INFORMATION TO USERS
This reproduction was m ade from a copy o f a docu m en t sent to us fo r microfilming.
While the m ost advanced technology lias been used to photograph and reproduce
this docum ent, the quality o f the reproduction is heavily dependent upon the
quality o f the material subm itted.
The following explanation o f techniques is provided to help clarify markings or
notations which m ay appear on this reproduction.
1 .T h e sign or “ targ et” fo r pages apparently tacking from the docum ent
photographed is “ Missing Page(s)” . If it was possible to obtain the missing
nage(s) o r section, they are spliced into the film along w ith adjacent pages. This
may have necessitated cu tting through an image and duplicating adjacent pages
to assure com plete continuity.
2. When an image on the film is obliterated w ith a round black m ark, it is an
indication o f either blurred copy because o f m ovem ent during exposure,
duplicate copy, or copyrighted materials th a t should n o t have been filmed. F o r
blurred pages, a good image o f the page can be found in the adjacent frame. If
copyrighted m aterials were deleted, a target n o te will appear listing the pages in
the adjacent frame.
3. When a m ap, drawing or ch art, etc., is p art o f the m aterial being photographed,
a definite m ethod o f “ sectioning” th e m aterial has been followed. It is
custom ary to begin filming at th e u pper left hand com er o f a large sheet and to
continue from left to right in equal sections with small overlaps. If necessary,
sectioning is continued again—beginning below the first row and continuing on
until com plete.
4. F o r illustrations th a t cannot be satisfactorily reproduced by xerographic
means, photographic prints can be purchased at additional cost and inserted
into y o u r xerographic copy. These p rints are available upon request from the
D issertations C ustom er Services D epartm ent.
5. Some pages in any docum ent may have indistinct print. In all cases the best
available copy has been filmed.

University
M icnJilm s

International
300 N. Zoob Road
Ann Arbor, Ml 48106

8224391

Abrams, M arc David

EARLY PLANT SUCCESSION ON CLEARCUT AND BURNED JACK PINE
SITES IN NORTHERN LOWER MICHIGAN

Ph.D. 1982

Michigan State University

University
Microfilms
International

300 N. Zeeb Road, Ann Arbor, MI 48106

PLEASE NOTE:

In all cases this material has been filmed in the best possible way from the available copy.
Problems encountered with this document have been Identified here with a check mark V

.

1.

Glossy photographs or p a g e s ______

2.

Colored illustrations, paper or print______

3.

Photographs with dark background

4.

Illustrations a re poor c o p y ______

5.

Pages with black marks, not original copy______

6.

Print shows through as there is text on both sides of p a g e ______

7.

Indistinct, broken or small print on several pages

8.

Print exceeds margin requirem ents______

9.

Tightly bound copy with print lost in spine______

10.

Computer printout pages with indistinct print_____

11.

Page(s)___________ lacking when material received, and not available from school or
author.

12.

Page(s)___________ seem to be missing in numbering only as text follows.

13.

Two pages num bered___________ . Text follows.

14.

Curling and wrinkled p a g e s ______

15.

^

Other______________________________________ „ _______ _____________________

University
Microfilms
international

EARLY PLANT SUCCESSION ON CLEARCUT AND
BURNED JACK PINE SITES IN
NORTHERN LOWER MICHIGAN
By
M a r c D a v i d Abrams

A DISSERTATION

S ubm itted to
M ich ig an S t a t e U n i v e r s i t y
in p a r t i a l f u lf illm e n t of th e req u irem en ts
f o r the degree o f

DOCTOR OF PHILOSOPHY

D epartm ent o f F o r e s t r y
1982

ABSTRACT
EARLY PLANT SUCCESSION ON CLEARCUT AND
BURNED JACK PINE SITES IN
NORTHERN LOWER MICHIGAN
By
M a r c D a v i d A b ra m s

E a r l y p l a n t s u c c e s s i o n was c h a r a c t e r i z e d on
unburned c l e a r c u t s
M ichigan.

an d b u r n e d s i t e s

i n n o r t h e r n low er

B urning r e s u l t e d i n th e e s ta b lis h m e n t o f

numerous s p e c i e s n o t fo u n d on u n b u rn ed a r e a s .

The

l o n g e v i t y o f m o st o f t h e s e s p e c i e s was s h o r t , r e s u l t i n g
i n a l a r g e d ro p i n s p e c i e s r i c h n e s s b e tw e e n one and
two y e a r s a f t e r

fire.

Unburned c l e a r c u t

s i t e s w ere

r a p i d l y c o n v e r t e d t o C a r e x meadows w i t h v e r y l o w s p e c i e s
d iv ersity .

Burned s i t e s

sh o w e d m u l t i p l e

su ccessio n al

p a t t e r n s w h ic h i n c l u d e d d o m in a tio n by s h r u b and e a r l y
s u c c e s s i o n a l h a rd w o o d s, c o n v e r s i o n t o C a r e x meadows,
and th e e s t a b l i s h m e n t o f j a c k p i n e .

The d o m i n a t i o n

o f C a re x p e n s y l v a n i c a on t h e s e s i t e s

in d icates

it

is

th at

an o p p o r t u n i s t i c s p e c i e s c a p a b le o f m o n o p o liz in g

resources

lib e rate d

f o l l o w i n g d i s t u r b a n c e s and

su p p re ssin g or excluding o th e r sp e c ie s.
P e r m a n e n t p l o t s w e r e e s t a b l i s h e d on a 3 - y e a r o ld j a c k p in e c l e a r c u t p r i o r t o b u r n in g from w h ich
v e g e ta tio n a l changes over th re e
r e c o r d e d and f i r e

grow ing s e a s o n s were

i n t e n s i t y was e v a l u a t e d .

Burned

Marc D a v id Abrams

b l o c k s , c o m p a r e d t o u n b u r n e d b l o c k s , sh o w e d g r e a t e r
cover o f g r a s s and t r e e

and s h r u b s p e c i e s ,

low er

c o v e r o f C a r e x , and a g r e a t e r number o f p e r e n n i a l h e rb
sp ecies.

F i r e i n t e n s i t y on th e b u rn e d b l o c k s v a r i e d .

H igher f i r e
sity ,

in te n sity resu lte d

low er t o t a l p l a n t c o v e r,

in low er s p e c ie s

d iv er­

increased s u rv iv a b ility

o f b l u e b e r r y compared t o C a r e x , and lo w e r f r e q u e n c y
of estab lish m en t of c e r ta in grass

sp ecies.

F e r t i l i z a t i o n e x p e r i m e n t s on m a t u r e a n d c l e a r c u t
ja c k pine s i t e s

sh o w e d t h a t b i o m a s s o f u n d e r s t o r y v e g e ­

ta tio n increased a f te r f e r t i l i z a t i o n ,

and t h a t C .

p e n s y l v a n i c a c o n s i s t e n t l y sh o w e d a n a b o v e a v e r a g e
response.
E xperim ents conducted w ith s u rfa c e
from d i f f e r e n t a g ed j a c k p i n e

sites

s o i l sam ples

in d icated th a t

t h e dom inance o f Geranium b i c k n e l l i i r e s t r i c t e d

to

1 - y e a r - o l d b u r n s was due t o t h e h e a t - s t i m u l a t e d g e r m i ­
n a tio n o f b u rie d seed.
C o m p a r i s o n s made b y t h e i n d e x o f s i m i l a r i t y
sh o w ed t h a t s u c c e s s i o n o n e a c h s i t e
T herefore,

is

(S)

h ig h ly unique.

the i n d i v i d u a l i s t i c n a tu re of each s i t e ,

r a t h e r t h a n age f o l l o w i n g d i s t u r b a n c e , becomes t h e
dom inant a s p e c t i n u n d e r s t a n d i n g th e s u c c e s s i o n a l
rela tio n sh ip s

i n th e s e com m unities.

X d e d i c a t e t h i s w o r k t o my p a r e n t s , who
p r o v i d e d c o n s t a n t e n c o u ra g e m e n t and lo v e
t h r o u g h o u t my g r a d u a t e s t u d i e s , a n d a l l
o t h e r f r i e n d s a n d r e l a t i v e s who k e p t me
g o in g when t h e g o in g g o t to u g h .

ACKNOWLEDGMENTS
X e x p r e s s d e e p a p p r e c i a t i o n t o D r . D o n a l d X.
D i c k m a n n , who w a s r e s p o n s i b l e f o r m uch o f my p e r s o n a l
a n d p r o f e s s i o n a l g r o w t h d u r i n g my g r a d u a t e
S incere

thanks i s

stu d ies.

a l s o e x te n d e d to Dr. D ouglas

S p r u g e l , who a l w a y s h a d t i m e a n d i n s i g h t f u l a n s w e r s
f o r my n u m e r o u s q u e s t i o n s ,
Dr.

J.

a n d D r . P e t e r Mur phy a n d

B. H a r t f o r s e r v i n g o n my g r a d u a t e c o m m i t t e e .
I t h a n k a l l my f r i e n d s

the

la s t fiv e years;

fo r t h e i r su p p o rt over

h o w e v e r , E r i c Menges d e s e r v e s

s p e c i a l m e n tio n f o r th e g u id a n c e and e n e r g y he p r o ­
v i d e d t o my w o r k , s t a r t i n g w i t h o u r u n d e r g r a d u a t e d a y s
a t S.U .N .Y .

Bingham ton.

I want to th a n k th e N o rth C e n tr a l F o r e s t E x p e r i­
m e n t S t a t i o n o f t h e U. S . F o r e s t S e r v i c e f o r t h e i r
f in a n c ia l support of th is re se a rc h ,
D epartm ent o f N a tu r a l R eso u rces
and f i e l d

for a d m in istrativ e

a s s is ta n c e they p ro v id ed .

G rie v e s , A lb e r t Sim ard,

and th e M ich ig an

Ron W i l s o n ,

G erald

and R ic h a r d B la n k from t h e s e

o r g a n iz a tio n s d e se rv e s p e c i a l m ention.
F in ally ,

X t h a n k my p a r e n t s

fo r sh arin g a l l

the

j o y s a n d h a r d s h i p s I e n c o u n t e r e d i n c o m p l e t i n g my P h . D .

TABLE OF CONTENTS
L IST OF TABLES........................................................................................ v i
L IST OF FIGURES.....................................................................................

xi

LIST OF

xv

PLATES........................................................................................

CHAPTER
I.
INTRODUCTION...........................................................................

1

I n t r o d u c t i o n .....................................................................

1

DESCRIPTION OF STUDY AREA AND S I T E S .................

4

G e n e r a l D e s c r i p t i o n o f S t u d y A r e a ................
D e s c r ip tio n o f S i t e s In c lu d e d f o r Study.

4
8

EARLY REVEGETATION OF BURNEDAND UNBURNED
JACK PINE CLEARCUTS IN NORTHERN LOWER
MICHIGAN....................................................................................

23

I n t r o d u c t i o n .....................................................................
M e t h o d s .................................................................................
R e s u l t s .................................................................................
D i s c u s s i o n ..........................................................................

23
24
25
39

II.

III.

IV.

V.

MULTIPLE PATHWAYS IN EARLY SUCCESSIONAL
DEVELOPMENT FOLLOWING DISTURBANCE TO JACK
PINE SITES IN NORTHERN LOWERMICHIGAN

48

I n t r o d u c t i o n .....................................................................
M e t h o d s ..................................................................................
R e s u l t s .................................................................................
D i s c u s s i o n ..........................................................................

48
50
50
87

FLORISTIC COMPOSITION BEFORE AND AFTER
FIRE ON A JACK PINE CLEARCUT SITE IN
NORTHERN LOWER MICHIGAN..............................................

98

I n t r o d u c t i o n .....................................................................
98
M e t h o d s .................................................................................
99
R e s u l t s ................................................................................. 101
D i s c u s s i o n .......................................................................... 119

iv

CHAPTER
VI.

RESPONSE OF UNDERSTORY VEGETATION TO
FERTILIZATION ON MATURE AND CLEARCUT
JACK PINE SITES IN NORTHERN LOWER
MICHIGAN....................................................................................

124

I n t r o d u c t i o n ..................................................................... 124
M e t h o d s a n d M a t e r i a l s .............................................. 125
R e s u l t s ................................................................................. 127
D i s c u s s i o n .......................................................................... 135
V II.

APPARENT HEAT STIMULATION OF BURIED SEEDS
OF GERANIUM BICKNELLII BRITT. ON JACK
PINE SITES IN NORTHERN LOWERMICHIGAN
141
I n t r o d u c t i o n ..................................................................... 141
E xperim ents w ith F re sh ly -M a tu re d
G e r a n i u m S e e d ............................................................. 143
E x p e r i m e n t s w i t h B u r i e d S e e d .............................. 144
D i s c u s s i o n .......................................................................... 153

V III.

SUMMARY AND CONCLUSION................................................. 157

APPENDICES
A.

R e la ti v e Cover and R e la tiv e Frequency
D a ta f o r S p e c ie s S u rv e y ed on Unburned
C l e a r c u t S i t e s ................................................................ 160

B.

R e l a t i v e Cover and R e l a t i v e F req u en cy
D a ta f o r S p e c ie s S u rv ey ed on S i t e s
B u r n e d b y P r e s c r i p t i o n o r W i l d f i r e ................... 167

C.

R e l a t i v e Cover and R e l a t i v e F re q u e n c y
D ata f o r U n d e rsto ry S p e c ie s Surveyed
on I n t e r m e d i a t e - a g e and M a tu re J a c k
P i n e S t a n d s .......................................................................

178

N a tio n a l C o o p e ra tiv e S o il Survey f o r th e
G r a y l i n g S e r i e s .............................................................

181

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

182

D.

v

L IS T OF TABLES
T able

Page

2.1

Summary o f S t u d y S i t e s ........................................

3.1

R e la tiv e Cover (C ) , R e la tiv e F requency ( F ) ,
a n d S p e c i e s R i c h n e s s (R) f o r V e g e t a t i o n a l
G r o u p s o n U n b u r n e d C l e a r c u t s .....................
26

3.2

R e l a t i v e Cover ( C ) , R e l a t i v e Freq uency ( F ) ,
a n d S p e c i e s R i c h n e s s (R) f o r V e g e t a t i o n a l
G r o u p s o n B u r n e d C l e a r c u t s ...........................
27

3.3

P l a n t S p e c ie s E n c o u n te r e d on
Burned and
Unburned C l e a r c u t S i t e s i n N o r t h e r n
L ow er M i c h i g a n .........................................................
31

4 .1

R e l a t i v e Cover (C ), R e l a t i v e Frequ en cy ( F ) ,
a n d S p e c i e s R i c h n e s s (R) f o r V e g e t a t i o n a l
G roups on M a tu re J a c k P i n e S t a n d s and
U n b u r n e d C l e a r c u t s ...............................................
53

4 .2

P l a n t S p e c ie s E n c o u n te re d on U nburned C l e a r c u t S i t e s a n d M a t u r e J a c k P i n e (_^ 55
Y e a r s ) S t a n d s i n N o r t h e r n L ow er M i c h i g a n .

54

R e l a t i v e C o v e r ( C ) , R e l a t i v e F r e q u e n c y (F) ,
a n d S p e c i e s R i c h n e s s (R) f o r V e g e t a ­
t i o n a l Groups on M a tu re J a c k P in e S ta n d s
a n d B u r n e d C l e a r c u t s ...................................................

61

P l a n t S p e c ie s E n c o u n te r e d on Burned C l e a r c u t S i t e s a n d M a t u r e J a c k P i n e (_^ 55
Y e a r s ) S t a n d s i n N o r t h e r n L ow er M i c h i g a n ,

63

R e la tiv e Cover (C ) , R e la tiv e Frequency ( F ) ,
a n d S p e c i e s R i c h n e s s (R) f o r V e g e t a t i o n a l
G roups on U nburned 3 5 - Y e a r - O l d J a c k P i n e
S ta n d s and B urned 3 5 -Y ear-O ld J a c k P in e
S t a n d s .......................................................................................

70

P l a n t S p e c i e s E n c o u n t e r e d on B u rn e d and
Unburned 3 5 -Y ear-O ld J a c k P in e S ta n d s
i n N o r t h e r n Lower M i c h i g a n ....................................

72

4 .3

4.4

4.5

4 .6

vi

9

T a b le

Page
C o m p o s i t e D a t a f r o m 1980 a n d 1 9 8 1 S u r v e y
o f 3 P e r m a n e n t C o v e r T r a n s e c t s (60 m)
E s t a b l i s h e d i n G eranium (G eranium
b i c k n e l l i i ) P a t c h e s o n S i t e 7A..........................

78

R e l a t i v e Cover ( C ) , R e l a t i v e F requency ( F ) ,
a n d S p e c i e s R i c h n e s s (R) f o r V e g e t a ­
t i o n a l G r o u p s on A r e a s . B u r n e d b y W i l d ­
f i r e s .........................................................................................

80

Y e a r - B y - Y e a r C o m p a r i s o n s o f t h e Mean I n d e x
o f S i m i l a r i t y (S) f o r U n b u r n e d C l e a r c u t
S i t e s C o m p risin g a 6 - Year S eq u en ce, and
t h e Mean I n t e r s i t e ( D i f f e r e n t S i t e s ) a n d
Mean I n t r a s i t e (Same S i t e ) I n d e x o f
S i m i l a r i t y o f S i t e s Compared a t C o n se c u ­
t i v e Y e a r s ............................................................................

84

4 . 1 0 Y e a r - B y - Y e a r C o m p a r i s o n s o f t h e Mean I n d e x
o f S i m i l a r i t y (S) f o r B u r n e d C l e a r c u t
S i t e s C o m p ris in g a 6 -Y ear S e q u e n ce , and
t h e Mean I n t e r s i t e ( D i f f e r e n t S i t e s ) a n d
Mean I n t r a s i t e (Same S i t e ) I n d e x o f S i m i ­
l a r i t y o f S i t e s Compared a t C o n s e c u t i v e
Y e a r s .........................................................................................

86

R e l a t i v e Cover and R e l a t i v e F re q u e n c y o f
E ach S p e c i e s , by S p e c ie s Group, S u r ­
v e y e d i n 1 9 7 9 , 1 9 8 0 , a n d 1981 ( 3 - ,
4 - , and 5 -Y ear-O ld C l e a r c u t , R e s p e c t i ­
v e l y ) , a s a C o m p o site o f t h e 3 Unburned
B l o c k s ......................................................................................

102

D i v e r s i t y T re n d s O ver t h e 1979, 1980, and
19 81 G r o w i n g S e a s o n s , E x p r e s s e d a s T o t a l
S p e c ie s R i c h n e s s , R ic h n e s s o f Each
S p e c i e s G r o u p , A v e r a g e R i c h n e s s p e r 1 m2
P l o t , a n d S h a n n o n I n d e x , A v e r a g e d Among
t h e U n b u r n e d C l e a r c u t B l o c k s ...............................

103

A d d i t i o n s a n d L o s s e s o f S p e c i e s , a s a Compo
s i t e o f t h e 3 Unburned B lo c k s , S u rv ey ed
i n 1979, 1980, and 1981.
V alues f o r
L o s s e s o f S p e c i e s Are E x p r e s s e d a s a
R a t i o o f Number o f P l o t s i n W h ic h a
S p e c i e s Was L o s t i n a S u r v e y , O v e r t h e
Number o f P l o t s I t Was P r e s e n t i n t h e
P r e v i o u s Y e a r S u r v e y ...................................................

107

4.7

4.8

4.9

5.1

5.2

5.3

v ii

T a b le
5 .A

5.5

5.6

P age
R e la ti v e Cover and R e la ti v e F requency o f
E a c h S p e c i e s , By S p e c i e s G r o u p , S u r ­
v e y e d i n 1979 ( 3 - Y e a r - O l d C l e a r c u t ) , and
1980 a n d 1981 ( 1 - a n d 2 - Y e a r - O l d P r e ­
s c r i b e d Burn, R e s p e c t i v e l y ) , as a
C o m p o s i t e o f t h e 3 B u r n e d B l o c k s .......................

109

D i v e r s i t y T ren d s Over t h e 1979, 1980, and
1 98 1 G r o w i n g S e a s o n s , E x p r e s s e d a s T o t a l
S p e c ie s R ic h n e s s , R ic h n e s s o f Each
S p e c ie s Group, A verage R ic h n e s s p e r
1 m2 P l o t , a n d S h a n n o n I n d e x , A v e r a g e d
Among t h e B u r n e d C l e a r c u t B l o c k s .........................

110

A d d i t i o n s and L o s s e s o f S p e c i e s , as a
Com posite o f th e 3 Burned B lo c k s , S u r ­
veyed i n 1979, 1980, and 1981.
V alues
fo r L osses o f S p e c ie s a re E xpressed as
a R a t i o o f Number o f P l o t s i n w h i c h a
S p e c i e s was L o s t i n a S u r v e y , O v e r t h e
Number o f P l o t s I t Was P r e s e n t i n t h e
P r e v i o u s Y e a r S u r v e y .....................................................

11A

5.7

R e l a t i v e (R el) Cover and R e l a t i v e F re q u en c y
o f Each S p e c ie s i n th e U n d e r s to r y o f a
M ature J a c k P in e S ta n d ( S i t e 1 3A ), A d ja ­
c e n t t o t h e E x p e r i m e n t a l B u r n A r e a ..................... 118

6 .1

B i o m a s s D a t a ( g / m 2) b y S p e c i e s G r o u p f o r
t h e 1 9 7 8 P r e s c r i b e d B u r n ...........................................

136

7.1

T o t a l Number o f G e r m i n a n t s f o r S p e c i e s
A p p e a r in g i n S o i l Sam ples from t h e 3 - Y e a r - ,
3 5 - Y e a r - , and 5 5 - Y e a r - O ld P o s t - F i r e S i t e s
D u rin g th e B u rie d Seed E x p e rim e n t.
T reat­
m e n t s Were a n U n h e a t e d C o n t r o l , 7 0 ° C - 3 0
M i n , 9 0 ° C - 3 0 M i n , a n d 7 0 ° C - 3 0 Min + S t r a ­
t i f i c a t i o n ............................................................................... 151

A .1

R e l a t i v e Cover and R e l a t i v e F requency o f
E ach S p e c i e s on S i t e 1B80-81 S u r v e y e d a s
a 0 - Y e a r a n d 1 - Y e a r - O l d C l e a r c u t .......................

160

R e l a t i v e Cover and R e l a t i v e F req u en cy o f
Each S p e c ie s on S i t e 2 S u rv ey ed as a
1 - Y e a r - O l d C l e a r c u t ........................................................

161

R e l a t i v e Cover and R e l a t i v e F re q u e n c y o f
E a c h S p e c i e s on S i t e 3A S u r v e y e d a s a
2 - Y e a r - O l d C l e a r c u t ........................................................

162

A .2

A .3

v iii

T a b le
A .4

A .5

A .6

A .7

B .l

B .2

B .3

B .4

B .5

B .6

R e l a t i v e Cover and R e l a t i v e F re q u en c y o f
Each S p e c ie s on S i t e 3B79-80-81 S urveyed
as a 2 -Y e a r-, 3 -Y e a r-, and 4-Y ear-O ld
C l e a r c u t ..................................................................................

163

R e l a t i v e Cover and R e l a t i v e F req u en cy of
Each S p e c ie s on S i t e 4A79-80-81 S u rv ey ed
as a 3 -Y e a r-, 4 - Y e a r - , and 5 -Y ear-O ld
C l e a r c u t .................................................................................

164

R e l a t i v e Cover and R e l a t i v e F re q u e n c y o f
E ach S p e c i e s on S i t e 5 A 79-80-81 S u r v e y e d
as a 4 - Y e a r - , 5 - Y e a r - , and 6-Y ear-O ld
C l e a r c u t .................................................................................

165

R e l a t i v e Cover and R e l a t i v e F re q u e n c y o f
Each S p e c ie s on S i t e 6 S u rv ey ed a s a 5Y e a r - O l d C l e a r c u t ..........................................................

166

R e l a t i v e Cover and R e l a t i v e F r e q u e n c y o f
E a c h S p e c i e s on S i t e 7 A 8 0 - 8 1 S u r v e y e d
as a 1-Y ear- and 2 -Y ear-O ld P r e s c r i b e d
B u r n ...........................................................................................

167

S p e c ie s Cover and F re q u e n c y D ata from S i t e
7 B (1 -, 2 - , and 3 -Y ear-O ld B urn). S i t e
12A ( 3 5 - Y e a r - O l d J a c k P i n e S t a n d ) , a n d
S i t e 13C ( 5 5 - Y e a r - O l d J a c k P i n e S t a n d )
U s e d i n t h e B u r i e d S e e d E x p e r i m e n t ...............

168

R e l a t i v e Cover and R e l a t i v e F re q u en c y o f
Each S p e c i e s , by S p e c ie s Group, S u rv e y ed
i n 1979 ( S i t e 1A, 0 - Y e a r C l e a r c u t ) , a n d
1980 a n d 1981 ( S i t e 7 C 8 0 - 8 1 , 1 - a n d 2 - Y e a r
P r e s c r i b e d B u r n ) .............................................................

170

R e l a t i v e Cover and R e l a t i v e F req u en cy o f
E ach S p e c ie s on S i t e 7D 79-80-81 S u rv e y e d
as a 1 -Y e a r-, 2 -Y e a r-, and 3-Y ear-O ld
P r e s c r i b e d B u r n ...............................................................

171

R e l a t i v e Cover and R e l a t i v e F re q u e n c y o f
E a c h S p e c i e s on S i t e 8 A 7 9 - 8 0 - 8 1 S u r v e y e d
a s a 2 - Y e a r - , 3 - Y e a r - , and 4 -Y e a r - O ld
W i l d f i r e .................................................................................
R e l a t i v e Cover and R e l a t i v e F re q u en c y o f
E ach S p e c i e s on S i t e 9A79-80-81 S u rv e y ed
as a 3 -Y e a r-, 4 -Y ear, and 5-Y ear-O ld
P r e s c r i b e d B u r n ...............................................................
ix

172

173

Page

T a b le
B.7

R e l a t i v e Cover and R e la tiv e Frequency of :
E ach S p e c i e s on S i t e 9B 79-80-81 S u rv e y ed
as a 3 -Y e a r-, 4 -Y e a r-, and 5-Y ear-O ld
W i l d f i r e ....................................................................... ..

174

R e l a t i v e Cover and R e l a t i v e F requency o f
E ach S p e i c e s on S i t e 10A 79-80-81 S u rv e y e d
a s a 4 - Y e a r - , 5 - Y e a r - , and 6 -Y e a r-O ld
P r e s c r i b e d B u r n ................................................................

175

R e l a t i v e Cover and R e la ti v e F requency of
E a c h S p e c i e s on S i t e 1 0 B 7 9 - 8 0 - 8 1 S u r v e y e d
as a 4 -Y e a r-, 5 -Y e a r-, and 6-Y ear-O ld
P r e s c r i b e d B u r n ................................................................

1 76

B.10 R e l a t i v e Cover and R e l a t i v e F re q u e n c y f o r
E a c h S p e c i e s o n S i t e 11 S u r v e y e d a s a
6 - Y e a r - O l d W i l d f i r e .....................................................

177

B .8

B .9

C .l

C .2

C .3

R e l a t i v e Cover and R e la ti v e F requency f o r
E a c h S p e c i e s o n S i t e 12B S u r v e y e d a s a n
In te rm e d ia te -A g e (35-Y ear-O ld) Ja ck Pine
S t a n d .........................................................................................

178

R e l a t i v e Cover and R e l a t i v e F req u en cy f o r
E a c h S p e c i e s o n S i t e 13B S u r v e y e d a s a
M a t u r e J a c k P i n e S t a n d ..............................................

179

R e l a t i v e Cover and R e l a t i v e F req u en cy f o r
E a c h S p e c i e s on S i t e 13D S u r v e y e d a s a
M a t u r e J a c k P i n e S t a n d ..............................................

180

x

*

L IS T OF FIGURES
F igure

Page

2.1

Map o f M i c h i g a n ............................................................. ......

3.1

A) S p e c i e s R i c h n e s s , B) A v e r a g e R i c h n e s s
p e r 1 m2 P l o t , a n d C) S h a n n o n I n d e x o f
D i v e r s i t y V ersus Y ears A f t e r T reatm ent
on B u r n e d a n d U n b u m e d S i t e s .
V e rtical
B a r s E q u a l One S t a n d a r d D e v i a t i o n ( S D ) .
O n l y One O b s e r v a t i o n a t Y e a r 3 B u r n e d
a n d Y e a r 6 U n b u r n e d ...................................................

29

T o t a l V e g e t a t i o n a l C o v e r (m) V e r s u s Y e a r s
A f t e r T r e a t m e n t on B urned and Unburned
S ites.
V e r t i c a l B a r s E q u a l One SD.
O n l y One O b s e r v a t i o n a t Y e a r 3 B u r n e d
a n d Y e a r 6 U n b u r n e d .....................................................

33

A) Number o f G r a s s S p e c i e s , a n d B) P e r c e n t
Cover o f G ra sses V ersus Years A f te r
T r e a tm e n t on Burned an d Unburned S i t e s .
V e r t i c a l B a r s E q u a l One SD.
O n l y One
O b s e r v a t i o n a t Y ear 3 Burned and Year 6
U n b u r n e d .................................................................................

35

P e r c e n t Cover o f Sedge V ersu s Y ears A f t e r
T r e a tm e n t on Burned and U nburned S i t e s .
V e r t i c a l B a r s E q u a l One SD.
O n l y One
O b s e r v a tio n a t Y ear 3 Burned and Y ear 6
U n b u r n e d .................................................................................

37

A) S p e c i e s R i c h n e s s ( T o t a l Number o f
S p e c i e s ) , B) A v e r a g e R i c h n e s s P e r P l o t ,
a n d C) S h a n n o n I n d e x o f D i v e r s i t y , o f
S i t e s A veraged W ith in Y e a rs , f o r M ature
J a c k P in e S ta n d s and Unburned C l e a r c u t
J a c k P in e S i t e s C om prising a 6-Y ear
Sequence F o llo w in g L ogging.
V ertical
B a r s E q u a l One S t a n d a r d D e v i a t i o n ( S D ) . . .

52

3.2

3 .3

3.4

4.1

xi

5

Page

F ig u r e
4.2

4 .3

4.4

4.5

4.6

4.7

4 .8

T o ta l V e g e t a t i o n a l Cover (m ), o f S i t e s
A verage W ith in Y e a r s , f o r M ature
J a c k P in e S ta n d s an d Unburned C l e a r c u t
Ja c k P in e S i t e s C o m prising a 6-Y ear
Sequence F o llo w in g L ogging.
V ertical
B a r s E q u a l One SD........................................................

56

R e l a t i v e Cover f o r Each S p e c ie s Group, o f
S i t e s A veraged W ith in Y e a r s , f o r M ature
J a c k P in e S ta n d s and Unburned C l e a r c u t
S i t e s C om prising a 6-Y ear Sequence
F o l l o w i n g L o g g i n g ..................................................... .

58

A) S p e c i e s R i c h n e s s ( T o t a l Number o f
S p e c i e s ) , B) A v e r a g e R i c h n e s s P e r P l o t ,
a n d C) S h a n n o n I n d e x o f D i v e r s i t y , o f
S i t e s A veraged W ith in Y e a rs , f o r M ature
J a c k P in e S ta n d s and B urned C l e a r c u t
S i t e s C om prising a 6-Y ear Sequence
F o llo w in g B urning.
V e r t i c a l Bars E qual
One SD....................................................................................

60

T o ta l V e g e ta tio n a l Cover (m ), o f S i t e s
A veraged W ith in Y e a rs , f o r M ature J a c k
P in e S ta n d s and B urned C l e a r c u t S i t e s
C o m p risin g a 6-Y ear Sequence F o llo w in g
B urning.
V e r t i c a l B a r s E q u a l One SD. . . .

65

R e l a t i v e Cover f o r Each S p e c ie s Group, o f
S i t e s A veraged W ith in Y e a rs , f o r M ature
J a c k P in e S ta n d s and Burned C l e a r c u t
Ja c k P in e S i t e s C om prising a 6-Y ear
S e q u e n c e F o l l o w i n g B u r n i n g .................................

67

A) S p e c i e s R i c h n e s s , B) A v e r a g e R i c h n e s s
P e r P l o t , a n d C) S h a n n o n I n d e x o f D i ­
v e r s i t y , o f S i t e s Averaged W ith in Y e ars,
f o r Unburned 3 5 -Y e a r-O ld J a c k P in e
S ta n d s and Burned 3 5 -Y e a r-O ld J a c k P in e
S tan d s C om prising a 5-Y ear Sequence
F o llo w in g B urning.
V e r t i c a l Bars Equal
One SD....................................................................................

71

T o ta l V e g e ta tio n a l Cover (m ), o f S i t e s
A v erag ed W it h i n Y e a r s , f o r Unburned 35Y e ar-O ld J a c k P in e S ta n d s and Burned
35-Y ear-O ld Ja c k P in e S ta n d s C om prising
a 5-Year Sequence F o llo w in g B u rn in g .
V e r t i c a l B a r s E q u a l One SD.................................

74

x ii

F ig u r e
4 .9

5.1

5 .2

6.1

6.2

6.3

Page
R e l a t i v e Cover f o r Each S p e c ie s Group,
o f S i t e s Averaged W ith in Y e a rs, f o r
Unburned and B urned 3 5 -Y e ar-O ld J a c k
P in e S tands C om prising a 5-Y ear
.......................
Sequence F o llo w in g B urning.

76

T o t a l C o v e r (m) a n d R e l a t i v e C o v e r (%) f o r
Each S p e c ie s Group, as a C o m posite o f
t h e 3 Unburned B l o c k s , S urveyed i n 1979,
1 9 8 0 , a n d 1 9 8 1 ..................................................................

105

T o t a l C o v e r (m) a n d R e l a t i v e C o v e r (%) f o r
Each S p e c ie s Group, as a C om posite o f t h e
3 Burned B l o c k s , S u rv e y ed i n 1979, 1980,
a n d 1 9 8 1 .................................................................................

112

T o t a l L i v e S h o o t B i o m a s s ( g / m 2) f o r t h e
F e r t i l i z e d and C o n tr o l P l o t s i n th e
M atu re and C l e a r c u t J a c k P i n e , and t h e
P r e s c r i b e d B u r n A r e a ...................................................

129

T o t a l L i v e S h o o t B i o m a s s ( g / m 2) o f S e d g e
(C arex p e n s y l v a n i c a ) f o r t h e F e r t i l i z e d
a n d C o n t r o l P l o t s i n (A) M a t u r e J a c k
P i n e a n d (B) C l e a r c u t J a c k P i n e .....................

130

T o t a l L i v e S h o o t B i o m a s s ( g / m 2) f o r B l u e ­
b e r r y (V accinium s p p .) f o r t h e F e r t i l i i z e d a n d C o n t r o l P l o t s i n (A) M a t u r e
J a c k P i n e a n d (B) C l e a r c u t J a c k P i n e . , . .

131

6.4

T o t a l L i v e S h o o t B i o m a s s ( g / m 2) o f
B racken F ern (F te rid iu m a q u ilin u m ) f o r
th e F e r t i l i z e d and C o n tr o l P l o t s in
(A) M a t u r e J a c k P i n e a n d (B) C l e a r c u t
J a c k P i n e .............................................................
132

6.5

F e r t i l i z e d / C o n t r o l Biom ass R a t i o s f o r
T o ta l L ive Shoot B iom ass, Sedge, B lu e ­
b e r r y , a n d B r a c k e n F e r n i n (A) M a t u r e
J a c k P i n e a n d (B) C l e a r c u t J a c k P i n e . . . .

134

C u m u l a t i v e a n d W e e k ly Number o f G e r m i n a n t s
f r o m t h e C o n t r o l , 7 0 ° C - 3 0 M i n , a n d 90 °C30 Min T r e a t m e n t s i n t h e 3 - Y e a r - O l d , 3 5 Y e a r - O l d , a n d 5 5 - Y e a r - O l d S i t e s (Com­
b in e d ) D u rin g th e B u rie d Seed E x p e r i ­
m e n t ......................

148

7 .1

x iii

P age

F ig u r e
7.2

W e e k l y T o t a l Number o f G e r m i n a n t s o f
G e r a n i u m , G r a s s e s ( A l l S p e c i e s Com­
b i n e d ) , an d Hawkweed ( H i e r a c i u m s p p . )
fro m t h e C o n t r o l , 70°C -30 Min, and
9 0 ° C - 3 0 Min T r e a t m e n t s i n t h e 3 - Y e a r O ld, 3 5 -Y e ar-O ld , and 55-Y ear-O ld
S i t e s (C o m b in e d ) D u r i n g t h e B u r i e d
S e e d E x p e r i m e n t .....................................................

x iv

150

L IS T OF PLATES
P lates
3 .1
3.2

4.1

4 .2
4 .3
5.1
5.2
5 .3

-

Page

C a r e x Meadow E s t a b l i s h e d o n a 5 - Y e a r
O l d U n b u r n e d C l e a r c u t S i t e .................................

44

S h r u b s an d E a r l y S u c c e s s i o n a l Hardwoods
D om inating a 5 -Y ear-O ld P r e s c r ib e d
B u r n S i t e ............................................................................

44

C o v e r T r a n s e c t (20m) T h r o u g h a G e r a n i u m
P a t c h on 1 -Y e a r-O ld P r e s c r i b e d Burn
S i t e .........................................................................................

79

Above C o v e r T r a n s e c t a t Y e a r 2 A f t e r
F i r e ; N ote D is a p p e a r a n c e o f G e ra n iu m .. . .

79

V ig o ro u s J a c k P i n e R e p r o d u c t i o n on a
4 -Y e a r-O ld S i t e Burned by W i l d f i r e

92

C arex p e n s y l v a n i c a D o m in atin g Unburned
B locks a t Year 4 A f t e r C l e a r c u t t i n g

106

L a r g e A r e a B a r e d o f V e g e t a t i o n One Y e a r
A f t e r F i r e ..........................................................................

113

O verview o f B urned B lo c k s a t Y ear 2 A f t e r
F i r e S h o w i n g R a p i d R e v e g e t a t i o n .....................

113

xv

CHAPTER I
INTRODUCTION
Jack pine
forest

( F i n u s b a n k s i a n a L a m b .)

is

a dom inant

ty p e i n t h e Lake S t a t e s and h a s b e e n t h e fo c u s

o f man y s i l v i c u l t u r a l
pine is
ju v en ile

of in te re s t
grow th,

and p a p e r ,

its

and e c o l o g i c a l s t u d i e s .

to f o r e s t e r s b e c a u se o f i t s

d e s i r a b l e wood p r o p e r t i e s

a b ility

because i t pro v id es

rap id

fo r pulp

t o grow on h a r s h s i t e s ,

and

t h e summer n e s t i n g h a b i t a t f o r

th e endangered K i r t l a n d 's w a rb le r
lan d ii) .

Jack

Jack pine is

(D endroica k i r t -

a sero tin o u s

s p e c ie s t h a t has

b e e n m a i n t a i n e d i n t h e Lake S t a t e s by p e r i o d i c f i r e
(H einselm an 1973).

In fa c t,

the p r e s e n t a re a of ja c k

p i n e i n t h e Lake S t a t e s

is

its

t h e w i l d f i r e s w hich fo llo w e d

p ro life ra tio n

e a r l y p ine lo g g in g

after

somewhat i n f l a t e d

due to

(B enzie 1977).

J a c k p i n e i s known a s a " f i r e

sp ecies",

and t r a ­

d i t i o n a l l y c l e a r c u t t i n g and b u r n in g have b e en u sed
for i t s

reg en eratio n .

m ethod o f c u t t i n g ,

However, v i r t u a l l y

s ite p rep aratio n ,

r e g e n e r a tio n has been t r i e d
(E yre and L eB arron 1944).

e v e r y known

and a r t i f i c i a l

i n m anaging t h i s
It

species

i s now a p p a r e n t t h a t

no one m e th o d w i l l p r o d u c e s a t i s f a c t o r y r e s u l t s

1

on a

2
co n sisten t b asis.
In n o r t h e r n low er M ichigan,

larg e areas of jack

p i n e a r e c u t and b u rn e d e a c h y e a r .
in itia l failu re

o f ja c k p in e re g e n e ra tio n fo llo w in g

t r e a t m e n t i s more t h e r u l e
fore,

th ere

c u ltu re
tu re,

On t h e s e a r e a s ,

than th e e x c e p tio n .

i s much i n t e r e s t i n t h e e c o l o g y a n d s i l v i ­

of th is

sp ecies.

A fte r a review o f th e l i t e r a ­

I r e a l i z e d t h a t m ost s tu d ie s of p r e s c r ib e d

b u r n i n g f o c u s e d on s l a s h r e d u c t i o n ,
tio n ,

T here­

and r e s p o n s e s o f s e e d l i n g s

m ental f a c t o r s

seedbed p r e p a r a ­

to v a rio u s e n v iro n ­

(E y re and L e B a rro n 1944, B e a u f a i t 1 9 6 2 ).

L im ite d d e t a i l e d w ork, how ever, h a s been co n d u cted
on p l a n t s u c c e s s i o n on j a c k p i n e
a n d / o r b u r n i n g i n t h e Lake S t a t e s
1970, K r e f t i n g and A h lg ren 1974),
of th is

type a re ,

sites

fo llo w in g lo g g in g

( A h l g r e n 1960, Vogl
In M ich igan,

stu d ies

t o my k n o w l e d g e , n o n e x i s t e n t .

T h e o b j e c t i v e o f my r e s e a r c h was t o c h a r a c t e r i z e
early p lan t

su c c e ssio n fo llo w in g c l e a r c u t t i n g an d /o r

b u rn in g on ja c k p in e

sites

i n n o r t h e r n low er M ichigan.

F i e l d d a t a w e r e c o l l e c t e d i n Roscommon, O s c o d a , C r a w ­
f o r d , a n d Ogemaw c o u n t i e s d u r i n g t h e sum m ers o f 1 9 7 9 ,
1980,

a n d 1 9 81 f r o m u n b u r n e d c l e a r c u t s i t e s

sites

com prising a s u c c e s s io n a l s e r i e s

years.

and bu rn ed

from 0 to 6

C o m m unity d a t a w e r e a l s o c o l l e c t e d f r o m i n t e r ­

m ediate-age

and m a tu r e j a c k p i n e

con sid ered r e p r e s e n ta tiv e

s ta n d s w hich were

of the p re d is tu rb a n c e

3
c o n d i t i o n o f many o f t h e s i t e s .

In a d d itio n a th re e -

y e a r i n v e s t i g a t i o n was u n d e r t a k e n ,
p lo ts,

u s in g perm anent

o f v e g e t a t i o n a l c h an g es on b u rn ed and u n b u rn ed

blocks in a ja c k p in e c le a r c u t.

When c e r t a i n t r e n d s

b e ca m e a p p a r e n t d u r i n g e a r l y s u c c e s s i o n

(e.g .

larg e

i n c r e a s e s i n C a re x dom inance and t h e f i r s t - y e a r
dom inance o f Geranium b i c k n e l l i i B r i t t ,

on b u r n e d

s i t e s ) , X became i n t e r e s t e d i n t h e m echanism s i n v o l v e d .
T herefore,
tatio n

stu d ies

of th e resp o n se of u n d e rsto ry vege­

to f e r t i l i z a t i o n

in a m ature ja c k p in e sta n d

and an a d j a c e n t c l e a r c u t ,

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

g e r m i n a t i o n o f b u r i e d s e e d from j a c k p i n e s t a n d s a lo n g
a s u c c e s s io n a l s e r i e s w ere i n i t i a t e d .

