere th e

p l a n t s grow a s s u f f r u t e s e e n t s .

32. c a l i f o r n i c a . c lim a tic

C anyon, an d P i c o C anyon n e a r N e w h a ll

12. c a l i f o r n i c a

r e g i o n o f S o u t h e r n C a l i f o r n i a a n d n o r t h e r n Low er

C a lifo rn ia .

As a d u n e a n d l i t t o r a l s p e c i e s t h e p l a n t w a s

d w a r f e d , m ore w oo d y , an d l e s s th e se a c l i f f s . 5*

c a lifo rn ic a

woody p l a n t . N ear th e

o c c u rs in e v e ry m a jo r

f r e q u e n t , t h a n when i t

As a com p on en t o f t h e

g r e w on

c o a s ta l sage b ru sh

o c c u rre d b o th a s a s u f f r u t e s c e n t and a s a I t w as n o t n o te d i n t h e

to p s of

c liffs ,

a t th e

h e rb a c e o u s g ro w th -fo rm .

c re sts of h ills ,

h e a d s o f c an y o n s i t w as a n a l l w oody s h r u b i f

and a t

th e

th e c o n d itio n s

o f s c a n t m o istu re and u p -d ra u g h t p r e v a ile d .

U nder a l l o th e r

c o n d itio n s th e s p e c ie s w as a s u f f r u t e s c e n t .

I t was fo u n d u p

to

3 ,0 0 0 f e e t ab o v e se a l e v e l in th e P a lo m a r, S a n ta I n e z ,

San G a b r i e l , and San J a c i n t o M o u n ta in s , a lth o u g h n o t fo u n d i n t h e m o n ta n e b e l t s

i n v e s t i g a t e d on an y o f t h e s e r a n g e s .

h ig h a l t i t u d e s

a p p e a r s m uch t h e

it

l i t t o r a l c o m m u n itie s .

sam e a s i n t h e d u n e a n d

U nder c h a p p a rra l c o n d itio n s th e p la n t

was u s u a l l y s u f f r u t e s c e n t .

W h ile a c t i n g a s a p i o n e e r a lo n g

r id g e s and c r e s t s a f t e r f i r e s w h en l e s s

In

t h e s p e c i e s show ed l i g n i f i c a t i o n

t h a n a y e a r o ld and r e m a in e d m ore o r l e s s

w oody u n t i l t h e c h a p a r r a l becam e e s t a b l i s h e d a g a i n .

c o m p le te ly B urned

53 a r e a s w ere

c h e c k e d , p a r t i c u l a r l y a f t e r th e San M arcos f i r e

19 3 9 , and m in o r f i r e s

of

s in c e t h a t d a t e , i n V e n tu ra , Los A n g e le s,

a n d S an D ie g o C o u n t ie s .

The s p e c i e s w as s e e n i n t h e

l a n d s , w o o d la n d s, s a v a n n a ,

rip a ria n ,

and canyon v e g e t a t i o n a s

a s u f f r u t e s c e n t o n ly and a s a woody p e r e n n i a l i n A lo n g t h e c o a s t o f Low er C a l i f o r n i a T a b le IV , page 5 9 , and t h e i r r e a c t i o n to th e

g rass­

a rid

re g io n s.

i t w as a s u f f r u t e s c e n t .

show s t h e d i f f e r e n t

g ro w th -fo rm s

v a rio u s e n v iro n m e n ts.

M any d e s e r t

fo rm s a c q u ir e a p e c u l i a r e x p r e s s i o n o f s u f f r u t e s c e n c e : th e h e rb a c e o u s ste m t i p s

d i e d b a c k i n som e d u r i n g t h e

c ritic a l

p e r i o d , m an y o f t h e m a s s u m i n g t h e f u n c t i o n o f s p i n e s . IV .