CHAPTER I I
DESCRIPTION OF STUDY AREA AND SITES
G e n e ra l D e s c r i p t i o n o f S tud y Area
S u rfa ce F o rm atio n s,

S o il,

and C lim ate

T h e r e s e a r c h s i t e s w e r e l o c a t e d i n Roscommon,
C r a w f o r d , O s c o d a , a n d Ogemaw c o u n t i e s i n t h e n o r t h e a s t
low er p e n i n s u la o f M ichigan

(Fig.

of th ese c o u n ties g e n e ra lly

l i e s b e t w e e n 275 a n d 365 m

above se a l e v e l .

2 .1 ),

The to p o g ra p h y o f t h i s a r e a i s

e c t r e s u l t o f th e W isconsin g l a c i a t i o n ;
tio n

T he e l e v a t i o n

t o o k p l a c e f r o m 14 8 00 B . P .

v a st areas of t i l l

the f in a l g la c ia ­

t o 9500 B . P . ,

v aria­

( V e a t c h eit al^. 1 9 2 4 ) .

M i c h i g a n sho w s h i g h e d a p h i c d i v e r s i t y ,
350 s o i l s e r i e s m a p p e d (Sommers 1 9 7 7 ) .
fo u r co unty a r e a used in
R ubicon, K a lk a sk a ,

leav in g

and outw ash p l a i n s w ith l i t t l e

tio n in surface r e l i e f

a d ir­

th is

study i s

w ith over

However,

the

d o m in a t e d by

and G ra y lin g s e r i e s o f th e H ap lo rtho d

an d Udipsamment g r e a t g r o u p s .
sand s e r i e s p red o m in ates,
stu d y w ere l o c a t e d on t h i s

In p a r t i c u l a r ,

and a l l

sites

so il series.

in terg rad in g of asso ciated s o il s e rie s ,

the G rayling

included fo r
However,
such as th e

G raycalm , w i t h t h e G r a y l i n g s e r i e s on th e s t u d y
s i t e s was p o s s i b l e ;

the e x te n t o f t h i s in te r g r a d in g
4

5

F ig.

2 .1 .

Map o f M i c h i g a n

6
is

unk n ow n b e c a u s e no f o r m a l s o i l a n a l y s e s w e r e c o n ­

ducted.
The G r a y l i n g s a n d s e r i e s
Udipsam m ent)

is

c h a r a c t e r i z e d by i t s

in coherent s tru c tu re ,
th ro u g h o u t,
m o istu re.

(a m ixed,

frig id ,

T ypic

lo o se c o n siste n c e ,

sandy o r s i n g l e - g r a i n e d t e x t u r e

and p e r v i o u s n e s s and n o n r e t e n t i v e n e s s
A verage m o is tu r e c o n te n t d u r i n g th e

to

grow ing

s e a s o n i s v e r y low t o a d e p t h o f 1 m o r m o r e a n d f e r ­
tility

is

c o r r e s p o n d i n g l y low.

R eactio n v a r ie s

from

medium t o s t r o n g l y a c i d t o a d e p t h o f 1 m o r m o r e
( V e a t c h e_t a l .

1 9 2 4 , A p p e n d i x D) .

The c l i m a t e o f t h e

fo u r cou nty a r e a i s

charac­

t e r i z e d by c o ld w i n t e r s , s h o r t m ild sum m ers, a l a r g e
number o f c lo u d y d a y s ,

lo w e v a p o r a t i o n ,

h i g h h u m i d i t y ( V e a t c h e_t a l .
tem perature is

1924) .

The m ea n a n n u a l

g e n e r a l l y 6° t o 7 ° C , w i t h m ean m o n t h l y

t e m p e r a t u r e s r a n g i n g f r o m - 8 ° t o 19 °C.
is

and m o d e r a te l y

P re cip ita tio n

f a i r l y w e ll d i s t r i b u t e d thro ugh out th e y e a r, w ith

m ea n a n n u a l r a i n f a l l a n d s n o w f a l l o f 77 a n d 180 cm,
resp ectiv ely .
100 t o

The l e n g t h o f t h e g ro w in g s e a s o n i s

120 d a y s , w i t h t h e l a s t p r o b a b l e

f r o s t in the

s p r i n g a t t h e e n d o f May, a n d t h e f i r s t p r o b a b l e f r o s t
in the f a l l

i n m id S e p te m b e r

(U .S.D .A .

1971).

I n 1924, V e atc h and a s s o c i a t e s w ro te t h e f o llo w in g
a c c o u n t o f t h e f o r e s t c o m m u n i t i e s i n Roscommon c o u n t y
(n early id e n tic a l

a c c o u n ts w ere w r i t t e n f o r Oscoda,

7
Ogemaw, a n d C r a w f o r d c o u n t i e s ) .
The w h o le l a n d a r e a a t t h e tim e o f
o c c u p a t i o n b y w h i t e men w a s a d e n s e f o r e s t
e x c e p t f o r a sm all a c re a g e o f bog o r m arsh
a n d p o s s i b l y some o p e n l a n d i n t h e d r i e r
sand p l a i n s .
The lu m b e r i n d u s t r y s t a r t e d
a b o u t 1870 a n d a t t h e p r e s e n t t i m e t h e
o r i g in a l f o r e s t has been c u t, except in
a f e w s m a l l s c a t t e r e d a r e a s a n d wood l o t s
a n d i n a f e w p l a c e s i n swamps.
S everal
ty p e s of f o r e s t o r t r e e a s s o c i a t i o n s w ere
rep resen ted :
(1 ) The h a r d w o o d f o r e s t s i n
w h i c h h a r d m a p l e o r s u g a r m a p l e (A c e*
6az c. haA. am) a n d b e e c h w e r e t h e d o m i n a n t
s p e c i e s and elm , a s h , bassw ood, y e llo w
b i r c h , and hem lock w ere s u b o r d i n a t e
s p e c i e s ; (2) t h e m ix ed hardw ood and
c o n i f e r o u s f o r e s t , i n w h i c h common h a r d ­
woods w e re i n t i m a t e l y a s s o c i a t e d w i t h
w h i t e p i n e , h e m l o c k , a n d some Norw ay p i n e
( r e d p i n e ) ; (3) t h e p i n e f o r e s t s i n w h ic h
w h i t e p i n e , Norway p i n e , o r Norway a n d
j a c k p i n e p r e d o m i n a n t e d ; a n d ( 4 ) t h e swamp
f o r e s t s i n w h ich t h e dom inant s p e c i e s were
arbor v i t a e , b la c k spruce, w hite spruce
( P Z z z a z a n a d z n s Z A ) , balsam f i r (AbZz6
b a Z 6 a m z a ) , and ta m a ra c k .
Much o f t h e
f o r e s t land is a t p re s e n t e it h e r d e s o la te
stump l a n d c o v e r e d w i t h a d e n s e g r o w th o f
b r u s h , b r i e r s , a n d g r a s s o r g r o w n up t o
scrubby aspen, o ak s, and r e d m aple, w ith
v e ry l i t t l e n a t u r a l second grow th o f th e
o r i g i n a l dom inant s p e c i e s .
The f o r e s t
r e m a i n s i n t a c t i n some p a r t s o f t h e
swamps, b u t t h i s la n d a l s o has b e e n p a r t l y
lo g g e d and d e s o l a t e d b y f i r e s .
Some o f
th e d ry sandy p l a i n s la n d i s c h a r a c t e r i z e d
b y a s c a t t e r e d g r o w t h o f j a c k p i n e , lo w
b l u e b e r r i e s (m ainly s p e c ie s o f V a c c in iu m ),
t h e sw e e t f e r n (Compto nZa a6pZznZ&oZZa) ,
and b ra c k e n , to g e th e r w i t h v a r io u s g r a s s e s ,
t h e m o s t common o f w h i c h a r e b l u e g r a s s
( P o a zompA.e.66 a ) , w i l d o a t g r a s s o r b u f f a l o
g r a s s (VanZkonZa s p .) , and b e a r d g r a s s
(A n d A . o p o g o n 6 z o p a A . Z u 6 ) .
I n t h e more open
swamps o r b o g s t h e r e i s a d e n s e c o v e r o f
h e a t h s h r u b s , such as b l u e b e r r i e s , C a ssa n d ra ,
and L ab ra d o r t e a
{ Lz dum gA.oznZandj.zum) , w i t h
some s c a t t e r e d b l a c k s p r u c e a n d t a m a r a c k .
I n p la c e s th e open m arsh i s covered w ith

8
various

se d g es and b l u e j o i n t (Catamag a o 6 t ^ 6 canade.n6.L6) .
I t is estim ated
t h a t a b o u t 10 p e r c e n t o f t h e c u t - o v e r
f o r e s t land has been c le a re d f o r farm ing
purposes.

At p r e s e n t ,

the s ta tu s

o f th e f o r e s t com m unities o f

the four c o u n tie s a re a is not th a t dism al.
n o way i m p l i e s , h o w e v e r ,

a retu rn

T h is in

to th e o r i g i n a l

d e n s e hardwood and c o n i f e r o u s f o r e s t s .

Today t h e s e

c o u n t i e s a r e a p a t c h w o r k o f s m a l l w o o d l o t s , so m e w h at
more e x t e n s i v e s e c o n d - and t h i r d - g r o w t h hardwood and
conifero u s f o r e s ts ,

farm land, b r a c k e n - g r a s s la n d s ,

a n d swamp f o r e s t s .

J a c k p in e r e p r e s e n t s an e x t e n ­

s iv e f o r e s t type in th e se c o u n t i e s , co v erin g a p p ro x i­
m a t e l y 20% o f t h e c o m m e r c i a l f o r e s t l a n d a r e a .
p resen t d is tr ib u tio n of jack p in e,

The

a ty p ic al p o st­

f i r e p io n e e r s p e c ie s , m ainly r e f l e c t s

its p ro life ratio n

f o llo w in g e a r l y p in e lo g g in g and s u b s e q u e n t w i l d f i r e s
( B e n z i e 1 9 77) .
cia tio n s

O th e r i m p o r t a n t f o r e s t t y p e s and a s s o ­

in th ese co u n ties

are o ak-hickory, n o rth e rn

w h ite c e d a r, m aple-beech-bassw ood,

D escrip tio n of S ite s
f o r Study

and aspen.

Included

A t o t a l o f 25 j a c k p i n e s i t e s w e r e u s e d f o r
d e ta ile d

study:

burned a re a s ,

8 u n b u r n e d c l e a r c u t s , 10 su m m er-

two 3 5 - y e a r - o l d s t a n d s ,

(55-years or g re a te r )

stands

and 4 m a tu re

(T able 2 .1 ) .

The 10

burned a re a s in c lu d e d 3 p r e s c r ib e d burns through

9

<13 r-*

01

'O ’•T

Li xi

H

O N sf

H

-T ^ n
i—I CM CM CM

i—*

O ' v£> r ^

*—1 cm r i

R] ^

u
C
01
MM e
D£
w
♦pH
tO rH
a> jd
>« to

a>ococor-^Ln<r
r^cor^r^r^r^f^r^
C'O'O'O'O'O'O'O'

0\
r%rin- in
^
r-

O' 00 vO
r*» r - r O' O' O'

0*» 0> O' O'

M O «A

\0 - t

f--, iO ro

i^3 vO \ f l <.0 *sO 'O

mm

r - .r - .r -

( 'I <<r a j n

• s t < T CM CM <M CM

O' O' O'

O 'O 'O ' O 'O ' O '

OO OO

i-i

(0
o>

u uu u
o o o o

*H ' r l *rl *H
U U M U

a.

cl a. cl

AJ
3
to
•
< x*
to

S IT E S

W
3

STUDY

3

3
to

W
^
to
-0 3
*
A J W - V ^
W
- W 3

u
yj
<u
o

r t f Jf»
W W A*
to co 3
-C O
UJ W A *
to to w

rM A *
a w
^
z g
g

3
5s
*
A J
M3

ti] W W ^ f *
3
3
-C O
.
* 0
O 'l N H m

* S
■
‘
*
*
- < m -j *
- r — r — *n
O N H < ^ n h - H H C S

u

3

*o
c
CO

A *

rM A *
S
- v to
3
K A J
A*
- W

OF
SUMMARY

m
CM

3
to

‘

u
u
u
u
u
u
u
o
< !> < L ia J Q J Q ItIld jO J
t f i t n t n u i u i v i t / i v i

a
qj

<n

u
qj
<0

u
qj

tf l

rH go r CM CM CM

u
<u
<fl

0
u
H» «
ID 0 )

u
Of
01

u
01
<Q
-h
-V
U1
CO
-A *
W

HI
CO

w
CO
-

A»
W

CO t O 3
«
*
«»
M3 r H M3
CM c n CM
0
0)
m

U)
3
CO

u
u
<U QJ
cn t o

A * rtf
W Ul A*
to 3
3
a » a *
*
- 3
*
w
z a w
^
- to 3
3
- t f rtf
- to
3
3
CO U 3 r) f *
*
*
-W
cm r - c - O '
CM H r H H r H

u
a
(j
QJ QJ a
CO to t o

u
u
u
0J O
OJ
CO CO t o

rH

<0
o:
QJ

3
3
U 3
W U I3
3 *
H N H H N N
C N N
DC DC C C D C DC 0 C D C ClC

3
u
s
n i
r H CM m CM
gc a i p i o s

w w
w
H H W
a s c tf a s

HI 3
3
r H CM CM
H i oc p :

w
H

3
3 ^ 3
3 * 3
3 ' 3
3
tN « n c o s O is N r N ( N

3 3
3 3
r-* r - t n < r
CM CM CM CM

3 "
-X
CM
H

3
< r
CM
H

3
- r
CM
£h

p

u a a w
a
H ( N CM p H n

H i PS OS P S p S Od

H-l

cm cm cm

H

h

h

cm cm cm c m

H

H

H

H

cm

H

H

H

H

H

3 *
■‘3 “
CM
H

3 "
-h t
CM
H

QJ
S’H
D
P
3
0
CJ

c
0
■rH
4J
<u (d
u P
be
tO »r4
CD
CJ
•O

V)
u
p
CJ
Li
ctj
<11
•—I
O
•O
CJ
p
Li
D

T3*0 TJ
Q0
fi 6
U Li
M
O O <0 O rd ra g g
UM4H tMT3 *o 0 O
a a O anj O 0 u U
(J a cn V)
<u Cfl U
Li Li V) U CDCD O 0
U U 0 CJ O O ad oc

pH

CO

O
CO
C < SQ
—H H
3 1

fH pH r i
00 00 00
4 t
1
O O O
CO CO GO
1 1 1
O 'O 'O '
r-* r-* r -< cq < <
CM c n c i <t i n

T3 *o
M D
0 cd 0 a
T3 LL-i to
a 0 a 6
<d u <u QJ
Li in M 00
U O CJ 0
cn
4J
D
O
Li
<n
Q>
pH
U
T3
<U
e
Li
3
PQ

p

«rH

au m
a <aa
h : <
a E E E
cO
0)
*r

T3

QJ

Q)

00 tiO 0 0
O O O

3 3
V D CM
CM CM
H H

P
T3 06
Li
aoj m0 g
O
b a O
QJ RJ (A

rH rH
rH 0 0 CO
00 1 1
t 0 0
rH O 0 0 OO
0 0 OO » |
1 1 d 'O '
o O 'f N h ,
CO f~->
EQ
0 0 0 0
r - r - r H r*H

3
—r
CM
H

3
< r
CM
H

3
in
CM
H

'O

Li
0
h
cti U
a

B

QJ

a

cd

00 U
O O c d PS 0 u

<n
TO

td

G
nj

a>

4J
Ul

>

rH
00
1
rH O
*13 OO OO
1
QJ
C O o>

1

m
ro

rH
CO
1
O
OO

1

O'
r-< A <
r-* O' O '

M 30

ca

3
< r
CM
H

c n
0□ g0
a ra
a g |3
n3
0 O
GB y U
QJ QJ <n cn
00 00 0 O

00 Ll O
O O Od

rH
0
1
U

3
- r
CM
H

rH r H
<n 30 CO
<u 1 1

Ll 0 0
»rl CO GO
UH 1 1
TJ O ' O'
rH r - r < tf> rH
s 0 0 a \ rH

QJ

a

*rH
PH

a < fO < 03 a Q
<d CM cm m m c n c n
* n pH rH pH i-H r—1 rH

10
stan d in g 3 5 -y ear-o ld ja c k p in e,
through c le a rc u t ja c k p in e,
jack p in e,
p ine.

and 1 w i l d f i r e

4 p re s c r ib e d burns

2 w ild fires

th ro u g h m ature

th ro u g h c le a r c u t m ature ja c k

A ls o i n c l u d e d was one s i t e u s e d f o r a c o n t r o l l e d

b u r n from w h ic h e a r l y s u c c e s s i o n on b u r n e d and u n b u r n e d
c l e a r c u t b l o c k s was m o n i t o r e d d u r i n g t h r e e c o n s e c u t i v e
seasons.

A ll unburned c l e a r c u t s w ere p r e v i o u s l y m ature

j a c k p i n e g r e a t e r t h a n 50 y e a r s o l d .
A l l s i t e s u s e d f o r s tu d y w ere l o c a t e d on th e
G ra y lin g sand s e r i e s , w ith as l i t t l e

v a r ia tio n in

to p o g ra p h y and o t h e r e n v iro n m e n ta l f a c t o r s
The f o l l o w i n g d i s c u s s i o n w i l l o u t l i n e

as p o s s i b l e .

the s i l v i c u l t u r a l

management and n a t u r a l d i s t u r b a n c e h i s t o r y o f e a c h
site.

U nique p h y s i c a l f e a t u r e s o r a s p e c t s o f t h e

a d ja c e n t v e g e t a t i o n w i l l a ls o be in c lu d e d .
The s t u d y s i t e s

a r e p r e s e n t l y b e i n g managed by

th e M ichigan D epartm ent o f N a tu r a l
and w ere a c q u i r e d i n t h e

1930's

Resources

or th e r e a f te r ;

(DNR),
no s i t e

h is to ry d ata are a v a ila b le b efo re th e ir a c q u is itio n .
T herefore,
for s ite s

stand o rig in in fo rm atio n is not a v a ila b le
50 y e a r s o r o l d e r .

p l a n t i n g s were c o n d u cted p r i o r

H o w e v e r , no w i d e s c a l e
t o 1930,

w i l d f i r e s w e r e common i n t h i s r e g i o n .

and f r e q u e n t
T h e r e f o r e , most

f o r e s t m anagers ag ree t h a t m atu re ja c k p in e sta n d s
of th is

a r e a o f M ichigan a re o f p o s t f i r e

w i l l be assumed u n l e s s o t h e r w i s e n o te d .

o rig in ;

th is

11
S t a n d age o f b u r n e d s t a n d i n g t i m b e r was d e t e r ­
m in ed by b o r i n g a t l e a s t

10 t r e e s

i n an a d j a c e n t

sta n d th a t appeared r e p r e s e n ta t iv e o f th e tr e a te d
a re a b efo re burning.

O v e rsto ry b a s a l a re a i n m ature

a n d 3 5 - y e a r - o l d j a c k p i n e s t a n d s was m e a s u r e d u s i n g
a 1 0 - f a c t o r p r i s m a t 10 r a n d o m l o c a t i o n s .
c learcu ts

Z ero-year

are d e fin e d as a re a s h a rv e ste d th e w in te r

a n d s p r i n g b e f o r e summer s a m p l i n g ;

1 -y e a r-o ld burns

a r e d e f i n e d a s a r e a s b u r n e d t h e summer b e f o r e sam­
p lin g ;

n o a r e a s w e r e s u r v e y e d t h e same g r o w i n g s e a s o n

th e y w ere b u rn ed .
Unburned c l e a r c u t s *
S ite

1A.

and a d ja c e n t
d uring the

T his a r e a i s n o r t h o f a t r a i l ro ad

(w est)

to r o u te F-97.

s p r i n g o f 1979,

a t w h ich tim e t h e dom inant

j a c k p i n e w a s 55 t o 6 0 - y e a r s - o l d .
r e d pine

I t was c l e a r c u t

(F inus r e s i n o s a ) w ere l e f t

W idely s c a t t e r e d
stan d in g .

T his

a r e a was s u r v e y e d a s a 0 - y e a r c l e a r c u t on 7 / 1 7 / 7 9
and l a t e r

t h a t d a y t h e a r e a was p r e s c r i b e b u r n e d .

D u r i n g t h e 1980 a n d 19 81 f i e l d

seasons th is

a r e a was

r e s u r v e y e d as a 1 - y e a r - o l d and 2 - y e a r - o l d b u r n
7C 80-81).

(site

No o t h e r d i s t u r b a n c e i n f o r m a t i o n i s known

*The n u m b e r 7 9 , 8 0 , a n d 81 f o l l o w i n g t h e a l p h a n u m e r ic a l s i t e d e s i g n a t i o n r e f e r s to th e 1979, 1980,
a n d / o r 19 81 g r o w i n g s e a s o n s when t h e s i t e s w e r e s u r ­
v e y e d , f o r s i t e s s u r v e y e d i n more t h a n one g ro w in g
season.

12
for th is

site.

S ite

1B8Q-81.

a larg er clearcu t

T h is a r e a s u r v e y e d was p a r t o f

lo cated a t th e so u th east ju n ctio n

o f 4 - M ile Road a n d S t a l e y Lake Road.
o l d j a c k p i n e on t h i s

The 5 7 - y e a r -

s i t e was c l e a r c u t d u r i n g t h e

w in te r and s p r in g o f 1980,

and t h e s i t e was s u r v e y e d

i n 19 80 a n d 1981 a s a 0 - y e a r - o l d a n d 1 - y e a r - o l d c l e a r cu t.

S c a tte re d in te r m e d ia te s iz e d oak (n o rth e rn

r e d g ro u p , Q uercus s p p .)

and b l a c k c h e r r y

s e r o t i n a ) w ere p r e s e n t .

No o t h e r p r e v i o u s d i s t u r ­

bance to

th is

S ite

2.

stand is

(Prunus

known.

I n 1 9 6 1 a n d 19 74 t h i s

tiv e ly cut fo r jac k pine

a r e a was s e l e c ­

l o g s e x c e e d i n g 20 cm d . b . h .

I n 19 7 8 t h e 55 t o 6 0 - y e a r - o l d j a c k p i n e was c l e a r c u t
a n d t h e s i t e w a s s u r v e y e d i n 1979 a s a 1 - y e a r - o l d
cle arc u t.

Subsequent to th a t survey,

p l a n t e d and n o t u se d f o r f u r t h e r

t h e s i t e was

study.

S cattered

3- t o 5 - m - t a l l j a c k p i n e and m a tu re r e d p i n e w ere
le ft

stan d in g a f t e r h a rv e stin g .

u n d u latin g ,

T h i s s i t e was g e n t l y

as opposed to n e a r t o t a l f l a t n e s s

o f m ost

o f the o th e r s i t e s .
S ite

3A.

T he s o u t h e r n b o r d e r o f t h i s a r e a l i e s

n o r t h o f 4 -M ile Road.

The s i t e was c l e a r c u t i n 1 9 7 8 ,

a n d s u r v e y e d i n 19 80 a s a 2 - y e a r - o l d c l e a r c u t .

The

a g e o f t h e d o m i n a n t s r a n g e d f r o m 48 t o 57 a t t h e t i m e
of c le a r c u ttin g .

S c a tt e r e d i n t e r m e d i a t e - s i z e d oaks

13
and b l a c k c h e r r y ,

a l o n g w i t h s u p p r e s s e d 3 - t o 5-m

j a c k p in e w ere p r e s e n t .
S ite

3B 79-80-81.

o f M u s k ra t Lake Road.
on t h i s

old c le a rc u t.

d ir e c tly w est

T h e 50 t o 5 5 - y e a r - o l d j a c k p i n e

s i t e w ere c l e a r c u t i n 1977,

v ey ed i n 1979, 1980,

tiv ely

T his a r e a l i e s

t h e s i t e was s u r ­

a n d 1981 a s a 2 - ,

I n 19 6 1 a n d 1 9 6 4 ,

3-,

t h e s i t e was s e l e c ­

logged f o r ja c k ^ p in e lo g s .

An o v e r g r o w n t r a i l

ro a d ra n through the c e n te r o f the s i t e .
o f 7 t o 10 m t a l ' l a s p e n

and 4 - y e a r -

A % ha c lo n e

( P o p u lu s t r e m u l o i d e s ) was a l s o

l o c a te d i n th e c e n te r o f the s i t e .
S ite

4A79-8Q -81.

e a s t o f R i c h a r d s o n Road.

T h is a re a i s

I n 1957 t h e r e was s e l e c t i v e

c u t t i n g f o r ja c k p in e pulp lo g s ,
was c l e a r c u t .

found d i r e c t l y

a n d i n 1976 t h e a r e a

At t h e tim e o f h a r v e s t i n g ,

t h e d o m i n a n t s r a n g e d f r o m 54 t o 7 2 y e a r s .
1980, and 1981 t h i s

th e age of
I n 1979,

a r e a was s u r v e y e d a s a 3 - ,

and 5 - y e a r - o l d c l e a r c u t .

4-,

S c a tte re d sa p lin g sized jack

and r e d p i n e and a few m a t u r e r e d p i n e w e re f o u n d .
T o p o g ra p h y o f t h e s i t e was g e n t l y r o l l i n g ,
tic

ch aracteris­

of a t i l l p la in .
S ite

5A79-80-81.

o f A i r p o r t Road,

T his a re a l i e s

d ir e c tly n o rth

and i s p a r t o f a l a r g e r r e s e a r c h

a r e a d i r e c t e d b y t h e U. S . F o r e s t S e r v i c e .

T h is a re a

w a s c l e a r c u t i n 1 9 7 5 , w i t h no p r e v i o u s d i s t u r b a n c e
to the sta n d r e p o rte d .

T he d o m i n a n t s r a n g e d f r o m 57

14
t o 66 y e a r s a t t h e t i m e o f c u t t i n g .

The a r e a was

s u r v e y e d i n 1 979, 1 9 8 0 , and 1981 a s a 4 - ,
6 -y ear-o ld c le a rc u t.

5-,

and

R e g u l a r l y s p a c e d d e n se oak and

c h e r r y s p ro u ts w ere found th ro u g h o u t.
S ite
Road.

6.

T his a re a l i e s

d irec tly

s o u t h o f Emery

No o t h e r d i s t u r b a n c e t o t h e s t a n d w a s r e p o r t e d

p rio r to c le a rc u ttin g

i n 1974.

Dom inant j a c k p i n e

r a n g e d f r o m 57 t o 63 y e a r s a t t h e

tim e o f h a r v e s t .

Dense oak and c h e r r y s p r o u t s w ere fo u n d th r o u g h o u t
the a re a ,

a n d t h i c k s t a n d s o f 20 m t a l l j a c k p i n e

surrounded t h is

site

on t h e e a s t , w e s t ,

and s o u th

borders.
Burned a r e a s
S ite
r o a d 13,

7A 80-81.

p in e,

lie s

d i r e c t l y w est of co un ty

and i s p a r t o f a l a r g e r e s e a r c h a r e a d i r e c t e d

b y t h e M i c h i g a n DNR.
th is

T h is

A larg e w ild fire

e n t i r e r e s e a r c h a r e a i n 1946.

swept th ro u g h

Some o a k ,

and j a c k p in e s u r v iv e d th e f i r e ;

red

those i n d iv i ­

d u a l s o f f i n a n c i a l m a t u r i t y w e r e l o g g e d i n 19 75 a n d
1976.

What r e m a i n e d w a s a v a r i a b l e

ja c k pine

age and d e n s i t y

stand, re p re s e n tin g p o s t f i r e re g e n e ra tio n

a n d e m e r g e n t s t h a t s u r v i v e d t h e 1946 f i r e

and sub-

s

sequent lo g ging.

The e m e r g e n t s r a n g e d f r o m 4 0 - t o

4 5 - y e a r s - o l d and i n t e r m e d i a t e s from 22- t o 3 5 - y e a r s old.

However,

t h e r e a l s o were p r e s e n t a s u b s t a n t i a l

number o f j a c k p i n e

i n th e 0-5 y e a r ,

5-10 y e a r ,

and

15
10-20 y e a r age c l a s s e s .
On 7 / 1 2 / 7 9 t h e s i t e w a s p r e s c r i b e b u r n e d w i t h
resid u al trees

left

stan d in g .

enough to k i l l n e a r l y a l l
parts

o f t h e hardw oods.

T h e f i r e was s e v e r e

th e p i n e s and a b o v e -g ro u n d
The h ard w o o d s h a v e s u b s e ­

q u e n t l y s p r o u t e d from t h e b a s e o r p r o d u c e d r o o t
suckers.

P a t c h y and s c a t t e r e d i n t e r m e d i a t e - s i z e d sn a g s

are d is tr ib u te d

throughout th e s i t e .

The a r e a was

s u r v e y e d i n 1980 a n d 1 9 8 1 a s a 1 - y e a r a n d 2 - y e a r - o l d
burn.

Burned s i t e s

7B79-80-81 and 9A79-80-81 and

3 5 - y e a r - o l d j a c k p i n e s t a n d s 12A a n d 12B a r e p a r t o f
th is

same r e s e a r c h a r e a .
S ite

7B79-80-81.

b u r n e d on 8 / 8 / 7 8 .

T h i s s i t e was p r e s c r i b e

See s i t e

7A80 -81 f o r s t a n d h i s t o r y .

A unique a s p e c t o f t h i s

site

the n o rth e a s t p o rtio n .

The s i t e

the bog, bu t i s

i s a 2 ha h e a t h bog i n
g e n t l y s lo p e s tow ards

l e v e l on t h e u p l a n d a r e a s .

O nly th e

l e v e l u p l a n d a r e a s away f r o m t h e b o g w e r e s u r v e y e d .
P a tc h y and s c a t t e r e d i n t e r m e d i a t e - s i z e d
found t h r o u g h o u t.
1980,

sn ag s were

T h e s i t e w a s s u r v e y e d i n 197 9,

a n d 19 8 1 a s a 1 - y e a r ,

2-year,

and 3 - y e a r - o l d

p r e s c r ib e d burn.
S ite
on 8 / 1 7 / 7 6 .
site

is

9A79-80-81.
See s i t e

T h i s a r e a was p r e s c r i b e b u r n e d
7A80 -8 1 f o r s t a n d h i s t o r y .

T his

somewhat u n i q u e due t o t h e l a r g e number o f

oak, b la c k c h e rry ,

and r e d and j a c k p i n e

th a t survived

16
the f i r e .

The r e m a i n i n g c a n o p y ,

alth o u g h p atch y ,

c r e a t e d an u n u s u a l m i c r o h a b i t a t f o r a r e c e n t l y
burned s i t e .

Surveys took p la c e

in 1979,

1980,

and

19 81 a s a 3 - ,

A - , and 5 - y e a r - o l d p r e s c r i b e d b u r n ,

resp ectiv ely .
S ite

7C8081.

year c learcu t

T h i s s i t e was s u r v e y e d a s a 0-

i n 1979 and a 1 - y e a r and 2 - y e a r - o l d

p r e s c r i b e d b u r n i n 1980 a n d 1 9 8 1 .

See u n b u r n e d s i t e

1A f o r s i t e h i s t o r y .
S ite

7D79-80-81.

T his a re a i s

lo cated d ir e c tly

s o u th o f Penn Road, and o r i g i n a t e d a s a j a c k p i n e
p lan tatio n

(80 0 t r e e s / a c r e ) , e s t a b l i s h e d

The a r e a w a s c l e a r c u t f r o m t h e
s p r i n g 1978.

s p r i n g o f 1976 t o

How ever, n e a r l y a l l h a r v e s t i n g was done

i n 197 7 , a n d t h i s
On 9 / 2 6 / 7 8 ,

i n 1930.

is

used as th e r e f e r e n c e d a te .

t h e a r e a w a s p r e s c r i b e d b u r n e d a s a 1-

y ear-o ld c le a rc u t.

S u r v e y s w e r e m ade i n 1 9 7 9 ,

a n d 19 81 a s a 1 - y e a r ,
sc rib e d burn.

2-year,

1980,

and 3 - y e a r - o l d p r e ­

The r e c o r d e d stu m p a g e a t t h e t i m e o f

c l e a r c u t t i n g r a n g e d f r o m 31 t o A9 y e a r s , w i t h m o s t
a g e s f a l l i n g b e t w e e n A0 a n d A7 y e a r s .

T h is a re a i s

s u r r o u n d e d by d e n s e s t a n d s o f m ixed r e d and j a c k p i n e
and a l a r g e c l e a r c u t to th e n o r t h and w e s t.

The

m i x e d r e d a n d j a c k p i n e s t a n d s h a d a much s p a r s e r
u n d e rsto ry than u su a lly
blueberry

seen in pure ja c k pine

stands;

(V a c c in iu m s p p . ) was c o n s p i c u o u s l y r e d u c e d .

17
S ite
fire

8A79-80-81.

on 7 / 2 / 7 7 ,

o f R o ll Road.

T h is a r e a was b u rn e d by w i l d ­

and i s

l o c a te d d i r e c t l y e a s t and w e st

The f o r e s t a t t h e t i m e o f b u r n i n g was

a w e l l s to c k e d j a c k p in e s t a n d , w i t h dom inants r a n g ­
i n g f r o m 40 t o 52 y e a r s - o l d .
w ere l e f t
d en sity ,

Most o f t h e i n d i v i d u a l s

s ta n d in g as c h a rre d sn ag s.

Because o f t h e i r

they g r e a t ly in c r e a s e d th e shading to

f o r e s t f l o o r com pared t o o t h e r b u rn e d a r e a s .
to p o g rap h y i s
areas.

g en tly r o l l i n g

3-year,

The

slopes w ith la rg e r f l a t

S urveys w ere co n d u cted in 1979,

1 98 1 a s a 2 - y e a r ,

the

1980,

and

and 4 - y e a r - o ld b u rn , r e s ­

p ectiv ely .
S ite
on 8 / 2 3 / 7 6 ,

9B79-8Q-81.
is

T h is a r e a , b u rn ed by w i l d f i r e

l o c a t e d d i r e c t l y n o r t h o f 4 -M ile Road.

T h e a g e o f t h e d o m i n a n t s a t t h e t i m e o f b u r n i n g was
h ig h ly v a ria b le ,

r a n g i n g f r o m 48 t o 66 y e a r s .

a r e a was a p p a r e n t l y w e l l - s t o c k e d ,

as in d ic a te d by the

r e m a i n i n g c h a r r e d s n a g s and s tu m p s ,
d e n s ity o f th e a d ja c e n t ja c k pine
to p o g r a p h y was g e n t l y r o l l i n g
was s u r v e y e d i n 1 9 7 9 ,

1980,

The

and th e h ig h

stands.

throu gh ou t.

The
T his a r e a

a n d 1981 a s a 3 - y e a r ,

4-

y e a r , and 5 - y e a r - o l d b u rn .
S ite

10A79-80-81.

of the r a i l r o a d
Road.

T h is a re a i s

lo c a te d south

tra c k s running a d ja c e n t to

5-M ile

The s i t e w a s c l e a r c u t i n t h e w i n t e r a n d s p r i n g

o f 1 97 3 a n d p r e s c r i b e b u r n e d a s a 2 - y e a r c l e a r c u t o n

18
10/7/75.

At t h e tim e o f c l e a r c u t t i n g

th e age o f

t h e d o m i n a n t s r a n g e d f r o m 42 t o 63 y e a r s .
is

The a r e a

s u rro u n d e d on t h r e e b o r d e r s by w e l l - s t o c k e d m atu re

ja c k p ine.

Hardwoods w e re a p p a r e n t l y an i m p o r t a n t

component o f th e p r e - e x i s t i n g v e g e t a t i o n ,

as in d ic a te d

by t h e i r c u r r e n t p r e s e n c e as d e n se s p r o u t s and s c a t t e r e d
su rv iv in g a d u lts.

T he a r e a w a s s u r v e y e d i n 1 9 7 9 ,

a n d 1981 a s a 4 - y e a r ,

5-year,

1980,

and 6 - y e a r - o l d b u r n ,

resp ectiv ely .
S ite

10B79-8Q-81.

e a s t o f F a i r v i e w Road.

T h is a re a is

lo cated d ir e c tly

I t w a s c l e a r c u t i n 1 97 4 a n d

b u rn e d as a 1 - y e a r - o l d c l e a r c u t on 8 /1 5 /7 5 .
to

th is,

t h e a r e a was s e l e c t i v e l y l o g g e d f o r m a t u r e

ja c k pine
1950,

Previous

( e x c e e d i n g 20 cm d . b . h . )

1956,

1958,

and 1965.

i n 1941,

1948,

The age o f t h e d o m in a n ts

a t t h e t i m e o f c l e a r c u t t i n g r a n g e d f r o m 44 t o 54 y e a r s .
T h i s a r e a was s u r r o u n d e d b y w e l l - s t o c k e d m a t u r e j a c k
p i n e w i t h a c a n o p y h e i g h t o f 15 t o 20 m.
o f d e n s e hardw ood s p r o u t s

(m ainly aspen s u c k e rs )

to 4 .5 m t a l l dom inated th e s i t e .
i n 1979,

Large a re a s
up

I t was s u r v e y e d

1 9 8 0 , a n d 1 9 81 a s a 4 - y e a r ,

5-year,

and 6-

y e a r-o ld burn.
S ite

11.

T h is a re a i s

to th e so u th w e st of a

t r a i l r o a d t h a t r u n s s o u t h e a s t o f Emery R o a d .
clearcu t

I t was

i n t h e w i n t e r a n d s p r i n g o f 1972 a n d 1973

and on 7 / 2 1 / 7 3 b u r n e d by w i l d f i r e .

The age o f t h e

19
d o m i n a n t s a t t h e t i m e o f c u t t i n g r a n g e d f r o m 51 t o
60 y e a r s .

Oak was a n i m p o r t a n t c o m p o n e n t o f t h e p r e ­

e x is tin g v eg etatio n ,
sprouts

as e x h i b i t e d by num erous d en se

thro u g h o u t th e a re a .

ja c k p in e b o rd er the s i t e .

M ature w e l l - s t o c k e d
The a r e a w a s s u r v e y e d

i n 1979 a s a 6 - y e a r b u r n .
E xp erim en tal burn s i t e .
T24N, R2W, s e c 8 ,

S% o f SE%.

Roscommon C o u n t y ,
T h is 7 .2 ha a r e a i s

l o c a t e d d i r e c t l y w e st o f A i r p o r t Road.
of th is

site

s t a r t e d i n 1976 a n d c o n t i n u e d t h r o u g h

t h e f a l l and w i n t e r o f 1977.
to o k p l a c e i n 1976,
d ate.

C learcu ttin g

and t h i s

When c l e a r c u t ,

f r o m 55 t o 65 y e a r s .

Most o f t h e h a r v e s t i n g
i s used as the r e fe re n c e

t h e age o f t h e d o m in a n ts ra n g e d
T his a r e a ,

used to

study e a r ly

p l a n t s u c c e s s i o n on b u r n e d a n d u n b u r n e d c l e a r c u t
b l o c k s , was m o n i t o r e d d u r i n g t h e 1979,
grow ing s e a s o n s

1980,

(see C hapter 5 f o r d e t a i l s

a n d 1981

o f the

stu d y ).
Interm ed iate-ag e
_j a c k p i n e s t a n d s
B oth s t a n d s i n t h i s
19 46 w i l d f i r e

c a te g o r y r e s u l t e d from a

i n Ogemaw C o u n t y .

d i s c u s s e d above (7A80-81,

Three stu d y s i t e s

7B79-80-81,

and 9A79-80-

8 1 ) a l s o o r i g i n a t e d f r o m t h e same f i r e .

A lthough

th e stand h is to r y d e scrib e d in d e t a i l fo r s i t e
7A80-81 a p p l i e s

to th ese

stan d s,

th e y were n o t

20
p r e s c r ib e burned.
S ite

12 A.

T his a r e a o f 3 5 - y e a r - o ld j a c k p in e

was s u r v e y e d i n 1981.

T he j a c k p i n e d i s t r i b u t i o n

was e x t r e m e l y v a r i a b l e ,
p atch es

r a n g i n g from im p e n e tr a b le

to i n t e r m e d i a t e - s i z e d openin g s devo id of t r e e s .

The m e a n b a s a l a r e a o f 6 . 0 m2/ h a , w i t h a s t a n d a r d d e v i a ­
tio n

(S .D .)

reflects

o f 7 . 4 6 m2 ( r a n g i n g f r o m 0 t o 2 7 . 6 s q . m ) ,

the v a r i a b i l i t y .

g rea tly affected

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

the d i s t r i b u t i o n

the f o r e s t flo o r.

of lig h t reach in g

The o p e n a r e a s a p p e a r e d h o t a n d

d ry and t h e sh a d e d a r e a s c o o l and m o i s t .

The open

a r e a s w ere dom inated by low -grow ing g r a s s e s ,

sedges,

and f o r b s , and t h e sh a d ed u n d e r s t o r y was d o m in a ted
by s m a l l t r e e s

and s h r u b s .

d is tr ib u tio n of ja c k p ine,
extrem ely v a r i a b le .

A long w i t h t h e p a t c h y
t h e c a n o p y h e i g h t was

T h is r e f l e c t e d

th e w ide ra n g e

o f age c l a s s e s r e p r e s e n t e d h e r e .
S ite

12B.

T h i s a r e a was a l s o s u r v e y e d i n 1981.

The s t a n d s t r u c t u r e w a s s i m i l a r t o

th at of s ite

h o w e v e r , m ean b a s a l a r e a w a s s o m e w h a t h i g h e r
h a ) , w i t h somewhat l e s s v a r i a b i l i t y
r a n g i n g from 0 t o

(S.D .

12;

( 9 . 7 m2/

= 6 . 2 m2 ,

16.1 s q . m ).

M ature j a c k p i n e s t a n d s
The m a t u r e j a c k p i n e s t a n d s

chosen f o r study

w e r e 55 y e a r s o r o l d e r a n d a p p e a r e d f r e e
d istu rb an ce.

from r e c e n t

These m a tu re s ta n d s were w e l l - s t o c k e d

21
w ith e v e n ly d i s t r i b u t e d dom inants.
fa irly reg u lar,

but l i g h t gaps,

b l o w d o w n , was a common f e a t u r e .
d o m in a n ts on a l l

sites

T he c a n o p y was

c r e a t e d from r e c e n t
The h e i g h t o f t h e

r a n g e d f r o m 15 t o 20 m.

In ter­

m e d ia te and s a p l i n g - s i z e d i n d i v i d u a l s w ere g e n e r a l l y
p resen t.
S ite

13A.

T his a r e a i s

d i r e c t l y so u th o f the

e x p e r i m e n t a l b u rn a r e a o f f o f A i r p o r t Road.
m ea n b a s a l a r e a w a s 1 7 . 0 m2/ h a

(S.D .

s t a n d a g e w a s c o n s i d e r e d 68 y e a r s ;
f r o m 55 t o
S ite

The

=4.4).

The

dom inants ra n g e d

68 y e a r s .
13B.

T his a r e a i s

o f A i r p o r t Roa d a n d i s

situ a te d d ire c tly north

l o c a t e d 1 . 6 km s o u t h o f s i t e

T h e m ea n b a s a l a r e a w a s 1 5 . 6 m2 / h a

(S.D . = 3 . 9 ) .

d o m i n a n t s r a n g e d i n a g e f r o m 55 t o 68 y e a r s ;

site

a n d s i t e 13A o r i g i n a t e d

b a s e d on t h e i r c l o s e p r o x i m i t y ,

The

a s on

s i t e 13A t h e s t a n d a g e w a s c o n s i d e r e d 68 y e a r s .
probably t h is

13A

M o st

from f i r e ,

age o f d o m in a n ts ,

and

stand s tru c tu re .
S ite

13 C ,

T his a r e a i s

o f c o u n ty r o a d 26.
(S.D .

f r o m 47 t o 55 y e a r s ,

The a g e o f t h e d o m i n a n t s r a n g e d
s t a n d a g e w a s c o n s i d e r e d 55 y e a r s .

The s i t e w a s s e l e c t i v e l y

1949,

south

The mean b a s a l a r e a was 2 2 . 1 m2/ h a

= 3.9 sq. m ).

t h a n 20 cm d . b . h .

lo cated d ir e c tly

logged

(diam eter c u ts g re a te r

as w e l l as s a lv a g e c u ts )

1956, and 1961.

i n 1946,

Twenty y e a r s l a p s e d b e tw e e n

22
th e l a s t c u t t i n g and th e p r e s e n t s u rv e y o f th e s i t e .
T his a p p a re n tly v ig o ro u s ,

h e a l t h y s t a n d i n n o way

a p p e a re d d i s t u r b e d or d e g ra d e d from lo g g in g .
S ite

13D.

T his s ta n d ,

d i r e c t l y e a s t of S taley

L a k e R o a d , was l e f t f r o m a l a r g e a r e a t h a t was c l e a r c u t i n 1980 ( s e e s i t e
w a s 1 9 . 8 m 2/ h a

(S.D .

1B 80-81).
= A .3 ).

r a n g e d f r o m h i t o 57 y e a r s ;
to be 5 7 - y e a r s - o l d .

The m e a n b a s a l a r e a

The age o f t h e d o m in a n ts
t h e s t a n d was c o n s i d e r e d

CHAPTER I I I
EARLY REVEGETATION OF BURNED AND
UNBURNED JACK PINE CLEARCUTS IN
NORTHERN LOWER MICHIGAN
INTRODUCTION
The i m p o r t a n c e o f f i r e
and i t s

a s an e c o l o g i c a l f a c t o r

u se as a management t o o l i n

t h e Lake S t a t e s

i s w e l l docum ented (Eyre and L eB arro n 1944, C a y fo rd
1970, Vogl 1970, H einselm an

1973, A hlgren 1974).

A

l a r g e p o r t i o n o f t h i s p r e v i o u s w ork f o c u s e s on t h e
e c o lo g y and management o f j a c k p i n e
L am b.).

Jack pine is a ty p ic a l p o s t f i r e p io n eer

s p e c i e s whose p r e s e n t d i s t r i b u t i o n
te d to w i l d f i r e s
1977).

(P i n u s b a n k s i a n a

cones and r e q u i r e s

optimum g e r m i n a t i o n .
manage t h i s

d irectly re la ­

fo llo w in g e a r l y pine lo g ging

Throughout m ost o f i t s

sero tin o u s

is

(B enzie

ra n g e , ja c k p in e produces
a m in e ra l seedbed fo r

T h e re fo re , m ost a tte m p ts

to

s p e c ie s have u t i l i z e d p r e s c r i b e d b u rn in g .

S tu d ie s o f p r e s c r i b e d b u rn in g have m ain ly fo cu sed
on s l a s h r e d u c t i o n ,
of seed lin g s

seedbed p r e p a r a t i o n ,

and r e s p o n s e

to v a rio u s e n v iro n m e n ta l f a c t o r s

and L eB arron 1944, B e a u f a i t 1962).

L im ited d e ta ile d

w o r k , h o w e v e r, h a s b e e n c o n d u c t e d on p l a n t
on j a c k p in e

sites

(Eyre

su ccessio n

fo llo w in g log ging a n d /o r burning

23

24
i n t h e Lake S t a t e s

(A h lg ren 1960, Vogl 1970, K r e f t i n g

a n d A h l g r e n 1 9 7 4 , Ohmann a n d G r i g a l 197 9 ) .

Thus a

s t u d y was u n d e r t a k e n t o c h a r a c t e r i z e e a r l y r e v e g e t a ­
t i o n o f burn ed and unburned c l e a r c u t s
M ich ig an w here j a c k p in e i s

i n n o r t h e r n low er

a p re d o m in a n t component

o f th e t r e e v e g e t a t i o n on sandy s o i l s .
tiv e of th is

chapter is

T he m a i n o b j e c ­

to d isc u ss the d iffe re n c e s

betw een b u rn e d v e rs u s unburned j a c k p in e c l e a r c u t s
and th e v a r i a t i o n

th at r e s u lts

from t h e s e d i s t u r b a n c e s .

A d i s c u s s i o n o f s u c c e s s i o n a l p a t t e r n s w i t h i n and betw een
b u rn e d and u n b u rn ed se q u en c es w i l l be p r e s e n t e d i n
C h a p t e r 4.

METHODS
Unburned c l e a r c u t s i t e s
3B 79-80-81,

1A,

1B80-81,

4 A 7 9 -8 0 -8 1 , 5A79-80-81 and 6,

clearcu t s ite s

7C79-80-81,

7D 79-80-81,

2,

3A,

and b u rn e d

10A79-80-81,

a n d 10B 7 9 - 8 0 - 8 1 w e r e u s e d f o r c o m p a r i s o n i n t h i s
(se e C hap ter 2 f o r s i t e
a l l v ascular p la n ts,
herbs,

shrubs,

d escrip tio n s).

in clu d in g g rasse s,

chapter

On e a c h s i t e
sedges,

ferns,

and t r e e s were c h a r a c t e r i z e d by fre q u e n c y

and co v er m easurem ents.

F re q u en c y was d e te r m in e d u s i n g

a i m 2 c i r c u l a r p l o t random ized alo n g t r a n s e c t s .

T ran­

s e c t s were o r i e n t e d to b e s t in c lu d e th e e n t i r e a r e a
to be su rv e y e d .

C o v e r d e t e r m i n a t i o n s w e r e made b y

summing t h e d i s t a n c e

i n t e r s e c t e d by e a c h s p e c i e s a l o n g

25
r a n d o m l y p l a c e d 20 m t r a n s e c t s .
and 3 c o v e r t r a n s e c t s
te r iz e each s i t e .
on s i t e

T h irty frequency p lo ts

(6 0 m t o t a l )

were used to c h a r a c ­

D e v i a t i o n from t h i s m ethod o c c u r r e d

1A w h e r e , d u e t o t i m e c o n s t r a i n t s , o n l y 20 f r e ­

quency p l o t s

a n d AO m c o v e r w e r e i n c l u d e d .

s i t e s a 10% o r l e s s

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

r e s u l t e d f r o m t h e f i n a l 10% i n c r e a s e
(G ain 1 9 3 8 ).

On a l l

i n sam ple a r e a

N o m e n c la tu r e f o l l o w s Voss

gymnosperms and m o n o c o ts and F e r n a l d

(1972)

(1950)

for

for a l l

o th er taxa.

RESULTS
The d a t a p r e s e n t e d a r e from s i t e s
once,

tw ice,

or th ree

1981 g ro w in g s e a s o n s

sam pled a n n u a l l y

tim es d u r in g th e 1979,
(T able 3 .1 and 3 . 2 ) .

1980, and

The a g e

o f t h e s e s i t e s make up a s e q u e n c e r a n g i n g f r o m 0 - y e a r
( c l e a r c u t t h e w i n t e r and s p r i n g p r e v i o u s
sam pling)

to 6 -y ear c l e a r c u ts

(burned th e
burns

(T able 3 .1 )

summer p r e v i o u s t o s a m p l i n g )

(T able 3 .2 ) .

th a t th ese s i t e s

summer

and 1 -y e a r
to 6-y ear

C au tio n is a d v ised in i n f e r r i n g

are t y p i c a l o r s u f f i c i e n t to r e p r e s e n t

average c o n d itio n s

fo r those ages.

w ere r i g o r o u s l y s e l e c t e d ,

A lthough s i t e s

th e s e v e r i t y and tim in g o f

t h e d i s t u r b a n c e on e a c h s i t e v a r i e d .
site

to

T herefore each

i s u n i q u e i n many a s p e c t s , m a k i n g d i r e c t c o m p a r i ­

sons d i f f i c u l t .

N onetheless,

I do f e e l

th at c e rta in

TABLE 3.1
RELATIVE COVER ( C ) , RELATIVE FREQUENCY ( F ) , AND SPECIES RICHNESS (R)
FOR VEGETATIONAL GROUPS ON UNBURNED CLEARCUTS

Years
S ince
Number Treatment
S it e 3

Area Annuals & B iennials
R
ha
C
F

Grasses & Sedges
C
F
R

Perennial Herbs
C
F
R

T otal
T o ta l
Trees & Shrubs_______ R______ Cover
R
C
F
per 60 m

.7
-

1
2

32.5
12.9

26.1
20.4

6
7

15.0
29.9

31.2
33 .3

9
9

52.5
56.7

42.0
46.3

10
13

26
31

36.0
61.5

.5
-

1
-

27.2
24.7

22.5
2 7.6

5
5

34.3
7.1

26.7
23.2

8
12

38.5
68.2

50.4
49.2

9
7

23
24

61.5
27.6

-

1 .9

1

49.6
36.5

4 3.8
24.8

9
5

21.6
13.6

21.6
32.2

9
10

29.1
49.6

34.6
41.1

9
9

27
25

50.7
37.1

.

_
-

1

53.1
83.4

32.0
52.7

7
7

4.3
5 .1

26.0
16.4

7
6

42.5
11.4

42.0
30.9

8
6

22
20

46.9
37.7

1 .5

1

45.7
68.8
77.2

30.4
50.0
35.7

6
6
2

1 .6
.3
2 .2

23.9
10.3
14.3

6
5
1

52.7
3 0 .9
20.7

44.2
39.5
50.0

9
8
7

22
19
10

64.4
64.5
41.7

51.6
37.1
45.6

6
5
3

.1
2 .9
.2

9.5
19.3
2 .9

5
4
3

18.0
29.8
22.5

39.0
43.5
51.5

7
5
10

18
14
16

58.9
44.7
30.1

35.0

4

5.7

18.0

4

28.7

47.0

7

15

46.8

1A
IB 80

0
0

16
16

.49

1B81
2

1
1

16
24

-

3A
3B79

2
2

16
16

3B83
4A79

3
3

16
32

.1

3B81
4A80
5A79

4
4
4

16
32
k

4A81
5A80
6

5
5
5

5A81

6

-

-

-

-

-

-

32
k
16

_

.

-

-

-

81.8
67.3
77.3

k

-

-

-

65.6

-

^ i t e ninbers s ta r tin g w ith the same conbination o f mirber and le t t e r represent the sane s i t e surveyed twice; the
nurbers 79, 80, and 81 in d ic a te th e year surveyed <1979, 1980, and 1981), for s i t e s surveyed in more than one growing season.

TABLE 3.2
RELATIVE COVER ( C ) , RELATIVE FREQUENCY ( F ) , AND SPECIES RICHNESS <R)
FOR VEGETATIONAL GROUPS ON BURNED CLEARCUTS
Years
S it e 3
S in ce
Nunber Treatment

Area
ha

Annuals & B ien n ials
C
F
R

7C80
7D79

1
1

16
24

7C81
7D80

2
2

16
24

.

__

-

7D81

3

24

-

10A79
10B79

4
4

26
23

_

10A80
10B80

5
5

10A81
10B81

6
6

^ e e Table 3 .1 .

Grasses & Sedges
F
R
C

Perennial Herbs
F
C
R

T otal
T otal
Trees £ Shrifcs_______ R______ Cover
R
C
F
per 60 n

4
3

35.8
73.5

32.0
4 4.6

9
9

2.8
8 .5

2 6.1
2 4.3

15
12

43.1
17 .8

34.8
28.1

7
10

35
34

26.9
30.6

-

-

45.7
71.6

38.4
56 .3

8
9

1.1
12.4

13.2
21.1

10
8

53.2
15.8

48.3
22.5

7
8

25
25

52.6
48.5

-

-

66.7

52.5

8

14.3

25.1

8

19.1

22.7

8

24

48.7

.2

1 .6
3 .9

1
1

71.8
53.5

35.7
40.0

13
14

15.2
22.9

39.3
2 3.3

13
10

13.0
23.4

2 3.3
3 2.8

9
14

36
39

58.2
45.4

26
23

.1
.8

1.7
5.4

2
3

57.3
57.3

38.2
37.3

11
10

18.5
8.0

3 2.9
23.5

16
8

24.1
33.8

27.2
33 .9

7
14

36
35

75.7
61.6

26
23

.2
.8

.6
2 .3

2
1

66.4
50.3

39.3
38.3

8
9

8 .9
12.2

34.9
21.4

14
10

24.5
36.7

25.2
38.0

8
14

32
34

74.4
74.8

18.3
.1

7 .1
2 .8

28
c o n siste n t d ifferen ces
c le arc u ts

i n burned v e rs u s unburned

can be d i s c e r n e d from t h i s

study,

A com parison o f t h r e e d i f f e r e n t m easures of d i ­
v e rsity ,

to ta l

sp ecies ric h n e ss

(com bined t o t a l

number o f s p e c i e s r e c o r d e d i n f r e q u e n c y a n d c o v e r
m e a s u re m e n ts ), a v e ra g e r i c h n e s s p e r p l o t , and Shannon
2
Index , of s i t e s averaged w ith in years fo r a 6-year
p ro g ressio n is

show n i n F i g u r e 3 . 1 .

g ro u p e d i n t o a n n u a l s and b i e n n i a l s ,
p e re n n ial herbs,
re la tiv e

and t r e e s

frequency,

S p e c ie s were
g r a s s e s and s e d g e s ,

and s h r u b s .

and r i c h n e s s

a r e a by v e g e t a t i o n a l group i s

d a ta fo r each surveyed

show n i n T a b l e s

3 .2 f o r un b u rn ed and b u rn e d c l e a r c u t s ,
R e l a t i v e c o v e r and r e l a t i v e

R e la tiv e cover,

3 . 1 and

resp ectiv ely .

frequency d a ta for each

s p e c i e s s u r v e y e d on u n b u rn ed c l e a r c u t and b u rn e d c l e a r cut sites

d u rin g 1979,

1 9 8 0 , a n d 1 9 81 a r e show n i n

A p p e n d i c e s A a n d B.
S pecies r ic h n e s s ,
rare

a d i v e r s i t y in d e x i n w hich

and d o m in a n t s p e c i e s

on b u rn e d c l e a r c u t s

c o n t r i b u t e e q u a l l y , was h i g h e r

compared t o u nburned c l e a r c u t s

f o r e a c h y e a r e x c e p t y e a r 2.

Average fre q u e n c y p e r

p l o t and Shannon I n d e x , w h ic h r e f l e c t number o f s p e c i e s
and e q u i t a b i l i t y , i n i t i a l l y w ere h ig h e r f o r th e unburned
c learcu ts.

L a te r in th e sequence

(years

4,

5,

and 6 ) ,

^ S h a n n o n I n d e x o f D i v e r s i t y (H) = I P i l o g P i ,
w h e r e P i = r e l a t i v e c o v e r (Odum 1 9 7 1 ) .

29

40

20

0

I

Z

3

4

5

6

0

1

2

3

4

5

fl

0

1

Z

3

4

S

6

0

4

2
0
3

H
R

M

*0J
N
I
N

D
E
X

0r
01
v
cR
S
[
r

0 .5

r

YERRS n r r E R

TflER t h e h t

F ig . 3.1
A) S p e c i e s r i c h n e s s , B) a v e r a g e r i c h n e s s
p e r 1 in2 p l o t , a n d C) S h a n n o n i n d e x o f d i v e r s i t y v e r s u s
y e a r s a f t e r t r e a t m e n t on b u r n e d a nd u n b u r u e d s i t e s .
V e r t i c a l b a r s e q u a l one s t a n d a r d d e v i a t i o n (SD).
Only
one o b s e r v a t i o n a t y e a r 3 b u rn e d and y e a r 6 u n b u rn e d .

30
how ever, v a lu e s of t h i s
th e burned a re a s ,

i n d i c e w e r e much h i g h e r o n

compared t o u n b u rn e d a r e a s .

A lthough

t h e a v e ra g e fre q u e n c y p e r p l o t and th e Shannon In d ex
w e r e l o w e r on c e r t a i n b u r n e d s i t e s ,
sites,

th ese s i t e s

m ostly r a r e ,

compared to u n b u rn ed

had a g r e a t e r number o f s p e c i e s ,

c o n tr i b u ti n g to th e s e i n d ic e s , w hereas

f e w e r , b u t m o r e common, s p e c i e s w e r e c o n t r i b u t i n g on
th e unburned c l e a r c u t s .
As sh o w n i n T a b l e 3 . 3 ,
re c o r d e d on bu rn ed c l e a r c u t s
cle arc u ts.

73 d i f f e r e n t s p e c i e s w e r e
compared to

49 o n u n b u r n e d

T w e n ty -e ig h t s p e c ie s were e x c lu s iv e to

b u rn ed a r e a s , w hereas o n ly 4 s p e c ie s w ere e x c lu s iv e
to unburned a r e a s .
a ls,

12 o f t h e 20 g r a s s e s

p eren n ial herbs,
r e c o r d e d on a l l
c u ts.

T hree o f th e 7 a n n u a ls and b i e n n i ­
18 o f t h e

30

a n d 16 o f t h e 23 t r e e s a n d s h r u b s
s i t e s w ere p r e s e n t on u n b u rn ed c l e a r ­

In c o n tr a s t,

and b i e n n i a l s ,

and s e d g e s ,

a ll

burned s i t e s

co n tain ed a l l

19 g r a s s e s a n d s e d g e s ,

28 p e r e n n i a l h e r b s , a n d a l l

23 t r e e s

7 annuals

24 o f t h e

and s h ru b s .

F i g u r e 3 . 2 sho w s t h a t c o v e r w a s m uch l o w e r on
burned s i t e s ,
By y e a r 2 ,

compared t o u n b u rn ed s i t e s ,

h o w e v e r , c o v e r on b u r n e d c l e a r c u t s was

h i g h e r th a n on unburned c l e a r c u t s .
sequence,

d u r i n g y e a r 1.

burned a re a s

Over t h e te m p o r a l

sh o w e d a g e n e r a l i n c r e a s e i n

c o v e r t h r o u g h y e a r 6, w h e r e a s c o v e r v a l u e s on u n b u r n e d
s ite s v aried l i t t l e .

However, n e a r l y a l l

sites

surveyed

31

TABLE 3 . 3
PLANT SPECIES ENCOUNTERED ON BURNED AND
UNBURNED CLEARCUT SITES IN
NORTHERN LOWER MICHIGAN
S p e c ie s E x c lu s iv e t o B urned A reas
(28 s p e c i e s t o t a l )
A nnuals and B i e n n i a l s
(4 s p e c i e s )

C-ix.4j.um Ap.
C o x y d a t i A A z m p z x v i x z n A L.
G zx a n iu m b-ic.knztti.-i B r i t t .
Ex.-igzx.on c a n a d z n A i A L.

G ra sse s and Sedges
(7 s p e c i e s )
A g x o p y x o n t x a c h y c a u t u m L ink
C a x z x X u g o A p z x m a Mack.
Fz & t u c a i p .
Muh. t znbzx. g- i a. m z x . i c a . n a L.
P a n i c u m x a n t h o p h y A u m Gray
P o a p x a t z n A i A L.
S o xghaAtxum nutanA L.

Hztiant hzmum canadznAiA
M ichx.

L z c k z a minox L.
P o t y g a t a p o ty g a m a W alt.
P o t y g o n u m c i t i n o d z M ichx.
P o t z n t i t t a a x g u ta Pursh.
P o t z n t i t t a A i m p t z x M ichx.
P o t z n t i t t a t x i d z n t a t a A it.
Rumex a c z t o A z t t a L.
S e nc c-to t o m z n t o A u A M i c h x .
I( i o t a a d u n c a Sm.

T re e s and Shrubs
(4 s p e c i e s )
A c z x xu.bx.um L.

P e r e n n i a l Herbs
(13 s p e c i e s )

CzanotkuA ovatuA D e s f .
GaytuAAacia b a c c a t a

A&tzx. A a g g i t i f i o t i u A

SympkoxicaxpoA atbuA L.

Wederaeyer

( W a n g .)

C. Koch

Co n v o t vu t u A ApithamazuA L.
Hizxacium canadznAz M i c h x .

S p e c ie s E x c l u s i v e t o Unburned A re as
(4 s p e c i e s )
P e r e n n i a l Herbs
A n t z n n a x i a n z g t z c t a G reene
A A c t z p i a A A y x i a c a L.

SpixantkzA

gxacitiA

( B i g e l . ) Beck
1f i o t a p z d a t i f a i d a G. Don.

32

TABLE 3 . 3

( C o n tin u e d )

S p e c i e s Common t o B o t h B u r n e d a n d
U n b u r n e d C l e a r c u t 's
(45 s p e c i e s t o t a l )
A nnuals and B i e n n i a l s
(3 s p e c i e s )
Aa a b t A gZabAa L.
La.ctu.ca canadcnAtA L.
McZampyAum Z t n c a A c Lam.
G ra ss e s and Sedges
(12 s p e c i e s )
AgAOAttA hycmaZtA W a l t .
AndAopogon gcA.aA.dtt V i t m a n
BAomuA k a Z m t t G r a y
CaA.CK p c n A y Z v a n t c a Lam.
V a n th o n ta Aptcata (L .)
Beauv.
V c a c h a m p A t a f iZcxuoAa L.
V t c h a n t h c Z Z u m dcpau.p c Aa t u m M u h l .
KocZcAta macAantha P e r s .
P a n t c u m c oZu mb t a n u m
S cribn.

OAyzopAtA a Ap cAt ^ o Z t a
M ichx.

OAyzopAtA pungcnA T o r r .
S c k t z a c h n c puApuAaAccnA
T orr.
P e r e n n i a l Herbs
(14 s p e c i e s )
Apocynum andAOAacmt fioZtum L.
Anemone q u Z n q u c & o Z t a L.
AaZca Z a c v t A L.
CampanuZa A o t u n d t (Jo Z t a L.

FAagaAta v t A g t n t a n a
Duchesne

G a u Z t h c A t a pAocumbcnA L.
H t c A a c t u m a u A a n t t a c u m L.
H t c A a c t u m vcnoAum L.
HcZtanthuA o c c t d c n ta Z tA R idd.
LtatAtA novac-angZtac
L unell
Mat a n t k c mu m c a n a d c n A t A
D esf.
PhyAaZt A v t A g t n t a n a M i l l .
PtcAtdtum aqutZtnum D esf.
S o Z t d a g o Ap.
T re e s and Shrubs
(1 6 s p e c i e s ^
Am c Z a n c h t c A Ap.
AActoAtaphyZoA uvauAAt L.
C o m p t o n t a p c A c g A t n a L.
CAat acguA Ap.
VtcAvtZZa ZontccAa M ill.
E p t g c a AcpcnA L.
PtnuA b a n k A t a n a Lamb.
PopuZuA t A c m u Z o t d c A M i c h x .
PAuntiA p u m t Z a L.
Pauhua A C A o t t n a E h r h .
PAunuA v t A g t n t a n a L.
Qulcacua App. ( r e d o a k
group)
RoAa b Zanda A i t .
Ru.ba-4 p c n A t Z v a n t c u A P o i r .
S a Z t x gZaucophyZZotdcA
Fern.
I/ a c c t n t u m App.

BURNED CLEARCUTS
UNBURNED CLEARCUTS

80

60
o cd > uj a i

40

20

0
0

1

2

3

4

5

YEARS AFTER TREATMENT

Fig. 3.2.
T o t a l v e g e t a t i o n a l c o v e r (m) v e r s u s y e a r s a f t e r t r e a t ­
m e n t on b u r n e d and u n b u r n e d s i t e s .
V e r t i c a l b a r s e q u a l o ne SD. O n ly
o ne o b s e r v a t i o n a t y e a r 3 b u r n e d and y e a r 6 u n b u r n e d .

34
d u r i n g two o r m o r e g r o w i n g s e a s o n s sho w ed i n c r e a s e d
c o v e r from t h a t o f th e p r e v i o u s y e a r .
Trends in r e l a t i v e
and b i e n n i a l s

o v e r t h e t e m p o r a l s e q u e n c e a r e shown

i n T a b le s 3.1 and 3 .2 .
in th is

c o v e r and f re q u e n c y o f a n n u a ls

On u n b u r n e d s i t e s ,

group a re v i r t u a l l y

rum l i n e a r e Lam.

ab sen t,

e x c e p t f o r M e lam py -

(cow w h e a t ) w h i c h i s common i n t h e

u n d e rs to ry o f m ature ja c k p in e s ta n d s .
the r e l a t i v e

sp ecies

cover of t h is

b e t w e e n t h e tw o s i t e s

species

On b u r n e d s i t e s

group v a r i e d g r e a t l y

surveyed as f i r s t

year burns.

Very h i g h l e v e l s o f c o v e r

( 1 8 .3 % ) o f a n n u a l s a n d b i ­

e n n i a l s w ere se e n on s i t e

7C80, w h e r e a s o n s i t e

o n l y 0.1% c o v e r f o r t h i s
(T able 3 .2 ) .
(geranium )
quin)

On s i t e

s p e c i e s g r o u p was r e c o r d e d

7C80, G eranium b i c k n e l l i i B r i t t ,

and C o r y d a l i s s e m p e r v i r e n s L.

dom inated t h i s

g erm ination of th ese

sp ecies

group.

sp ecies is

d r a s t i c r e d u c t i o n from y e a r s
on s i t e

7D79

(ro ck -h arle­

Heat s tim u la te d

i n d i c a t e d by t h e i r

1 to 2 fo llo w in g f i r e

7C80-81 ( s e e C h a p t e r 7 ) .

P e r e n n i a l g r a s s e s and s e d g e s w ere d o m in a n t on
b o th burned and unburned c l e a r c u t s
3 .2 ).

The r i c h n e s s

of t h is

( T a b le s 3 .1 and

g r o u p , h o w e v e r, was due

t o t h e many s p e c i e s o f g r a s s e s .

Burned s i t e s

y e a r s 1 t o 6 showed g r e a t e r s p e c i e s r i c h n e s s
than

an alo g ou s unburned s i t e s

from
of grasses

( F i g . - 3 . 3 A ) . The b u r n e d

c le a rc u ts r e ta in e d high le v e ls of ric h n e s s

of grass

35

4

BURNED CLEARCUTS
UNBURNED CLEARCUTS

2

0
B
6
4

2

0
0

R
A
S

2

3

4

5

B

SO
0

1

6

BURNED CLEARCUTS

2 2 UNBURNEO CLEARCUTS

40

.

20

_

S
c
0

V
E
R

i

I

pL

1
3

4

S

B

YEARS AFTER TREATMENT
F ig. 3 .3 .
A) Number o f g r a s s s p e c i e s a n d B) p e r ­
c e n t c o v e r o f g r a s s e s v e r s u s y e a r s a f t e r t r e a t m e n t on
b u r n e d and un b u rn ed s i t e s .
V e r t i c a l b a r s e q u a l on e SD.
O nly one o b s e r v a t i o n a t y e a r 3 b u r n e d and y e a r 6 u n ­
burned .

36
species

th roughout th e tem p o ral sequence, w hereas

u n b u r n e d c l e a r c u t s sh o w e d a d e c l i n e i n r i c h n e s s
th e l a t t e r p a r t of th e sequence.
g r a s s c o v e r on t h e b u rn e d s i t e s

From y e a r s 2 t o 6
g r e a tly exceeded th a t

o f t h e u n b u r n e d s i t e s o f t h e same a g e
D om inant g r a s s

in

(F ig.

3 .3B ).

s p e c ie s occupying th e b u rn ed s i t e s

i n c l u d e O r y z o p s i s a s p e r i f o l i a M i c h x . , 0_^ p u n g e n s T o r r . ,
and S c h iza ch n e p u r p u r a s c e n s T o r r.

In c o n tra s t,

the

c o v e r o f g r a s s e s o n u n b u r n e d s i t e s was n e g l i g i b l e .
The r e l a t i v e
c u ts

is

co ver o f sedges on unburned c l e a r ­

shown i n F i g .

3 . A.

O n l y two s p e c i e s o f s e d g e ,

C a r e x p e n s y l v a n i c a Lam. a n d C^ r u g o s p e r m a M a c k . , w e r e
o b s e r v e d on th e s tu d y s i t e s ,
as C arex spp.

w hich w ere c h a r a c t e r i z e d

a n d g i v e n a r i c h n e s s v a l u e o f 1.

Sedges

c o m p r i s e d t h e m a j o r i t y o f t h e c o v e r o f t h e g r a s s and
sedge s p e c ie s group,
8 0 - 8 1 ( y e a r s 4,

a n d o n l y on b u r n e d s i t e

5 , a n d 6) a n d u n b u m e d s i t e

( y e a r 1) d i d t h e r e l a t i v e
of sedges.

10A791B81

cover of g rasse s exceed th a t

On u n b u r n e d c l e a r c u t s C a r e x c o v e r i n i t i a l l y

( y e a r s 0 a n d 1) w a s l o w .

By y e a r

2, h o w e v e r , C a r e x

c o v e r d r a m a t i c a l l y i n c r e a s e d and by y e a r 5 C arex
c o v e r e d a n a v e r a g e 70% o f t h e u n b u r n e d s i t e s .
tia lly ,

C arex r e l a t i v e

In i­

cover in c re a s e d to a g r e a te r

e x t e n t on b u r n e d c l e a r c u t s c o m p a r e d t o u n b u r n e d c l e a r ­
c u ts.
tiv e

In c o n t r a s t to th e unburned s i t e s ,
cover did not in c re a s e in th e

Carex r e l a ­

l a t t e r p a r t of the

^

m o o m c/>

60

a im < o n

BURNED CLEARCUTS
CLEARCUTS

7ZA UNBURNED

60

40

20

0
0

1

2

3

4

S

6

YEARS AFTER TREATMENT

F i g . 3 . h.
P e r c e n t cover o f sedge v e r s u s y e a r s a f t e r tr e a tm e n t
on b u r n e d and u n b u r n e d s i t e s .
V e r t i c a l b a r s e q u a l one SD. Only
o ne o b s e r v a t i o n a t y e a r 3 b u r n e d and y e a r 6 u n b u r n e d .

38
burned sequence.

In fa c t,

on t h e b u r n e d s i t e s ,

showed a g e n e r a l d e c r e a s e i n r e l a t i v e

C arex

c o v er betw een

y e a r s 3 a n d 6'.
Bracken f e r n
w in terg reen

(P te rid iu m aquilinum D e sf.)

( G a u l t h e r i a procum bens L . ) ,

p e re n n ia l herbs in t h i s
trib u to rs

i n c l u d e d as

s t u d y , w ere th e dom inant c o n ­

to th e r e l a t i v e

cover of t h is

A ll o th e r p e re n n ia l herbs c o n trib u te d
tiv e cover of th is

and

sp ecies

s p e c ie s group.

little

to r e l a ­

group b u t added g r e a t l y

to s p e c i e s r i c h n e s s on a l l s i t e s ,

e s p e c ia lly those

w h ic h had b e e n b u r n e d ( T a b le s 3 . 1 and 3 . 2 ) .

A steady

d e c r e a s e i n t h e n u m b e r o f h e r b s was s e e n o n t h e u n b u r n e d
sequence, w hereas burned s i t e s
(years

sh o w e d a n i n i t i a l d e c l i n e

1 t h r o u g h 3) b u t l a t e r e x h i b i t e d h i g h r i c h n e s s

(years 4 through 6).
e n n ia l herbs e x c lu s iv e

Some f r e q u e n t l y o c c u r r i n g p e r ­
to burned c le a r c u ts

H ie ra c iu m c a n a d e n s e M ichx.
a c e t o s e l l a L.
Michx.

include

( C a n a d i a n h a w k w e e d ) , Rumex

(sh e ep s o r r e l ) , and S e n e c io to m e n to su s

(g ro u n d sel).
The n u m b e r o f s p e c i e s o f t r e e s a n d s h r u b s d i f f e r e d

little

betw een b u rn ed and unburned c l e a r c u t s o f th e

same a g e
species

(T ables
groups,

3 . 1 and 3 . 2 ) .
trees

Of t h e f o u r m a j o r

a n d s h r u b s sh o w e d t h e l e a s t i n ­

crease in species rich n e ss a f te r

fire.

R e la tiv e cover

o f t h e s e s p e c i e s w a s on t h e a v e r a g e h i g h e r o n u n b u r n e d
c l e a r c u t s d u r i n g y e a r s 1 t h r o u g h 4, b u t s l i g h t l y

low er

39
d u r i n g y e a r s 5 a n d 6.

Unburned c l e a r c u t s

general decrease in r e la tiv e
burned s i t e s

sh o w e d a

cover over tim e , w hereas

sh o w e d a d e c l i n e f r o m y e a r 2 t o y e a r s

3 and 4, b u t i n c r e a s e d d u r i n g y e a r s 5 and 6.
s p e c ie s d om inating t h i s
unburned s i t e s

g ro u p on b o t h b u r n e d and

i n c l u d e d V accinium s p p .

P o p u lu s t r e m u l o i d e s M ichx.

(b lu eb erry ),

( a s p e n ) , Q uercus spp.

( o a k ) , Prunus s e r o t i n a Ehrh.
spp.

The

( b l a c k c h e r r y ) , a n d R u b us

(dew berry),

DISCUSSION
A m ajor e f f e c t o f f i r e

on v e g e t a t i o n i s

the

a d d i t i o n and l o s s o f s p e c i e s th r o u g h m echanism s i n v o l v ­
ing

both s i t e

burn v e g e ta tio n

facto rs

and c h a r a c t e r i s t i c s

(in clu d in g propagules).

of the p re ­

These

m echanism s i n c l u d e c r e a t i o n o f l a r g e o p e n in g s i n th e
forest

canopy and f o r e s t

a v aila b ility

flo o r,

(A hlgren 1960, S t.

W e lls 1979) , i n c r e a s e d s o i l
m o istu re

in creased n u trie n t
J o h n and R undel 1976,

t e m p e r a tu r e s and s o i l

(A h lg re n and A h l g r e n 1960, W e lls 1 9 7 9 ),

stim u­

l a t i o n o f s p r o u t i n g o f p r e e x i s t i n g v e g e t a t i o n and
g erm in atio n o f b u rie d seed
al.

(Sweeney 1956, M u l l e r e t

1968, C h r i s t e n s e n and M u lle r 1975, T r e d i c i 1 977),

v o l a t i l i z a t i o n o f a l l e l o p a t h i c chem icals
a l.

1968, C h r i s t e n s e n and M u l l e r 1975)

s o il m icro b ial a c tiv ity

(M uller e t

and i n c r e a s e d

(A h lg ren and A h lg re n 1960,

40
W ells 1979).
The i n c r e a s e i n s p e c i e s r i c h n e s s a n d d o m i n a n c e
o f c e r t a i n p la n t groups o c c u rrin g a f t e r
th is

study is

fire

seen in

a phenomenon w h ic h h a s b e e n d e s c r i b e d

in a v a r i e t y o f ecosystem s

(L ittle

a n d M oor e 1 9 4 9 ,

A h l g r e n a n d A h l g r e n I 9 6 0 , D y r n e s s 1 973, S h a f i and
Y a r r a n to n 1973, C h r i s t e n s e n and M u lle r 1975, P u rd ie
and S l a t y e r 1976).
tem p o rary ,
cally

T h is in c r e a s e d d i v e r s i t y i s

l a s t i n g o n ly a few y e a r s .

A sh ift

u su ally

is

ty p i­

s e e n i n p l a n t c o m p o s i t i o n a n d d o m i n a n c e away

from i n v a d i n g s p e c i e s , w h ic h e s t a b l i s h
seed i n i t i a l l y

after fire ,

them selves by

to th o s e s p e c i e s p r e s e n t

b e f o r e th e f i r e w hich r e e s t a b l i s h th e m s e lv e s l a r g e l y
by v e g e t a t i v e m eans.
The p o s t f i r e

su ccessio n al trends p rev io u sly

r e p o r t e d from t h e Lake S t a t e s ,

h o w e v e r , do n o t a l w a y s

follow

th is p a ttern .

(1979)

stu d ied e a rly re v e g e ta tio n of th e L i t t l e

w ild fire

For exam ple,

Ohmann a n d G r i g a l

i n n o r t h e a s t e r n M innesota b u t d id n o t f in d

an abundance o f d is tu r b a n c e s p e c ie s .
p r e s e n c e o f a few " f i r e
and r o c k - h a r le q u i n ,
after

Sioux

fo llo w ers",

Except fo r the
su c h as geranium

s p e c i e s c o m p o s i t i o n b e f o r e and

f i r e were n e a r l y i d e n t i c a l .

Vogl (1 9 7 0 ), w o rk in g

i n th e n o r t h e r n W isco n sin p i n e b a r r e n s , found t h a t
th e frequency of occurrence o f b a rren sp ecies did
n o t show a s i g n i f i c a n t r e s p o n s e t o b u r n i n g .

Changes

41
i n u n d e rs to ry balsam f i r

and p a p e r b i r c h v e g e t a t i o n

f o l l o w i n g tim b e r h a r v e s t i n n o r t h e r n M in n e so ta were
r e p o r t e d by O u t c a l t and W hite

(1981).

T h e y sh o w e d

d e c r e a s e d d i v e r s i t y o f n e a r l y a l l s p e c i e s g r o u p s on
lo g g ed and b u rn ed s i t e s
sites;

compared to u n b u rn ed lo g g ed

how ever, h ig h d e n s i t i e s o f geranium and r o c k -

h a r l e q u i n a f t e r f i r e w ere found.
v e g e t a t i o n a l d a t a f r o m 11 s e r i e s
n o rth e rn co n ifero u s

fo rests

A h lg re n (1959)
of p lo ts

These d a ta i n d i c a t e

a n d 35 s p e c i e s

4 species ex clu siv e
common t o b o t h a r e a s .

th a t larg e in creases in species

ric h n e s s fo llo w f i r e ,

K r e f t i n g and A h lg re n (1 9 7 4 ) ,

how ever, r e p o r t e d changes i n s p e c ie s
th is

in the

o f M i n n e s o t a a n d f o u n d 60

sp e c ie s e x c lu siv e to burned a re a s ,
to unburned a r e a s ,

grouped

same s t u d y a r e a ,

th a t burning r e s u lte d

sp ecifically

c o m p o s it io n from

by s i t e ,

a n d sh o w e d

i n e i t h e r no s i g n i f i c a n t

change

o r d e c r e a s e d s p e c i e s r i c h n e s s compared t o u n b u rn ed
c o n tro ls.
The ab o v e s t u d i e s r e p o r t d a t a fro m p r e s c r i b e d
b u rn s and w i l d f i r e s
(A h lg ren 1960,

th a t took p la c e d u rin g th e sp rin g

1 9 7 4 , V o g l 1 9 7 0 , Ohmann a n d G r i g a l 1979 )

a n d summer ( A h l g r e n 1 9 6 0 ,

1974, O u t c a l t and W hite 1 9 8 1 ),

b u t none r e p o r t e d a la r g e

site -sp e c ific

species
I found,

d iv ersity a fte r
as d id A hlg ren ,

fire

in crease in

as seen in t h is

study.

t h a t b u r n in g c l e a r l y prom otes

the e sta b lish m e n t o f a la rg e v a r ie ty of sp e c ie s not

42
t y p i c a l to unburned a r e a s .

A t o t a l o f 73 d i f f e r e n t

s p e c i e s w ere re c o r d e d on bu rn ed s i t e s
on unburned s i t e s .

c o m p a r e d t o 49

However, n e a r l y a l l b u r n e d s i t e s

i n t h i s s t u d y sh o w e d g r e a t e r
to an alo go u s unburned s i t e s

species ric h n e ss
(F ig.

3 .1 ).

t h e number o f a n n u a l s and b i e n n i a l s ,

In c re a se s in

grasses,

p e r e n n i a l h e rb s f o llo w in g f i r e w ere found.
l i s h m e n t o f new s p e c i e s
v iab le

seed sto re d

S t u d i e s h a v e shown t h a t
s u c c e s s io n a l ages are

in th e f lo o r under f o r e s t

s t o n and A l l e s s i o

stands

(L iv ing ­

1968, T r e d i c i 1977, A h lg re n 1979a).

T h e l o n g e v i t y o f many f i r e - f o l l o w e r s
two y e a r s a f t e r

The e s t a b ­

in th e f o r e s t flo o r or tra n sp o rte d

s e e d from s p e c i e s o f a l l

u b iq u ito u s

and

f o l lo w in g f i r e r e s u l t s from

by w ind an d a n im a l v e c t o r s .
v iab le

compared

f i r e many a r e g o n e

is

short;

(note d ecreased

s p e c i e s r i c h n e s s i n b u r n e d s i t e s b e tw e e n y e a r s 1 and 2,
T able

3 .2 and F ig .

3 .1 ).

In the l a t t e r y ears a f t e r f i r e ,

s p e c i e s t h a t w e re p r o b a b l y members o f t h e p r e - e x i s t i n g
v eg etatio n

a nd w h i c h e s t a b l i s h a n d p e r p e t u a t e

m a i n l y by v e g e t a t i v e m ea n s d o m i n a t e .
y e a r-o ld burned s i t e s

(sites

Four,

them selves

5 , an d 6-

10A 79-80-81 and 10B79-80-81)

r e t a in e d a hig h le v e l o f d i v e r s i t y com prising a m ix tu re
o f f i r e - s t i m u l a t e d g r a s s e s and h e r b s , whose p r e s e n c e
a p p e a r s t o be w a n in g ,

and t r e e s

and s h r u b s , whose

dom inance a p p e a r s t o be i n c r e a s i n g ( T a b l e 3 . 2 ) .
These f in d in g s r e p r e s e n t

im portant c o n tr a s ts

in

43
t h e e a r l y s u c c e s s i o n a l d e v e lo p m e n t on t h e b u r n e d and
unburned c l e a r c u t s i n t h i s s tu d y .

M ajor d i f f e r e n c e s

dom inance and d i v e r s i t y o f a l l s p e c i e s
the e a rly re v e g e ta tio n p ro cess i s
and u n b u rn ed c l e a r c u t s .

in

groups d u rin g

s e e n on t h e b u rn e d

The m o st i m p o r t a n t c o n t r a s t ,

and c e r t a i n l y t h e m o st o b s e r v a b l e

in the f i e l d ,

is

the

d i f f e r e n t p athw ays o f v e g e t a t i o n a l d evelopm ent c h a r a c t e r ­
i z e d on t h e s e s i t e s .

On a l l b u r n e d a n d u n b u r n e d c l e a r c u t

a reas ja c k pine re g e n e ra tio n is
lish ed ,
tio n .

and t h e s e s i t e s

f a i l i n g t o b ecom e e s t a b ­

a re dom inated by o t h e r v e g e t a ­

The u n b u r n e d c l e a r c u t s a r e b e i n g c o n v e r t e d t o

e s s e n t i a l l y C a r e x " m ea do w s" ( P l a t e

3 .1 ).