RESULTS OF THE ANATOMICAL WORK

T he woody g r c w th - f o r m o f S a l v i a a p i a n a h a d a g e n e r a l l y l i g n i f i e d x y le m a n d x y lem p a re n c h y m a , l i g n i f i e d p h lo e m , lig n ifie d ste m s,

p ith

c e l ls , h e a v ily

s ta rc h sto ra g e

c u b i'n ilz e d e p i d e r m i s , h o llo w

in th e r a y c e l l s ,

lig n ifie d

sc le re n c h y m a ,

v e ry d en se tric h o m e s, no i n t e r c e l l u l a r s p a c e s , p i t t e d and a c o n tin u o u s s t e l e .

som e

v e sse ls,

T he s u f f r u t e s c e n t g r o w t h - f o r m o f t h e

sam e s p e c i e s s h o w e d a p a r t i a l l y o r s c a t t e r e d l i g n i f i e d x y l e m a n d o t h e r s p e c im e n s h a d l i g n i f i e d x y lem p a re n c h y m a 9 a few h ad lig n ifie d

p ith c e lls ,

some o t h e r s h a d h e a v i l y c u t i n i z e d e p i ­

d e r m i s , m an y h a d h o l l o w s t e m s , s m a l l t r a c e s o f s t o r e d s t a r c h , lig n ifie d

sc le re n c h y m a ,

in te rc e llu la r

d en se tric h o m e s , w h ile

spaces, p itte d

som e o t h e r s h a d

v e s s e ls , and a c o n tin u o u s s t e l e .

54 T he h e r b a c e o u s g r o w t h - f o r m o f t h e lig n ific a tio n

sam e s p e c i e s sh o w e d n o

o th e r th a n tr a c e s o f th e

s u b s ta n c e n e a r th e b ase

o f t h e p l a n t i n b o t h t h e p h lo em an d x y le m . lig n ific a tio n

i n sc le re n c h y m a f i b e r s ,

e p i d e r m i s , no h o l lo w s te m , no s t a r c h v e sse ls, ste le .

in te rc e llu la r

T h ere w as s l i g h t

no h e a v i l y c u t i n i z e d sto ra g e ,

spaces in c o rte x ,

som e p i t t e d

and a c o n tin u o u s

T h i s i n f o r m a t i o n w a s c o m p i l e d i n T a b l e IV , p a g e 5 9 . T h e w o o d y g r o w t h - f o r m o f S . l e u c o p h y l l a sh o w e d a g e n ­

e ra lly lig n ifie d f i e d p h lo e m ,

x y le m a n d x y lem p a re n c h y m a ,

i r r e g u l a r e p id e rm is,

v e ry dense tric h o m e s , no i n t e r c e l l u l a r ste le .

lig n ifie d

p itte d v e sse ls,

lig n ifie d

p h lo em , p a r t i a l l y few l i g n i f i e d

v e sse ls,

a few h o llo w

th a n s l i g h t tr a c e s

sam e s p e c i e s

ste m s,

som e i n t e r c e l l u l a r

a

spaces,

The h e r b a c e o u s g r o w th in any o rgan o th e r

i n t h e x y le m a t t h e b a s e o r b e lo w t h e g ro u n d , no a b n o rm a lly c u t i n i z e d e p id e r m is ,

n o h o l l o w s t e m s , m any i n t e r c e l l u l a r

c o rte x and p i t h ,

lig n ifie d

som e i r r e g u l a r e p i d e r m i s ,

show ed no l i g n i f i c a t i o n

no i r r e g u l a r e p i d e r m i s , tric h o m e s,

sc le re n c h y m a ,

some p a r t i a l l y

and a c o n tin u o u s s t e l e .

sp e c ie s

e p id e rm is,

x y lem an d p h lo em p a re n c h y m a ,

sc le re n c h y m a f i b e r s ,

dense tric h o m e s ,

fo rm o f t h i s

lig n ifie d

x y lem ,

lig n i­

and a c o n tin u o u s

T he s u f f r u t e s c e n t g r o w t h - f o r m o f t h e

sh o w e d p a r t i a l l y

p itte d

h e a v ily c u tin iz e d

h o llo w ste m s ,

spaces,

p a rtia lly

som e p i t t e d v e s s e l s ,

few

sp aces in th e

and a c o n tin u o u s s t e l e .

S e e T a b l e V, p a g e 6 0 . The w oody g r o w th - f o r m o f E .

c a l if o r n i c a had a g e n e r a lly

55 lig n ifie d

x y le m , x y lem p a r e n c h y m a , p h lo e m and s c l e r e n c h y m a .

They had a ls o

i r r e g u l a r and h e a v ily c u tin iz e d e p id e rm is ,

h o l l o w s t e m s , m an y t r i c h o m e s , n o i n t e r c e l l u l a r - a d isse c te d

ste le .