The o n l y o t h e r

v e g e t a t i o n o f a n y c o n s e q u e n c e a r e o a k a n d c h e r r y stu m p
s p r o u t s and b l u e b e r r y grow ing i n and aro u n d t h e shade
of slash p ile s

(logging d e b r i s ) .

In c o n tra s t,

burned

c le a r c u ts a re b ein g converted to e a r ly su c c e ss io n a l
hardw oods su c h as a s p e n , o a k ,
sp ecies,

Shrub

su c h as b l u e b e r r y , w i l lo w , C e a n o t h u s , and

A m elanchier a re a l s o
(P late

and b l a c k c h e r r y .

3 .2 ).

im p o r ta n t on t h e s e s i t e s

C a r e x i s a l s o a d o m i n a n t member o f t h e

o ld e r burned s i t e s ,

but th ere is

little

in d icatio n th a t

t h e s e s i t e s w i l l b e c o n v e r t e d t o a C a r e x ’'m e a d o w . "
fact,

In

t h e dom inance o f C a re x i s b e i n g c h e c k e d by t r e e

and s h r u b s p e c i e s m e n tio n e d above and g r a m i n o id s s u c h
as O r y z o p s i s a s p e r i f o l i a and S c h iz a c h n e p u r p u r a s c e n s .
An i n c r e a s e i n s e d g e s f o l l o w i n g f i r e

has been

44

P l a te 3 .1 .
C a r e x meadow e s t a b l i s h e d o n a
5 - y e a r - o ld unburned c l e a r c u t s i t e .

P la te 3 .2 .
S h ru b s and e a r l y s u c c e s s i o n a l
hardwoods d o m in a tin g a 5 - y e a r - o l d p r e s c r i b e d
burn s i t e .

45
p r e v i o u s l y docum ented (A h lg re n 1960, Vogl 1970, O u t­
c a l t and W hite 1 9 8 1 ).

However,

t h e overw helm ing

dom inance o f C arex o b s e r v e d i n t h i s
be uniq u e to ja c k p in e s i t e s
gan.

s t u d y seems to

i n n o r t h e r n low er M ic h i­

A C a r e x "meadow", o n ce e s t a b l i s h e d ,

cap able of e x clu d in g tr e e
d u c t i o n f o r many y e a r s
Noble 1 980).

appears

and s h r u b s e e d l i n g r e p r o ­

( N i e r i n g a n d Go od w in 1 9 7 4 ,

The d o m in a n ce o f C a r e x o n u n b u r n e d

c le a rc u ts p rev en ted the e stab lish m en t of a m u lti­
la y e re d canopy,

as o b s e rv e d on th e b u rn e d c l e a r c u t s .

The m o n o la y e re d C a re x c an o p y on u n b u rn e d c l e a r c u t s
was r e s p o n s i b l e f o r t h e l a c k o f i n c r e a s e
over the tem poral sequence.
treatm en t,

in t o t a l cover

By y e a r s 5 a n d 6 a f t e r

b u r n e d c l e a r c u t s h a d 47% a n d 63% m o r e v e g e ­

t a t i o n cover, r e s p e c tiv e ly ,
There a re

th a n unburned c l e a r c u t s .

s e v e r a l p o s s i b l e m echanism s t h a t e x p l a i n

th e dom inance o f C a re x p e n s y l v a n i c a .
may b e C a r e x ' s a b i l i t y
m ad e a v a i l a b l e

One p o s s i b i l i t y

to e x p lo it n u tr ie n ts

and sp a ce

fo llo w in g a p e r t u r b a t i o n such as c l e a r -

c u ttin g an d /o r b urning.

Recent s tu d ie s

in v o lv in g

d i s t u r b a n c e s h a v e show n t h a t c e r t a i n e x p l o i t a t i v e
sp ecies,

due to t h e i r l i f e - h i s t o r y

can m onopolize r e s o u r c e s

ch aracteristics,

l i b e r a t e d by d i s t u r b a n c e

and s u p p re s s o r e x c lu d e o t h e r s p e c i e s .

M a r k s 's work

( 1 9 7 4 ) o n New H a m p s h i r e c l e a r c u t s h a s shown p i n
cherry

(Prunus p e n s y l v a n ic a L .)

t o be s u c h an

46
ex p lo itativ e

sp ecies.

B a k e l a a r a n d Odum ( 1 9 7 8 )

c o n c l u d e d t h a t a few o p p o r t u n i s t i c s p e c i e s a l r e a d y
e s t a b l i s h e d on abandoned f i e l d s
to expand t h e i r n ic h e s a f t e r
em pting c e r t a i n
d iv ersity .

i n G e o rg ia were a b le

f e r t i l i z a t i o n by p r e ­

s u b o rd in a te s , thus red u cin g o v e ra ll

A hlgren

(1960)

showed t h a t n u t r i e n t s

r e l e a s e d fo llo w in g f i r e have f e r t i l i z i n g
s t i m u l a t e th e grow th o f c e r t a i n s p e c i e s .
o f a study i n i t i a t e d

i n May t o t e s t

a r e d i s c u s s e d i n C h a p t e r 6.

effects

The r e s u l t s

t h i s h y p o th esis

A second e x p la n a tio n

f o r t h e dom inance o f C a r e x , a l t h o u g h u n t e s t e d ,
p o s s i b l e r e l e a s e o f a chem ical i n h i b i t o r
At t h i s

th at

is

its

(allelochem ) .

tim e I can n o t f u l l y e x p la in th e c o n t r a s ­

t i n g s u c c e s s i o n a l p a th w a y s f o l l o w e d on b u r n e d and u n ­
burned ja c k p in e c le a r c u ts .

However, fro m t h e f o r m a l

c h a ra c te riz a tio n of e a rly re v e g e ta tio n of th ese s i t e s
a n d my own p e r s o n a l o b s e r v a t i o n s ,

the fo llo w in g

p o in ts a re p ro b a b ly im p o rta n t in answ ering t h i s q u es­
tio n .

C a no p y r e m o v a l w i t h o u t s u b s e q u e n t b u r n i n g r e s u l t s

i n th e r a p i d fo rm a tio n o f a n e a r m o no cu ltu re o f Carex
p en sy lv an ica.

Canopy r e m o v a l f o l l o w e d by f i r e

prom oted th e s p re a d o f se d g e s ,

also

b u t i n a d d i t i o n prom oted

t h e e s t a b l i s h m e n t o f a l a r g e v a r i e t y o f new s p e c i e s
a n d t h e r a p i d s p r e a d o f many s p e c i e s t h a t w e r e p r o b a b l y
members o f t h e p r e e x i s t i n g v e g e t a t i o n .
effects of fire

These d ram atic

on s p e c i e s c o m p o s i t i o n and s p e c i e s

47
re la tiv e

dom inance a l t e r e d

the c o m p etitiv e a b i l i t i e s

o f C arex.

F ollow ing f i r e

t h e r e was t h e e s t a b l i s h m e n t

and c o n tin u e d developm ent o f a m u l t i l a y e r e d canopy
c o n s i s t in g o f gram in o id s,
tree s.

herbs,

shrubs,

and sm a ll

The s h a d e p r o v i d e d b y t h e c a n o p y a p p e a r e d t o

check the f u r th e r spread of C arex .
th e c o m p e titiv e n e s s o f C arex i s
situ atio n s

is

its

F u rth er proof th a t

low ered i n shaded

h i g h f r e q u e n c y b u t v e r y lo w r e l a t i v e

c o v er under a m ature ja c k p in e canopy.

A lso,

a con­

s i s t e n t p a t t e r n o b s e r v e d on u n b u r n e d c l e a r c u t s was
t h e v i g o r o u s g ro w th o f b l u e b e r r y i n and a ro u n d t h e
shade o f th e s l a s h p i l e s , w h ile C arex dom inated th e
l a r g e r open a r e a s .

CHAPTER IV
MULTIPLE PATHWAYS IN EARLY SUCCESSIONAL
DEVELOPMENT FOLLOWING DISTURBANCE
TO JACK PINE STANDS IN NORTHERN
LOWER MICHIGAN
In tro d u ctio n
Of a l l t h e a n t h r o p o g e n i c a n d n a t u r a l d i s t u r ­
bances , f i r e has been th e m ost w id e ly s tu d ie d .
S e c o n d a ry p l a n t s u c c e s s i o n has b e e n a m a jo r theme
in the f i r e ecology l i t e r a t u r e ,

and num erous te rm s

and m odels have b een p ro p o s e d t o d e s c r i b e
(1929)

used " d e f le c te d

it.

Godwin

su ccessio n " to d e sc rib e

s u c c e s s i o n a l developm ent f o llo w in g b u rn in g ,
and c u t t i n g , w hich d i f f e r e d

from t h e

sio n" in the absence o f d is tu rb a n c e .
h y p o th e s iz e d an " i n i t i a l f l o r i s t i c

g razing,

"norm al s u c c e s ­
E gler

(1954)

co m position"

(IFC)

model for. o l d - f i e l d v e g e t a t i o n developm ent w hich has
b e e n s i m i l a r l y u s e d t o d e s c r i b e comm unity d e v e lo p m e n t
fo llo w in g f i r e
to E g le r,

( P u r d i e a n d S l a t y e r 1 9 76 ) .

A ccording

a f i e l d r e c e i v e s many d i f f e r e n t p r o p a g u l e s

up t o t h e t i m e o f a b a n d o n m e n t ; a f t e r
dev elop m ent u n fo ld s from t h i s
ad d itio n al

abandonm ent

i n i t i a l f lo r a , w ith o u t

i n c r e m e n t s by f u r t h e r

in v asio n s.

E g ler

r e c o g n i z e d t h a t s e v e r a l e q u i l i b r i a may e x i s t on a
given s i t e ,

and s u c c e s s i o n c o u ld be a r r e s t e d i f
48

the

49
woody p l a n t s p e c i e s t h a t w o u l d d o m i n a t e t h e m a t u r e
c o m m u n i t y w e r e k i l l e d a t some i n t e r m e d i a t e s t a g e .
R ichards

(1955)

term ed d i f f e r e n t se c o n d a ry s u c c e s s io n s

"p rogressive" i f ,

fo llo w in g c le a rin g ,

a young se c o n d a ry

f o r e s t r a p i d l y e s t a b l i s h e s w hich e v e n t u a l l y le a d s to
a fo re st lik e

the o r ig in a l,

or "reg ressiv e " i f

v a n n a d e v e l o p s w i t h few o r no t r e e s .

Loucks

a sa­

(1970)

co n clu d ed t h e r e i s a n a t u r a l ten d e n c y i n f o r e s t system s
tow ard p e r i o d i c p e r t u r b a t i o n t h a t r e c y c l e s

th e system

and m a i n t a i n s a p e r i o d i c wave o f p e a k d i v e r s i t y .
C o n n e ll and S l a t y e r

(1977) o u t l i n e d th e

"in h ib itio n

m odel", w hich d e s c r i b e s a p o s t d i s t u r b a n c e s u c c e s s io n a l
p r o c e s s w here e a r l i e r

co lo n ists

secure

space and/or

o t h e r r e s o u r c e s and i n h i b i t th e i n v a s i o n o f s u b s e q u e n t
co lo n ists,

o r s u p p re ss o r e x c lu d e th e grow th o f th o se

already p re se n t.

T h e ir model o u t l i n e s

the ap p aren t

mechanism i n v o l v e d i n a r r e s t e d o r r e g r e s s i v e s u c c e s s i o n .
I n t h e L ak e S t a t e s ,

l i m i t e d d e t a i l e d work has

b e e n c o n d u c te d on t h e c h a r a c t e r i z a t i o n o f p l a n t
sio n fo llo w in g f i r e

succes­

(A h lg re n 1960, Vogl 1970, K r e f t i n g

a n d A h l g r e n 1 9 7 4 , Ohmann a n d G r i g a l 1 9 7 9 , O u t c a l t a n d
W h ite 1 981, S c h e i n e r and T e e r i 1981)
( O u t c a l t and W hite 1 9 8 1 ).
on j a c k p i n e s i t e s
In th is

However,

and lo g g in g
sim ila r stu d ies

i n M ichigan have n o t b e en p u b l i s h e d .

chapter,

d e t a i l e d com parisons o f s p e c ie s

c o m p o s i t i o n a n d s p e c i e s d o m i n a n c e a r e made b e t w e e n

50
d istu rb ed s ite s

( c l e a r c u t a n d /o r burned)

ja c k pine sta n d s.
d irect effects

and u n d is tu rb e d

C o n c lu s io n s a r e drawn c o n c e r n i n g t h e

o f c l e a r c u t t i n g a n d / o r b u r n i n g on t h e s e

j a c k p i n e com m unities and s u b s e q u e n t e a r l y s u c c e s s i o n a l
tren d s.

C o m p a r i s o n s among s i t e s

show t h a t m u l t i p l e

s u c c e s s i o n a l p athw ays a r e e v i d e n t soon a f t e r d i s t u r b a n c e .

M ethods
On e a c h s i t e ,

a l l v a s c u la r p l a n t s were c h a r a c t e r ­

i z e d by fre q u e n c y and c o v er m easurem ents a s d e s c r ib e d
i n C h a p t e r 3.
stan d s,
tall)

In th e m atu re and 3 5 - y e a r - o l d j a c k p in e

only th e u n d e rs to ry v e g e ta tio n

( p l a n t s £ 1.5 m

w ere c h a r a c t e r i z e d .

R esu lts
Unburned C l e a r c u t S i t e s
V ersu s M ature J a c k F in e
Stands
A ll unburned c l e a r c u t
fo rm e rly m ature ja c k p in e .
s ta n d s su rv e y ed w ere u se d ,

sites

The f o u r m a t u r e j a c k p i n e
th erefo re,

of the p re d is tu rb a n c e c o n d itio n .
rig o ro u sly

s e l e c t e d , was u n i q u e ,

some e x t e n t i n p r e d i s t u r b a n c e
dom inance,

tu rb an ce.

However,

as r e p r e s e n t a t i v e

Each s i t e ,

alth o u g h

in th a t they v a rie d to

sp ecies

seed pool com position,

invading p ro p ag u les,

i n c l u d e d f o r stu d y were

c o m p o s itio n and

nearby sources of

and s e v e r i t y and t i m i n g o f t h e d i s ­
c e r ta in o v e ra ll su c c e ss io n a l tren d s

51
can be d i s c e r n e d .
R e l a t i v e c o v er and r e l a t i v e

frequency d a ta fo r

e a c h s p e c i e s on u n b u rn ed and b u rn e d c l e a r c u t s i t e s , as
w e ll as burned 3 5 -y e a r-o ld ja c k p in e s i t e s ,

in term ed i­

a t e - a g e a n d m a t u r e j a c k p i n e s t a n d s a r e show n i n
A p p e n d i c e s A, B, a n d C, r e s p e c t i v e l y .
C learcut s ite s
species ric h n e ss
stands.

i n i t i a l l y had in c re a s e d t o t a l

compared to t h e u n d i s t u r b e d f o r e s t e d

However,

on c l e a r c u t s i t e s

sp ecies ric h n e ss g e n e ra lly decreased
over the s ix year sequence,

i n m ost o ld e r c l e a r c u t s i t e s
ness

h a v in g low er s p e c i e s r i c h ­

th a n m ature j a c k p in e u n d e r s t o r i e s

T able 4 .1 ) .

S im ilar tren d s

resu ltin g

(F ig.

4 . 1 and

in average ric h n e s s p er p lo t

and th e Shannon in d e x w ere found.
As shown i n T a b l e 4 . 2 ,
r e c o r d e d on c l e a r c u t s i t e s

51 d i f f e r e n t

s p e c ie s were

c o m p a r e d t o 41 s p e c i e s on

m ature ja c k p in e s i t e s .

I n c r e a s e s i n g r a s s and p e r e n n i a l

herb

sites

s p e c i e s on c l e a r c u t

creases

in fa c t,

in ­

i n th e s e s p e c ie s gro u p s were r e s p o n s i b l e f o r th e

in itia l rise
sites

are e v id en t;

(F ig.
It

in sp ecies rich n ess

on r e c e n t c l e a r c u t

4 .1 and T a b le 4 . 1 ) .

a p p e a r s t h a t t o t a l v e g e t a t i o n a l c o v e r on c l e a r -

cut s ite s v aried l i t t l e
fo r e s te d stands

(Fig.

exam ined i n d i v i d u a l l y ,

from t h a t o f t h e u n d e r s t a r y o f

4 .2 ).

How ever, when s i t e s

c le a r c u ttin g did,

in fa c t,

are
cause

c h an g e s i n th e t o t a l p l a n t c o v e r and c o v e r o f i n d i v i d u a l

52
SO

s

>55

R

D

1

?

3

4

&

6

0

I

2

3

«

s

G

10

a

3

Z
1 .s

1

0
? e r g s r f ' cb m n m t N T

F ig. 4 .1 .
A) S p e c i e s r i c h n e s s ( t o t a l n u m b e r o f
s p e c i e s ) , B) a v e r a g e r i c h n e s s p e r p l o t , a n d C) S h a n n o n
index o f d i v e r s i t y , o f s i t e s averaged w i t h in y e a r s ,
f o r m atu re ja c k p in e sta n d s and unburned c l e a r c u t
ja c k p in e s i t e s com prising a 6-year sequence fo llo w in g
lo g g in g .
V e r t i c a l b a r s e q u a l one s t a n d a r d d e v i a t i o n
(SD) .

TABLE 4 .1
RELATIVE COVER ( C ) , RELATIVE FREQUENCY ( F ) , AND SPECIES RICHNESS (R)
FOR VEGETATIONAL GROUPS ON MATURE JACK PINE STANDS
AND UNBURNED CLEARCUTS
Years
S ite a
Since
Nirber Treatment

Area Annuals & B ien n ials
“p
R
ha
C

>55
>55
>55
^55

12
4
8
3

1A
1B80

0
0

16
16

LB81
2

1

_

1

16
24

3A
3B79

2
2

16
16

_

3B80
4A79

3
3

16
32

3B81
4A80
5A79

4
4
4

16
32
4

4AB1
5A80
6

5
5
5

32
4
16

5A81

6

4

13A
13B
13C
13D

2.1
1.2
.8

.6

11.6

-

-

Trees & Shrubs
C "F
R
5 1.3
41 .9
4 6.9
43.7

9
7
11
14

18
21
23
27

30.7
32-1
42.7
71.6

1

3
6
4
4

.3
8 .5
28.8
41.2

11.6
14.9
25.8
30.5

8

94.1
75.0
56.7
48.4

1
2

32.5
12.9

26.1
20.4

6

-

7

15.0
29.9

31.2
33.3

9
9

52.5
56.7

42.0
4 6 .3

10
13

26
31

30.0
61.5

.5

1

-

-

27.2
24.7

22.5
27.6

5
5

34.3
7 .1

26.7
23.2

8
12

38.5
68.2

50.4
49.2

9
7

23
24

61.5
27.6

1 .9

1

49.6
36.5

43.8
24.8

9
5

21.6
13.6

21.6
32.2

9
10

29.1
49.6

34.6
41.1

9
9

27
25

50.7
37.1

32.0
52.7

7
7

4 .3
5.1

26.0
16.4

42.5
11.4

42.0
30 .9

8

1

53.1
83.4

7

-

6

22
20

4 6.9
37.7

1 .5

1

45.7
68.8
77.2

30.4
50.0
35.7

6
6
2

1.6
.3
2 .2

23.9
10.3
14.3

6
5
1

52.7
30.9
20.7

44.2
39.5
50.0

9
8
7

22
19
10

64.4
60.5
41.7

51.6
37.1
45.6

6
5
3

.1
2 .9
.2

9.5
19.3
2 .9

5
4
3

18.0
29.8
22.5

39.0
43.5
51.5

7
5
10

18
14
16

58.9
44.7
30.1

35.0

4

5 .7

18.0

4

28.7

47.0

7

15

46.8

1

-

-

-

-

-

-

-

-

-

-

-

-

81.8
67.3
77.3

-

-

65.6

-

Perennial Herbs
C
F
R

25.6
28.4
17.5
18.4

.

.1

T otal
Cover

3.5
15.7
13.7
9.7

14.9
9.8
7.4
.7

.49

1
1

Grasses & Sedges
C '
R
P

Total
R

5
7
7

6

aS ite numbers s ta r tin g w ith the same combination o f number and le t t e r represent the sane s i t e surveyed twice; the
nutber 79, 80, and 81 in d ic a te the year surveyed (1979, 1980, and 1981), fo r s it e s surveyed in more than one growing season.

54

TABLE 4.2
PLANT SPECIES ENCOUNTERED ON UNBURNED CLEARCUT
SITES AND MATURE JACK PINE (>55 YEARS)
STANDS IN NORTHERN LOWER MICHIGAN
S p e c i e s E x c l u s i v e t o Unburned C l e a r c u t S i t e s
(18 sp e c ie 's t o t a l )
A n n u als and B i e n n i a l s
(2 s p e c i e s )
HZz4.acZum a a a a n Z Z a c a m L.
A4.abZA gZabn. a L.
LZaZflZa n o v a e - angZZae.
L a c Z u a a canad&nAZA L.
L unell
PhyAaZZA vZ4. gZnZa. na M i l l
G ra sse s and Sedges
SpZ4.anZh&A g4.ac.ZZZA
(5 s p e c i e s )
( B i g e l . ) Beck
Ag4.oAZZA hye.maZZA W a l t .
VZoZa p e d a Z Z ^ Z d a g . Don.
V&AchampAZa ^ Z z x u o A a L.
0Zc . k anZhz ZZa m de. patipz4. a~
turn M u h l.
0 4. y z opAZA aApe.4.Z&oZZa

T re e s and Shrubs
(4 s p e c i e s )

M ichx.
Paa-cccm co£umb^.anam
S chribn.

VZe f i v ZZ Za Z o n Z e e 4 . a M i l l .
P4.unuA v Z a g Z n Z a n a L.
RoAa b Z a n d a A i t .
SaZZx gZaueophyZZoZdzA

P e r e n n i a l Herbs
(7 s p e c i e s )

Fern.

AnZe.nna4.Za m g Z z c Z a

Greene
AAC.Ze.pZa6 A y a . Z a c a L.

S p e c ie s E x c lu s iv e t o M ature J a c k P in e S ta n d s
(8 s p e c i e s t o t a l )
G r a s s e s and Sedges
(1 s p e c i e s )
Mu.hZe.nbe.4.gZa me.x.Zc.ana L.

P e r e n n i a l Herbs
(3 s p e c i e s )
ChZmaphZZa a mb e Z Z a Z a

(L .) N utt
P o Z e n Z Z Z Z a Zn.Zde.nZaZa A i t .
VZoZa a d u n c a Sm.

T r e e s and Sh ru b s
(4 s p e c i e s )
Aceh. A.U.b4.um L.
Ce.anoZhuA o v a Zu A D e s f .
PZnuA AZh.o bu.A L.
RubuA hZApZduA L.

55

TABLE 4.2 (Continued)
S p e c i e s Common t o B o t h U n b u r n e d C l e a r c u t s
and M ature J a c k P in e S ta n d s
(33 s p e c i e s t o t a l )
A nnuals and B i e n n i a l s
(1 s p e c i e s )
iAeZamptjAum Z Z n e a i t e Lara.
G r a s s e s and Sedges
(7 s p e c i e s )
AndAopogo n geAaAdZZ V itm a n
Baomu6 kaZmZ-i G r a y
CaJiex. p e n 6 y Z v a n Z c a Lam.
VanZhonZa 6 pZ c aZ a

(L .)

Beauv.

K0 e.Z2.AZa m a c A a n Z h a P e r s .
0 A y z o p 6 Z 6 pu.nge.n6 T o r r .
Sc . h Z z a . c h n e p u A p u A a 6 c e n 6

T orr.
P e r e n n i a l Herbs
(1 1 s p e c i e s )
Ap o c y n u m a n d A 0 6 aemZ&oZZum L.
A n e m o n e q u Z n q u e f i o Z Z a L.
A6 t e A Z a e v Z 6 L.
C a mp a n u Z a Ao Z u n d Z &o Z Z a L.
pAagaAZa vZngZnZana

Duchesne
G a u Z Z h e A Z a pAo c u m b e n 6 L.
He ZZanZhu6 o c c Z d e n Z a Z Z 6

R idd.
HZeAacZum v e no Aum L.
MaZanZhemum c a n a d e n 6 e D e s f .
PZeAZdZum aquZZZnum D e s f .
SoZZdago spp.

T re e s and Shrubs
(14 s p e c i e s )
Am e Z a n c h Z e A s p .
A A c Z o 6 Z a p h y Z o 6 u, v a - u A6 Z L.
C o mp Z o n Z a p e A e g A Z n a L.
CAaZaegu6 s p .
B p Z g e a A e p e n 6 L.
PZnu6 b a n k 6 Z a n a Lamb.
P o p u Z u 6 Z A e mu Z o Z d e 6 M i c h x .
PAunu6 pumZZa. L.
PAunu6 6 e A o Z Z n a E h r h .
QueAcu6 s p p . ( r e d oak )
Ru.ba4 p e n 6 Z Z v a n Z c u 6 P o i r .
Va c c ZnZum a ng u 6 Z Z ^ o Z Z u m

A it.
V a c c Zn Zu m my AZZZZo Zd e 6

Michx.
\ / ac c ZnZum v a c Z Z Z a n 6 T o r r .

MATURE JACK PINE
UNBURNED CLEARCUTS

80 _

60
C

0
V
E
R

40 M

20

T -

11 I

YEARS AFTER TREATMENT
F i g . A. 2.
T o t a l v e g e t a t i o n a l c o v a r (m), o f s i t e s a v e r a g e d w i t h i n
y e a r s , f o r m a t u r e j a c k p i n e s t a n d s and u n b u r n e d c l e a r c u t j a c k p i n e
s i t e s c o m p risin g a 6 -y e a r sequence f o llo w in g lo g g in g . V e r t i c a l b a rs
e q u a l one SD.

57
species

(T able 4 . 1 ) .

cle arc u t)
to s i t e

F o r e x a m p le , on s i t e

a l a r g e drop i n cover i s

was a l s o

s e e n on s i t e

site

13B l i e s

c le arc u t).

stand)

due to

A l a r g e drop i n b l u e b e r r y

1B80, b u t t h i s was o f f s e t b y an

i n c r e a s e i n A m elanchier sp.
pine

(0-year

s e e n when c o m p a r e d

13D ( a n a d j a c e n t m a t u r e j a c k p i n e

decreased bracken fe rn cover.

1B80

(se rv ic eb e rry ).

a d ja c e n t to s i t e

5A79 ( 4 - y e a r - o l d

I n c r e a s e d c o v er on s i t e

e x trao rd in ary increases

M ature j a c k

5A79 was d u e t o t h e

i n C a r e x c o v e r , w h i c h more t h a n

com pensated f o r th e d r a s t i c d e c r e a s e i n b l u e b e r r y .
gen eral,

In

c o v e r i n c r e a s e d from one y e a r t o t h e n e x t ,

b a s e d on d a t a fro m s i t e s

s u r v e y e d o v e r more t h a n one

grow ing se a s o n .
C o v er o f a n n u a l s and b i e n n i a l s was n e g l i g i b l e
the m ature f o r e s t s

and a l l c l e a r c u t s i t e s

H o w e v e r , cow w h e a t

(Melampyrurn l i n e a r e L a m .) w a s q u i t e

f r e q u e n t on m a tu re j a c k p i n e s i t e s
grass

and sedg e s p e c i e s

(F ig.

in

4 .3 ).

(T able 4 .1 ) .

The

g r o u p sh o w ed e x t r a o r d i n a r y i n ­

c r e a s e s i n dom inance a f t e r c l e a r c u t t i n g .

T h is group

d o m i n a t e d b y C a r e x p e n s y l v a n i c a L a m . , r e p r e s e n t e d 10%
o f t h e c o v e r i n t h e m a t u r e j a c k p i n e u n d e r s t o r y a n d 75%
o f th e cover by th e f i f t h y e a r a f t e r

c le arc u ttin g .

e n n ia l herb cover in c re a s e d s h a rp ly i n i t i a l l y
c le arc u ttin g ,

fo llo w in g

b u t was d r a s t i c a l l y r e d u c e d a s t h e d o m i ­

nance o f Carex i n c r e a s e d .
blueberry)

Per­

T ree and sh ru b s p e c i e s

dom inated m atu re j a c k p in e

(e.g .

s t a n d s , b u t sh o w ed

Unburned clearcuts
100

.

oc u j

*— ^ANNUALS flND 8 1ENNIflLS
Q— 13 GRASSES FIND SEDGES
° °PERENNIAL HERBS
* * TREES AND SHRUBS

80 _

i n — »—< r>

Os.
60 _

uj

* ------c_> cd :» iij ac

40 _

If

,..-0

s

20

\

a

\

V..
T

>55

T
0

t
1

t
2

f~
3

"T "

4

~~T

t -

S

YEARS AFTER TREATMENT

F i g . A . 3.
R e la tiv e cover fo r each s p e c ie s group, o f s i t e s
av erag e d w i t h i n y e a r s , f o r m ature j a c k p in e s ta n d s and unburned
c le a r c u t s i t e s com prising a 6 -y ea r sequence fo llo w in g logging.

59
a ra p id decrease fo llo w in g c le a r c u ttin g .
Burned C l e a r c u t s
V ersu s M ature
Ja ck P in e Stands
A l l burned c l e a r c u t s i t e s w ere of m ature j a c k
pine o rig in .

The f o u r m a t u r e j a c k p i n e s t a n d s s u r ­

veyed in t h i s

stu d y were c o n s id e r e d r e p r e s e n t a t i v e

of the p r e d is tu rb e d c o n d itio n .
D uring th e f i r s t
cu ts

year fo llo w in g b u rn in g ,

sh o w e d g r e a t l y i n c r e a s e d

le v e ls of sp ecies r i c h ­

n e s s compared t o m a tu re j a c k p in e

stands

b u t s p e c i e s r i c h n e s s on t h e s e s i t e s
7D 79-80-81)
b urning.
to

dropped d r a s t i c a l l y

(F ig.

t h a t a t y e a r one, w ere a ls o

(T able 4 . 3 ) .

4,

5,

2 and 3 y e a r s a f t e r
sim ilar

s e e n on s i t e s

1QA79-

and 6 y e a r s a f t e r b u r n i n g

D i f f e r e n t tre n d s developed i n th e average

r i c h n e s s p e r p l o t and Shannon i n d e x .
burned s i t e s

4.4),

(7C 80-81 and

H igh l e v e l s o f s p e c i e s r i c h n e s s ,

8 0 - 8 1 a n d 10B79 - 8 0 - 8 1 ,

c le ar-

O ne-year-old

and m a tu re j a c k p i n e s i t e s w ere s i m i l a r

i n a v e r a g e r i c h n e s s , b u t showed c o n s i d e r a b l e v a r i a t i o n
in th is

index.

A g r a d u a l drop i n a v e ra g e r i c h n e s s

p e r p l o t o c c u r r e d d u r i n g y e a r s 2 a n d 3.
a n d 10B t h e h i g h e s t l e v e l s o f t h i s
d u rin g y e a r s 4,

5 and 6.

On s i t e s

in d ex w ere seen

A ll burned c le a r c u ts ,

a v e ra g e d w i t h i n age c l a s s e s , had h ig h e r

le v e ls o f the

Shannon in d e x compared to m a tu re j a c k p i n e s i t e s .
Trends in t h i s

10A

i n d e x d i d n o t show a n y p a t t e r n , b u t

60

nrttuRC JACK ^ I he
BURNED C ltf lR C U rs

i

SO .

30

.

Z$ ,

X

10 „

>55

10

e
€

4
2

0
t

3

5

4

NATURE JACK PINE
BURNED CLEARCUrs

I

TEARS A M E R

TREATnEHl

F i g . A . 4. A) S p e c i e s r i c h n e s s ( t o t a l n u m b e r
o f s p e c i e s ) , B) a v e r a g e r i c h n e s s p e r p l o t , a n d C)
Shannon in d e x o f d i v e r s i t y , o f s i t e s a v erag e d w i t h i n
y e a r s , f o r m a t u r e j a c k p i n e s t a n d s and b u r n e d c l e a r c u t
s i t e s com prising a 6 -y ear sequence fo llo w in g b u rn in g .
V e r t i c a l b a r s e q u a l one SD.

TABLE 4.3
RELATIVE COVER ( C ) , RELATIVE FREQUENCY ( F ) , AND SPECIES RICHNESS <R)
FOR VEGETATIONAL GROUPS ON MATURE JACK PINE STANDS
AND BURNED CLEARCUTS
Years
Since
S ite
Nunber Treatment

Area Annuals & B ien n ia ls
F
ha
C
R

Grasses & Sedges
F
R
C

Perennial Herbs
C
F
R

Trees & Shrubs
C
F
R

T otal
R

Total
Cover
per 60 m

>55
>55
">55
>55

12
4
8
3

2 .1
1 .2
.8
.6

11.6
14.9
9.8
7.4

1
1
1
1

3 .5
15.7
13.7
9.7

25.6
28.4
17.5
18.4

3
6
4
4

.3
8 .5
28.8
41.2

11.6
14.9
25.8
30.5

5
7
7
6

94.1
75.0
56.7
48.4

5 1.3
4 1 .9
4 6 .9
43.7

9
7
11
14

18
21
23
27

30.7
32.1
42.7
71.6

7C80
7D79

1
1

16
24

18.3
.1

7.1
2.8

4
3

35.8
73.5

32.0
44.6

9
9

2.8
8 .5

26.1
24.3

15
12

43.1
17.8

3 4 .B 7
2 8.1 10

35
34

26.9
30.6

7C81
7D80

2
2

16
24

-

-

-

45 .7
71.6

38.4
56.3

8
9

1 .1
12.4

13.2
21.1

10
8

53.2
15.8

4 8.3
22.5

7
8

25
25

52.6
48.5

7D81

3

24

-

•

-

66 .7

52.5

8

14.3

25.1

8

19.1

22.7

8

24

48.7

1QA79
10B79

4
4

26
23

.

.2

1 .6
3 .9

1
1

71 .8
53 .5

35.7
40.0

13
14

15 .2
22 .9

39.3
23.3

13
10

13.0
23.4

23.3
32.8

9
14

36
39

58.2
45.4

10A80
10B80

5
5

26
23

.1
.8

1.7
5.4

2
3

57.3
5 7 .3

38.2
37.3

11
10

18.5
8 .0

32.9
23.5

16
a

24.1
33.8

27.2
3 3 .9

7
14

36
35

75.7
61.6

10A81
10B81

6
6

26
23

.2
.8

.6
2 .3

2
1

66.4
5 0 .3

39.3
38.3

8
9

8 .9
12.2

34.9
21.4

14
10

24.5
36.7

25.2
38.0

8
14

32
34

74.4
74.8

134
13B
IX
13D

^ e e Table 4 .1 .

_

62
h i g h e s t l e v e l s were a t t a i n e d

6 years a f t e r burning.

B urning s tim u la te d th e e s ta b lis h m e n t o f a la rg e
v a r i e t y o f sp e c ie s n o t p r e s e n t in m ature ja c k p in e
stands

(T able

4 .4 ).

S ev en ty -six d if f e r e n t sp ecies

w ere r e c o r d e d on b u rn e d c l e a r c u t

sites

compared to

41 s p e c i e s o n m a t u r e j a c k p i n e s i t e s .
s p e c ie s w ere e x c l u s i v e

T h irty -n in e

to burned c l e a r c u t s i t e s ,

as o n ly 4 s p e c ie s w ere e x c l u s i v e

w here­

to m ature j a c k p in e

sites.
One y e a r a f t e r f i r e ,

b u rn e d c l e a r c u t s i t e s had

g r e a t l y d e c r e a s e d v e g e t a t i o n a l c o v er compared to m a tu re
ja c k p ine s i t e s

(F ig.

4 .5 ).

However,

larg e

in creases

i n c o v e r o c c u r r e d by y e a r 2, w i t h f u r t h e r i n c r e a s e s
d u r i n g y e a r s 5 a n d 6.

G en erally ,

one y e a r t o t h e n e x t on s i t e s
one grow ing s e a s o n .
to th e se in c re a s e s

c o v e r i n c r e a s e d from

s u r v e y e d o v e r more t h a n

D iffe re n t sp ecies c o n trib u te d
in cover.

For exam ple,

7 C 8 0 - 8 1 , C a r e x , V a c c i n i u m , P r u n u s p u m i l a L.
cherry),

on s i t e
(sand

and O ry z o p s is pungens T o r r. w ere i m p o r ta n t;

w h e re a s on s i t e

7 D 7 9 - 8 0 - 8 1 , F t e r i d i u m a q u i l i n u m L.

(bracken fern )

a n d D i c h a n t h e l i u m d e p a u p e r a t u r n M u h l.

c o n t r i b u t e d m ost to th e c o v e r.

On s i t e

10A 79-80-81

l a r g e i n c r e a s e s w ere n o te d i n C a r e x , V a cc in iu m ,
F te rid iu m a q u ilin u m , Schizachne p u rp u ra sc e n s T o r r . ,
and P ru n u s
site

s e r o t i n a Ehrh.

( b l a c k c h e r r y ) , w h i l e on

1 0 B 7 9 - 8 0 - 8 1 i n c r e a s e d c o v e r was d ue t o C a r e x ,

63
TABLE 4 . 4
PLANT SPECIES ENCOUNTERED ON BURNED
CLEARCUT SITES AND MATURE JACK PINE
<> 55 YEARS) STANDS IN NORTHERN
LOWER MICHIGAN
S p e c ie s E x c l u s i v e to Burned C le a r c u t S i t e s
(" 3 9 ™ sp e c ie s t o t a l )
A n n u a l s and B i e n n i a l s
(7 s p e c i e s )

P e r e n n i a l H erb s
( 1 2 sp e c ie s)

AA.ab-t.4 g Z a b n a L.
CtA.4 -turn s p .
C o n y d a Z Z i i e m p e n v Z n e n i L.
ca n a d ctii-i-i L.
Ge-tantum
B r itt.
Lac-tuea caiiadeaA-iA L.
L-i-t/ioape/imum a n v e m e . L.

A iten ia g L ttlio Z ia i
Wederaeyer
C o n u o Z v u Z u i i p i t t i a m a e u i L.
HcC iait*(temum caiia rfen 4e
M ic h x .
U t e n a c L a m a u x a iU x a c u m L.
Hte^iactum c a iia d e iiA t M ic h x .
L e c h e a m i n o n L.
L Z a tn ii novae-angZLae
L u n e ll
Po Zt jgaZa p o t t j g a m a W a l t .
P o l y g o n u m c i Z Z n o d e M ic h x .
PhtjiaZii vin g in L a n a M ill.
Rumex a c e t o i t t Z a L.
S e a e c x o t o m e n t o i u i M ic h x .

G r a s s e s and S e d g e s
(12 s p e c i e s )
Agnoptjnon t n a c h y c a u Z u m
L in k
Agn.oit.LA hijo.ma.ZLi W a le .
B.tumui kaZmLL G ray
Peac/iampA- ia i l t x u o i a L.
DichanthQtLum depaup&naturn M uhl.
F e iia c a sp.
Koe.Ze.nLa m a e n a n t h a P e r s .
O m jzo p iii aipcnZfioH a
M ic h x .
P aiiicum co£ambAa>:um
S crib n .
Pan-tcum x a n i / i u p b i / i u r a

T r e e s and S h r u b s
(B s p e c i e s )
O Le.nvLtta Z o tU c e n a M i l l .
G aty lu 4 4 a c x a b a c e a i a
(W an g.) C. Koch
P o t e n t l Z Z a a n g u ta Pursh.
P'iunu4 v i n g i n L a n a L.
Ro,ia b Z a n d a A i t .
Si/m p/io /iica^ po 4 adbuA L.

Gray
Poa p n a . t e . n i L i L.
Sci'ig/tai-tftum t i u t a n i L.
S p e c ie s E x c lu siv e

c o M a tu re J a c k P i n e S t a n d s
(A s p e c i e s )

G r a s s e s and S e d g e s
(1 s p e c i e s )
An d n a p o g o n i e o p a n i u i M ic h x .

T r e e s and S h r u b s
(2 s p e c i e s )
PitmA
RubuA

P e r e n n i a l H erbs
(1 s p e c i e s )

CliLmaptiLZa u m b e ZZ a t a
(L .)

N u tt.)

j-t/iebuA L.
L.

biipxdui

64
TABLE A. A ( C o n t i n u e d )

S p e c i e s Common t o B o t h B u r n e d C l e a r c u t s
and M a tu r e J a c k P i n e S t a n d i
(37 s p e c i e s t o t a l )
A n n u a l s an d B i e n n i a l s
(1 s p e c ie s )

P e r e n n i a l H erbs
(13 s p e c i e s )

Mefampi/Aum l i n e a t ie. Lam.

Anemone g u - t u g u e &oZia L.
Apoci/num artdAOAaem-ifioZium L.
AaEca Z a e v i i L.
C am panula / l o t u n d i & o Z i a L.
FAagaA-ia u iA g - in ia n a
D uchesne
G au f-t/icA ia pA ocum bcnl L.
He.ZZauZ.hui o c c . Z d e n t a . Z i i

G r a s s e s and S e d g e s
(7 s p e c i e s )
AudAopogon geAaAd-i-i
V it m a n
BAomuA kaZmZZ Gray
CaAtx p e m y Z o a t i Z c a Lam.
V anthonia i p i c a t a
( L . ) Beauv.
MuhZznbe.ft.QZa m e x i c a n a L.
O t i y z o p i i i pungcnA T o r r .
S c h E z a c f in e puApuAaAceui
Torr.

Ridd.

H ie .f i ac iu m ueiiojum L.
Maianthemum c a n a d e m e
D esf.
PotcntZZZa t t i i d c n t a t a
A it.

Pte.tiidiu.rn aq u ZZZ n u m D e s f .
S o f - id a g o s p .
VZoZa a d u n c a Sm.

T r e e s an d S h r u b s
("l5""ap"ecies)_"
Acca AubAum L.
AmcEanc/ti ca s p .
A t i c t o i t a p h y Z o i uva-uAA-i L,
Ceano-t/iuA cua-tuA D e s f .
C o m p t o n Z a p c n e g / i i n a L.
Ctiataegui sp.
EpEgea AcpcnA L.
PEnuA b a n f e iia t ia Lamb.
P o p u l u i t t i e m u Z o Z d e i M ic h x .
Pauuua pumiEa L.
Pauhua 4£Ao-ti.na E hrh.
Qucacua s p p . ( r e d o a k )
RubuA pou-tE oaH -icuA P o i r .
Vaccinium anguiti& oZium A it.
yaec.in-t.um m y r t t Z Z Z o i d a M ic h x .
Vaccinium vacZZZam Torr.

100

_
HRTURE JACK PINE
2 2 BURNED CLEARCUTS
80

0

*

60

V
E
R
40

20

0
1

2

3

4

S

6

YEARS AFTER TREATMENT
F i g . A . 5.
T o ta l v e g e t a t i o n a l cover (m ), o f s i t e s av erag ed
w i t h i n y e a r s , f o r m a tu re j a c k p i n e s t a n d s and b u rn e d c l e a r c u t
s i t e s com prising a 6 -y ear sequence fo llo w in g b u rn in g .
V ertical
b a r s e q u a l one SD.

66
V a c c i n i u m , and F o p u l u s t r e m u l o i d e s M ichx.

(trem b lin g

aspen).
W h e r e a s c o v e r o f a n n u a l s a n d b i e n n i a l s was
n eg lig ib le

under m ature j a c k p i n e ,

in cover of t h is
(Fig.

4 .6 ).

a sharp

group o c c u r r e d one y e a r a f t e r

age c l a s s .

(7C80 a n d

The c o v e r o f t h e s e s p e c i e s ,

m a in ly G eranium b i c k n e l l i i B r i t t ,
v i r e n s L.

fire

The p r e s e n c e o f a n n u a l s a n d b i e n n i a l s ,

h o w e v e r , w a s d i f f e r e n t o n t h e two s i t e s
7D79) i n t h i s

in crease

and C o r y d a lis sem per-

( r o c k - h a r l e q u i n ) , was e x c e p t i o n a l l y h i g h

( 1 8 .3 % )

on s i t e

on s i t e

7D79.

7C 80, w h e r e a s i t was n e g l i g i b l e
S p ecies in t h i s

(.1% )

group w ere e i t h e r

absent

o r r e d u c e d t o s c a t t e r e d i n d i v i d u a l s on t h e 2 - t o 6y ear o ld burned c le a r c u ts .

D ram atic i n c r e a s e s

in

g r a s s e s and se d g e s w ere s e e n on t h e b u rn e d c l e a r c u t s .
For e x a m p le , on 3 - y e a r - o l d

sites,

r e p r e s e n t e d a n a v e r a g e 66% o f t h e
cover.
tire ly

s p e c i e s group

to ta l v eg etatio n al

The g r a s s a n d s e d g e s p e c i e s g r o u p was n o t e n ­
d o m in a te d b y C a r e x , a s s e e n on t h e u n b u r n e d

clearcu t s ite s .
burned s i t e s
grasses

th is

(e.g .

T h i s was e s p e c i a l l y t r u e on t h e o l d e r

(10A 79-80-81 and 1 0 B 7 9 -8 0 -8 1 ), w here
Q r y z o p s i s a s p e r i f o l i a M ichx.

Schizachne p u rp u ra s c e n s) r e p r e s e n te d
th e s p e c i e s group c o v e r .

and

38% t o 51% o f

In c o n tra s t,

grasses re p re ­

s e n t e d o n l y 1% t o 5% c o v e r o f t h e g r a s s a n d s e d g e
s p e c i e s group on u n b u rn ed c l e a r c u t s .

Burned clearcu ts

100

R

00

_

60

_

A * ANNUALS AMD BIENNIALS
Q— 3 GRASSES AND SEOGES
«— ^PERENNIAL HERBS
*" * TREES AND SHRUBS

E
L

R
T
I
V
E

.-

0'

t3 -

P'

C

0
V
E
R

V '

V

40

;\

-■*

V -~
20

_

►—
! \

0

\
—

v

t-------t-----T------ 1
>55

2

3

4

5

6

YEARS AFTER TREATMENT

F ig . 4.6.
R e la tiv e cover fo r each sp e c ie s group, o f s i t e s
a v e r a g e d w i t h i n y e a r s , f o r m a t u r e j a c k p i n e s t a n d s and b u r n e d
c le a r c u t ja c k pine s i t e s com prising a 6-yaar sequence fo llo w in g
burning.

68
A sharp d e c lin e in cover o f p e re n n ia l h e rb s,
m a i n l y d u e t o d e c r e a s e d c o v e r o f b r a c k e n f e r n , was
s e e n when c o m p a rin g m a tu r e j a c k p i n e s i t e s
o ld burned c le a r c u ts
crease

(years

1 t o 4)

(Fig.

4 .6 ).

to

1-year-

The s u b s e q u e n t i n ­

and d e c r e a s e

(years

4 t o 6)

p e r e n n i a l h e r b c o v e r was a l s o due t o v a r i a t i o n
bracken fe rn cover.

in

in

O ther p e r e n n ia l herb s p e c ie s ,

alth o u g h c o n trib u tin g s u b s ta n tia lly

to s i t e

co n trib u ted l i t t l e

A v e r y l a r g e drop

in tre e

to t o t a l cover.

and sh ru b c o v e r,

rich n ess,

due to t h e l o s s o f b l u e ­

b e r r y , was a l s o n o t e d i n 1 - y e a r - o l d b u r n e d c l e a r c u t s .
Cover o f t h i s

s p e c ie s group g e n e r a l l y d e c l i n e d th ro u g h

y e a r 4, b u t s h a r p l y i n c r e a s e d a t y e a r 5 due t o i n c r e a s e s
in aspen,

black ch erry , b lu eb e rry ,

a n d d e w b e r r y (R u b u s

p en silv an icu s P o ir .) .
B urned V ersus
Unburned 35- Y e a rO ld J a c k P in e
A l l b u rn e d and unburned a r e a s were p a r t of a
l a r g e r a r e a consumed by w i l d f i r e

in

1946.

The f o r e s t

t h a t s u b s e q u e n t l y d e v e l o p e d w a s l o g g e d i n 19 75 a n d
1976 o f f i n a n c i a l l y m a t u r e j a c k p i n e ,
oak.
pine

T h is a re a ,

red p in e,

d o m in a ted by i n t e r m e d i a t e - a g e d j a c k

( r e p r e s e n t i n g p o s t f i r e r e g e n e r a t i o n ) , was d i v i d e d

in to blo ck s;

some o f t h e s e b l o c k s w e r e p r e s c r i b e

b u r n e d and o t h e r s have re m a in e d u n b u rn e d
site

and

7A80-81 f o r c o m p l e t e d e t a i l s ) .

(s e e C h a p te r 2,

Five b lo c k s ,

69
3 b u r n e d and 2 u n b u rn e d , have b een s u r v e y e d and a r e
u se d to c o n t r a s t d i f f e r e n c e s i n unburned and b u rn ed
s t a n d s , m a k i n g up a p o s t f i r e

sequence o f 1 to 5 y e a rs

(T able 4 .3 ) .
B urned j a c k p in e

s ta n d s w ere c o n s id e r a b ly r i c h e r

i n s p e c i e s one y e a r a f t e r f i r e
(F ig.

4 .7 ),

b u t a l a r g e drop in s p e c ie s r i c h n e s s

o c c u r r e d betw een y e a r s
7B 79-80).
years

S ite

1 and 2 ( s i t e s

7B d r o p p e d f r o m 45 t o

1 and 2, a l t h o u g h s i t e

unchanged

th a n th e unburned sta n d s

(T able 4 .5 ) .

7A80-81 and
30 s p e c i e s b e t w e e n

7A r e m a i n e d e s s e n t i a l l y

A steady in c re a se in species

r i c h n e s s o c c u r r e d from y e a r s 2 t o 5, m a i n l y due t o
site

9A79-80-81

(3-,

4-,

and 5 - y e a r o ld b u r n ) .

A drop

in average ric h n e s s p er p lo t occurred d u rin g years
1 and 2,

f o l l o w e d by i n c r e a s e d l e v e l s o f t h i s

d uring y ears

3,

4,

a n d 5.

the f i r s t year a f t e r f i r e ,
3,

index

The S h a n n o n i n d e x w a s h i g h e r
b u t low er d u r in g y e a r s 2,

a n d 4 , f o l l o w e d by a s h a r p i n c r e a s e a t y e a r 5.

s e v e n s p e c i e s w e re r e c o r d e d on t h e b u r n e d s i t e s
pared to

43 s p e c i e s o n t h e u n b u r n e d s t a n d s

S ixty -

com­

(T able 4 .6 ) .

A l a r g e drop i n c o v e r o c c u r r e d as a r e s u l t o f
burning

(Fig.

4 . 8 ) , b u t t h e o p e n in g s c r e a t e d by f i r e

w ere v e ry r a p i d l y re o c c u p ie d ,
creases

re s u ltin g in larg e in ­

i n c o v e r d u r i n g y e a r s 2 a n d 3.

y e a r l y on e a c h s i t e
ing season.

Cover in c r e a s e d

s u r v e y e d o v e r more t h a n one grow­

TABLE 4.5
RELATIVE COVER ( C ) , RELATIVE FREQUENCY ( F ) , AND SPECIES RICHNESS (R)
FOR VEGETATIONAL GROUPS ON UNBURNED 35-YEAR-OLD JACK PINE STANDS
AND BURNED 35-YEAR-OLD JACK PINE STANDS

S itea

Years
Since

Total
ha

C

F

R

C

F

R

C

F

R

C

F

R

Total
Cover

L2A
12B

35
35

16
24

.1
-

.9
2.6

1
1

20.5
26.1

33.1
36.5

10
9

1.8
3 .5

24.1
21.9

14
11

77.6
70.4

42.0
39.1

11
8

36
29

52.7
51.4

7A80
7B79

1
1

19
26

14.2
25.6

13.3
12.0

4
5

30.5
27.0

34.4
34.8

10
13

2 .5
8 .1

13.3
18.0

12
13

52.8
38.7

39.0
34.8

10
14

36
45

35.8
28.7

7A81
7B80

2
2

19
26

.1
-

2 .3
1 .1

3
2

47.6
60.8

37.4
44.0

10
10

.9
12.9

12.4
14.7

11
10

51.4
26.3

47.8
40.1

10
8

34
30

49.8
43.7

7B81
9A79

3
3

26
31

.1
.6

2.2
-

3
2

58.8
63.7

48.9
41.0

11
12

4 .8
1 .2

12.4
14.9

10
12

36.2
34.4

36.5
44.2

9
12

33
38

60.9
54.9

9A80

4

31

-

.5

1

62.7

40.7

11

1.2

18.0

13

36.0

40.7

11

36

56.7

9A81

5

31

.15

.51

3

49.2

37.2

10

3.6

20.4

13

47.1

41.8

13

39

65.6

o

aSee Table 4 .1

71
00
3S- TEA* JACK MKE
BURNED 3&+YCAA JACK PJNE

50

30

20
0
0
12

35

I

2

3

B

4

5

□
3S-TEAR JACK PfKE
E 3 BURNED 3 5 - T f f l * JACK PtNC

35

i

2

3

4

5

«
3$-<EAft JACK P M C
BURNED 33-TEAK JACK MNE
J

2

0
35

I

2

3

4

S

YCAA3 ATTC* tHEflmCNT

F ig. 4 .7 .
A) S p e c i e s r i c h n e s s , B) a v e r a g e r i c h n e s s
p e r p l o t , a n d C) S h a n n o n i n d e x o f d i v e r s i t y , o f s i t e s
averaged w i t h in y e a r s , f o r unburned 3 5 - y e a r - o ld ja c k
p i n e s t a n d s and b u rn e d 3 5 - y e a r - o l d j a c k p i n e s t a n d s
c o m p risin g a 5 -y e a r sequence fo llo w in g b u rn in g .
V e r t i c a l b a r s e q u a l o n e SD.

72

TABLE 4 . 6
PLANT SPECIES ENCOUNTERED ON BURNED AND
UNBURNED 3 5 -YEAR-OLD JACK PINE STANDS
IN NORTHERN LOWER MICHIGAN
S p e c ie s E x c lu s iv e t o Burned A reas
(28 s p e c i e s t o t a l )
A nnuals and B i e n n i a l s
(7 s p e c i e s )
CZn.&Zum s p .
Con. ydaZZ& &z m p z n . v Z n . z n i
En. Zgzn. on c a n a d z n & Z i L.
Gzn. anZum b Z a k n z Z Z Z Z

Ante.nnaA.Za ne.gZe.ata

Greene
L.

B ritt.
Kf l Zg Za
Q4.n-Lc.GL L.
Lac.tac.ci c a n a d z n i Z i L.
S-t£ e n e a n t 4 . f i n . h t n a L.

G rasses and Sedges
(5 s p e c i e s )
Agn. opy n. on t n . a a h y c . a a Z a m

L ink
C a f i z x f i u g o &p z f i m a Mack.

F e ^ -tu c a s p .
Koe. Ze. f ita m a a n . a n t k a P e r s .
S o n . g h a i t n . u m n u t a n i L.

P e r e n n i a l Herbs
(1 1 s p e c i e s )

C a m p a n u Z a n o t u n d Z f i o Z Z a L.
ConvoZvuZui ip Z th a m a zu i

L.
E q u Z i z t u m h y m a Z z L.
HZz n. acZum a u n . a n t Z a c . u m L.
L e c /ie a mZnon. L.
MaZanthzmum c a n a d z m z

D esf.
PoZygonum aZZZnodz

M ichx.
V Z o Z a p z d a t Z & Z d a G.

T re e s and Shrubs
(5 s p e c i e s )
Cfiatazgui sp.
Ro-ia b Z a n d a L.
SaZZx gZauaophyZZoZdzi

Fern.
Si/mp/io^t.Lca^po4 a Z b u i L.

Anemone. q u Z n q u z f i o Z Z a L.
Anemone n.Zpan.Za F e r n .

S p e c ie s E x c l u s i v e to Uhburned A re as
(4 s p e c i e s )
P e r e n n i a l Herbs
(4 s p e c i e s )
CypfiZpzdZum a a a u Z z A i t .
G a u Z t h z f i Z a ph.o a u m b z n A L.
PoZygaZa poZygama W alt.
S p Z n a n t h z i gfiacZZZi

(B ig el.)

Beck

Don.

73

TABLE 4 . 6

(C ontinued)

S p e c i e s Co mmon t o B o t h B u r n e d a n d
U n b u rn ed 3 5 -Y ear-O ld J a c k Pine
(39 s p e c i e s t o t a l )
A nnuals and B i e n n i a l s
(1 s p e c i e s )
Me.ZampyH.um Z Z n z a n z Lam.

G ra sse s and Sedges
(12 s p e c i e s )
A n d n . o p o g o n gzn.aH.dZZ

HZz n. acZum v z n o A u m L.
LZatnZA n o v a z - a n g Z Z a z

L unell
PoZznZZZZa Zn.ZdznZaZa A i t .
PZznZdZum aquZZZnum D e s f .
S z n z c Z o ZomznZoAUA M ic h x .
SoZZdago sp.
V Z o Z a a d u n c a Sm.

V itm an
A n d a o p o g a n Ac. opan.Zu6

M ichx.
Bn.omu6 k a t m Z Z G r a y
Ca n . z x p z n A y Z v a n Z c a Lam.
VanZhonZa ApZcaZa

( L . ) Beauv.
V z a c h a m p A Z a f i Z z x u o A a L.
VZckanZhzZZum d zp a u p zn a t urn M u h l.
M u h Z z n b z n . g Z a me. xZc. ana L.
On. yzopAZA a A p z n . Z &o Z Z a

M ichx.
On. y zopAZA p u n g z n A T o r r .
PanZcum coZumbZanum

S cribn.
S c h . Z z a c . k n z pun.pun.aAcznA

T orr.

A p o c y n u m an d n. o6 a e . mZfioZZum L.
AAZzn. j un c Z&o n. mZA R ydb.
Aa Z&h. Z a z v Z A L.
Fn.agan.Za v Z n g Z n Z a n a

Duchesne
HzZZanZhzmum c a n a d z n A Z

M ichx.
R idd.

An. c Zo AZap h . y ZoA u v a - uh.a Z
Amz Z a n c h Z z n . s p .
CzanoZhuA ovaZuA D e sf.
C o m p Z o n Z a p z n . z g n . Z n a L.
PZnuA b a n k A Z a n a Lamb.
P n u n u A p u m Z Z a L.
Pn u n u A a z n o Z Z n a E h r h .
Pn u n u A v Z n g Z n Z a n a L.

QuzncuA s p p . ( r e d o a k )
Rubui p z n A Z Z v a n Z c u A P o i r .
VaccZnZum

anguAZZ&oZZum

A it.
VaccZnZum mynZZZZoZdzA

P e r e n n i a l Herbs
(13 s p e c i e s )

HzZZanZhuA

T rees and Shrubs
(13 s p e c i e s )

occZdznZaZZA

M ichx.
1/ ac cZn Zu m v a c Z Z Z a n A T o r r .

L.

100

^
7ZA

0
V
E
R

35-YEAR JACK PINE
BURNED 35-YEAR JACK PINE

60 _

40 _

20

.

0

35

1

2

3

4

5

YEARS AFTER TREATMENT
Fig. 4 .8 .
T o ta l v e g e t a t i o n a l cover (m), o f s i t e s averaged
w i t h i n y e a r s , f o r u n b u r n e d 3 5 - y e a r - o l d j a c k p i n e s t a n d s and b u r n e d
3 5 -y e a r -o ld ja c k p in e sta n d s com prising a 5 -y e a r sequence fo llo w in g
burning.
V e r t i c a l b a r s e q u a l one SD.

75
A la r g e in c r e a s e i n cover o f a n n u als and b i ­
e n n i a l s o c c u r r e d d u r i n g th e f i r s t y e a r on b u rn ed
sites

(F ig.

4 .9 ).

T h is sp e c ie s

geranium and ro c k h a r l e q u i n ,
the

group,

d o m in a te d by

a v e r a g e d 20% c o v e r on

1 - y e a r - o l d b u m s , w i t h a maximum o f 25% o n s i t e

7B79.

A d r a s t i c r e d u c t i o n i n t h e s e s p e c i e s was s e e n

by y e a r 2.

Large i n c r e a s e s

in grasses

o ccu rred in the e a r ly p o s t f i r e
7A80-81 and 7B 79-80,
burns).

and s e d g e s

sequence

(e.g .

sites

s u rv e y e d as 1- and 2 - y e a r - o l d

G r a s s e s and s e d g e s r e p r e s e n t e d a p p r o x i m a t e l y

60% o f t h e v e g e t a t i o n a l

co ver 3 and 4 y e a r s a f t e r

fire,

b u t t h e i r c o v er dropped betw een y e a r s 4 to 5 ( s i t e
9A 80-81).

P e re n n ia l herbs c o n trib u te d l i t t l e

t o t a l v e g e t a t i o n a l c o v e r on t h e s i t e s .
t r e e and sh ru b s p e c i e s ,
and sw eet f e r n

The c o v e r o f

such as b l u e b e r r y ,

sand c h e rry ,

(C o m p to n ia p e r e g r i n a L . ) was d r a s t i c a l l y

r e d u c e d t h r o u g h y e a r 4 on t h e b u r n s i t e s .
f i f t h year in creases

in th is

cherry,

oak, w illow

sand c h e rry ,

lo id e s F e rn .)

to th e

group,

By t h e

attrib u ted

to b lac k

(S a lix g lau co p h y l-

and dew berry, had o c c u r r e d .

G e r a n i u m was a d o m i n a n t c o n t r i b u t o r t o t h e c o v e r
on f i r s t

y e a r b u r n s , b u t was a b s e n t o r r e d u c e d to

s c a t t e r e d i n d i v i d u a l s on t h e s e s i t e s
ch aracterize
years

th ereafter.

To

th e s p e c i f i c v e g e t a t i o n a l changes betw een

1 and 2 on p a t c h e s d o m in a te d by g e ra n iu m ,

3 perm a­

n e n t 20 m c o v e r t r a n s e c t s w e r e e s t a b l i s h e d o n s i t e

Burned 3 5 -y e a r jack pine

100

S F i< o n

m e —i —i D r * m 3 3

a—

^ANNUALS AND BIENNIALS
Q—3 GRASSES ANO SEDGES
°
^PERENNIAL HERBS
*■ * TREES AND SHRUBS

80

—-B .

60

40

'ik-

20

-----

0
35

1

2

3

4

5

YEARS AFTER TREATMENT

Fig. 4.9.
R e la tiv e cover f o r each s p e c ie s group, o f s i t e s
a v e r a g e d w i t h i n y e a r s , f o r u n b u r n e d and b u r n e d 3 5 - y e a r - o l d j a c k
p in e sta n d s com prising a 5 -y e a r sequence fo llo w in g bu rn in g .

77
7A80.

T a b l e 4 . 7 show s t h e t o t a l a n d r e l a t i v e

to ta lle d

cover,

f o r 60 m, d u r i n g 1980 ( 1 - y e a r - o l d b u r n )

1981 ( 2 - y e a r - o l d b u r n ) .

and

I n 1 9 8 0 , a t o t a l o f 14 s p e c i e s

c o m p rise d th e 5 0 .6 m c o v e r on th e 3 t r a n s e c t s

(P late

4 .1 ).

fo llo w ed

G e r a n i u m r e p r e s e n t e d 74% o f t h e c o v e r ,

by b l u e b e r r y
In 1981,

(10%),

(P late

(6%),

and C a rex

a la rg e d ecrease in v e g e ta tio n cover

o c c u r r e d on t h e s e
a larg e

sand c h e rry

tran sec ts;

( 3 3 . 8 m)

t h i s was a c c o m p a n ie d by

in cre ase in species ric h n e ss
4 .2 ).

(3%).

(24 s p e c i e s )

G e r a n i u m w a s r e d u c e d t o u n d e r 1% c o v e r ,

w h e r e a s s e d g e a n d b l u e b e r r y i n c r e a s e d t o 41% a n d 34%
cover,

resp ectiv ely .

a l b u s L.

Three s p e c i e s ,

Sym phoricarpos

( s n o w b e r r y ) , P r u n u s v i r g i n i a n a L.

(choke

c h e r r y ) , a n d r o c k h a r l e q u i n w e r e p r e s e n t i n 1980 b u t
w ere m i s s i n g from th e

1981 s u r v e y .

i n 19 8 1 on t h e s e t r a n s e c t s
5 p ere n n ial herbs,

New s p e c i e s p r e s e n t

in clu d ed 1 annual,

and 2 s h r u b s ,

5 grasses,

b u t none of th e s e

s p e c i e s r e p r e s e n t e d m o r e t h a n 2% o f t h e t o t a l c o v e r .
A r e a s B u r n e d By
W ild fires
T he s i t e s

in clu d ed fo r t h i s

stu d y were 2 a re a s

o f s t a n d i n g m atu re j a c k p in e and 1 c l e a r c u t a r e a
( 1 - y e a r - o l d o f m ixed j a c k p i n e and oak) b u r n e d by w i l d ­
fire.

These 3 s i t e s

d i s p l a y e d a w ide ra n g e i n s p e c ie s

c o m p o s i t i o n and dom inance
site

(T able 4 . 8 ) .

8 A 7 9 - 8 0 - 8 1 sh o w ed t h e h i g h e s t

For exam ple,

sp ecies ric h n e ss

TABLE 4 . 7
COMPOSITE DATA FROM 19 80 AND I 9 8 I SURVEY
OF 3 PERMANENT COVER TRANSECTS (60m)
ESTABLISHED IN GERANIUM (GERANIUM
BICKNELLII) PATCHES ON SITE 7A
Cover
19 80
T o ta l(m )
R e la tiv e (Z )

S p e c ie s

Cover
1981
T o ta l(m )
R e la tiv e (Z)

S p e c ie s p r e s e n t in
1980 and 1981
Gcnanium b i c k n c t t i i
P^iunuA p u m i l a „
Vacc-inium i pp.
Ca/t ex 4 P P - ^
Ccaiio-Cliua o v a -tu i
Aiten. t a c v i i
Amctanchicn. i p .
Vic.hanthe.ilum
d c p a u p e . u a turn
L ith o ip e n m u m aa.ve.mc
C im ium ip .
C a m p a n u l a n o t u n d i (Jo l i a

37.6
3.0
5.3
1.7

.2
.1

74.2

6.0

.2
2.7

10. A

11.2

3.3
.4

13.5
.2

.2

.1

.2

.4

.5

.3

.7
.4
1.9

1.2

.2
1. 0

.6

8.2
33 . 9
41.0
.7
.2
1.6

.1

3.7
.4

.5

1. 6

.2

S p e c i e s p r e s e n t i n 1980
b u t a b s e n t in 1981
Symphon.ican.poi a t b u i
Paunui v i n g i n i a n a
C a n i j d a t i i ic mpen.vin.cni
S p e c ie s p r e se n t
1981 o n ly

.6
.3
.1

1.2
.6
.2

in

S tite tio to m c n to iu i
Qnyzcrpiii p u n g c m
K o c i c n i a mac^a*ii/ia
Comp^otua p e 'ie g - t t n a
F 'la g a 'iia v i n g i n i a n a .
Soiidago ip .
A ndoitaphtjioi u va -u m i
Andnopogon gcn.an.dii
H i e n a c i u m v e n o iu m
L c c h c a minon
Panicum caiu m b ia n u m
Vanthonia ip i c a t a
Mctampijnum t i n e a n c

1.4

1.1
.4

T o ta ls
50.6

-

in d ic a te s

a co ver v a lu e

33.8

( t o t a l or r e l a t i v e )

le ss

th an . 0 5 .

" I n c l u d e s V. a n g u s t l f o l i u m . V. m y r t i l l o i d e s . and V. v a c i l l a n s .
■ ^Includes C.

p e n s v l v a n i c a and C. r u g o s p e r m a .

79

P l a t e A .I.
C o v e r t r a n s e c t (20 m) t h r o u g h a
g eran iu m p a t c h on 1 - y e a r - o l d p r e s c r i b e d b u rn s i t u .

P la te 4 .2 .
Above c o v e r t r a n s e c t a t y e a r 2
a f t e r f i r e ; n o te d is a p p e a ra n c e o f geranium .

TABLE 4 . 8

RELATIVE COVER ( C ) t RELATIVE FREQUENCY ( F ) , AND SPECIES RICHNESS (R)
FOR VEGETATIONAL GROUPS ON AREAS BURNED BY WILDFIRE

S ite3
Nunber
8A79

Years
Since

2

T otal
ha

C

F

R

C

F

R

C

R

R

C

32

.1

1.8

4

44.0

39.9

14

2 .8

17.6

16

53.0

F

R

40.5

13

47

Total
Cover

47.4
00

8A80
«79

3
3

32
30

.1
-

2.2

2
-

39.4
60.2

33.2
49.6

12
7

6 .6
1 .1

23. L
3 .5

15
5

54.2
38.5

41.5
47.6

11
6

40
18

52.8
42.9

8A81
9B80

4
4

32
30

.2
-

1.8

3
-

43.7
71.8

38.6
50.0

14
7

7 .6
.7

19.8
9 .9

15
8

48.5
27.1

39.9
40.1

12
6

44
21

61.7 ‘
54.7

9BS1

5

30

-

-

62.7

4 2 .9

6

-

2.5

2

37.3

54.6

6

14

56.2

11

6

6

-

-

91.8

59.2

6

.5

10.3

1

7.7

30.8

6

13

31.2

^ e e Table 4 .1 .

o

81
o f any b u rn e d s i t e ,

in c lu d in g p r e s c r ib e burned c l e a r -

c u ts and 3 5 - y e a r - o ld s t a n d s ,
4.

In c o n tr a s t,

site

d u r i n g y e a r s 2,

9B 79-80-81 and s i t e

l e a s t s p e c i e s r i c h o f any b u rn e d s i t e
3,

t h a t com paring t h e s e s i t e s

sequence, w ith
less

It

seem s,

along a tem poral

d a t a a v e r a g e d w i t h i n y e a r s , w o u l d be

in fo rm a tiv e than lo o k in g a t each s i t e
S ite

11 w e r e t h e

during y e ars

4, and 5 , and y e a r 6 , r e s p e c t i v e l y .

th erefo re,

3, and

in d iv id u a lly .

8A 79-80-81 was u n iq u e b e c a u s e o f t h e h i g h

d e n s ity o f sta n d in g snags th a t in c re a s e d shading to
the f o r e s t

flo o r.

A lso,

th is

site

came c l o s e r t o

ap p ro ach in g adequate s to c k in g of ja c k p in e r e g e n e r a tio n
th a n any o t h e r b u rn ed o r c l e a r c u t s i t e
th is

study.

I n 1 9 8 0 , j a c k p i n e r e g e n e r a t i o n was f o u n d

i n 13 o f 30 f r e q u e n c y p l o t s .
c o v e r on t h e s i t e
years

1 to

3;

sp ecies c o n trib u tin g to t h is

bracken fe rn .

As s e e n o n o t h e r b u r n e d s i t e s

of sim ilar

and p e r e n n i a l h e r b s c o n ­

to th e v e g e t a t i o n c o v er.

However,

s i t e was e s p e c i a l l y r i c h i n p e r e n n i a l h e r b s .

q u en tly o ccu rrin g sp e cie s in
f e r n , A s t e r l a e v i s L.
tosus

increase

O ry z o p sis p u n g e n s , and

a n n u a ls and b i e n n i a l s

trib u ted l i t t l e
th is

The t o t a l v e g e t a t i o n a l

i n c r e a s e d from 4 7 .4 m t o 6 1 .7 m from

in clu d ed C a rex , b lu e b e rry ,

ages,

surveyed in

Fre­

t h i s group were b r a c k e n

(sm o o th a s t e r ) , S e n e c io tomen-

( w o o l y r a g w o r t ) , a n d Apocynum a n d r o s a e m i f o l i u m L.

(spreading dogbane).

G rasses and sedges com prised

82
b e t w e e n 39% a n d 44% o f t h e c o v e r o n t h i s
the

3 grow ing s e a s o n s .

site

during

D om inant g r a s s s p e c i e s on t h i s

s i t e w e r e A n d r o p o g o n s c o p a r i u s M i c h x . , A. g e r a r d i i V i t man, S o r g h a s tr u m n u t a n s L . , and O r y z o p s i s p u n g e n s .
t e r e s t i n g l y , C a r e x c o v e r on t h i s
lo w e s t o f any b u rn e d s i t e

s i t e w a s among t h e

o f analogous age.

s h r u b s r e p r e s e n t e d 48% t o 53% c o v e r on t h i s
th e 1979,

1 9 8 0 , a n d 1981 g r o w i n g s e a s o n s ;

sp e cie s included b lu e b e rry ,
On s i t e

sand c h e r r y ,

9B79-80-81 and s i t e

In­

T r e e s and
site

during

t h e dom inant
and sw eet f e r n .

11, C arex p e n s y lv a n ic a

r e p r e s e n t e d f r o m 60% t o 86% c o v e r , r e s p e c t i v e l y .

For

a l l p r a c t i c a l p u r p o s e s t h e s e s i t e s became C a re x "mead­
o w s " , o n w h i c h o t h e r v e g e t a t i o n was s u p p r e s s e d o r
excluded.
S u c c e s s i o n a l T r e n d s on
Unburned C l e a r c u t S i t e s ,
B urned C l e a r c u t Site~iT
and Burned 3 5 -Y ear-O ld
S ites
The i n d e x o f s i m i l a r i t y

is

used h ere to

test

d if­

f e r e n c e s i n s p e c i e s c o m p o s itio n betw een s i t e p a i r s ,
i n c l u d i n g t h e same s i t e

s u r v e y e d more t h a n o n c e a s w e l l

a s d i f f e r e n t s i t e s w i t h i n t h e same a g e c l a s s a n d b e t w e e n
d i f f e r e n t age c l a s s e s .

The i n d e x o f s i m i l a r i t y

(S)

com puted as f o l l o w s :
2C
A + B
w here A i s

t h e number o f s p e c i e s

i n s a m p l e A, B i s

is

83

t h e number o f s p e c i e s

i n s a m p l e B, a n d C i s

o f s p e c i e s common t o b o t h s a m p l e s

t h e number

(Odum 1 9 7 1 ) .

in d e x o f s i m i l a r i t y r a n g e s from 0 to

The

1; 1 i n d i c a t e s

i d e n t i c a l s p e c i e s c o m p o s i t i o n oh t h e s i t e s .
T a b l e 4 . 9 shows t h a t t h e d e g r e e o f s i m i l a r i t y
among u n b u r n e d c l e a r c u t
o ften less
c la sses.
2 -y ear-o ld

th a n t h a t o f s i t e s w i t h i n d i f f e r e n t age
For exam ple,
sites

is

t h e a v e r a g e S v a l u e among t h e

0 .6 5 , w hereas the

for 2 -y ear-o ld s it e s
0 .7 0 .

s i t e s w i t h i n an a g e - c l a s s was

compared to 3 - y e a r - o l d s i t e s

E ven more s t r i k i n g

among 4 - y e a r - o l d

sites

f o r 4- and 5 - y e a r - o l d

average S value

is
site

is

is

th a t th e average S valu e

0 .5 6 , w hereas i t

is

0.64

co m p ariso n s and 0 .6 9 f o r

4- and 6 -y e a r o ld c o m p a riso n s.

T h is p a t t e r n a ls o occurs

i n t h e 3- a n d 5 - y e a r s e q u e n c e .
When c o m p a r i s o n s w e r e made b e t w e e n c o n s e c u t i v e
years,

in tra site

(sa m e s i t e )

sim ila rities

among u n b u r n e d

c l e a r c u t s i t e s w e r e much g r e a t e r t h a n i n t e r s i t e
(d ifferen t
site

sites)

sim ila rities

(T able 4 .9 ) .

For exam ple,

1 B 8 0 - 8 1 was s u r v e y e d a s a 1 - a n d 2 - y e a r - o l d u n ­

b u r n e d c l e a r c u t a n d had an S v a l u e o f 0 .7 4 when t h e
d a t a f r o m two y e a r s w e r e c o m p a r e d .
d ifferen t site s

In c o n tra st,

com pared w i t h i n 1- and 2 - y e a r age

c l a s s e s had an a v e ra g e S v a lu e o f 0 .5 9 .
t i o n s h i p was fo u n d f o r a lm o s t a l l
site

T his r e l a ­

in tra site

com parisons betw een c o n s e c u tiv e y e a r s .

and i n t e r ­
O v erall,

TABLE 4 . 9
YEAR-BY-YEAR COMPARISONS OF THE MEAN INDEX
OF SIMILARITY (S) FOR UNBURNED CLEARCUT
SITES COMPRISING A 6-YEAR SEQUENCE, AND
THE MEAN INTERSITE (DIFFERENT SIT E S) AND
MEAN INTRASITE (SAME SITE) INDEX OF
SIMILARITY OF SITES COMPARED AT
CONSECUTIVE YEARS
1

sar

0

0

.74

1

.62

.64

2

.67

. 63

.65

3

.60

.61

.70

. 62

4

.5 6

.59

.63

.62

. 56

5

.53

.57

.60

. 61

. 64

. 59

6

. 53

.60

.58

.61

. 69

. 63

2

3

4

I n t e r s i t e versus i n t r a s i t e d iffe re n c e s in
s i m il a r it y a t consecutive y e ars.
Year 0 - 1
m ean i n t e r s i t e
m ean i n t r a s i t e

= .59
= .74

Year 1 - 2
m ean i n t e r s i t e
m ean i n t r a s i t e

= .63
= -

Year 2 - 3
m ea n i n t e r s i t e
mean i n t r a s i t e

= . 71
=* . 6 8

5

the index

Year 3 - 4
mean i n t e r s i t e
m ea n i n t r a s i t e
Year 4 - 5
mean i n t e r s i t e
m ean i n t r a s i t e

= .56
= .75

= .61
=* . 77

Year 5 - 6
m ean i n t e r s i t e = . 5 3
mean i n t r a s i t e =* . 8 3

85
t h e m ea n S v a l u e f o r a l l

in tra site

and i n t e r s i t e

com pari­

s o n s b e tw e e n c o n s e c u t i v e y e a r s was 0 .7 5 and 0 . 6 0 ,
resp ectiv ely .

T herefore,

from d i f f e r e n t s i t e s

o v e r a t e m p o r a l s e q u e n c e w i l l show

a g reater d isp a rity
of s ite

su ccessio n a l d ata c o lle c te d

i n s p e c i e s c o m p o s itio n by v i r t u e

d i f f e r e n c e s t h a n would s i m i l a r d a t a c o l l e c t e d

o v e r t i m e o n t h e same s i t e .
T h e s e d a t a show t h a t e a c h s i t e ,
its

d istu rb an c e age,

The S v a l u e o f t h e

re g a rd le ss of

has a unique s p e c ie s

same s i t e c o m p a r e d a t d i f f e r e n t

was g r e a t e r t h a n t h a t o f d i f f e r e n t s i t e s
d istu rb an ce ag e.
v alue fo r years

co m position.
ages

o f t h e same

To i l l u s t r a t e , t h e mean i n t r a s i t e
3-4 i s

S

0 .7 5 , w hereas th e w ith in - y e a r

S v a l u e f o r y e a r 3 a n d y e a r 4 was 0 . 6 2 a n d 0 . 5 6 , r e s p e c ­
tiv e ly .
sites

T h is r e l a t i o n s h i p

e x i s t e d on u n b u r n e d c l e a r c u t

fo r every y ear o f the sequence.

m ean w i t h i n y e a r S v a l u e
c o m p a r e d t o a m ea n

In f a c t ,

for th ese s ite s

the

was 0 . 6 3 ,

i n t r a s i t e v a lu e o f 0 .75.

Burned c l e a r c u t s i t e s

o f t h e same a g e w e r e g e n ­

e r a l l y no t as s im ila r to each o th e r as burned c le a r c u t
sites

o f d i f f e r e n t ages

can be seen in y e a rs

1,

(T able 4 .1 0 ) .
2, 4,

and 5.

T his r e l a t i o n s h i p
A lso,

in tra site

s i m i l a r i t y was c o n s i s t e n t l y h i g h e r t h a n i n t e r s i t e
s i m i l a r i t y w he n s i t e

c o m p a r i s o n s w e r e made b e t w e e n

co n secu tiv e y e ars.

The m e a n i n t r a s i t e

S v alu e

(0 .73)

fo r burned c le a r c u t

s i t e s w a s much h i g h e r t h a n t h e

86

TABLE 4 . 1 0
YEAR-BY-YEAR COMPARISONS OF THE MEAN INDEX
OF SIMILARITY ( S ) FOR BURNED CLEARCUT SITES
COMPRISING A 6-YEAR SEQUENCE, AND THE MEAN
INTERSITE (DIFFERENT SIT ES) AND MEAN
INTRASITE (SAME SITE) INDEX OF
SIMILARITY, 'OF SITES COMPARED
AT CONSECUTIVE YEARS

ar

1

1

.5 8

2

.58

.60

3

.60

.67

-

4

.5 4

.53

,49

. 68

5

.51

.47

.47

.69

.54

6

.52

.5 5

.49

.67

. 65

2

3

4

5

6

. 65

I n t e r s i t e v e rsu s i n t r a s i t e d if f e r e n c e s in th e in d ex o f
s i m i l a r i t y a t co n secu tiv e y e a rs.
Year 1 - 2
m ean i n t e r s i t e
m ean i n t r a s i t e
Year 2 - 3
m ean i n t e r s i t e
m ean i n t r a s i t e

= .52
.64

Year 4 - 5
mean i n t e r s i t e
m ean i n t r a s i t e

= . 62
= .76

.57
.78

Year 5 - 6
me a n i n t e r s i t e
me a n i n t r a s i t e

= . 55
= .74

“

Year 3 - 4
m ean i n t e r s i t e
m ean i n t r a s i t e

= . 49
=

-

87
m ean w i t h i n y e a r S v a l u e

(0 .6 0 ).

A l t h o u g h n o t sh o w n , i n d e x o f s i m i l a r i t y r e l a ­
t i o n s h i p s among b u r n e d 3 5 - y e a r - o l d s i t e s w e r e s i m i l a r
to t h a t o f b u rn ed and unburned c l e a r c u t s .

D iscu ssio n
F ir e has long been a r e c u r r in g f a c to r in shaping
f o r e s t c o m m u n ities i n t h e Lake S t a t e s
Swain 1 9 7 3 ).

(H e in selm an 1973,

Even b e f o r e t h e a p p e a ra n c e o f t h e abun­

dant d istu rb an ce

literatu re

g enerated sin ce

the e a r ly

1 9 7 0 's , p e r i o d i c

f i r e was r e c o g n i z e d a s a n o r m a l e v e n t

in flu e n c in g the s tr u c tu r e

and m a i n t e n a n c e o f Lake

S t a t e s f o r e s t com m unities

( C o o p e r 1 9 1 3 , Graham 1 9 4 1 ,

M aissurow 1941, S t e a r n s

1949).

W hereas f i r e

h a s b e en w e l l docum ented i n a r e a s
W a te r s Canoe A re a i n M i n n e s o t a ,
Sim ard and B lan k
co unted f i r e

(1981),

it

su c h as t h e Boundary
is

s c a r c e i n M ichigan.

i n one o f t h e few s u c h s t u d i e s ,

s c a r s on r e d p i n e t r e e s k i l l e d by t h e

Mack L a k e f i r e , w h i c h o c c u r r e d on May 5 ,
co un ty M ichigan,
sp ecific

histo ry

and found t h a t f i r e

1980,

i n Oscoda

o c c u r r e d on a

s i t e w i t h a n a v e r a g e i n t e r v a l o f 23 y e a r s

s i n c e 1830.
Trends in s p e c ie s d i v e r s i t y d u rin g e a r l y p o s t f i r e
s u c c e s s i o n have b e e n r e p o r t e d from w i d e ly v a r y i n g
com m unities.

C e r t a i n a u th o r s have r e p o r t e d t h a t f i r e

did not g r e a tly a l t e r

the sp e cie s

co m position or

88
ric h n e s s of the s i t e

(Dix 1960, Vogl 1970, K r e f t i n g

a n d A h l g r e n 1 9 7 4 , P u r d i e 1 9 7 7 , Ohmann a n d G r i g a l 1 9 7 9 ) .
In c o n tra s t,

th ere

is

a d o cu m entation t h a t f i r e

g re a tly in cre ases sp ecies ric h n e ss
s p e c ie s com p o sitio n )

of the s i t e

(i.e .

a lters

( G a s h w ile r 1970,

D y r n e s s 1 9 7 3 , P a r s o n s 1 9 7 6 , B e l l a n d Koch 1 9 8 0 , T r a b a u d
a n d L e p a r t 1 9 8 0 , Abr ams a n d D i c k m a n n 1 9 8 2 ) .
from t h i s

The d a t a

s t u d y show t h a t s p e c i e s r i c h n e s s a n d t o t a l

n u m b e r o f u n i q u e s p e c i e s , w i t h a f e w e x c e p t i o n s , was
g r e a t e r on t h e b u r n e d s i t e s

th a n e i t h e r unburned c l e a r ­

c u ts o r unburned j a c k p in e s t a n d s .
Denslow (1980)

s u g g e s ts t h a t p l a n t com m unities

w hich e v o lv ed w ith r e c u r r in g l a r g e - s c a l e c le a r i n g
d istu rb an ces
tia lly

such as f i r e ,

sh o u ld be m ost d i v e r s e i n i ­

fo llo w in g th e d is tu rb a n c e ,

but species d iv e rs ity

should d e c re a se d u rin g s u c c e ss io n .

D ocum entation

s u p p o rtin g t h i s h y p o th e s is does e x i s t

(C hristensen

and Kimber 1975, P a r s o n s 1976, P u r d i e and S l a t y e r
1976, Trabaud and L e p a rt 1980).

Much o f t h e

literatu re

s u p p o r t i n g D e n s l o w ' s i d e a s a l s o s u p p o r t E g l e r 1s IFC
m odel,

i n t h a t n e a r l y a l l m em b ers o f t h e p r e d i s t u r b a n c e

com m unity w e r e p r e s e n t d i r e c t l y
It

is

d iffic u lt

fo llo w in g f i r e .

to g e n e ra liz e

about the p a tte r n

o f s u c c e s s i o n f o l l o w i n g b u r n i n g on j a c k p i n e s i t e s
used in t h i s

study.