See T a b le V I,

T a b le s V II to X I I I , th e

page. 61.

p a g e s 62 t o 6 6 ,

in c lu siv e ,

s p e c i f i c new g r o w th c h a n g e s n o t e d i n t h e

sp e c im e n s.

B rie fly ,

sp a c e s, and

show

tra n s p la n te d

th e c r i t i c a l c o n d itio n s a t th e

sta tio n s

w e re m et b y m o r p h o lo g i c a l c h a n g e s o r e c o n o m ie s w i t h i n t h e p la n t tis s u e s .

I n a r e a s w h ere w o o d in e ss o c c u r r e d i n a l l

in d iv id u a ls o f th e s p e c ie s ,

th e t r a n s p l a n t e d sp e cim e n s soon

a p p ro p ria te d th e

c h a ra c te ristic .

d iv id u a ls o f th e

s p e c i e s w e re m ore o r l e s s h e r b a c e o u s ,

tra n s p la n te d in th e

ste m s.

th e

I n a r e a s w here a l l t h e

sp e c im e n s f a i l e d t o l i g n i f y

In ­

th e

c e r t a i n new g ro w th

56

TABLE I CERTAIN ENVIRONMENTAL CONDITIONS AT THE STATIONS A ltitu d e (fe e t)

E x p o su re

S o il

2 ,8 0 0

S o u th e rn

S an d ­ S te e p sto n e

800

S o u th e a st

Adobe

G e n tle

S lig h t

50 0

W e ste rn

Sand C lay Loam

G e n tle

W oody S . le u c o .

C o n tin u e d u p -d rau g h t

500

O pen

Sand

L evel to g e n tle

S u ff. S. le u c o .

V a ria b le

200

S o u th e a st

C lay Loam

G e n tle to le v e l

H erb . S . le u c o .

S lig h t

600

S o u th e a st

C lay S h a le Loam

g e n tle

Vfoody S . c a lif.

C o n tin u e d u p -d rau g h t

300

S o u th w e st

S h a le

S te e p

S u ff. c a lif.

V a ria b le

S ta tio n

A ir m o v em en ts

W oody S . a p ia n a

C o n tin u e d u p -d rau g h t

S u ff. S. a p ia n a

V a ria b le

H erb . S . a p ia n a

20-30 N o rth w e st

D e c liv ity

L im e ­ S te e p sto n e S h a le

57

TABLE I I PERCENTAGES OF S O IL MOISTURE AT THE STATIONS IN OCTOBER W et w e i g h t (k ilo s )

S ta tio n

W oody S .

D ry w e ig h t (k ilo s )

Per cent of m o is tu re ^

a p ia n a

5

4 .9 0 1

2 .0 2

S u f f * _S. a p i a n a

5

4 ,8 5 6

2 .9

H erb. S . a p ia n a

5

W oody S .

le u c o •

5

4 .8 5

2 .9

S u ff. S.

le u c o .

5

4 .8 1

3 .8

H erb . S. le u c o .

5

W oody. W. c a l i f .

5

4 .8 5

2 .9

5

4 .9 1

1 .8

S u ff. E.

c a lif.

4-8

4 —8

a F i g u r e d on t h e d r y w e ig h t b a s i s .

58

TABLE I I I DISTRIBUTION OF EXPERIMENTAL PLANTS P la n ts in tro d u c e d

S ta tio n

N um ber tra n s p la n te d

S u rv iv a ls

W oody S . a p i a n a

S u f f . S. a p ia n a H erb . S . a p ia n a

6 6

2 1

S u f f . S . a p ia n a

W oody S . a p i a n a H e r b . _S. a p i a n a

6 6

3 2

H erb . S . a p ia n a

Woody S . a p i a n a S u f f . S. a p i a n a

6 6

1 6

W oody S . l e u c o .

S u f f . S . le u c o . H erb. S . le u c o .

6 6

3 2

S u ff. S. le u c o .

Woody S . l e u c o . H e r b . £>. l e u c o .

6 6

2 1

H erb . S . le u c o .

Woody S . l e u c o • S u f f . S. le u c o .

6 6

2 1

W oody E . . c a l i f .