The r a p i d c o n v e r s i o n a n d d o m i n a ­

t i o n o f a l l b u t p o s s i b l y one b u rn e d s i t e by v e g e t a t i o n

89
o th e r than ja c k p in e is

th e m ain d i f f i c u l t y .

However,

t h e f o r m a t i o n o f C a r e x meadows o n m any o f t h e d i s t u r b e d
s i t e s c a n be l a b e l l e d a " r e g r e s s i v e

su ccessio n ",

th e term in o lo g y o f R ich ard s

because of the

(1955);

in

a p p a r e n t s t a b i l i t y o f t h e s e m e a d o w s , t h e y may b e
invo lv ed in a r r e s tin g or i n h ib i t in g su c cessio n
1954, C o n n e ll and S l a t y e r
trends

1977).

(E gler

E arly su ccessio n al

i n s p e c i e s r i c h n e s s a n d c o m p o s i t i o n o n many

o f the s i t e s

a r e c o n s i s t e n t w i t h D e n slo w 's h y p o t h e s is

a n d E g l e r ' s IFC m o d e l .

T hat burned s i t e s

had n e a r l y

e v e r y s p e c i e s found on th e u n d i s t u r b e d s i t e s

(T ables

4.2,

E vidence

4.4,

and 4 . 6 ) ,

supports

t h e IFC m o d e l .

s u p p o r t i n g D e n slo w 's h y p o t h e s is
rich n ess

in clu d es

the d e crea sin g

t h a t g e n e r a l l y o c cu rs i n a sequence going

from r e c e n t l y b u rn e d a r e a s
m ature sta n d s

to

3 5 -y e a r-o ld stan d s to

(T ab les 4 .3 and 4 . 9 ) .

However,

tw o a r e a s

b u rn e d by w i l d f i r e s ,

w hich w ere dom inated by C a r e x ,

had s p e c ie s r i c h n e s s

lev els

low er th a n t h a t of i n t e r ­

m e d ia te -ag e d or m ature ja c k p in e s ta n d s .

A lso,

the

f a c t t h a t 3 5 - y e a r - o l d and m a tu r e j a c k p i n e s t a n d s e a c h
had a s u b s t a n t i a l number o f s p e c i e s n o t s u r v e y e d i n
th e o t h e r age c l a s s

is

i n c o n s i s t e n t w i t h t h e IFC m o d e l .

To s u p p o r t t h i s m o d e l , m a t u r e s t a n d s

sh o u ld have

e s s e n t i a l l y no s p e c i e s n o t p r e s e n t i n an e a r l i e r
su ccessio n al stag e.

How ever,

t h i s d i s c r e p e n c y may

be due t o t h e s m a l l number o f s i t e s

surveyed an d /o r

90
u s i n g a s a m p l i n g t e c h n i q u e t h a t was n o t d e s i g n e d t o
u n c o v er e v e r y s p e c i e s on th e s i t e .
The d o m i n a t i o n o f d i s t u r b e d s i t e s by v e g e t a t i o n ,
u su ally early

su ccessio n al sp ecies,

o th e r th an the

p r e v io u s dom inants has b e en r e p o r t e d f o r a v a r i e t y
o f p l a n t com m unities.

W i l l i a m s et: a l .

(1969) r e p o r t e d

t h a t L an tan a t h i c k e t s w hich e f f e c t i v e l y

su p p re ss norm al

s u c c e s s i o n can form f o l l o w i n g c l e a r c u t t i n g o f r a i n
fo rests

in Q ueensland.

A h lg re n (1976)

w ith o u t ad eq u ate seed s o u rc e s ,
pine s i t e s

showed t h a t

logged r e d and w h ite

i n n o r t h e a s t e r n M in n eso ta w ere c o n v e rte d

to bru sh lan d .

R e c e n t l y , C a t t e l i n o et: a l .

(1979) r e c o g ­

n i z e d t h a t man y d i s t u r b e d c o m m u n i t i e s do n o t f o l l o w
a s i n g l e r e g e n e r a t i o n pathw ay.

The c o n v e r s i o n o f a

p l a n t com m unity t o a c e r t a i n d o m in a n t v e g e t a t i o n
r e p r e s e n t s one p ath w ay i n a m u l t i p l e p ath w ay s u c c e s i o n a l
s c h em e t h a t a p p a r e n t l y e x i s t s
com m unities.
f ir e p erio d
1981),

in fre q u e n tly d istu rb ed

M odels have b e e n d e v e l o p e d b a s e d on i n t e r ­
( C a t t e l i n o et: a l .

scorch h e ig h t

1979, K e s s e l l and F i s c h e r

( K e s s e l l and P o t t e r

1980),

and

v i t a l a t t r i b u t e s o f i n d i v i d u a l s p e c i e s and t h e comm unity
(N oble and S l a t y e r 1980)

to p r e d i c t s u c c e s s io n a l changes

i n p l a n t com m unities s u b j e c t to r e c u r r e n t d i s t u r b a n c e s .
The d a t a p r e s e n t e d i n t h i s
h y p o th esis

study su p p o rt th e

t h a t v a r i o u s d e v e lo p m e n ta l pathw ays a r e

p o s s ib le during e a r ly p o s t-d is tu rb a n c e

su ccessio n .

A

p a r t i a l ex p lan atio n fo r t h is
is

th a t ja c k pine f a ile d

su c c e ss io n a l divergence

to re p r o d u c e on t h e s e s i t e s

w i t h any c o n s is te n c y f o llo w in g d i s t u r b a n c e .

Unburned

c l e a r c u t s i t e s v e ry q u ic k ly c o n v e rte d to Carex
mea dows ( s e e P l a t e

3 .1 ).

T h i s same p a t h w a y was

f o l l o w e d b y two a r e a s b u r n e d b y w i l d f i r e s

a n d two p r e -

scrib e-b u rn ed a re as.

apparently

O ther burned a r e a s ,

f o l lo w in g a d i f f e r e n t d e v e lo p m e n ta l p ath w ay , were
b e in g c o n v e r te d to sh ru b s and e a r l y
woods s u c h as a s p e n ,
P late

3 .2 ).

oak,

cherry,

su ccessio n al h ard­

and w illo w

(see

A n o t h e r p a th w a y was t h e r e e s t a b l i s h m e n t

o f a j a c k p in e dom inated s i t e
c o v e r on t h i s

(P late

4 .3 ).

C arex

s i t e was among t h e l o w e s t o f a n y o f t h e

d istu rb ed s ite s .

P a r t i a l s h a d e , p r o v i d e d by th e h i g h

d e n s ity of stan d in g snags,

a p p a re n tly provided s u ita b le

c o n d i t i o n s f o r t h e s u r v i v a l and grow th o f j a c k p in e
r e p r o d u c t i o n and c o n t r o l l e d

the sp read o f C arex , a

p ro v en c o m p e tito r i n open s i t u a t i o n s .
The s p e c i f i c

f a c to r s d eterm in in g su c c e ss io n a l

p a t h w a y s f o l l o w e d on b u r n e d s i t e s
tain ly

sources,

flo ra,
so ils

b u rie d seed p o o ls,

d ifferen ces,

a d ja c e n t seed

burn i n t e n s i t y ,

turbance h is to r y a re im p o rtan t.

is

C er­

s p e c i e s c o m p o s i t i o n and dom inance o f t h e p r e ­

d istu rb ed

little

a r e unk n ow n .

and d i s ­

U n fo rtu n ately ,

very

in fo rm a tio n about the s i t e s p r i o r to burning

av ailab le.

B ut,

even a l i t t l e

i n f o r m a t i o n may

92

P la te 4 .3 .
V ig o r o u s j a c k p i n e r e p r o d u c t i o n on a 4 - y e a r o l d s i t e b u r n e d by w i l d f i r e .

93
p r o v i d e some v a l u a b l e c l u e s a b o u t t h e r e s p o n s e o f
a c o m m u n i ty t o d i s t u r b a n c e .
and burned s i t e
in the

For exam ple,

10B h a d s c a t t e r e d a r e a s o f a s p e n w i t h ­

s i t e b e f o r e lo g g in g and b u r n in g .

appears th a t t h i s s i t e
dom inated e a r l y

clearcu t

I t now

i s b e in g c o n v erted to aspen-

s u c c e s s i o n a l hardw oods.

A lso, o ld e r

s t a n d i n g i n d i v i d u a l s o r stu m p s o f hardw ood s p e c i e s
w ith sp ro u tin g or suckering c a p a b ilitie s a re o fte n
p r e s e n t on n ew ly d i s t u r b e d s i t e s .

These r e l i c t

species

o f t e n have a pro fo u n d l o c a l i z e d or w id e sp re a d e f f e c t
in shaping the p o s t- d is tu r b a n c e
I n man y i n s t a n c e s
age were l e s s
sites

comm unity.

s i t e s o f t h e same d i s t u r b a n c e

s im ila r to each o th e r th an to d i f f e r e n t

o f a n o th e r age.

T h is,

too,

can be e x p la in e d

by m u l t i p l e s u c c e s s i o n a l p a th w a y s , b e c a u s e s i t e s
i n g t h e same p a t h w a y

(e.g .

fo llo w ­

C a r e x meadows o r d o m i n a t i o n

by e a r l y s u c c e s s i o n a l h a rd w o o d s) w e re more s i m i l a r
to th em selv es th a n to s i t e s
ways.

How ever, ev en s i t e s

fo llo w in g d if f e r e n t p a th ­
f o l l o w i n g t h e same s u c c e s ­

s i o n a l pathw ay w ere h i g h l y u n iq u e ;
t e n t l y more s i m i l a r t o i t s e l f ,
p eriod ,
age.

than to o th e r s i t e s

a s i t e was c o n s i s ­

even over a th re e year

o f t h e same d i s t u r b a n c e

I t was a l s o n o t e d t h a t a u n i q u e s e t o f s p e c i e s

w ere r e s p o n s i b l e
T herefore,

f o r th e in c r e a s e d c o v e r on e a c h s i t e .

the i n d i v i d u a l i s t i c

of each s i t e ,

( G le a s o n 1926) n a t u r e

r a t h e r th a n age f o l l o w i n g d i s t u r b a n c e ,

94
becomes t h e d o m in a n t a s p e c t i n d i s c e r n i n g s u c c e s s i o n a l
rela tio n sh ip s
is

i n th e com m unities.

T h is c o n c lu sio n

c o n s i s t e n t w i t h B e l l a n d Koch ( 1 9 8 0 ) , who f o u n d

t h a t g e o g r a p h i c a r e a was more i m p o r t a n t t h a n t i m e s in c e - la s t- b u r n in d eterm inin g s i m i l a r i t y

of s i t e s

i n th e j a r r a h f o r e s t s o f W estern A u s t r a l i a .
The u n i q u e c o m m u n i t y o n e a c h s i t e m a k e s i t v e r y
d iffic u lt,
sites

o r e v e n i n a p p r o p r i a t e t o compare t h e s e

along tem p o ral seq u en ces.

7B81 ( b u r n e d c l e a r c u t )

For exam ple,

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

b e i n g c o n v e r t e d t o a C a r e x meadow;

th is

s p e c i e s r i c h n e s s o f 24 a t y e a r 3.

S ites

(4-,

5-,

site

site

had a

10A a n d 10B

and 6 - y e a r - o l d p r e s c r i b e d b u rn s ) w ere b e in g

c o n v e r t e d t o s h r u b and e a r l y s u c c e s s i o n a l hardw oods
and w ere r i c h e r i n

species

(36 a n d 39 s p e c i e s , r e s p e c - ■

tiv e ly a t year 4).

T h e r e i s n o way s i t e

w i l l resem ble s i t e s

10A79 a n d 10B79.

processes

on t h e s e s i t e s

a n d 3 a n d two d i f f e r e n t s i t e s

sequence.

T he d e v e l o p m e n t a l

a re too v a r i e d .

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

and 6 f o l lo w in g f i r e

7B a t a g e 4

A lso,

d ata

co m p risin g y e a r s 1,
com prising y e ars

2,

4,

5,

to e s t a b l i s h th e burned c le a r c u t

T h is e x p l a i n s t h e anom alous i n c r e a s e i n

s p e c i e s r i c h n e s s and a v e ra g e r i c h n e s s p e r p l o t t h a t
o c c u r r e d betw een y e a r s
It

is

3 and 4 ( F ig .

4 .4 ).

g e n e r a lly tho u g h t t h a t d is tu rb a n c e

b a c k comm unity s u c c e s s i o n t o an e a r l i e r

sets

developm ental

95
stag e

( C l e m e n t s 1 9 1 6 , M a r g a l e f 1 9 6 3 , Odum 1 9 7 1 ) .

Jack pine is
S tates

among t h e m o s t s h a d e i n t o l e r a n t L a k e

sp ecies,

and h i s t o r i c a l l y n eed ed p e r i o d i c f i r e

to p e rp e tu a te i t s e l f .

In th e absence of f i r e ,

jack

p i n e s t a n d s w i l l e v e n t u a l l y be r e p l a c e d by more
t o l e r a n t hardwoods and c o n i f e r s , w h ic h a r e i n t u r n
f o llo w e d by l a t e r

su ccessio n al a ss o c ia tio n s

L eB arron 1944, H einselm an 1 9 7 3 ).
e v e r , b u r n i n g on c e r t a i n s i t e s
ment o f j a c k p i n e t o e a r l y

In th is

(E yre and

s t u d y , how­

a ccelerated

the r e p la c e ­

s u c c e s s i o n a l hardw oods.

T h i s r e s u l t e d b e c a u s e t h e j a c k p i n e o v e r s t o r y was
d estroyed, ja c k pin e rep ro d u c tio n f a i l e d ,
such as aspen,
fire
after

cherry,

oak,

and s p e c i e s

and w illo w s u r v iv e d th e

and s p r e a d v e g e t a t i v e l y o r became e s t a b l i s h e d
the f i r e

by s e e d .

I have n o t i d e n t i f i e d o l d e r burn ed o r c l e a r c u t
a r e a s on w h ic h j a c k p i n e d i d n o t become a d e q u a t e l y
e s t a b l i s h e d im m ediately fo llo w in g the d is tu r b a n c e .
X, t h e r e f o r e ,

cannot say what th e u ltim a te f a t e o f

th e c l e a r c u t an d /o r burned a re a s used in t h i s
w i l l be.
every s i t e

Jack p in e,

a l t h o u g h r a r e , was p r e s e n t on

used in t h i s

study.

I f th ese in d iv id u a ls

and a d j a c e n t s o u r c e s p r o v i d e enough s e e d ,
f in d adequate " s a fe s i t e s "
tio n ,

jack p in e,

study

( H a r p e r 1977)

and t h e

seed

fo r germ ina­

o v e r a l o n g t i m e p e r i o d , may a g a i n

dom inate th e s i t e .

There i s

docum entation t h a t ja c k

p i n e c a n become r e e s t a b l i s h e d i n a r e a s i t o n c e d o m in a ­
ted but fa ile d
turbance

to rein v a d e i n i t i a l l y

fo llo w in g d i s ­

(Thomson 1 9 4 3 , V o g l 1 9 6 4 , M a r s h a l l 1 9 8 0 ) .

T his s i t u a t i o n ,

if

i n f a c t p o s s i b l e on t h e s e s i t e s ,

a p p e a r s more l i k e l y

t o o c c u r on t h e s i t e s

coverted

t o C a r e x m e a d o w s.

These s i t e s

are n o t d ev elo p in g

a s tr a tif ie d p lan t

canopy and,

th erefo re,

flo o r,

even 6 y e a rs a f t e r

lo g g in g or b u rn in g ,

h ig h ly exposed to f u l l s u n l i g h t .
u n lik e ly th a t jack pine,

the f o re s t
is

In c o n tr a s t,

because of i t s

still

it

is

e x t r e m e l y low

s h a d e t o l e r a n c e , w i l l be c o m e e s t a b l i s h e d u n d e r a c a n o p y
o f e a r l y s u c c e s s i o n a l hardw oods.
s i t e s may b e a r r e s t e d
o r may p r o g r e s s
However,

later

S u c c e s s io n on t h e s e

a t a s h r u b - e a r l y hardw ood s t a g e ,

to l a t e r

su ccessio n al a sso c ia tio n s .

a s s o c i a t i o n s d e v e l o p in g on t h e s e s i t e s

w i l l be l i m i t e d by t h e e x tr e m e e d a p h i c c o n d i t i o n s
im p o sed by t h e G r a y l i n g sa n d s e r i e s , w h ic h r e p r e s e n t s
th e m ost d e p a u p e ra te o f th e sandy s o i l ty p es
low er M ichigan,
c ap a b ilities.
a sso ciatio n ,

in n o rth ern

i n term s o f n u t r i e n t and m o i s t u r e h o l d in g
A su g a r m ap le- bassw ood o r s p r u c e - f i r

said

to e v e n t u a l l y r e p l a c e j a c k p in e in

the absence o f f i r e

(E yre and L eB arron 1944),

u n l i k e l y on t h e s e s i t e s ;
and b la c k c h e r r y ,
p o ssib ilities.

n o rth e rn re d oak,

se e m s

re d m aple,

or w h ite p in e re p re s e n t stro n g e r

A lso,

any i n t e r m e d i a t e o r

long-range

s u c c e s s i o n a l developm ent on t h e s e s i t e s w i l l depend

97
on f u t u r e d i s t u r b a n c e e v e n t s , w h ic h o c c u r f r e q u e n t l y
i n n o r t h e r n low er M ichigan,

and a r e

the f a c to r s

u l t i m a t e l y shape the v e g e ta tio n in the a re a.

th at

CHAPTER V
FLORXSTIC COMPOSITION BEFORE AND AFTER
FIRE ON A JACK PINE CLEARCUT SITE
IN NORTHERN LOWER MICHIGAN
In tro d u ctio n
There e x i s t t h r e e b ro a d s t r a t e g i e s

in d ata

c o lle c tio n used to c h a ra c te riz e p o s t f i r e
a n d / o r compare d i s t u r b e d s i t e s
not been r e c e n tly d istu rb e d .
m any s i t e s

( u s u a lly once)

to o th e r

su ccessio n
sites

One s t r a t e g y

is

to survey

th a t to g eth er rep resen t a

te m p o r a l se q u en c e r a n g i n g from new ly d i s t u r b e d
lo n g -sin ce d istu rb ed

t h a t have

to

(V ogl 1 9 7 0 , Hanes 1971, S h a f i and

Y a r r a n t o n 1 9 7 3 , P a r s o n s 1 9 7 6 , B e l l a n d Koch 1 9 8 0 ) .
second s t r a t e g y ,

g e n e r a l l y l i m i t e d to e a r l y

A

succes­

s i o n a l s t u d i e s , in v o lv e s e s t a b l i s h i n g perm anent p l o t s
soon a f t e r a f i r e

and t h e n s u r v e y i n g t h e s e s i t e s ,

u su a lly an n u ally ,

fo r v a ry in g le n g th s o f tim e

(A hlgren

1 9 6 0 , L y on a n d S t i c k n e y 1 9 7 6 , Ohmann a n d G r i g a l 1 9 7 9 ,
T rabaud and L e p a r t 1980).

These ty p e s o f s t u d i e s

may i n c l u d e d a t a f r o m a d j a c e n t s t a n d s

th at

s e n t a t i v e o f the p r e - d i s t u r b e d c o n d itio n
Ohmann a n d G r i g a l 1 9 7 9 , U h l et: a l .
stra te g y ,

a ls o used in e a r ly

to e s t a b l i s h perm anent p l o t s
98

1981).

are r e p r e ­

(A h lg ren 1960,
A th ird

su ccessio n al s tu d ie s ,
i n an a re a p r i o r to

is

99
burning.

D ata i s

t h e n c o l l e c t e d from th e p r e - d i s t u r b e d

comm unity and from t h e d i s t u r b e d com m unity f o r v a r y i n g
l e n g t h s o f tim e f o l l o w i n g d i s t u r b a n c e .

T h is s t r a te g y

e l i m i n a t e s t h e c o n f o u n d i n g e f f e c t s o f many d i f f e r e n t
aged s i t e s

in e sta b lish in g

a u th o r s have u t i l i z e d

su ccessio n al tre n d s.

th e l a t t e r approach (G ashw iler

1970, D yrness 1973, P u r d ie and S l a t y e r
knowledge t h i s
used in f i r e

th ird

th e t h i r d

i n t h e Lake S t a t e s .

(DNR) l a n d w a s d e s i g n a t e d t o

f ir e - in d u c e d v e g e ta tio n a l changes u sin g

strateg y .

On t h i s a r e a i t w a s p o s s i b l e

t o 1) m o n i t o r b l o c k s b e f o r e a n d a f t e r b u r n i n g ,
ate f i r e

To my

a c l e a r c u t a r e a on M i c h ig a n D e p a r tm e n t

o f N a tu ra l R esources
ch aracterize

1976).

s t r a t e g y has n o t been p re v io u s ly

su ccessio n s tu d ie s

I n 1979,

Fewer

in te n sity

t a t i o n a l changes,

to d ete rm in e i t s

effects

2) e v a l u ­

on v e g e ­

3) c h a r a c t e r i z e v e g e t a t i o n a l d e v e l o p ­

m e n t on u n b u r n e d c l e a r c u t b l o c k s ,

a n d 4) c o m p a r e b u r n e d

and u n burned c l e a r c u t b lo c k s w i t h an a d j a c e n t m a tu re
ja c k p in e stand.

M ethods
T h e e x p e r i m e n t a l b u r n a r e a w a s c l e a r c u t i n 1976
from m atu re j a c k p i n e
th ereafter.

( 6 5 - y e a r s - o l d ) , and was u n t r e a t e d

See C h a p te r 2 f o r a more d e t a i l e d d e s c r i p ­

t i o n o f t h e e x p e r i m e n t a l b u rn a r e a and m a tu re j a c k p i n e
site

13A u s e d i n t h i s

study.

100
I n 1979 t h e e x p e r i m e n t a l b u r n s i t e was d i v i d e d
in to
(1,

6 b lo c k s r a n g i n g from 1 .0
3,

to 1.5 ha.

Three b lo c k s

6) w e r e r a n d o m l y d e s i g n a t e d t o b e b u r n e d a n d

th e th re e rem ain in g b lo ck s
unburned.

(2,

D u rin g J u l y 1979,

quency p l o t s

k,

5) w e r e t o b e l e f t

15 p e r m a n e n t 1 m2 f r e ­

a n d 2 p e r m a n e n t 20 m c o v e r t r a n s e c t s w e r e

random ly l o c a t e d i n e ach b lo c k ;
t a t i o n was r e c o r d e d .

at

th at

On S e p t e m b e r 1 9 ,

tim e th e v e g e ­
1979,

the th re e

d e s ig n a te d b lo c k s were burned by p e r s o n n e l o f th e M ich i­
g a n DNR.

However, b l o c k 6 was n o t c o m p l e t e l y b u r n e d ;

th is resu lte d

in only 6 freq u en cy p l o t s

and 1 c o v e r

t r a n s e c t b ein g in c lu d e d in th e burned a re a .

T herefore,

o n ly t h e s e b u rn e d p l o t s from b l o c k 6 w ere in c l u d e d i n
the d a ta a n a ly s e s .

The b u r n e d a n d u n b u r n e d b l o c k s w e r e

resu rv ey ed d u rin g the

1980 a n d 1 9 81 g r o w i n g s e a s o n s .

B urn i n t e n s i t y was e v a l u a t e d u s i n g a m eth o d m o d i­
f i e d from t h a t d e s c r i b e d by A h lg r e n ( 1 9 6 0 ).

D uring

J u l y 1 9 7 9 , a s t e e l d u f f p i n was p l a c e d a t a r a n d o m l y
c h o s e n c a r d i n a l d i r e c t i o n a d j a c e n t t o 10 r a n d o m l y c h o s e n
frequency p lo ts

i n b l o c k s 1 a n d 3.

s u c h t h a t t h e i r c o l l a r was e x a c t l y
o f the l i t t e r

lay er

l e v e l w i t h t h e top

( u n a l t e r e d o r o n l y s l i g h t l y decom­

posed p la n t m a t e r i a l ) .
burned,

The p i n s w e r e l o c a t e d

The d a y a f t e r t h e a r e a w a s

t h e d u f f p i n s w ere l o c a t e d and t h e amount o f

c o n s u m e d o r g a n i c m a t t e r was m e a s u r e d .
e a c h d u f f p i n w a s w h e t h e r 1)

the l i t t e r

A lso n o te d a t
l a y e r was s t i l l

101
present

(i.e .

lo w i n t e n s i t y b u r n ) , 2)

the l i t t e r

l a y e r was consumed b u t p a r t o f t h e d u f f l a y e r

(m oder-

a t e - t o w e l l d ecom posed o r g a n i c m a t t e r ) was s t i l l
in ta ct

(i.e .

m oderate i n t e n s i t y b u r n ) ,

o r 3) a l l o r ­

g a n ic m a t t e r was consumed, e x p o s in g m i n e r a l s o i l
(i.e .

high i n t e n s i t y b u rn ).
The u n d e r s t o r y

the a d ja c e n t,
d u rin g the
th ree

( v a s c u l a r v e g e t a t i o n <_ 1 . 5 m) o f

m ature,

u n c u t j a c k p i n e s i t e was s u r v e y e d

summer o f 19 8 1 u s i n g 30 f r e q u e n c y p l o t s a n d

20 m c o v e r t r a n s e c t s

a s d e s c r i b e d i n C h a p t e r 3.

R esu lts
T able 5 .1

shows t h e r e l a t i v e

c o v er and r e l a t i v e

fre q u e n c y o f e a c h s p e c i e s , by s p e c ie s group
and b i e n n i a l s ,

g r a s s e s and s e d g e s ,

and t r e e s

and s h r u b s ) ,

1981 ( 3 - ,

4-,

p e re n n ia l herbs,

su rv e y e d i n 1979,

and 5 - y e a r - o l d c l e a r c u t )

o f th e 3 unburned b lo c k s .

1980,

and

as a com posite

T r e n d s i n d i v e r s i t y on t h e

u n b u r n e d b l o c k s show t h e mea n t o t a l r i c h n e s s
f r o m 1979 t o

(annuals

in creased

1980 a n d d e c r e a s e d i n 19 81 ( T a b l e 5 . 2 ) .

The i n c r e a s e d r i c h n e s s

s e e n i n 1980 w a s m a i n l y d u e

t o t h e i n c r e a s e i n t h e number o f g r a s s s p e c i e s , i n c l u d ­
in g D anthonia s p i c a t a

( L .) B e a u v ., D ichanthelium

d e p a u p e ra tu m M u h l . , Andropogon g e r a r d i i V itm an, and
O ryzopsis pungens T o rr.

(T able 5 . 1 ) .

The m ea n a v e r a g e

r i c h n e s s p e r p l o t on t h e b l o c k s d i d n o t c h a n g e ,

and

102

TABLE 5.L
RELATIVE COVER AND RELATIVE FREQUENCY OF
EACH SPECIES, BY SPECIES GROUP, SURVEYED
IN 1 9 7 9 , 1 9 8 0 , AND 1 9 8 1 ( 3 - , 4 - , AND 5 YEAR-OLD CLEARCUT, RESPECTIVELY), AS A
COMPOSITE OF THE 3 UNBURNED BLOCKS

Year o f S u rv ey
S p e c ie s

1979
C over(T T F req (Z )

1980
C o v e r ( T T F r e g ( Z)

1981
C over(T T F req (Z )

A n n u a ls and B i e n n i a l s
Me,£ampy4um £ine.aA £

-

-

-

1 ,4

.7

7 0 .3

2 9 .0
1.3
2.7
.5
1.1

76.7
.2
. 1

30. A
1.1
1.5
3.6
1.1

.3

1.3

.1

2.0

.3
-

6.2
3.6
1.1

-

5.7
3.0
-

-

1.2

-

-

-

-

-

G r a s s e s an d S e d g e s
C a i e x pc.«4^£va*i£ca
Andxapogan g c x a x d i i
D au£(ioHia i p i z a t a
O xtjzopiii p u n g zm
O xtjzopiii a ip z x i& o lia
Vichanthzlium
dzpaupzxatum

6 5 .3
,1
.1
-

3 0 .0
,5
.5
.5
1 .0

.

1
.2

-

-

P e r e n n i a l H erb s
Ptzx.-id4.tim a q u i l i n u m
G a u l t h z x i a p /io c u m b a n i
P otzntiZZa tx id z n ta ta
A p oc y n u m a n d x o i a z m i -

iJo££um
Campanula x o t u n d i i o Z i a
A itzx Z azvii

3 .3
,3
-

1 0 .6
4. A
1 .0

.1

1.2

.1

.5
. 1

T r e e s and S h r u b s
1/acc-i.n-tum 4 p p . ^
Pdunua putni£d
C otnpiotiia p z x z g x i n a
Axz to i i a p i t t j l o i u v a - u X i i
P£hu4 b a n k i i a n a
Epigza X z p z m
Rubui h i i p i d u i
Que'tca-i 4 p p . ( r e d o a k )
PduMuJ i z x o t i n a
Am zlanchizx ip .

2 2 .7

2 8 .3

1. 8

8.8

.2
5 .1
,1
1 .0
-

1 .2
3 .9
3 .7
1 .0
.5
2 .A

23.8
1.3
. 1
2. A
.2
-

.7
.5
-

28. A
10. 9
1.2
3.9
2.7
1.1
.5
.5
1.5
-

18.3
.9
-

2.0
.5
-

.7
.5
.1

^ I n c l u d e s V. a n g u s c i f o l i u m . V. m v r t i l l o l d e s . a n d V. v a c i l l a n s .

29.2
12.0
.8
3.0
2.8
1.1
1.1
-

103

TABLE 5 . 2
DIVERSITY TRENDS OVER THE 1 9 7 9 , 1 9 8 0 , AND 1 98 1
GROWING SEASONS, EXPRESSED AS TOTAL SPECIES
RICHNESS, RICHNESS OF EACH SPECIES GROUP,
AVERAGE RICHNESS PER 1 m 2 PLOT, AND
SHANNON INDEX, AVERAGED AMONG THE
UNBURNED CLEARCUT BLOCKS
Year o f Survey
1979

1980

1981

T o ta l S p ecies R ichness

12.4

14.7

12.4

R ic h n e s s - A nnuals and
B ien n ials

-

.3

.7

D iv ersity

- G r a s s e s and
Sedges

2.7

4.7

4.0

- P erennial
H erbs

3.7

3.0

1.7

- T r e e s and
Shrubs

6.0

6.7

6.0

A verage R ic h n e ss p e r
iP lo t

3.4

3.4

3.4

Shannon In d ex

.95

.81

.70

104
a s l i g h t d e c l i n e i n t h e Shannon I n d e x was s e e n o v e r
th e 1979,

1 9 8 0 , a n d 19 81 s u r v e y s .

in th ese d iv e r s ity in d ic es

However,

a ll

changes

o v e r th e t h r e e grow ing

se aso n s w ere n o n s i g n i f i c a n t .
S ig n ific a n t in creases

(P < . 0 1 )

in t o t a l cover

o c c u r r e d over th e t h r e e grow ing se a s o n s

(F ig.

1) du e

t o t h e s p r e a d o f C a r e x p e n s y l v a n i c a Lam. o n a l l u n b u r n e d
blocks

(P late 5 .1 ).

A nnuals and b i e n n i a l s p ro d u ced

no c o v e r w h i l e t h e r e l a t i v e

cover of p e re n n ia l herbs

was n e g l i g i b l e on a l l u n b u rn e d b l o c k s .

R e la tiv e cover

o f g r a s s e s and se d g es i n c r e a s e d s i g n i f i c a n t l y
w hereas t r e e s

(P <.01>

and sh ru b s d e c l i n e d s i g n i f i c a n t l y

(P c . O l )

over the th re e y e a rs .
T a b le 5 .3 sum m arizes a d d i t i o n s

and l o s s e s o f

s p e c i e s , as a co m posite o f th e t h r e e unburned b l o c k s ,
d u r in g 1979,

1980,

and 1981.

Seven s p e c i e s d is a p p e a r e d

from p l o t s b etw een t h e 1979-1980 g ro w in g s e a s o n s .

-

B r a c k e n f e r n was m o st f r e q u e n t l y l o s t from t h e u n b u rn e d
p lo ts.

A t o t a l o f 9 s p e c i e s were added to unburned

p l o t s b e t w e e n 19 7 9 a n d 1 9 8 0 , P r u n u s p u m i l a L .
cherry)

and D ic h a n th e liu m depauperaturn b e in g t h e m ost

common a d d i t i o n s .

From 1980 t o 1 9 8 1 ,

l o s t from th e p l o t s , w h ereas

10 s p e c i e s w e r e

9 s p e c ie s w ere added.

O r y z o p s i s p u n g e n s a n d Melampyrum l i n e a r e Lam.
w h e a t ) w e r e t h e m o s t common a d d i t i o n s
1981.

(sand

( co w -

to the p l o ts

in

Unburned B lo ck s

120

100

*—
Q—
«—
*—
0—

* TOTAL COVER (tt)
SRNNURLS AND BIENNIALS CXI
» GRASSES AND SEDGES (X)
* PERENNIAL HERBS (X)
0 TREES RNO SHRUBS C*/.)

80

-fr
$------

60

40

e—
20

0
1979

1960

1981

YEAR OF SURVEY
Fig. 5 .1 .
T o t a l c o v e r (m) a n d r e l a t i v e c o v e r (%) f o r e a c h
s p e c i e s g ro u p , as a com posite o f th e 3 unburned b l o c k s , surveyed
i n 19 79 , 1 9 8 0 , a n d 1931.

P la te 5 .1 .
C arex p e n s y lv a n lc a d o m in a tin g unburned
blocks a t year 4 a f t e r c le a r c u ttin g .

107

TABLE 5.3.
ADDITIONS AND LOSSES OF SPECIES, AS A COMPOSITE
OF THE 3 UNBURNED BLOCKS, SURVEYED IN 1 9 7 9 ,
1 9 8 0 , AND 1 9 8 1 .
VALUES FOR LOSSES OF SPECIES
ARE EXPRESSED AS A RATIO OF NUMBER OF PLOTS
IN WHICH A SPECIES WAS LOST IN A SURVEY,
OVER THE NUMBER OF PLOTS IT WAS PRESENT
IN THE PREVIOUS YEAR SURVEY

S p e c ie s L o s t B etw een
197 9 - 1 9 8 0 S u r v d y
P tc ild lu m CLqultlnum
P/tunui p urni t a
Campanula l o - t u n d l ^ a l l a
Vaccinlum i p p ,
O u jzo p ili a& pcil$olia
PJtunui i c i o l l n a
Gautttici-ia piocum bcni

S p e c i e s L o st B etw een
19 80-1981 Survey
QueA cui i p p . ( r e d o a k )
Vanthonia i p i c a t a
P o tc n tllla tild& ntata
G a u lth c ila piocum bcni
A ic to ita p h g lo i uva-ui& l
P^luHUi i c i o t i n a
Andiopogon g e i a i d i i
M e t a mp t j iu m l i n c a i c
P tcild iu m aquitinum
Apoctj num a n d i o i a c m l fiolium

Number o f P l o t s
_______L o s t _______
T otal P resen t
8 /1 7
1 /1 3
1/1
1 /4 2

1/2
1 /3
1 /7

Number o f P l o t s
L o s t _______
T otal P resent
1 /1
1 /3
1 /2
1 /6
1 /6
1 /2
1 /2
2 /2
2 /8
1 /2

S p e c i e s Added B e t w e e n
1979-1980 Survey
Q ueA cui i p p . ( r e d oa k )
P a u n u i p u m if a
01 chan t h e H u m
dcpaupciatum
Vacc-infum i p p .
P te iid iu m aqultinum
Oan-thonia. ip - ic a -t a
Andiapogon g c i a i d l l
M c ta mp i jl u m t l n c a i c
Ap o c y n u m a n d i o & a c m l tfo l l u m

S p e c i e s Added B e t w e e n
1 9 8 0 - 1 9 8 1 S u r v e y ______
And^LOpogoH g e i a i d i l
O iijz o p iiq pungeni
Comptonia p c ie g iln a ,
M clampyium t i n z a i c
Rubui k - L i p i d u i
P teiid iu m aqultlnum
G a u l t h & l i a p /io cu m b en i
p/iunui p u m l l a
CaiCK p e m t j l v a n i c a

Number o f P l o t s

1
3
3
2

1

2

1
2

Number o

P lo ts

108
T a b l e 5 . 4 show s t h e r e l a t i v e

c o v e r and r e l a t i v e

fre q u e n c y o f e a c h s p e c i e s , by s p e c ie s group,

surveyed

i n 19 79 ( 3 - y e a r - o l d c l e a r c u t ) , a n d 1980 a n d 1 9 8 1 ( 1 and 2- y e a r - o l d p r e s c r i b e d b u r n , r e s p e c t i v e l y ) , as a
com posite o f th e
in the r e l a t i v e

3 burned b lo ck s.
frequency of a l l

in d u ced by b u rn in g .

(P < . 0 5 )

grow ing se a s o n s

and s h ru b s d e c r e a s e d

in r e la tiv e

bracken fe rn , w in terg reen
and k i n n i c k k i n i c k

s p e c i e s g ro u p s were

Over t h e t h r e e

b o t h p e r e n n i a l h e rb s and t r e e s
sig n ific an tly

S u b s ta n tia l changes

frequency, w ith

( G a u l t h e r i a procum bens L . ) ,

(A rc to s ta p h y lo s u v a - u r s i L .)

ing the l a r g e s t d e crea se s d uring t h i s

show­

in terv al.

In

s p i t e o f th e low ered fre q u e n c y o f C arex f o llo w in g
burn in g ,

th e .g ra s s and sedge s p e c i e s group s i g n i f i ­

c an tly in creased
to in c re a se s

(P < . 0 1 )

in r e la tiv e

f r e q u e n c y due

i n D ic h a n th e liu m d e p a u p e ra tu m , Panicum

colum bianum S c r i b n . , and O r y z o p s i s p u n g e n s .
T h e r e was a n o n s i g n i f i c a n t d e c r e a s e i n m ean
t o t a l r i c h n e s s on t h e b u rn e d b l o c k s t h e f i r s t y e a r
a f t e r burning, but a s ig n if ic a n t
d u rin g y e a r 2 (T able 5 .5 ) .

in crease

(P < . 0 5 )

D e cre ased r i c h n e s s betw een

1 9 7 9 a n d 1980 was m o s t l y d u e t o t h e l o s s o f many
tree

and sh ru b s p e c i e s on b lo c k s

3 and 5, and th e

lo ss

o f 3 p e r e n e n n i a l h e r b s p e c i e s o n b l o c k 3.

m ea n r i c h n e s s p e r p l o t d e c r e a s e d s l i g h t l y
fo llo w in g burning,

The

at year 1

b u t i n c r e a s e d a t y e a r 2.

The

109

TABLE 5.4
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES,
BY SPECIES GROUP, SURVEYED IN 1 9 7 9 (3-YEAR-OLD
CLEARCUT), AND 1 9 8 0 AND 1 9 8 1 ( 1 - AND 2-YEAR-OLD
PRESCRIBED BURN, RESPECTIVELY), AS A COMPOSITE
OF THE 3 BURNED BLOCKS
Year o f S u rv ey
S p e c ie s

1979
C over(Z )
F reg(Z )

1980
C over(T T F r e g (Z )

1981
C over(7T
Freg(Z )

G r a s s e s and S e d g e s

P e r e n n i a l H erb s
PtzA idium aquiZinum
AtteA Z aevii
SoZidago i p .
G auZthzAia pAocumbeni
LiaZAii novaz-angZ iaz
Apocynum a ndAoiazm i.(5o ££um
A i-fa* i a g i t Z Z ^ o t i u i
PoZi jgaZa p o Z y g a m a
die.Aac.Zum v z n o i u m
CoA.ijipzA.mum h t j i i o p Z (,o H u m
VZot a adunca

28.9
4.3
4.7
1 .4

21.8

17.9

23.7

.3

2.2
.6

2.6

2.9
7.8

-

.6

-

-

14.3
3.0

24.1
9.1

-

-

20.3
1.1
.1

20.7
5.5
.5

2 .3

8 .8
1 .4

57.2
.2
.6
.1

-

.1
-

-

-

-

-

—

.6

-

-

-

-

.3

3.

.2
.1
.3

.5
1.5

.6
.6

-

-

-

2.5
3.8

5 .3
—

-

..

19.6

.6
1.6

Ln Ln Ln Ui L n

Crt'iet p e n a y £ v f u t £ c a
A i t df i op o go n g t A a A d i i
0 Ai j z o p i i i a i p z A i & o t i a
O A tjz o p iii pungent
VichantheZium dzpaup ZA a t u m
Panicum coZumbianum
O anihonia i p i c a t a

.

.1
1.5

T r e e s and S h r u b s
Vacc-cnium -ipp.
P-tunai p u m i t a
P^iuHui Aa'io^-itta
AActoitaphijZoi u va -u A ii
Q u e t c u i Jpp. ( r e d o a k )
Sa Zi z gZaucopUtfiZoidei
Comptonia peA zgA ina
Ru 6 ua peiiA ,t£v an icuA
Rufaui h i i p i d u i
E p Z g z a ^icpenA
Poputui tA e m u to id zi
Re l a bf <inda
C o'tyfa* <ime*.£cana
Cta£aegu<i i p .

30.7
2. 5
1 .3
.8
1 .3
-

.2
2 .6
-

.2

28.3
2 .2
.6
4 .0
.6
-

.6
2 .7
2 .6
.6
-

1 .3
-

44.2
5.6

29.1
1.2

36. 7
2.8

1.0

_

1.3

-

1.5

_

.6
1.5

-

9.6

3.2

27.1
1.1

.1

.7
.5
.7

10.1

3.6

—
-

.1

~

m

-

110

TABLE 5 . 5
DIVERSITY TRENDS OVER THE 1 9 7 9 , 1 9 8 0 , AND 1 98 1
GROWING SEASONS, EXPRESSED AS TOTAL SPECIES
RICHNESS, RICHNESS OF EACH SPECIES GROUP,
AVERAGE RICHNESS PER 1 m2 PLOT, AND
SHANNON INDEX, AVERAGED AMONG THE
BURNED CLEARCUT BLOCKS
Year o f Survey
D iv ersity

1979

1980

1981

12.0

9.7

1 3. 3

- G r a s s e s and
Sedges

3.3

3.7

4 .3

- P eren n ial
Herbs

2.7

2.7

5.0

- T r e e s and
Shrubs

6.0

3.3

4.0

A verage R ic h n e ss p e r
P lo t

3.4

3.1

3 .6

1.2

1.4

T o ta l S p e c ie s R ichness
R ic h n e s s - A n n u als and
B iennials

Shannon In d ex

1.1

Ill
S h a n n o n I n d e x sh o w ed s l i g h t i n c r e a s e s
t h r e e grow ing s e a s o n s .

during the

T hese c h a n g e s , h ow ever, were

not sig n ific an t.
A s i g n i f i c a n t drop

(P < . 0 5 )

in t o t a l

o c c u r r e d f o l l o w i n g b u r n in g on th e b l o c k s ;
y e a r 2,

cover in cre ased s ig n if ic a n tly

n e a r ly the p reb u rn le v e l
5 .3 ).

(Fig.