S u f f . JU c a l i f .

6

6

c a lif.

6

5

S u f f . JL

c a lif.

W oody E .

59 TABLE IV MORPHOLOGICAL COMPARISONS OF THE GROWTH-FORMS OF S . APIANA W oody S . a p ia n a

L i g n i f i e d x y lem P a rtia lly x y le m

lig n ifie d

P a rtia lly p h lo em

lig n ifie d

S u ff. S . a p ia n a

H erb . S . a p ia n a

N one

A lla A ll

None

A ll

A ll

None

L i g n i f i e d x y lem p aren ch y m a

A ll

8o m ea

None

L ig n ifie d p ith c e lls

A ll

Some

N on e

H e a v ily c u tin iz e d e p id e rm is

A ll

Some

N one

H o llo w ste m

A ll

Some

N on e

S ta rc h sto ra g e

A ll

Some

None

L ig n ifie d sc le re n c h y m a

A ll

A ll

Some

D ense

S p arse

T ric h o m e s

V ery d e n se

In te rc e llu la r sp aces in c o rte x

N on e

Some

A ll

P itte d v e sse ls

A ll

A ll

Some

S te le

A ll

A ll

A ll

c o n tin u o u s

a Found, i n a l l p l a n t s

stu d ie d ,

NOTE: Some r e f e r s t o q u a n t i t y , w h i c h d i f f e r e d w i t h in d iv id u a ls . N on e i n d i c a t e s n o t f o u n d .

60

TABLE V MORPHOLOGICAL COMPARISONS OF. THE. GROWTH-FORMS OF S . LEUCOPHYLLA Woody S . le u c o . L i g n i f i e d x y lem P a rtia lly x y lem

A ll

S.

S u ff. le u c o .

H erb . S . le u c o .

Some

None

A ll

None

lig n ifie d

I r r e g u l a r e p id e rm is

A ll

Some

N o ne

H e a v ily c u tin iz e d e p id e rm is

A ll

N o ne

N o ne

L ig h tly c u tin iz e d e p id e rm is

N on e

Some

A ll

V ery d e n se

D ense

S p arse

T ric h o m e s H o llo w ste m s

A ll

Some

N one

L i g n i f i e d x y lem p aren ch y m a

A ll

Some

N one

L ig n ifie d

A ll

Few

Some

A ll

A ll

Some

A ll

A ll

Some

A ll

A ll

A ll

sc lere n ch y m a

In te rc e llu la r P itte d S te le

spaces

v e sse ls c o n tin u o u s

61 TABLE V I MORPHOLOGICAL COMPARISONS OF THE GROWTH-FORMS OF E . CALIFORNICA

E.

Woody c a lif.

S u ff. E. c a l if .

L i g n i f i e d x y lem

A ll

Some

P a rtia lly x y lem

lig n ifie d None

A ll

P a rtia lly p h lo em

lig n ifie d Some

None

Irre g u la r

e p id e rm is

A ll

Some

H e a v ily c u tin iz e d e p id e rm is

A ll

None

S lig h tly c u tin iz e d e p id e rm is

Some

A ll

L i g n i f i e d x y lem p aren ch y m a

A ll

N one

L ig n ifie d

Some

None

Some

None

N one

A ll

A ll

A ll

sc le re n c h y m a

H o llo w s te m s In te rc e llu la r

spaces

S te le d isse c te d

62

TABLE V I I NEW-GROWTH CHANGES IN PLANTS TAKEN FROM WOODY S . APIANA STATION To S u f f . S . a p i a n a sta tio n

To H e r h . S . a p i a n a sta tio n

L i g n i f i e d x y lem

S lig h t

N one

L i g n i f i e d p h lo em

Some

None

S ta rc h

Some

N one

None

None

in ra y c e l l s

L ig n ifie d p ith

L arg er c e lls

L arg er c e l ls

H eavy

S lig h t

T ric h o m e s

S h o rte r

S p arse

L i g n i f i e d x y le m p aren ch y m a

Some

None

L ig n ifie d sc le re n c h y m a

Some

None

Some

M any

H o llo w ste m s

Some

None

P itte d

Some

None

C o rte x C u tin iz e d

e p id e rm is

In te rc e llu la r

v e sse ls

spaces

63 TABLE V I I I NEW-GROWTH CHANGES IN PLANTS TAKEN FROM THE SUFFRUTESCENT S . APIANA STATION To Woody S . a p i a n a s ta tio n

To H e r b . S . a p ia n * sta tio n

L i g n i f i e d x y lem

A ll

None

L i g n i f i e d p h lo em

Some

None

S ta rc h

A ll

None

Some

N on e

in ra y c e l l s

L ig n ifie d p ith

S m a lle r c e l l s

C o rte x

L arg er c e l l s

C u tin iz e d e p id e rm is

H eavy

S lig h t

T ric h o m e s

L onger

S p arse

L i g n i f i e d x y lem p aren ch y m a

A ll

N one

L ig n ifie d

A ll

None

L i g n i f i e d p h lo em p aren ch y m a

Some

None

In te rc e llu la r

N one

Many

H o llo w s te m s

A ll

None

P itte d v e sse ls

Some

None

sc le re n c h y m a

spaces

64

TABLE IX NEW-GR0 WTH CHANGES IN PLANTS TAKEN FROM THE HERBACEOUS S . APIANA STATION To Woody S ^ a p l a n a sta tio n

To S u f f . S . a p i a n a sta tio n

L i g n i f i e d x y lem

A ll

A ll

L i g n i f i e d p h lo em

Some

Some

S ta rc h

None

Some

Some

None

in ra y c e lls

L ig n ifie d p ith C o rte x

S m a lle r c e l l s

S m a lle r c e l l s

C u tin iz e d e p id e rm is

H eavy

S lig h t

T ric h o m e s

Longer

L onger

L i g n i f i e d x y le m p aren ch y m a

Some

Some

L i g n i f i e d phloem p a re n c h y m a

Some

Some

H o llo w s te m s

A ll

Some

P itte d

A ll

Some

v e sse ls

65

TABLE X NEW-GROWTH CHANGES IN PLANTS TAKEN FROM THE WOODY- S . . LEUC OPHYLLA STATION To S u f f . S . l e u c o . sta tio n

To H e r b . S . ' ' l e u c o sta tio n

L i g n i f i e d x y lem

Some

N one

L i g n i f i e d p h lo em

Some

None

S ta rc h in ra y c e lls

R are

N on e

L ig n ifie d p ith

None

N o ne

L arg er c e lls

L arg er c e l l s

C u tin iz e d e p id e rm is

H eavy

S lig h t

T ric h o m e s

S h o rte r

S parse

L i g n i f i e d x y lem p aren ch y m a

A ll

N on e

L i g n i f i e d p h lo em p aren ch y m a

Some

N one

L ig n ifie d

A ll

N one

I n te r c e llu la r spaces

Some

Many

H o llo w s te m s

A ll

N one

P itte d v e sse ls

Some

N on e

C o rte x

sc le re n c h y m a

66

TABLE XI NEW-GROWTH IN PLANTS TAKEN FROM THE SUFFRUTESCENT S . LEUCOPHYLLA STATION

To Woody le u c o ♦ sta tio n

To H e r b . S . l e u c o sta tT o n

L i g n i f i e d x y lem

A ll

N o ne

L i g n i f i e d p h lo em

Some

None

S ta rc h in ra y c e lls

A ll

None

L ig n ifie d p ith

Some

None

S m a lle r c e l l s

C o rte x

L arg er c e lls

O u tin iz e d e p id e rm is

H eavy

S lig h t

T rie h o m e s

Longer

S p arse

L i g n i f i e d x y lem paren ch y m a

A ll

None

L i g n i f i e d p h lo em paren ch y m a

Some

None

L ig n ifie d

A ll

None

N o ne

M any

H o llo w s te m s

A ll

None

P itte d v e sse ls

Some

None

sc lere n eh y m a

In te rc e llu la r

spaces

67

TABLE XII NEW-GROWTH CHANGES IN PLANTS TAKEN FROM THE HERBACEOUS S ^ LEUCOPHYLLA STATION To Woody S . l e u c o . s ta tio n