5 .2 )

cover

how ever by

(P < .0 5) t o

( P l a t e s 5 .2 and

No c o v e r o f a n n u a l s a n d b i e n n i a l s w a s r e c o r d e d ,

and r e l a t i v e

c o v e r o f p e r e n n i a l h e r b s was n e g l i g i b l e

fo r a l l th ree

grow ing s e a s o n s .

g r a s s e s and se d g e s,

and t r e e s

The d o m inance o f
a n d s h r u b s was d r a m a ­

t i c a l l y a l t e r e d by b u rn in g (T able 5 .4 ) .
species

T ree and sh ru b

( m a i n l y b l u e b e r r y a n d b r i s t l y d e w b e r r y , R ubus

h is p i d u s L .)
in r e la tiv e
and se d g e s

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

in crease

(P < . 0 5 )

cover fo llo w in g b u rn in g , w hereas g ra s se s
(m ain ly A ndropogon g e r a r d i i , O ry z o p s is

a s p e r i f o l i a M i c h x ., and C arex p e n s y l v a n i c a ) d e c r e a s e d
sig n ific an tly

(P < . 0 1 )

during th a t

th e second y e ar a f t e r b u rn in g ,
cover of tr e e s

sedges;

th ese changes,

o f th e th re e burned b lo c k s ,

cover o f g r a s s e s and

of sp e c ie s,

as a com posite

o v e r th e t h r e e grow ing

shown i n T a b l e 5 . 6 .

w ere l o s t from p l o t s

corresponding

how ever, w ere n o n s i g n i f i c a n t .

A d d i t i o n s and l o s s e s

seasons is

D uring

a decrease in r e la tiv e

and sh ru b s o c c u rre d ,

to an i n c r e a s e i n th e r e l a t i v e

in te rv al.

A t o t a l o f 17 s p e c i e s

during th e f i r s t year a f t e r f i r e .

Burned B locks
80 _

*— * TOTAL COVER (H)
Q' ' ‘Q ANNUALS AND BIENNIALS £2)
0 0GRASSES ANO SEDGES 17.)
*" * PERENNIAL HERBS 17.)
°— 0 TREES ANO SHRUBS (*/.)

60

y.

o
R
40

112

M

20

0

T~~r
1979

—

*--------------1980

j
1981

YEAR OF SURVEY

Fig. 5 .2 .
T o t a l c o v e r (m) an d r e l a t i v e c o v e r (%) f o r
each s p e c ie s group, as a com posite o f th e 3 burned b l o c k s ,
s u r v e y e d i n 1 9 7 9 , 19C0, a nd 1981.

113

T able 5 .2 .
Large a r e a b a re d o f v e g e t a t i o n
one y e a r a f t e r f i r e .

T able 5 .3 .
O verview o f b u rn e d b l o c k s a t
y e a r 2 a f t e r f i r e show ing r a p i d r e v e g e t a t i o n .

114

TABLE 5.6
ADDITIONS AND LOSSES OF SPECIES, AS A COMPOSITE
OF THE 3 BURNED BLOCKS, SURVEYED IN 1 9 7 9 ,
1 9 8 0 , AND 1 9 8 1 .
VALUES FOR LOSSES OF SPECIES
ARE EXPRESSED AS A RATIO OF NUMBER OF PLOTS
IN WHICH A SPECIES WAS LOST IN A SURVEY,
OVER THE NUMBER OF PLOTS IT WAS PRESENT
IN THE PREVIOUS YEAR SURVEY

S p e c i e s L ose B etw een
197 9 - 1 9 8 0 S u r v e y

Number o f P l o t s
______ L o s t _______
T o ta l P resent

P Au nu i p u m i Z a
PiC ftidium a q u i t i n u m
Caftex p en 4</ £v ar ti ca
Atidftopogon g z A a A d i i
O Ayzopiii aipzA i& otia

1 / acc in iu m

1 /3
8 /1 1
1 3 /3 6

6/6

4pp.

A ii e . t o &t a p h y Z o h u v a - u A i i
Rubu4 p £ n 4 i £ v a n i c u 4
E p i g z a Acpe«4
G auZthzxia pAocumbzm
0ftyzop4i4 p u n g z m
Hicft.ac-i.um ucitoium

A4tcft i a c u i . 4
P o p u fu 4 t A z m u t o i d z i
Pftunui i z A . o t . i n a
(JuCft CU4 4 p p .
(red oak group)
CoA.yi.ui a m z A i c a n a

S p e c ie s L o s t B etw een
1980-1981 Survey
P a n i c u m c o Z u m b i a n um
O Ayzopiii pungzni
AitZA 4a g x t t i ^ o Z i u i
V a z z i n i u m 4pp.
Apoctjnum a n d A o i a z m i ftoZium
G auZthzAia pAocumbzm

3/A
5 /3 6
A /6
1 /1
A/A
2/A
1 /2
1 /1
1 /2
1 /1
1 /1
1 /1

S p e c i e s A dded B e t w e e n
1979-1980 Survey
V ic k a n th ztiu m dzpaupzAatum
P«nicum c o t umbianum
SaZix gZaucophyZZoidzi
L ia tA ii novaz-angtiaz
Camptonia pzAzgA ina
SoZidago i p ,
Caftex p z m y t v a n i c a
A itzx ia g ittiio Z iu i
PtZAidium a q u itin u m
Apoct jnum a n d A O i a z m i ioiium

Number o f P l o t s
23
1A

1
1

2

1
1
1
1

1 /1

Number o f P l o t s
______ L o s t
T o ta l P resen t6 /1 A
1 /1
1 /1
1 /3 6
1 /1
2 /2

S p e c i e s A dded B e t w e e n
1 9 7 9 - 1 9 8 0 S u r v e y _____

Number o f P l o t s

Apoctjnum a n d x o i a z m i loZium
O x y z o p i i i p u ng en 4
Caftcx p z m y t v a n i c a .
PtzAidium aquitinum
Vaccinium i pp.
V anthonia i p i c a t a
A 4icft i a g i t t i i o t i u i

6
3
A
1
1

P au i cu m c o f u mb i a nu m
Rubu4 Hi.4pi.du4
O A y z o p i i i a4pcfti|(o£ia
If i o t a a d u n c a

1
1
1
1

1

8

115
The s p e c i e s m o s t f r e q u e n t l y l o s t w e r e b r a c k e n f e r n ,
C a re x , Andropogon g e r a r d i i , O ry z o p s is a s p e r i f o l i a ,
blu eb erry ,

and k i n n i c k k i n i c k .

On a p e r c e n t a g e b a s i s ,

h o w e v e r , b l u e b e r r y sh o w e d t h e l o w e s t l o s s e s among t h e
group.
th is

Ten s p e c i e s w ere added t o b u r n e d p l o t s

in te rv al.

during

D ic h a n th e liu m d e p a u p e ra tu m and Panicum

c o l u m b i a n u m c o m p r i s e d 37 o f t h e 46 i n s t a n c e s o f s p e c i e s
a d d itio n s
fire

to the p l o ts

(1981),

a f t e r burning.

Two y e a r s a f t e r

6 s p e c i e s were l o s t from th e p l o t s ;

colum bianum c o m p r i s e d h a l f o f t h e s e l o s s e s .

P.

D uring

t h i s p e r i o d 11 s p e c i e s w e r e a d d e d t o t h e p l o t s , w i t h
C arex, bracken fe rn ,

blu eb erry ,

and O ry z o p sis a s p e r i -

f o l i a m ost o f t e n added.
I n 1 9 8 1 , many d i f f e r e n c e s b e t w e e n u n b u r n e d
blocks

(5 -y ear-o ld c le a rc u t)

and b u rn ed b lo c k s

y e a r - o l d p r e s c r i b e d b u rn ) were o b se rv e d
and 5 .4 ) .

burned b lo ck s
blocks

(T ables 5 .1

C a re x c o v e r on t h e u n b u rn e d b l o c k s

was s i g n i f i c a n t l y

g reater

(P < . 0 5 )

(24%) , w h e r e a s g r a s s

(1 % ).

(77%)

t h a n t h a t on t h e
cover on th e bu rn ed

(23%) w a s s i g n i f i c a n t l y g r e a t e r

t h a t on t h e u n b u rn e d b l o c k s

(2-

(P < . 0 5 )

than

A t o t a l o f 10

p e r e n n i a l h e r b s p e c i e s w e r e p r e s e n t on t h e b u r n e d
b l o c k s , b u t o n l y 2 w ere r e c o r d e d on t h e u n b u r n e d
b lo ck s.

T ree and sh rub s p e c i e s co m p rise d s i g n i f i c a n t l y

more c o v e r

(P < . 0 5 )

on t h e b u r n e d b l o c k s

t h a n on u n b u rn ed b lo c k s

(2 3 % ).

(51%)

i n 1981

1X6
F i r e i n t e n s i t y o n b l o c k 3 , w i t h a m ea n o r g a n i c
r e d u c t i o n o f 30 ± 1 2 . 5 mm w a s h i g h e r t h a n b l o c k 1 w h e r e
o r g a n i c m a t t e r was r e d u c e d 20 ± 9 . 0 mm.
5 p lo ts
p lo ts

of high f i r e

of high f i r e

in te n sity ,

B lock 3 had

compared t o o n ly 3

i n t e n s i t y o n b l o c k 1.

P o stfire

s p e c i e s d i v e r s i t y was a p p a r e n t l y a f f e c t e d by f i r e
sity .

The 10 p l o t s

in block 1 p r io r

in ten ­

t o b u r n i n g had

9 d i f f e r e n t s p e c ie s p r e s e n t , w ith an average r i c h n e s s
o f 3 . 2 s p e c i e s p e r 1 m2 p l o t .

One y e a r a f t e r

fire,

t o t a l r i c h n e s s o n t h e p l o t s was 11 s p e c i e s , w i t h a n
average ric h n e s s p e r p l o t o f 3 .9 .

A lthough b lo c k 3

p r i o r to b u rn in g had s i m i l a r d i v e r s i t y to b lo c k 1
(10 s p e c i e s w i t h a n a v e r a g e r i c h n e s s
p e r p l o t ) , one y e a r a f t e r

fire

o f 3.2 s p e c ie s

only 6 s p e c ie s o c cu rred

on t h e s e p l o t s w i t h an a v e r a g e r i c h n e s s o f 2 . 2 .

Two

y e a r s a f t e r b u r n i n g , b l o c k 1 r e m a i n e d more d i v e r s e
( s p e c i e s r i c h n e s s o f 12 a n d a v e r a g e r i c h n e s s p e r p l o t
o f 4.4)

t h a n b l o c k 3 ( s p e c i e s r i c h n e s s o f 8 and a v e r a g e

rich n e ss per p lo t of 2 .9 ).

P l a n t c o v e r on b l o c k 3 was

r e d u c e d b y 77% o n e y e a r a f t e r f i r e ,

whereas p l a n t cover

o n b l o c k 1 w a s r e d u c e d by 50%, p r o v i d i n g i n d i r e c t e v i ­
dence t h a t f i r e

i n t e n s i t y on b l o c k 3 w a s h i g h e r t h a n

b l o c k 1.
I n d iv id u a l sp e c ie s responded d i f f e r e n t l y
in ten sity .

to f i r e

For exam ple, C arex and b l u e b e r r y w ere

p r e s e n t o n a l l 10 p l o t s o n w h i c h f i r e

i n t e n s i t y was

117
e v a lu a te d in b lo ck s 1 and 3 p r i o r
after

to burning.

One y e a r

f i r e C a r e x w a s m i s s i n g f r o m 6 o f 10 p l o t s o n

b l o c k 3 (m o re s e v e r e l y b u r n e d ) , w h e r e a s b l u e b e r r y was
m i s s i n g from o n ly 2 p l o t s .

I n b l o c k 3 C a r e x was l o s t

from a l l f i v e p l o t s o f h i g h f i r e

in te n sity .

In b lo c k 1

(less

s e v e r e l y b u r n e d ) , C a r e x w a s l o s t f r o m 3 o f 10

p lo ts

( a l l h i g h i n t e n s i t y ) , w h e r e a s b l u e b e r r y was l o s t

from o n ly 1 p l o t

(high i n t e n s i t y ) .

appeared to a f f e c t

the a b i l i t y

become e s t a b l i s h e d on a b l o c k .
e s t a b l i s h e d on a l l
fire,

10 p l o t s

0.

sp e c ie s to

D. d e p a u p e r a t u m b e c a m e

i n b lo c k 1 one y e a r a f t e r

becam e e s t a b l i s h e d on 6 p l o t s

fire,

in te n s ity also

of c e rta in

b u t o n l y 6 p l o t s i n b l o c k 3.

a fter fire ,

F ire

b u t only 2 p l o t s

A lso, P.

colum bianum

i n b l o c k 1, one y e a r
i n b l o c k 3.

p u n g e n s was p r e s e n t i n 4 p l o t s

Year 2 a f t e r
on b l o c k 1 a n d

o n l y 1 p l o t i n b l o c k 3.
T a b l e 5 . 7 sh o w s t h e r e l a t i v e

cover and r e l a t i v e

frequency o f each s p e c ie s in the u n d e rs to ry of the
m ature ja c k p in e f o r e s t
e x p erim e n tal burn s i t e .

(site

13A ), a d ja c e n t to th e

T h e s u r v e y r e c o r d e d 18 s p e c i e s ,

w i t h an a v e ra g e r i c h n e s s p e r p l o t o f 4 .0 .

The Shannon

I n d e x o f d i v e r s i t y w a s e x t r e m e l y lo w ( . 4 9 )

because

b l u e b e r r y r e p r e s e n t e d o v e r 90% o f t h e v e g e t a t i v e c o v e r .
C a r e x p e n s y l v a n i c a o c c u r r e d i n 25 o f 30 f r e q u e n c y p l o t s ,
b u t r e p r e s e n t e d o n l y 3% o f t h e t o t a l p l a n t c o v e r .
A lso,

every sp ecies

surveyed in th e m ature ja c k p in e

118

TABLE 5 . 7
RELATIVE (REL) COVER AND RELATIVE FREQUENCY
OF EACH SPECIES IN THE UNDERSTORY OF A
MATURE JACK PINE STAND (S IT E 1 3 A ) ,
ADJACENT TO THE EXPERIMENTAL BURN AREA

S p ecies

Cover
(R el)

Frequency
(R el)

A nnuals and B i e n n i a l s
MzZampynum IZnzaJie.

2 .1

11.6

.2
3.2

3.3
20.7
1.7

G ra sses and Sedges
An d a o p o g o n g e n a n d Z Z
Catizx p z m y Z v a n Z z a
OnyzopAZA p u n g zn A

.2

P e r e n n i a l Herbs
Ap o c y n u m a n d n o i a z m Z f i o l Z u m

-

Campanula. n o l u n d Z ^ o l Z a
GauZZhznZa paocumbznA
HZznazZum v zn o iu m
PZznZdZum aquZZZnum

-

.3
-

.8
.8
8 .3
.8
.8

T re e s and S hrubs
AmzlanzhZzn 4p.
AnzZo&ZapkyloA uva-unAZ
C o m p Z o n Z a pe.ae.gn.Zna
EpZgea azpznA
PZnuA b a n k A Z a n a
PaunuA p u m Z l a
PnunuA A z n o Z Z n a
QuzncuA App. ( r e d oak)
VaccZnZum A p p .

.3
1.0
1.0
.4
-

.7
90 .7

.8
9.9
3.3
.8
1.7
5.0
.8
4.1
24.8

■ ^ I n c lu d e s V. a n g u s t i f o l i u m , V. m y r t i l l o i d e s .
a n d V. v a c i l l a n s .

119
u n d e r s t o r y w a s p r e s e n t on e i t h e r t h e a d j a c e n t u n b u r n e d
o r burned c le a r c u t b lo c k s.

But th e unburned b lo c k s had

5 s p e c i e s and th e b u rn e d b lo c k s

14 s p e c i e s n o t f o u n d

in th e m ature ja c k p in e sta n d .

D iscu ssio n
P revious p o s t f i r e

su ccessio n al stu d ies

in the

Lake S t a t e s h a v e n o t c h a r a c t e r i z e d t h e p l a n t community
on a s p e c i f i c

s i t e b e f o r e and a f t e r b u r n in g .

stu d ies of th is
system s

type have been conducted in o th e r eco ­

(G a sh w ile r 1970, D y rn ess 1973, P u r d i e and S l a t -

y e r 1976).
fir

However,

D y r n e s s , w orking i n o ld -g ro w th D ouglas-

f o r e s t s i n th e C ascade M ountains o f Oregon, r e p o r ­

ted th a t

c le a r c u ttin g reduced sp e cie s ric h n e s s ,

compared t o o l d - g r o w t h f o r e s t s ,
in in creased rich n ess

b u t burning r e s u l te d

through y ear 5 a f t e r

fire.

Woody

t r e e v e g e t a t i o n was r e d u c e d b u t s t i l l p r e s e n t a f t e r
fire,

a n d s p e c i e s common i n t h e o l d - g r o w t h f o r e s t s ,

w h i c h sh ow ed l o w e r e d d o m i n a n c e i n i t i a l l y a f t e r f i r e ,
r e g a i n e d d o m in a n ce by y e a r 5 a f t e r

fire.

G ashw iler,

a ls o w orking i n o ld -g ro w th D o u g la s - f ir s ta n d s in the
C a s c a d e M o u n ta in s i n O r e g o n , showed t h a t a l a r g e number
o f s p e c i e s n o t found i n
present a fte r
the s i t e

fire.

th e o ld - g r o w th f o r e s t s were

A lso,

h e rb a c e o u s p l a n t s dom inated

through y ear 3 a f t e r f i r e ;

p l a n t s p e c ie s dom inated.

th ereafter,

From s t u d i e s

woody

i n w oodland

120
co m m u n ities i n s o u t h e a s t e r n A u s t r a l i a , P u r d i e and
S laty er
tia lly

(1976) r e p o r t e d t h a t a l l

species p re s e n t i n i ­

a f t e r f i r e w ere p r e s e n t as

b u rie d seed p r i o r to f i r e ,
invaded a f t e r b u rn in g .

liv in g p la n ts or

a n d t h a t n o new s p e c i e s

They c o n c l u d e d t h a t r e g e n e r a ­

t i o n o f t h e c o m m u n i ty c l o s e l y r e s e m b l e d t h e i n i t i a l
flo ristic

c o m p o s it i o n m odel p r o p o s e d by E g l e r

The p r e s e n t s t u d y sh o w e d t h a t :

(1954).

1) c l e a r c u t t i n g

a m a tu re j a c k p in e s t a n d , w i t h an u n d e r s to r y dom inated
by b l u e b e r r y ,

resu lted

meadow b y y e a r 3 a f t e r
5 after

i n th e fo rm a tio n o f a C arex
lo g g in g ,

2) b e t w e e n y e a r s 3 a n d

c l e a r c u t t i n g , C a re x dom inance i n c r e a s e d from

65% t o 77% r e l a t i v e

c o v e r on t h e u n b u r n e d b l o c k s ,

3)

a t t h e c o n c l u s i o n o f th e e x p e r im e n t, b u rn e d b l o c k s were
d ram atically d if f e r e n t,
and dom inance,
flo ristic

i n term s of s p e c i e s c o m p o s itio n

from u n b u rn ed b l o c k s ,

c o m p o s itio n model i s

4) E g l e r ' s

in itia l

s u p p o r te d by t h e f a c t

t h a t every s p e c ie s in th e m ature j a c k p in e u n d e rs to ry ,
p l u s many u n i q u e s p e c i e s , w e r e p r e s e n t o n t h e u n b u r n e d
and b u rn e d c l e a r c u t b l o c k s ,

5)

t h e same f i r e m o v i n g

t h r o u g h two a d j a c e n t b l o c k s o f t h e same s o i l t y p e a n d
topography, w ith s im ila r p o s t f i r e
resu lte d

in v a ria b le f i r e

b etw een th e b lo c k s ,

com position,

i n t e n s i t i e s w i t h i n and

6) f i r e

a d d i t i o n and l o s s o f s p e c i e s
the blo ck s,

flo ristic

in te n sity

affects

and s p e c i e s

a n d 7) h i g h e r f i r e

the

d o m i n a n c e on

in te n sity resu lted

in

121
low er d i v e r s i t y

and lo w er p l a n t c o v e r .

V e g e t a t i o n a l d evelopm ent on th e s tu d y a r e a th e
first

two y e a r s a f t e r f i r e

d i f f e r e d from th e g e n e r a l

p a t t e r n d e s c r i b e d on o t h e r r e c e n t l y b u r n e d j a c k p i n e
sites

(Abrams a n d D i c k m a n n 1 9 8 2 ; C h a p t e r s 3 a n d 4 ) .

F o r e x a m p le , no a n n u a l o r b i e n n i a l s p e c i e s w ere
p r e s e n t a t y e a r one a f t e r f i r e .

A lso,

compared to p r e ­

b u rn l e v e l s , t o t a l s p e c i e s r i c h n e s s and r e l a t i v e c o v er
o f C arex p e n s y l v a n i c a w ere r e d u c e d on th e b u rn e d b l o c k s .
T h e s e f i n d i n g s may b e r e l a t e d

to

th e extrem e s e v e r i t y

o f t h e b u r n , w h ic h re d u c e d t h e f u e l lo a d i n g on th e s i t e
f r o m 23 t o 3 m t / h a a n d c o n s u m e d a l l f u e l s 6 . 4 cm o r
less

in diam eter

( p e r s o n a l c o m m u n i c a t i o n , Roy M i l n e s ,

DNR A r e a F i r e S u p e r v i s o r ) .

No m e a s u r a b l e p r e c i p i t a t i o n

o c c u r r e d f o r 28 d a y s p r i o r t o t h e b u r n , r e s u l t i n g i n
an e x tre m e ly h o t f i r e .
R esearch e v a lu a tin g the e f f e c t s o f f i r e
on p o s t f i r e p l a n t s u c c e s s i o n i s

scarce

(A h lg ren 1960,

D y rn e s s 1 9 7 3 , C h r i s t e n s e n and Kimber 1 9 7 5 ) .
concluded th a t f i r e

in ten sity

v e g e ta tiv e rep ro d u ctio n ,
q u ality ,

in te n sity

A hlgren

a ff e c te d seed g erm in atio n ,

p la n t co m p etitio n ,

seedbed

a n d n u t r i e n t c o n c e n t r a t i o n s on t h e s o i l s u r f a c e .

A l l t h e s e f a c t o r s may a c t i n d e t e r m i n i n g t h e e s t a b l i s h ­
m ent, p e r p e t u a t i o n ,

and s u b s e q u e n t dom inance o f s p e c i e s

during e a rly p o s t f i r e

su ccessio n .

D y rn e ss showed t h a t

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

122
p lo ts

compared t o l i g h t l y b u r n e d p l o t s .

D yrness a ls o

fo u n d t h a t c o v er on s e v e r e l y b u rn e d p l o t s
lagged b e h in d th e l i g h t l y burned p l o t s
fire.

co n sisten tly

fo r 5 years a f te r

C h r i s t e n s e n and Kimber r e p o r t e d t h a t h o t f i r e s ,

i n a d d it i o n to prom oting seed g e rm in a tio n o f c e r t a i n
sp ecies,
seed.

d e s t r o y e d c o n s i d e r a b l e am ounts o f s o i l - s t o r e d

A lso,

s o i l t e m p e r a t u r e d u r i n g a f i r e was f o u n d

to v ary g r e a tly w ith in th e s i t e ;

so il

tem perature

r e c o r d e d a t 56 p o i n t s a t t h e 1 cm a n d 2 cm d e p t h v a r i e d
f r o m < 66°C t o

> 399°C.

i n New S o u t h W a l e s ,
n ific a n tly

Floyd

(1966,

1976), w orking

concluded t h a t f i r e

in te n sity

a ffe c te d th e germ ination ( i . e .

of ty p ic a l p o s tfire

estab lish m en t)

sp ecies.

The d e p t h o f t h e u n d e r g r o u n d p e r e n n a t i n g
of understory sp ecies is

d irec tly rela te d

s u r v iv a b ility during f i r e
W ein 1 9 7 7 ) .

sig ­

system

to th e ir

(M clean 1968, F l i n n and

The g r e a t e r f i r e

resistan ce

shown by b l u e ­

b e r r y compared t o sed g e a n d o t h e r s h a l lo w r o o t e d s p e c i e s
is

c o n siste n t w ith th e ir in d iv id u a l ro o tin g h a b its .

H o w e v e r , b r a c k e n f e r n , known a s a d e e p - r o o t e d s p e c i e s ,
was p a r t i c u l a r l y
(lo st

su scep tib le

to b u rn in g in t h i s

study

f r o m 8 o f 11 p l o t s ) .
In th e p re s e n t study f i r e

i n t e n s i t y was h i g h l y

v a r i a b l e , w i t h two a d j a c e n t b l o c k s h a v i n g m a r k e d l y
d i f f e r e n t o rg a n ic m a tte r r e d u c t io n and m in e r a l s o i l
exposure.

I f e e l t h e s e v e r i t y o f b u r n i n g was

123
resp o n sib le
p la n t cover,
sp ecies,

for v a r ia b ility

in d iv e rs ity ,

to ta l

s u r v iv a b ility of v e g e ta tiv e ly reproducing

an d t h e number o f n ew ly e s t a b l i s h e d i n d i v i d u a l s

on t h e b u r n e d b l o c k s .

O t h e r s t u d i e s h a v e shown t h a t

v e r y d i f f e r e n t s u c c e s s i o n s c an r e s u l t on s i m i l a r p l a n t
com m unities f o l lo w in g d i s t u r b a n c e .
sites

i n n o r t h e r n low er M ic h ig a n ,

On o t h e r j a c k p i n e
early rev e g eta tio n

f o l lo w in g b u rn in g can be h i g h ly v a r i a b l e i n term s of
sp ecies ric h n e ss,

p r e s e n c e o f a n n u a ls and b i e n n i a l s ,

and t h e dom inance o f g r a s s e s
shrubs

and se d g e s and t r e e s

and

(A bram s a n d D i c k m a n n 1 9 8 2 ; C h a p t e r s 3 a n d 4 ) .

M u l t i p l e s u c c e s s i o n a l p a t h w a y s a r e a p p a r e n t on t h e s e
sites

in as l i t t l e

as 5 years a f te r

The c o n c l u d i n g p o i n t s

t o b e made f r o m t h e d a t a

p re s e n te d here is

th a t each s i t e

event is

Even when s i t e s

unique.

s e l e c t e d f o r hom ogeniety,

fire.

and e a c h d is tu r b a n c e
are rig o ro u sly

d i f f e r e n c e s w i l l m ost c e r ­

ta in ly e x is t in p re-d istu rb an ce

s p e c ie s co m p o sitio n

and dom inance, e d a p h ic f e a t u r e s , b u r i e d s e e d - p o o l s ,
and a d ja c e n t seed s o u r c e s .
to compare t h e e f f e c t s
but each event
w i t h i n an a r e a .

(in th is

Not on ly i s i t

d iffic u lt

of d if f e r e n t d istu rb an ce e v e n ts ,
case f ir e )

is h ig h ly v a ria b le

CHAPTER VI
RESPONSE OF UNDERSTORY VEGETATION
TO FERTILIZATION ON MATURE AND
CLEARCUT JACK PINE SITES IN
NORTHERN LOWER MICHIGAN
In tro d u ctio n
I n many f o r e s t e d e c o s y s t e m s ,
i n t h e dom inance o f s e d g e s
bances such as f i r e

dram atic in c re a se s

(C a r e x s p p . )

(L ittle

fo llo w d i s t u r ­

a n d Moore 1 9 4 9 , B u e l l a n d

C a n t I o n 1 9 5 3 , A h l g r e n 1 9 6 0 , V o g l 1 9 7 0 , W ein a n d
B liss

1973),

atom ic r a d i a t i o n e x p o su re

f e r t i l i z e r ap p licatio n

(W o o d w e ll 1 9 6 7 ) ,

(N oble e t a l . 1979)

and t r a n s ­

p la n tin g f o r e s t s o i l to a greenhouse (Z av itk o v sk i
1976)

have been r e p o r t e d .

I n o u r own s t u d i e s

(Abrams

a n d D i c k m a n n 1 9 8 2 ) , C a r e x p e n s y l v a n i c a Lam. o b t a i n e d
re la tiv e

c o v e r v a l u e s o f 86% a n d 71% o n r e c e n t b u r n e d

and u n burned c l e a r c u t s , r e s p e c t i v e l y .

We h y p o t h e s i z e d

t h a t C a r e x , a c t i n g as an o p p o r t u n i s t i c

sp ecies,

p lo its

r e s o u r c e s made a v a i l a b l e

an d /o r bu rn in g ,

ex­

fo llo w in g c le a r c u ttin g

thereby in c re asin g i t s

dom inance w h i l e

r e d u c i n g t h e dom inance o f n e i g h b o r i n g v e g e t a t i o n .
Large n u t r i e n t r e s e r v e s
a r e m ade a v a i l a b l e

s to r e d in the v e g e ta tio n

fo llo w in g c le a r c u tt i n g

and b u r n i n g

( A h l g r e n 1 9 6 0 , Bo rm an n e £ a l . 1 9 6 8 , D e b a n o a n d C o n r a d
1 9 7 8 , L i k e n s ej; a l . 197 8)
124

and r e s u l t

in profound

125
c h a n g e s i n t h e dom inance and d i v e r s i t y o f p o s t - d i s turbance v e g e ta tio n .

N u tr ie n t enrichm ent s tu d ie s

have been r e p o r t e d from w id e ly v a r y i n g e co sy ste m s
(W illis

1963,

J e f f e r y and P i g o t t 1973, Stephenson

1 9 7 3 , H a rco m b e 1 9 7 7 , B a k e l a a r a n d Odum 1 9 7 8 ) .
fin d in g of th ese a u th o rs is
sp ecies,

t h a t "A f e w o p p o r t u n i s t i c

a l r e a d y e s t a b l i s h e d i n t h e com m unity, expand

t h e i r n i c h e s b y p r e e m p t i n g some n i c h e
n ates,

A general

space o f su b o rd i­

thus red u cin g o v e ra ll d iv e r s ity "

( B a k e la a r and

Odum 1 9 7 8 ) .
T h i s s t u d y was e s t a b l i s h e d o n j a c k p i n e s i t e s
i n n o r t h e r n low er M ichigan to i n v e s t i g a t e
of fe r tiliz a tio n

on t h e r e l a t i v e

the e f f e c ts

grow th o f u n d e r s to r y

v e g e t a t i o n on c l e a r c u t and u n d i s t u r b e d s i t e s .

Methods and M a t e r i a l s
A 5 5 -y ea r-o ld ja c k pine
M ichigan

(C raw ford C ounty) was c l e a r c u t d u r i n g t h e

w i n t e r and s p r in g o f 1979-80.
lin g sand;
frig id ,

stand n ear G rayling,

a n u tr ie n t poor,

ty p ic,

udipsam m ent.

The s o i l

ty p e i s a Gray­

s t r o n g l y a c i d , m ixed,
A 70 m e t e r - w i d e b o r d e r

o f j a c k p i n e o f s i m i l a r a g e , q u a l i t y a n d s t o c k i n g was
left

alo n g an a d ja c e n t ro a d .
Four b lo ck s

(10 m x 11 m) w e r e e s t a b l i s h e d i n

b o th th e s ta n d in g j a c k p in e and th e a d ja c e n t c l e a r c u t .
Each b lo c k c o n ta in e d an u n f e r t i l i z e d h a l f

(control)

126
and a f e r t i l i z e d h a l f ,
tio n s tr ip .

s e p a r a t e d by a 1 - m e t e r i s o l a ­

The b l o c k s w e r e e s t a b l i s h e d b e f o r e g r e e n -

up d u r i n g t h e s p r i n g o f 1 9 8 0 , a n d w e r e l o c a t e d i n
a r e a s o f s e e m i n g ly homogeneous v e g e t a t i o n .

The

b lo c k s were l a i d o u t nonrandom ly to red u c e v a r i a b i l i t y
w ith in the b lo ck s,

but the f e r t i l i z a t i o n

treatm en t

w i t h i n e a c h b l o c k was ra n d o m ly a s s i g n e d .
The f e r t i l i z a t i o n

r a t e was 1 0 0 - 6 0 - 6 0 k g / h a

N -P -K p l u s 200 k g / h a Ca a s d o l o m i t i c l i m e s t o n e .

Osmo-

c o t e , a slo w r e l e a s e f o r m u l a t i o n , was u s e d f o r 6 0 -6 0 60 k g / h a N - P - K .
um n i t r a t e

The r e m a i n i n g 40 k g / h a o f N was ammoni­

(quick r e l e a s e ) .

e v e n l y b y h a n d on May 3 ,

1980.

t i o n was d e te r m in e d a t t h r e e
o f 1980

F e r t i l i z e r was d i s t r i b u t e d
Response to f e r t i l i z a ­

t i m e s d u r i n g t h e summer

( w e e k s e n d i n g J u n e 1 4 , J u l y 2 1 , a n d A u g u s t 28)

a n d o n e t i m e i n 1 98 1 ( w e e k e n d i n g J u l y 2 4 ) , b y s a m p l i n g
bio m ass from e a c h b l o c k i n th e s t a n d i n g j a c k p i n e and
clearcu t areas.

A l l v e g e t a t i o n u n d e r 1 . 5 m t a l l was

c l i p p e d a t ground l e v e l i n 5 random ly l o c a t e d
qu ad rats

w ith in each tre a tm e n t.

s o r t e d by s p e c i e s ,

The v e g e t a t i o n was

c le a n e d and s e p a r a t e d i n t o

y e a r s h o o t biom ass and r e s i d u a l biom ass
years b io m ass).

cu rren t-

(th e previous

T h e s a m p l e s w e r e d r i e d a t 8 0 °C f o r

48 h o u r s a n d w e i g h e d .

D e t e r m i n a t i o n o f a n n u a l and

r e s i d u a l b io m ass was n o t p o s s i b l e
green fo rb s.

, 2 5 m2

for c e rta in ever­

T h e i r b i o m a s s was i n c l u d e d i n t h e

127
r e s i d u a l p o r t io n o f th e t o t a l s ta n d in g biom ass.

These

s p e c i e s u s u a l l y r e p r e s e n t e d l e s s t h a n 10% o f t h e t o t a l
b iom ass.
Biomass d a t a w ere a l s o c o l l e c t e d from a j a c k p i n e
a r e a t h a t h a d b e e n p r e s c r i b e b u r n e d i n 1978 a s p a r t
of th is

study.

T his a r e a ,

l o c a t e d i n Ogemaw C o u n t y ,
study a re a .

a l s o on G r a y l i n g s a n d ,

is

31 k i l o m e t e r s f r o m t h e a b o v e

I n 1975 a n d 1 97 6 t h e m a t u r e j a c k a n d r e d

p i n e was lo g g e d , b u t s c a t t e r e d im m ature j a c k p i n e
(a b o u t 3 5 - y e a r s - o l d ) w ere l e f t

stan d in g .

T w e n t y . 2 5 m2

biom ass sam p les, random ized a lo n g t r a n s e c t s , w ere ta k e n
d u r i n g t h e same f o u r t i m e i n t e r v a l s ,
as in d ic a te d above.
i n 1980 a l l

D uring th e t h r e e

i n 1980 a n d 1 9 8 1 ,
sam p lin g p e rio d s

tw e n ty b iom ass sam p les were g ro u p ed t o g e t h e r .

In 1981, however,
of co llec tio n ,

th e sam ples w ere g ro u p ed ,

i n t o 4 g ro u p s o f 5 so t h a t a s t a n d a r d

d e v i a t i o n c o u ld be c a l c u l a t e d from th e d a t a .
w ise,

in order

O ther­

th e sa m p lin g and h a n d l i n g o f t h e v e g e t a t i o n were

conducted as d e s c r ib e d above.
A ll te s ts

f o r s t a t i s t i c a l d i f f e r e n c e s were con­

ducted u sin g B a r t l e t t 's

test

f o r hom ogeneity o f v a r i a n c e

and a n a l y s i s o f v a r i a n c e .

R esu lts
T o tal liv e

sh o o t biom ass f o r th e p r e s c r i b e d b u rn

a r e a a nd t h e f e r t i l i z e d a n d c o n t r o l p l o t s

in th e m ature

128
a n d c l e a r c u t j a c k p i n e a r e sho w n i n F i g .
sig n ifican t

fertiliz e r

6 .1 .

No

e f f e c t was d e t e c t e d d u r i n g

sam ple p e r i o d s 1 and 2, b u t t o t a l

liv e

sh o o t biom ass

i n t h e f e r t i l i z e d p l o t s was s i g n i f i c a n t l y g r e a t e r
(P < . 0 5 )

than the c o n tr o ls ,

m ature j a c k p in e ,
T o tal liv e

in b o th th e c l e a r c u t and

d u r i n g sam ple p e r i o d s

s h o o t b i o m a s s o n t h e b u r n e d s i t e was

g re a te r than f e r t i l i z e d

and c o n t r o l p l o t s

and c l e a r c u t j a c k p in e d u r i n g a l l
how ever,

3 a n d A.

th is

d ifferen ce

in m ature

f o u r sam ple p e r i o d s ;

c o u ld n o t be t e s t e d

sta tis­

tic ally .
In F ig u res 6 .2 ,

6 .3 ,

a n d 6 . A,

the t o t a l

liv e

sh o o t biom ass o f t h r e e dom inant s p e c i e s , sedge
(C arex p e n s y l v a n i c a ) , b l u e b e r r y
bracken fe rn

(P terid iu m aqu ilinu m D esf.)

fo r the f e r t i l i z e d
c le a rc u t jack p ine.
biom ass i s
th is

( V accin iu m s p p .) , and

g reatly

is

shown

and c o n t r o l p l o t s i n m a tu re and
F ig.

6 . 2 A a n d 6 .2 B show t h a t

s t i m u l a t e d by f e r t i l i z a t i o n ,

d i f f e r e n c e was o n l y s i g n i f i c a n t

(P < . 0 1 )

sedge

but

a t sam ple

p e r i o d A, i n b o t h m a t u r e a n d c l e a r c u t j a c k p i n e .

L ive

sh o o t biom ass o f b l u e b e r r y

(F ig,

i n c r e a s e d by f e r t i l i z a t i o n

i n m a tu re and c l e a r c u t j a c k

pine;

th is

6 . 3 A a n d 6 . 3 B ) was

d i f f e r e n c e was s i g n i f i c a n t

(P < . 0 5 )

during

sam ple p e r i o d A i n m a tu re j a c k p i n e and p e r i o d 3 i n
c le a rc u t jack p in e.

S ig n ific a n tly in creased

(P < . 1 )

b i o m a s s o f b r a c k e n f e r n was m e a s u r e d o n f e r t i l i z e d

400

350

CONTROL
FERTILIZED
PRESCRIBED BURN

_
m

300

_

5
H
0
0

200

_

cu
c
‘a

_

u
.«
a
3
(U
£

T

6

150

I
R

s
s

3
U
>_

4—»

0

n

I

T

250

100

_QJ

u

d

50 _

G
/„
0

I
1

6/14/80

T
7 /2 1 /6 0

T

T

8/2 8 /B O

7 /2 4 /8 1

BIOMASS SAMPLING DATE

Fig. 6 .1 .
T o t a l l i v e s h o o t b i o m a s s ( g / m 2) f o r t h e f e r t i l i z e d
an d c o n t r o l p l o t s i n t h e m a t u r e a n d c l e a r c u t j a c k p i n e , and t h e p r e
s c rib e d burn a re a .
V e r t i c a l b a r s e q u a l o n e s t a n d a r d d e v i a t i o n (SD)

130

Sedge

r

0
T
R

CONTROL

7ZAf e r t i l i z e d

L
L

1

V
E
S

H
0
0

zo

T

B
I
0
n
H
S
s
D
i2
6 /1 4 /8 0

7 /2 1 /8 0

7 /2 4 /8 1

8 / 2 0 /0 0

175

m
CONTROL
Z 3 FERTILIZED
150
125
5

H
0
0

100 .

T

B
I

0
M
R
S
S

2
6 /1 4 /8 0

7 /2 1 /8 0

0 /2 8 /8 0

7 /2 4 /0 1

SIQflfiSS S A M P L I N G D ATE

F ig . 6 .2 .
T o t a l l i v a s h o o t b i o m a s s ( g / m 2) o f
s e d g e (G a rex p e n s y l v a n i c a ) f o r t h e f e r t i l i z e d and
c o n t r o l p l o t s i n A) m a t u r e j a c k p i n e a n d B) c l e a r cut ja c k pine.
V e r t i c a l b a r s e q u a l o n e SD.

131

T

Blueberry

too

0
T
A

L
L

VZk
80

I
V
E
S

SO

H
0
0
T

a

40

i
o

n
A

s
s
g/m'

20

0
7 /2 1 /8 0

6 /1 4 /8 0

r
r

0
A
L

8 /2 8 /8 0

7 /2 4 /8 1

120

_
S
100

CONTROL
FERTILIZED

-

L

1

V
E

BO

S
H

0
0
T

8
I
0
n
A
s
s
0
/
MZ

40

6 /1 4 /0 0

7 /2 1 /8 0

BIOHflSS S R M P L I N O OflTE

F ig. 6 .3 .
T o t a l l i v e s h o o t b i o m a s s ( g / m 2) f o r
b l u e b e r r y (V accinium s p p . ) f o r t h e f e r t i l i z e d and
c o n t r o l p l o t s i n A) m a t u r e j a c k p i n e a n d B) c l e a r c u t
jac k p ine.
V e r t i c a l b a r s e q u a l o n e SD.

132

Bracken fern

T
0
T
R

C O N T RO L
FERTILIZED

L

L
[
V

E

100

.

S

H
0
0
r
e
i
o
n
a
s
s
o
L2
7 /2 1 /8 0

T

8 /2 8 /6 0

7 /2 4 /8 1

8 /2 0 /8 0

7 /2 4 /8 1

100

0
T
R
L

_

CONTROL

7ZA F E R T I L I Z E D

L

I

V

E
S

H
0
0
T

8

I
0
M
R

S
S

20

0

i2
6 /1 4 /8 0

7 /2 1 /8 0

B I O M R S S S R M P L I N O O RTE

Fig. 6 .4 .
T o t a l l i v e s h o o t b i o m a s s ( g / m 2) o f
bracken fe rn (P teridium a q u ilin u m ) fo r th e f e r t i l i z e d
a n d c o n t r o l p l o t s i n A) m a t u r e j a c k p i n e a n d B) c l e a r cut ja c k pine.
V e r t i c a l b a r s e q u a l o n e SD.

133
p lo ts

i n m a tu re j a c k p i n e d u r i n g sam ple p e r i o d s

a n d 4 , w h e r e a s no s i g n i f i c a n t f e r t i l i z e r
found i n th e c l e a r c u t a r e a

(F ig.

F e r t il i z e r response,

e f f e c t was

6.4A and 6 . 4 B ) .

c a l c u l a t e d as a r a t i o

l i v e biom ass on f e r t i l i z e d p l o t s

as w e ll as fo r t o t a l

6.5A and 6 .5 B ) .

liv e

in d icativ e

to f e r t i l i z e r

(F ig.

of the th re e

t h e i r t r e a tm e n t r e s p o n s e and can

be com pared to t h e t o t a l l i v e
w hich i s

and b ra c k e n

sh o o t biom ass

The b io m a s s r a t i o s

dom inants i n d i c a t e

s h o o t biom ass r a t i o ,

o f t h e a v e r a g e comm unity r e s p o n s e

treatm en t.