To Woody Sj_ l e u c o « sta tio n

L i g n i f i e d x y le m

A ll

Some

L i g n i f i e d p h lo em

Some

N one

S ta rc h in ra y c e lls

Some

Some

L ig n ifie d p ith

Some

N one

S m a lle r c e l l s

C o rte x

S m a lle r c e l l s

C u tin iz e d e p id e rm is

H eavy

Some

T ric h o m e s

L onger

Longer

L i g n i f i e d x y le m p aren ch y m a

Some

N on e

L i g n i f i e d p h lo em p aren ch y m a

Some

None

In te rc e llu la r

N one

Some

H o llo w s te m s

Some

None

P itte d

Some

Some

v e sse ls

spaces

68

TABLE XIII NEW-GROWTH CHANGES IN PLANTS TAKEN FROM THE WOODY E . CALIFORNICA STATION TO THE SUFFRUTESCENT E . CALIFO&NICA STATION AND VICE VERSA F ro m t h e Woody E. c a lif , s ta tio n to S u ff. E. c a lif , sta tio n

F ro m t h e S u f f . E ^ c a lif , sta tio n t o Woody E . c a l i f , sta tio n

L i g n i f i e d x y lem

Some

A ll

L i g n i f i e d p h lo em

Some

A ll

S ta rc h in ra y c e lls

Some

N one

L ig n ifie d p ith

N one

Some

L arg er c e l ls

C o rte x

S m a lle r c e l l s

C u tin iz e d e p id e rm is

Some

Some

T rie h o m e s

M o re

Few er

L i g n i f i e d x y lem p aren ch y m a

None

Some

L i g n i f i e d p h lo em p aren ch y m a

None

Some

None

Some

Some

N one

H o llo w s te m s

None

A ll

P itte d

Some

A ll

L ig n ifie d

sc le re n c h y m a

In te rc e llu la r

v e sse ls

spaces

69

TABLE XIV TYPES OF ENVIRONMENT AND THEIR EFFECTS UPON SUFFRUTESCENT GROWTH-FORHS

E n v iro n m e n t

E ffe ct

L i t t o r a l and dune

R e d u c tio n in

s ta tu re

Sea c l i f f

L i g n i f i c a t i o n o f a l l x y lem

C o a sta l sag eb ru sh S o u th e rn ex p o su re W este rn e x p o s u re N o rth e rn ex p o su re

S u ffru te sc e n t v a ria tio n S u ffru te sc e n t v a ria tio n H erbaceous to s u f f r u t e s c e n t

C h ap arral S o u th e rn ex p o su re W e ste rn e x p o s u re N o rth e rn ex p o su re U p -d ra u g h t 'c o n d it i o n

S u f f r u t e s c e n t t o woody G e n e r a l l y w oody G e n e ra lly h erb aceo u s A lw a y s w o o d y

W o o d la n d a n d s a v a n n a

H erb aceo u s to

G ra s s la n d and p l a i n

G e n e ra lly s u f f r u te s c e n t

R i p a r i a n and canyon

G e n e ra lly h erb aceo u s i f p r o t e c te d fro m d ra u g h t

S e m ia rid h i l l s i d e s

S u f f r u t e s c e n t t o w o o dy

M o ja v e D e s e r t

S u f f r u t e s c e n t t o w oody

C o lo ra d o D e s e r t

S u f f r u t e s c e n t t o woody

M o n tan e

Too v a r i a b l e t o c l a s s i f y w ith o u t f u r t h e r stu d y

su ffru te sc e n t

70

P late I

Woody f o r m o f S a l v i a a p l a g a .

T ran sv erse

se c tio n

71

Plate II Woody fora of Salvia apiana.

Longitudinal section.

72

P la te I I I

S a f f y u t e s e e n t fo rm o f S a l v i a

a p ia n a .

T ransverse

se c tio n .

73

^

X

X

r -

•V.° .* •*» : .*.*V.* .* /..* : .• .* . %//.*,*.• . > * / / * v .> % v /.*

^ — ( -ivy^____ ____ ___ - i

r" ■

r

i

- J L _____1

f n ^ r T I

k '- ''7

.*.» .* o . . „

i

□ ^p-p*f

r __,) L

i f

J L

T

______ 1

l

i

J L

Plate IV

Suffrutescent form of Salvia apiana.

Longitudinal section.

I 1

74

•Mft]

P la te V

Herbaceous form ef Salvia, apiana.

Transverse section..

F Ia te VI H erb aceo u s T o rn o f S a lv ia a p ia n a .

k o n ^ liu d in a l s e c tio n .

P l a t e VII Woody f o r m o f S a l v i a l e u c o p h y l l a .

T ran sv erse

se c tio n .

? la -te V III Wo o ily form o f S a lv ia leuco ph y l 1 a .

Lo-ngltsud in a l s a c tio n

78

P l a t e IX S u f f r u t e s c e n t fo rm o f S a lv ia l e u c o p h y l la ,

T ran sv erse

se c tio n .

79

S u f f r u t e s o e n i fo rm o f S a l v ia l e u c o p h y ll a .

L o n g itu d in a l s e c tio n .

F ia t© XI

H e rb a c e o u s fo rm o f S aX vla l e u c o p h y l l a .

T ran sv e® ?^ s « 9 f f o n .

81

Plate XII Herbaceous form of Salvia leucophylla.

Longitudinal section.

? la te X IIt Woody fo rm , o f E n c e X i a c a l t f o r n l c a .

T ran sv erse s e c tio n .

S3

Plate XIV Woody form o f Encelia californica* Longitudinal section

8 4-

F la te XV S u f f r u t e s c e n t fo rm e f E n c e li a

c a lifo ra lc ft.

T ran sv erse S e c tio n .

85

Suf\f;rut©scent form o f

Plate XVI Elncella callfornlca*

longitudinal section.

S7 d e ta ile d in

th e t e x t o f t h i s p a p e r.

D isc u ssio n o f r e s u l t s

h a s been b a s e d on d a t a c o m p ile d i n t h e form o f t a b l e s

in

C h a p t e r V. II. 1.

CONCUJSIONS

C o n tin u e d u p - d r a u g h t o f a i r

in flu e n c e d th e l i g n i f i -

c a t i o n o f p h lo em , x y lem , p aren ch y m a, an d s c le re n c h y m a e l e ­ m e n ts,

r e s u ltin g in th e

o f th e

th re e

a s s u m p tio n o f t h e w oody g ro w th -fo rm

s p e c ie s under stu d y .

2. V a r ia b le w in d s i n f lu e n c e d th e l i g n i f i c a t i o n ta in

of cer­

c e l l s w i t h i n t h e p h lo e m , x y le m , p a re n c h y m a , an d s c l e r e n ­

chym a e l e m e n t s , r e s u l t i n g c e n t g ro w th -fo rm o f th e

in th e a ssu m p tio n o f th e s u f f r u t e s ­

th re e

sp e c ie s.

3* A s o i l m o i s t u r e o f f r o m U t o te c tio n

from w in d ,

8 per c e n t,

d rau g h t c u rre n ts o f a i r ,

g ro w in g c o n d i t i o n s ,

in flu e n c e d th e

p lu s p ro ­

and o v ercro w d ed

a ssu m p tio n o f th e h e r b a ­

c eo u s g ro w th -fo rm by S a lv ia a p i a n a , J e p .

and S a lv ia

le u c o p h y lla , G reen e. 4 . The in d e x o f a i r seem s d i r e c t l y r e f l e c t e d

d rau g h t i n te n s i t y and d u ra tio n

i n th e am ount o f l i g n i f i e d t i s s u e

p ro d u ced in S a lv ia a p ia n a , J e p . , S a lv ia le u c o p h y lla , G re e n e ., and E n c e lia

c a l if o r n i c a . N u tt.

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(5 4 )

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,

d e se rts

C o m p a ra tiv e a n a to m y o f dune 5 4 :2 6 3 -3 0 5 .

and F . E . C le m e n ts, 1 9 2 9 . New Y o r k a n d L o n d o n .

sh ru b s.

il m o is tu re , ch ap arral T he A n g e le s.

P la n t

(53)

W ie g an d , K. M ., 1 9 1 0 . T he r e l a t i o n o f h a i r y a n d c u t i n i z e d c o v e r i n g s t o t r a n s p i r a t T o n . B’o t . G a z . 1+9:4 3 6 —4 4 4 •

(5 6 )

W i l s o n , C. L . , 1 9 2 2 . L i g n i f i c a t i o n o f m a tu r e p h lo em i n h e r b a c e o u s t y p e s . A m e r. «Tour* B o t* $ 12 3 9 -2 4 3 •