O nly se d g e s c o n s i s t e n t l y

sh o w e d a n a b o v e - a v e r a g e r e s p o n s e t o f e r t i l i z a t i o n
m ature ja c k p in e

(F ig.

6.5A ).

t o t a l annual p ro d u ctio n ,
f e r n in c r e a s e d w ith tim e.

Biom ass r a t i o s

sedge, b lu e b e rry ,

o f 1 .7 ;

at

in

for

and b ra c k e n

T h i s was m o s t e v i d e n t w i t h

s e d g e s , w h ic h a t sam ple p e r i o d 1 had a f e r t i l i z e d
co n tro l r a t i o

of

t o t h a t on c o n t r o l

p l o t s , was com puted f o r s e d g e , b l u e b e r r y ,
fern,

3

sam ple p e r i o d 4 t h i s

to

ratio

had i n c r e a s e d m arkedly to 4 .9 .
Biomass r a t i o s
a r e shown i n F i g .

fo r c le a rc u t ja c k pine p lo ts

6.5B .

response to f e r t i l i z a t i o n

S e d g e s sh o w ed a n a b o v e - a v e r a g e
during a l l

f o u r sam ple

p e r i o d s , b u t t h i s r e s p o n s e was m ost i m p r e s s i v e d u r i n g
p eriods

3 and 4.

T rends i n th e biom ass r a t i o s w ith

tim e as se e n i n th e m atu re j a c k p in e p l o t s a r e n o t
as a p p aren t in the c le a r c u t p l o t s .

134
6
5 .

m

TOTAL L IVE SH O O T B I OM A S S
SEO O E
BLUEBERRY
B R A C K E N FERN

8

I

0
M
n
s
s

3 .

R
a

r
i
o

2 .
1 .

7 /2 4 /8 1

6 /1 4 /S O

5

4

TO TA L L I V E SHOO T B I O M A S S
SEDDE
BLUEBERRY
B R A C K E N FERN

3

2

I

0
6 /1 4 /8 0

7/2 1 /0 0

8 /2 8 /0 0

7 /2 4 /8 1

B I O M A S S S A M P L I N O DATE

F ig. 6 .5 .
F e r t i l i z e d / c o n t r o l biom ass r a t i o s
f o r t o t a l l i v e s h o o t b i o m a s s , s e d g e , b l u e b e r r y , a nd
b r a c k e n f e r n i n A) m a t u r e j a c k p i n e a n d B) c l e a r c u t
ja c k pine.

135
Biomass d a t a from th e p r e s c r i b e d b u r n a r e a
a re p re s e n te d in T able 6 .1 .

M onocots, p a r t i c u l a r l y

t h e s e d g e s C. p e n s y l v a n i c a a n d C. r u g o s p e r m a M a c k . ,
c o n trib u te d g r e a tly to the t o t a l l iv e
of th e s i t e .

sh o o t biom ass

G ra ss e s w ere a l s o v e ry p r o d u c t i v e ,

t h e dom inant s p e c i e s i n c l u d i n g D a n th o n ia s p i c a t a
(L.)

B e a u v . , O r y z o p s i s a s p e r i f o l i a M ichx.

and 0.

pungens T o r r . , P ic h a n th e liu m depauperatum M u h l.,
and S c h iz a c h n e p u r p u r a s c e n s T o r r.

T re e s and sh ru b s

w ere a l s o an im p o r ta n t group i n term s o f p r o d u c t i v i t y
on t h i s b u r n , w i t h V a c c in iu m s p p . , P r u n u s p u m i l a L . ,
a n d C o m p t o n i a p e r e g r i n a - L.
than bracken fe rn ,

co n trib u ted l i t t l e

biom ass o f th e s i t e ,
co m p o sitio n .

dom inating.

F o rb s, o th er
to

the t o t a l

but g re a tly enriched species

Forb s p e c ie s

commonly f o u n d t h r o u g h o u t

th e b u rn w ere A s te r l a e v i s L . , H ieraciu m a u ra n tia c u m
* S e n e c i o t o m e n t o s u s M i c h x . , and C am panula r o t u n d i f o l i a L.

D iscu ssio n
B oth c l e a r c u t t i n g and b u r n in g in d u c e th e
rele ase of n u trie n ts

sto red in v e g etatio n ,

and s t i m u l a t e

o th e r n u t r i e n t c y c lin g p r o c e s s e s w hich " f e r t i l i z e "
the

site.

In the f i r s t

grow ing s e a s o n f o l lo w in g

c l e a r c u t t i n g , L i k e n s £t: a l .
so ils

as both " i r r i g a te d "

(1978)

d escribed fo re s t

(in c re a se d s o i l m o istu re)

136

TABLE 6 . 1
BIOMASS DATA ( g / m 2) BY SPECIES GROUP
FOR THE 1978 PRESCRIBED BURN

Sam ple D a te
6 /1 4 /8 0

7/2 1 /80

2.3

9.4

13.3

23.1
77.6

41.6
92.5

4 2.6
134.1

T re e s and
Shrubs
Annual
R esidual

45.6
9.5

101.8
28.5

123.7
24.7

T o ta l L ive
Biomass

148.6

245.3

313.7

Forbs

8/28/80

7/24/81
8.2 ±

2.2

M onocots
G rasses
Sedges

8 8 .3 ± 5.4
16 5 .9 ± 28.2

86.4 ±
50 .3 ±

7.3
5.2

348.8 ± 1 0.6

137
and " f e r t i l i z e d "

(elev ated c o n cen tratio n s of d isso lv e d

su b stan ces in s o i l s o l u t i o n ) .

A h lg r e n (1960) a t t r i ­

b u t e s th e r a p i d r e g r o w th o f v e g e t a t i o n and m arked
lu sh n e ss d u rin g the f i r s t

few p o s t - f i r e

to the f e r t i l i z i n g a c tio n of n u t r i e n t s
Borm ann £ t a l .

grow ing se a so n s
from t h e a s h .

( 1 9 6 8 ) h a v e show n s t i m u l a t e d n i t r i f i ­

c a t io n fo llo w in g c l e a r c u t t i n g , w hich th ey a t t r i b u t e
to w arm ing o f t h e s o i l and re m o v a l o f v e g e t a t i o n
w hich p o s s ib l y i n h i b i t e d

the n i t r i f i c a t i o n p ro cess.

I n c r e a s e d n u t r i e n t a v a i l a b i l i t y and e l e v a t e d l e v e l s
o f n i t r o g e n f i x a t i o n and n i t r i f i c a t i o n have a l s o been
r e p o rte d fo llo w in g f i r e

( J o rg e n s e n and W ells 1971,

C h r i s t e n s e n 1 9 7 3 , D e b an o a n d C o n r a d 1 9 7 8 ) .
N u tr ie n t enrichm ent w i l l in c r e a s e prim ary
p r o d u c t i v i t y o f b i o l o g i c a l com m unities.
F o ster

M o r r is o n and

(1977), w orking i n ja c k p in e s ta n d s in n o r th e r n

O ntario,

found a t h r e e - f o l d i n c r e a s e i n dry m a t t e r

increm ent fo llo w in g f e r t i l i z a t i o n .

In th is

a n e a rly tw o-fold in c re a s e in t o t a l liv e

study,

sh o o t biom ass

o c c u r r e d by th e m id d le o f t h e seco n d grow ing se a so n
due to f e r t i l i z a t i o n
pine stan d s.
tie s

in b o th m ature and c l e a r c u t ja c k

The d o m i n a n t s p e c i e s i n t h e s e com m uni­

(sedge, b lu e b e rry ,

and b r a c k e n f e r n )

co n sisten tly

sh o w ed i n c r e a s e d p r o d u c t i v i t y i n r e s p o n s e t o f e r t i l i ­
zatio n .

Sedge ^C. p e n s y l v a n i c a ) , h o w e v e r, was t h e

o n l y s p e c i e s t h a t c o n s i s t e n t l y sh o w ed a r e s p o n s e a b o v e

138
t h e comm unity a v e r a g e .
P r e f e r e n t i a l uptake of f e r t i l i z e r s
sp e cie s has p rofoundly a lte r e d

by c e r t a i n

th e d i v e r s i t y and

s p e c i e s dom inance i n w i d e l y v a r y i n g c o m m u n itie s .
W orking i n an 8 - y e a r - o l d f i e l d
a n d Odum ( 1 9 7 8 )

in G eorgia, B a k e la a r

found t h a t f e r t i l i z a t i o n

t h e dom inance o f g o ld e n r o d

(Solidago s p . ) ,

o r e l i m in a t e d o t h e r codom inant s p e c i e s .
fo rests

in creased

i n C o s t a R i c a , H a rco m b e ( 1 9 7 7 )

bu t reduced

In tro p ic a l
sh o w ed t h a t

f e r t i l i z a t i o n r e t a r d e d s u c c e s s io n by e n h a n c in g th e
c o m p etitiv e a b i l i t y

of a single forb

lacca riv in o id e s ) .

Stephenson

o ld -fie ld

species

(Phyto­

(1973), w orking in

com m unities, o b se rv e d a t r e n d tow ard o v e r ­

a l l r e d u c t i o n i n d i v e r s i t y and e v e n n e ss f o l lo w in g
fe rtiliz a tio n .

On c l e a r c u t n o r t h e r n h a r d w o o d f o r e s t s

i n New H a m p s h i r e ,
th at f e r tiliz a tio n

S t a f f o r d and F i l i p
g reatly

(1974) r e p o r t e d

s tim u la te d p in c h e rry to

t h e d e tr i m e n t o f com m ercial s p e c i e s su ch as y e llo w
b i r c h and p a p e r b i r c h .

S p e c h t ej: a l .

on s a n d - h e a t h v e g e t a t i o n i n A u s t r a l i a ,

(1977) , w o r k in g
sh o w e d t h a t

e ig h t years a f t e r f e r t i l i z a t i o n h eath sp ecies d eclin ed
a n d t h e n a t i v e g r a s s , Them eda a u s t r a l i s , e x p a n d e d
in to v acated gaps.
Sedges a re w e ll-a d a p te d to d is tu r b a n c e s
as f i r e ,
tio n .

r a d i a t i o n e x p o su re and f e r t i l i z e r

Abra ms a nd D i c k m a n n ( 1 9 8 2 ) r e p o r t e d

such

a p p lica­

139
e x tra o rd in a ry in creases

in sedge 3 to

6 years a f te r

c l e a r c u t t i n g a n d /o r b u r n in g i n j a c k p in e com m unities.
The lo w d i v e r s i t y o n many s i t e s w a s a d i r e c t r e s u l t
o f th e d o m in a tio n o f Carex p e n s y I v a n i c a .
through a v a r ie ty of avenues,
the a c tiv e

grow th o f th e l a r g e

buds a c c u m u la te d by C arex

D isturbance,

ap p aren tly stim u lates
"bank" o f dorm ant

( N o b l e et: a l .

W ithin th e i n t e r v a l of t h i s

1979) .

study,

I did not

s e e t h e e x p a n s i o n o f C a r e x , o r any o t h e r comm unity
member,

to th e p o i n t w here i t

excluded n e ig h b o rin g s p e c ie s .

g re a tly suppressed or
In f a c t ,

no s i g n i f i c a n t

o r even n o tic a b le d if f e r e n c e s e x is te d in sp e c ie s
r i c h n e s s betw een f e r t i l i z e d

and c o n t r o l p l o t s .

What

w a s shown w a s t h a t t h e b i o m a s s o f v e g e t a t i o n i n c l e a r c u ts and u n d er m a tu re j a c k p in e i n c r e a s e d a f t e r f e r ­
tiliz a tio n ,

a n d t h a t C. p e n s y I v a n i c a c o n s i s t e n t l y

showed a n above a v e r a g e r e s p o n s e .
however,

It

is

apparent,

t h a t th e e x p an sio n o f Carex fo llo w in g c l e a r -

c u t t i n g a n d b u r n i n g i n v o l v e s many m o r e f a c t o r s
sim ply i t s

than

resp o n se to n u t r i e n t enrichm ent o f th e s i t e .

For exam ple,
sig n ific an tly

in th is

stu d y Carex biom ass in c r e a s e d

in the c le a r c u t c o n tro l p l o ts w ith l a t e r

sam p lin g d a te , b u t d id n o t in c r e a s e w ith tim e in th e
c o n tro l p lo ts
was a l s o

under m ature ja c k p in e .

g reatly

C arex biom ass

s t i m u l a t e d on t h e b u r n e d s i t e

compared t o th e m a tu re j a c k p i n e s i t e

(F ig.

6 . 2A a n d

140
T able 6 .1 ) ,

a l t h o u g h c o m p a r i s o n s b e t w e e n t h e s e two

s i t e s m u s t b e made w i t h c a u t i o n .

O ther o v e rt changes

r e s u l t i n g from c l e a r c u t t i n g and b u r n i n g , b e s i d e s
n u tr ie n t enrichm ent,
m ade a v a i l a b l e

in c lu d e the in c re a s e d space

from t h e d e s t r u c t i o n o f v e g e t a t i o n ,

and i n c r e a s e d l i g h t ,
a t th e r o o t i n g zone.

s o i l tem peratures,

and m o is tu re

Most l i k e l y a l l t h e s e f a c t o r s

c o n tr i b u te to th e re s p o n se o f C arex fo llo w in g d i s ­
tu rb an ce.

CHAPTER V I I
APPARENT HEAT STIMULATION OF BURIED SEEDS
OF GERANIUM BICKNELLII BRITT. ON
JACK PINE SITES IN
NORTHERN LOWER MICHIGAN
In tro d u ctio n
The g e r m i n a t i o n o f b u r i e d s e e d s i s

an im p o rta n t

m echanism o f s p e c i e s e s t a b l i s h m e n t d u r i n g s e c o n d a r y
p lan t

s u c c e s s i o n i n man y e c o s y s t e m s

(Sweeney 1956,

M ajo r and P y o t t 1966, A h l g r e n 1 9 7 9 a , H a l l and Swaine
1 9 8 0 , McGraw 1 9 8 0 ) .

The a p p e a r a n c e o f a l a r g e v a r i e t y

o f s p e c i e s , n o t s e e n on u n b u rn e d a r e a s ,
fire

has been a t t r i b u t e d

o f b u rie d seed

to

(Went e t a l .

fo llo w in g

s tim u la tio n of g erm ination
1952,

Sweeney 1956,

F loyd 1966,

1976, C h r i s t e n s e n and M u lle r 1975,

S h e a a t aJ..

1979).

fire

V a r io u s f a c t o r s r e s u l t i n g from

have b e e n i m p l i c a t e d i n t h i s phenomenon.

e t a)..

Went

(1952) r e p o r t e d t h a t rem o v a l o f c o m p e t i t i o n

was a m a j o r f a c t o r

i n th e Abundant g e r m i n a t i o n o f

c h ap a rra l sp ecies the f i r s t

season a f t e r f i r e .

M cPherson and M u lle r

c o n c lu d e d t h a t h e a t from

fire

(1969)

d e g r a d e s some s u b s t a n c e i n t h e s o i l w h i c h o t h e r ­

w ise su p p re sses

germ ination o f c h a p a rra l sp e c ie s.

The a b u n d a n c e o f t h e a n n u a l S e n e c i o s y l v a t i c u s on
o n e -y e a r-o ld burned D o u g la s -fir
141

(P seudotsuga

142
m e n z i e s i i ) c l e a r c u t s and i t s
has been a t t r i b u t e d
ted w ith burning

disappearance t h e r e a f te r

to s o i l n u t r ie n t changes a s s o c ia ­

(W est a n d C h i l c o t e

1968).

R upture

or a l t e r a t i o n of th e w ater-im perm eable seed co at of
h a rd -s e e d e d s p e c ie s a llo w in g g e rm in a tio n has been
a ttrib u te d

to f i r e

M a r t i n e_t a l .

( F l o y d 1 9 6 6 , Cushwa e t a l .

1968,

1 9 7 5 , P u r d i e a n d S l a t y e r 1 9 7 6 , McDonough

1977).
G e r m i n a t i o n o f man y s p e c i e s
the f i r s t y e ar a f t e r

fire

to

(H orton and K ra e b e l 1955,

Sweeney 1 9 5 6 , West and C h i l c o t e
S l a ty e r 1976).

is re s tric te d

1968, P u r d i e and

On j a c k p i n e s i t e s

i n n o r t h e r n low er

M i c h i g a n , Abr ams a n d D i c k m a n n ( 1 9 8 2 )

r e p o r t e d t h a t many

s p e c i e s o n o n e - y e a r - o l d b u r n s w e r e n o t p r e s e n t on t h e
site

the fo llo w in g y ear.

The m o s t s t r i k i n g e x a m p l e

was G eranium b i c k n e l l i i B r i t t . , c l a s s i f i e d a s an a n n u a l
o r b i e n n i a l by F e r n a l d

(1950).

T his s p e c ie s r e p r e s e n t s

a s much a s 22% o f t h e v e g e t a t i o n a l c o v e r o n f i r s t - y e a r
b u r n s , b u t b y y e a r two i t w a s n o t p r e s e n t o r w a s r e d u c e d
to

sc a tte re d in d iv id u als.

fore,

I t was h y p o t h e s i z e d ,

t h a t h e a t fro m f i r e was r e s p o n s i b l e

appearance o f t h i s
To t e s t

th ere­

fo r the

s p e c i e s on o n e - y e a r - o l d b u r n e d s i t e s .

th is h y p o th esis,

tw o s e r i e s

o f experim ents

w ere i n i t i a t e d u s in g f r e s h ly - m a tu r e d geranium seeds
and se e d s b u r i e d i n t h e s o i l o f d i f f e r e n t - a g e j a c k p i n e
sites.

The p r i m a r y o b j e c t i v e o f t h e s e e x p e r i m e n t s

143
was t o d e t e r m i n e i f

h e a t was i n v o l v e d i n t h e g e r m i n a ­

t i o n o f geranium seed.

However,

i t was a l s o p o s s i b l e

t o d i s c e r n how w e l l t h e g e r m i n a n t s f r o m b u r i e d s e e d
m atched th e e x i s t i n g f l o r a o f th e s i t e .

E x p e rim e n ts W ith F r e s h ly - M a tu re d
Geranium S eea
F r e s h l y - m a t u r e d s e e d s c o l l e c t e d from p a r e n t
p lan ts

i n m i d t o l a t e A u g u s t 1980 f r o m s i t e s

7B80 w e r e u s e d i n t h i s

s e r i e s of experim ents.

r e p r e s e n t e d 13% o f t h e p l a n t
old p re sc rib e d burns.
m ixed t o g e t h e r ,

7A80 a n d
Geranium

c o v er on t h e s e o n e - y e a r -

T he s e e d s f r o m a l l p l a n t s w e r e

a ir d ried ,

th e n r e f r i g e r a t e d a t 1-2°C.

A se rie s of sm all-scale

g e rm in a tio n t e s t s were

u n d e r t a k e n b e t w e e n M a r c h a n d May, 1 9 8 1 t o d e t e r m i n e
th e f a c t o r s im p o rta n t in geranium seed g e rm in a tio n .
A l l t e s t s w e r e c o n d u c t e d i n p e t r i d i s h e s o n Whatman
//I f i l t e r p a p e r .

T hree o r four r e p l i c a t i o n s ,

co n sistin g

o f 10 t o 20 s e e d s e a c h , w e r e u s e d i n e a c h t r e a t m e n t .
P e t r i d i s h e s were p la c e d u n d e r " c o o l w h i t e " f l o r e s c e n t
l i g h t s w i t h a 14 h o u r p h o t o p e r i o d a n d a l i g h t i n t e n s i t y
o f a p p r o x i m a t e l y 20 m i c r o E i n s t e i n s m"2 s e c - ^-; t e m p e r a ­
t u r e s f l u c t u a t e d b e t w e e n 18 a n d 2 2 °C.

M o i s t h e a t was

u se d i n a l l t r e a t m e n t s by p l a c i n g s e e d on top o f 3 o r
4 lay ers of f i l t e r

paper s a tu ra te d w ith d i s t i l l e d w ater

d i r e c t l y b e f o r e h e a t in g i n a l a b o r a t o r y d r y in g oven.
S e e d s w e re s c a r i f i e d by n i c k i n g t h e r a d i c l e en d o f

144
th e seed w ith a ra z o r b la d e .

S tra tific a tio n

treatm en t c o n siste d of r e f r i g e r a t in g

(1-2°C)

(strat)
s e e d s on

m o i s t f i l t e r p a p e r f o r an 8 week p e r i o d i n d a r k n e s s .
The a l t e r n a t e w e t - d r y t r e a t m e n t ,

l a s t i n g 3 weeks,

in v o lv ed keeping seeds m oist fo r 5 days,
h o ld in g w ater fo r 5 days.
a ll

treatm e n ts,

then w ith ­

D uring th e t e s t s

seeds in

e x c e p t t h e w e t-d r y re g im e , w ere k e p t

c o n tin u a lly m oist w ith d i s t i l l e d w a te r.

Seeds were

c o n s i d e r e d g e r m i n a t e d when t h e r a d i c l e em erged from
th e seed c o a t.
The f o l l o w i n g t r e a t m e n t s w e r e a p p l i e d t o f r e s h
s c a r i f i e d and u n s c a r i f i e d geranium se ed :
( c o n t r o l ) ; 40°C-5 m in;
60 m i n ;

80°C-60 m in;

+ stra t;

70°C-5 m in;

no h e a t

80°C-10 m in;

7 0 ° C - 1 0 r a in + s t r a t ;

60°C-

70°C -20 min

70°C -60 m in + s t r a t ;

70°C-10 m in + f r e e z e

(1 week) + a l t e r n a t e w e t - d r y ;

7 0 ° C - 3 0 m i n 4- f r e e z e

(1 week) + a l t e r n a t e w e t - d r y .
R esu lts
None o f t h e u n s c a r i f i e d s e e d s g e r m i n a t e d .
c o n trast,

80-100% o f t h e s c a r i f i e d

In

seeds germ inated

in a l l tre a tm e n ts .

E x p e rim e n ts w i t h B u rie d Seed
From t h e a b o v e r e s u l t s ,

i t was a p p a r e n t t h a t i f

h e a t was i n v o l v e d i n g e r a n iu m s e e d g e r m i n a t i o n ,

seeds

r e q u i r e a p e r i o d o f dormancy i n th e s o i l b e f o r e h e a t

U 5

exposure.

T herefore,

s u r f a c e s o i l sam p les were

c o l l e c t e d from t h r e e d i f f e r e n t - a g e d j a c k p i n e s i t e s
alon g a p o s t - f i r e
u se d w ere s i t e
site

su c c e s s io n a l sequence.

The a r e a s

7B81 ( 3 - y e a r - o l d p r e s c r i b e d b u r n ) ,

12A ( 3 5 - y e a r - o l d j a c k p i n e s t a n d ) ,

and s i t e

13C

(5 5 -y ea r-o ld ja c k pine s ta n d ) .
On May 1 9 ,

1981,

12 r a n d o m l y l o c a t e d s o i l s a m p l e s

w ere c o l l e c t e d from e ac h s i t e .

The s a m p l e s w e r e

25 cm x 25 cm t o a d e p t h o f 2 . 5 cm i n t o m i n e r a l s o i l .
T herefore,

t h e volum e o f e a c h sam ple v a r i e d w i t h t h e

d ep th of th e o v e rly in g o rg a n ic m a te r ia l.
bagged s e p a r a t e l y and a i r - d r i e d
May 3 0 ,

1981.

in the

A f t e r rem oving a l l

e a c h s a m p l e was s i f t e d

lab o rato ry u n til

green v e g e ta tio n ,

t h r o u g h 60 mm m e s h s c r e e n i n g

t o rem ove l a r g e r t w i g s , r o o t s and r o c k s .
s a m p l e a 13 52 cm3 s u b s a m p l e w a s t a k e n .
(unheated c o n tr o l ,

Sam ples w ere

From e a c h
Four tre a tm e n ts

h e a t e d 7 0 ° C - 3 0 m i n . , h e a t e d 90 °C-

30 m i n . , a n d h e a t e d 7 0 ° C - 3 0 m i n + s t r a t ) , r e p l i c a t e d
3 tim e s , w ere random ly a s s ig n e d to
each s i t e .

H eated sam ples w ere p la c e d i n m e ta l t r a y s ,

m o isten ed s l i g h t l y ,
oven.

To r a i s e

70°C f o r

sam p les t a k e n from

30 m i n ,

and p la c e d in a f o r c e d - a i r d ry in g

t h e s a m p l e s t o a n d m a i n t a i n th e m a t
th e y were f i r s t

e x p o s e d t o 150°C

f o r 20 m i n ( p r e h e a t i n g ) , f o l l o w e d b y h e a t i n g a t 1 0 0 °C
f o r 30 m i n .

D uring th e p r e h e a t i n g ,

sam ples w ere p e r i ­

o d i c a l l y s t i r r e d and m o is te n e d to a llo w even h e a t i n g

146
and m o is te n in g th ro u g h o u t-

No a d d i t i o n a l s t i r r i n g

o r m o i s t e n i n g was done d u r i n g t h e f i n a l h e a t i n g p e r i o d .
A t h e r m o m e te r was i n s e r t e d

in to

the c e n te r o f each

sam ple t o m o n it o r t e m p e r a t u r e d u r i n g h e a t i n g .

The

h e a t i n g regim e u se d to r a i s e and m a i n t a i n sam ples to
90 °C f o r 30 m i n .
f o r 30 m i n .

i n v o l v e d p r e h e a t i n g s a m p l e s a t 1 8 0 °C

f o l l o w e d b y h e a t i n g a t 1 2 0 °C f o r 30 m i n .

The s a m p le s w e r e s t i r r e d

and m o is te n e d d u r in g p r e h e a t ­

in g , b u t n o t d u rin g the f i n a l h e a tin g .

In b o th the

70°C a n d 90°C h e a t i n g r e g i m e s t h e s a m p l e s r e m a i n e d m o i s t
through th e f i n a l h e a tin g p e rio d .
A l l s a m p l e s w e r e p l a c e d o v e r a 3 . 5 cm l a y e r
of s te r iliz e d

sand

( a u t o c l a v e d - 121°C f o r 60 m i n . )

i n 26 cm x 52 cm x 6 . 5 cm p l a s t i c

flats.

The sa m p le s

w e r e s p r e a d e v e n l y o v e r t h e s a n d t o f o r m a l a y e r 1 cm
deep.

The f l a t s

d esig n ated fo r

p laced in r e f r i g e r a t io n

(1-2°C)

s t r a t i f i c a t i o n were
f o r 8 weeks.

A ll o th er

f l a t s were p la c e d under " c o o l w h ite " f l o r e s c e n t
i n a c o m p le te ly random ized d e sig n .

lig h ts

The f l a t s w e r e

e x p o s e d t o a 14 h o u r p h o t o p e r i o d w i t h a n a p p r o x i m a t e
-2
-1
l i g h t i n t e n s i t y o f 36 m i c r o E i n s t e i n s m
sec
and to
t e m p e r a t u r e s t h a t f l u c t u a t e d b e t w e e n 20 a n d 2 5 °C.
flats

c o n ta in in g only s t e r i l i z e d

te c t p o ssib le

Two

sand w ere u sed to d e ­

c o n tam in an ts to th e e x p erim e n t.

The

f l a t s w ere s y s t e m a t i c a l l y r o t a t e d once a week t o r e d u c e
any p o s s i b l e p o s i t i o n e f f e c t s .

G erm ination counts

147
w e r e made e v e r y tw o d a y s f o r t h e f i r s t m o n t h o f t h e
e x p e r im e n t and once o r t w ic e a week t h e r e a f t e r .
A fter

s t r a t i f i c a t i o n was c o m p le te d ,

were p l a c e d under th e
flats.

th o se f l a t s

lig h t tra y s w ith the o th er

Sam ples w e re k e p t m o i s t t h r o u g h o u t t h e e x p e r i ­

ment by a d d in g ta p w a te r e v e r y 1 t o 2 d a y s .
stra tific a tio n ,
days.

D uring

s a m p l e s w e r e w a t e r e d e v e r y 7 t o 10

G erm inants w ere re c o r d e d as th ey ap p ea re d .

I n d i v i d u a l s n o t r e a d i l y i d e n t i f i a b l e w ere rem oved,
p o tte d ,

an d grown u n t i l t h e y c o u ld be i d e n t i f i e d .

The

e x p e r i m e n t w a s c o n c l u d e d a f t e r 18 w e e k s .
At t h e c o n c l u s i o n o f t h e b u r i e d s e e d e x p e r i m e n t ,
the c o n tro l f l a t s

from t h e 3 - y e a r and 3 5 - y e a r - o l d

s i t e s w ere h e a t e d t o 70°C-30 m in.
procedure)

to determ ine i f

( u s in g th e above

geranium se e d s i n th o s e

f l a t s w o u ld g e r m i n a t e when e x p o s e d t o h e a t .
R esu lts
No s e e d s g e r m i n a t e d i n t h e f l a t s
ste riliz e d
sp.

sand.

However,

came up i n o t h e r f l a t s

tam inants;

th ree

c o n ta in in g only

s e e d lin g s o f O x alis

and w ere c o n s i d e r e d c o n ­

t h i s s p e c i e s h a s n o t b e e n f o u n d on a n y

of the ja c k pine s i t e s

s t u d i e d i n n o r t h e r n low er M ic h i­

gan.
Most o f t h e g e r m i n a n t s a p p e a r e d d u r i n g t h e f i r s t
2 weeks o f th e e x p e r im e n t

(F ig.

7 .1 ).

A m oderate

amount o f g e r m i n a t i o n o c c u r r e d from w eeks 3 t o 8 and

148

* — ^ C U M U L A T I V E TOT AL
Q - - - B W E E K L Y T OT AL

200

150

100

50

1

2

3

4

5

G

7

6

9

10 11 12 13 14 15 16 17 16
WEEKS

Fig. 7 .1 .
C u m u l a t i v e an d w e e k l y n u m b e r
o f g e r m in a n ts fro m t h e c o n t r o l , 70°C-30 m in ,
a nd 90°C-30 m in t r e a t m e n t s i n t h e 3 - y e a r - o l d ,
3 5 - y e a r - o l d , and 5 5 - y e a r - o l d s i t e s (com bined)
d u rin g the b u rie d seed experim ent.

149
very l i t t l e

th ereafter,

among s p e c i e s

(F ig.

7 .2 .).

peaked th e f i r s t week,
geranium a p p e a re d .

b u t t h e r e were d i f f e r e n c e s
G eranium g e r m i n a t i o n

a n d b y t h e f o u r t h w e e k n o new

A s u b s t a n t i a l number o f g r a s s

s e e d s g e rm in a te d d u r i n g t h e f i r s t week, b u t peak
g erm in atio n occurred d u rin g the

s e co n d week.

g e r m i n a t i o n o f hawkweed ( H i e r a c i u m s p p . )

Peak

occurred

b etw een weeks f i v e and e i g h t .
When h e a t e d t r e a t m e n t s
+ stra t)
site,

(70°C,

9 0 ° C , a n d 70°C

a r e compared t o t h e u n h e a te d c o n t r o l s a t e a c h

some s t r i k i n g d i f f e r e n c e s i n

species

and number o f g e r m in a n ts a r e e v i d e n t
Most i m p o r t a n t l y ,

com position

(T able 7 .1 ) .

geranium s e e d l i n g s a p p e a re d o n ly

i n th e h e a te d t r e a t m e n t s from th e 3- and 3 5 - y e a r - s i t e s .
The d i f f e r e n c e i n t h e number o f g e ra n iu m g e r m i n a n t s
in the

9 0 °C a n d 7 0 oC + s t r a t

t r e a t m e n t s compared t o

th e u n h eated c o n tr o l s were s i g n i f i c a n t

(P < . 0 5 )

W i l c o x o n ' s n o n p a r a m e t r i c two s a m p l e t e s t
T o r rie 1960).

55-year s i t e .

and s e d g e s

from any s i t e .

( S t e e l and

No s e e d s o f g e r a n i u m g e r m i n a t e d i n

sam ples from th e
grasses

using

O nly a few s e e d s o f

germ inated in th e h e a te d tre a tm e n ts

In c o n tr a s t,

58 a n d 22 g r a s s a n d s e d g e

g e rm in a n ts were r e c o r d e d i n th e c o n t r o l f l a t s

from

th e 3 5 -y e a r and 5 5 -y e a r s i t e s ,

The

resp ectiv ely .

d i f f e r e n c e i n t h e number o f g r a s s

and sed g e g e r m in a n ts
4

betw een th e h e a te d and c o n t r o l t r e a t m e n t s

from a l l

150

so
*— “ GERANIUM
B 0 GRASSES
HRWKWEED
40

30

20

WEEKS

Fig. 7 .2 .
W ee k ly t o t a l n u m b e r o f g e r m i n a n t s
o f g e ra n iu m , g r a s s e s ( a l l s p e c i e s c o m b i n e d ) , and
hawkweed ( H i e r a c i u m s p p . ) f r o m t h e c o n t r o l , 70 °C30 m i n , a n d 9 0 ° C -3 0 m in t r e a t m e n t s i n t h e 3 - y e a r o l d , 3 5 - y e a r - o l d , and 5 5 - y e a r - o l d s i t e s (com bined)
d u rin g th e b u r ie d seed ex p erim e n t.

TABLE 7.1
TOTAL NUMBER OF GERMINANTS FOR SPECIES APPEARING IN SOIL SAMPLES
FROM THE 3-YEAR, 3 5 -YEAR-, AND 55-YEAR-OLD POST-FIRE SITES
DURING THE BURIED SEED EXPERIMENT.
TREATMENTS WERE AN
UNHEATED CONTROL, 7 0 aC - 3 0 MIN, 9 0 ° C - 3 0 MIN, AND
7 0 ° C - 3 0 MIN + STRATIFICATION

3-year
Species

cone

70 °C

90 “C

55-year

35-year
70°C +
strat

cont

70°C

90 “C

70°C +
strat

cont

70°C

21

15

90°C

70°C +
strat

Annual h e r b s
32

G e i a i t - i u m ii -ccf cucCI ' t . t

10

23

P e r e n n ia l herbs

151

Ante.nna.riia n e g l e c t a

F ia g a 't.ta v i.tg in -ta iia
Hic^ac-tum 4 pp. 1
V io la adunca

3

Woody p e r e n n i a l s

Ch-imap/i-tf a a n b e . l la . ta

10

G a u t t h c n i a prioc.umbe.nl>
G r a s s e s and s e d g e s
C tuet

2

4pp.

Vanthonia i p i c a t a
V i c h a n t k e t i u m depaupe>iatum
Q n t j z o p i i i pUJt£j£H4
Paiucum c a p itla ric
Poa p^a-t£H4t4
U nidentified grasses
Treatment t o t a l s

2
1

23
4
7
3
17

17

1

2

21

37

H i e r a c i u m a u r a n t i a c u m and H. venosum

24

65

11

56

22

152
s i t e s was s i g n i f i c a n t
sam ple t e s t .

(P < . 0 1 )

u s i n g W i l c o x o n ' s two

The t o t a l n u m b e r o f g e r m i n a n t s i n t r e a t ­

m en ts from e a c h s i t e r a n g e d from 0 t o 65, b u t t h e
d i f f e r e n c e s w ere n o t s i g n i f i c a n t .

However,

th e most

f r e q u e n t l y o c c u r r i n g s p e c i e s i n e ac h t r e a t m e n t from
th e s i t e s

d iffered .

F o r exam ple,

on t h e 3 - y e a r s i t e w ere g e ra n iu m ,
th e 3 5 -y e a r s i t e were g r a s s e s .

68% o f t h e g e r m i n a n t s
a n d 71% o f t h o s e o n

Hawkweed, w i t h 36 g e r m i ­

n a n t s , was th e d o m in an t s p e c i e s on t h e 5 5 - y e a r s i t e .
G rass s p e c ie s and w in te r g r e e n

( G a u l t h e r i a procum bens

L . ) a l s o w e r e i m p o r t a n t m em bers o f t h e b u r i e d s e e d
p o o l on t h i s

site.

Many i m p o r t a n t m e m b e r s o f t h e p l a n t c o m m u n i ty
on t h e s i t e s

d id n o t a p p e a r as e m e rg e n ts from b u r i e d

s e e d (A ppendix B . 2 . ) .
seedlings
u b iq u ito u s

N otew orthy i s

the s c a r c ity of

o f C a re x and a b s e n c e o f b l u e b e r r y ,
com ponents o f j a c k p i n e s i t e s

low er M ichigan.

both

in n o rth ern

A l s o , many s p e c i e s t h a t g e r m i n a t e d

from b u r i e d s e e d w ere n o t su rv e y e d on t h e r e s p e c t i v e
sites.

E xam ples o f t h i s

um bellata
year s ite ,

(L .)

B art,

a re w in te r g r e e n and C him aphila

(common p i p s i s s e w a )

geranium , A n te n n a ria n e g le c t a

from t h e 3Greene

(field

p u s s y t o e s ) , w i n t e r g r e e n , Fanicum c a p i l l a r e L . , and
common p i p s i s s e w a f r o m t h e
weed, V io la adunca Sm .,
sp icata

(L .)

35-year s i t e ,

and hawk-

(h o o k ed -sp u r v i o l e t ) , D anthonia

B e a u v ., D ic h a n th e liu m depauperaturn M u h l.,

153
and P.

c ap illa re

from th e 55- y e a r s i t e .

When t h e c o n t r o l f l a t s
y e a r - o l d s i t e s w ere h e a t e d ,

fro m t h e 3 - y e a r and 35one f l a t

from t h e 3 - y e a r

s i t e p r o d u c e d 7 g e r a n i u m g e r m i n a n t s a n d two f l a t s
the

from

3 5 -y e a r s i t e prod u ced 2 and 3 geranium g e rm in a n ts .

D iscu ssio n
Many c h a p a r r a l s p e c i e s common t o r e c e n t l y b u r n e d
a r e a s p ro d u c e s e e d s w h ich re m a in dorm ant i n th e s o i l
b etw een f i r e s
1975b).

(Sw eeney 1 956, C h r i s t e n s e n and M u l l e r

H e a t t r e a t m e n t h a s b e e n shown t o r e l e a s e

seeds

o f s e v e r a l c h a p a r r a l s p e c i e s from en d o g en o u sly e n fo r c e d
d o r m a n c y ( S t o n e a n d J u h r e n 1 9 5 1 , Sw e e n e y 1 9 5 6 , C h r i s t e n ­
s e n and M u l l e r 1 9 7 5 a,
g erm ination in t h is

1975b).

study,

C o n s is te n t w it h geranium

g e rm in atio n o f b u rie d seed

of c e rta in ch ap arral species

occur a f t e r h e a tin g s o il

sam ples, w hereas f r e s h l y m atured seed did n ot germ in ate
a f t e r h e at treatm en t

(M cPherson and M u lle r 1969,

C h r i s t e n s e n and M u lle r 1 9 7 5 a ).
(1975a)

C h ris te n s e n and M uller

sp e cu la te th a t germ ination of th ese sp ecies

may be d e p e n d e n t o n t h e g r a d u a l d e t e r i o r a t i o n o f s e e d
tissu e

im p o s in g m e c h a n i c a l r e s t r i c t i o n on t h e embryoj

dormancy o f s e e d s s t o r e d i n t h e s o i l f o r l o n g p e r i o d s
may r e s u l t f r o m c h e m i c a l i n h i b i t i o n b y n e i g h b o r i n g
v eg etatio n .

R esearchers

source o f to x in s

(e.g .

s u g g e s t f i r e rem oves th e

shrub f o lia g e )

and d e n a tu re s

154
r e s i d u a l chem icals in th e s o i l ,

th ereb y allo w in g i n ­

c r e a s e d g e r m i n a t i o n o f many s p e c i e s

( M u l l e r ej: a l .

1968, M cPherson and M u lle r 1969, C h r i s t e n s e n and
M u l l e r 1975a*

1975b).

However,

o f many h a r d - s e e d e d s p e c i e s

abundant g erm in atio n

fo llo w in g f i r e has been

a ttrib u te d d irec tly

t o a l t e r a t i o n o f t h e s e e d c o a t by

heat

1968, P u r d ie and S l a t y e r 1976,

(C u sh w a e £ a l .

G ill

1977).
The a p p e a r a n c e o f G e r a n i u m b i c k n e l l i i f o l l o w i n g
fire

( A h l g r e n 1 9 6 0 , Ohmann a n d G r i g a l 1 9 7 9 , O u t c a l t

a n d W h i t e 1 9 8 1 , A bram s a n d D i c k m a n n 1 9 8 2 )
distu rb an ces

such as c u lt i v a t io n ,

and ro a d c o n s t r u c t i o n

b u i ld i n g rem oval,

(A h lg re n , p e r s o n a l com m unication)

h a s b e e n d o cu m en ted i n t h e Lake S t a t e s .
sites

and o t h e r

On j a c k p i n e

i n n o r t h e r n Lower M i c h ig a n , p e a k o c c u r r e n c e o f

geranium i s r e s t r i c t e d

to the f i r s t y e a r a f t e r

e i t h e r no o r v e r y r a r e

in d iv id u a ls of t h is sp ecies

a r e p r e s e n t on o l d e r b u r n e d s i t e s .
geranium b e h a v io r ,
Lake S t a t e s

how ever,

s i t e s where t h i s

fire;

T his p a t t e r n of

does n o t o c c u r on a l l
s p e c i e s grow s.

For

e x a m p l e , A h l g r e n ( 1 9 7 9 c ) r e p o r t e d t h a t G. b i c k n e l l i i
occurred in

90 Z o f h i s

sam ple p l o t s

th ree years

(equiva­

l e n t t o 2 y e a r s u s i n g my c r i t e r i a o f d i s t u r b a n c e a g e )
a f t e r an o ld -g ro w th r e d p in e s ta n d i n n o r t h e a s t e r n
M i n n e s o t a was b u r n e d .

K r e f t i n g and A h lg re n

a ls o w orking in n o r t h e a s t e r n M innesota,

(1974),

found geranium

155

i n 27% o f t h e p l o t s

su rv e y e d on a 5 - y e a r - o l d b u rn

( 4 - y e a r s u s i n g my c r i t e r i a ) .
A hlgren (1979a,
seeds in s o i l

1979b,

1979c)

e x tra c te d b uried

s a m p l e s f r o m many s i t e s

f o r e s t ty p e s and b u r n in g h i s t o r i e s ,

of d ifferen t

and c o n s i s t e n t l y

fo u n d g era n iu m t o be an i m p o r t a n t com ponent.
i n g l y , w hen A h l g r e n

In tere st­

(1979c) m o n ito r e d s e e d l i n g emergence

f r o m i n t a c t s o i l b l o c k s c o l l e c t e d f r o m a 270 y e a r - o l d
r e d p in e s t a n d b u rn ed 3 y e a r s p r e v i o u s l y and from an
a d ja c e n t unburned a re a ,
th e burned s o i l .

g e ra n iu m a p p e a r e d o n l y from

A h lg re n (1979a) p l a n t e d s o i l - e x t r a c t e d

geranium seed i n s t e r i l e

g ree n h o u se s o i l and o b ta in e d

30% g e r m i n a t i o n .
These d a ta a re
my e x p e r i m e n t s .

in c o n tr a s t to th e fin d in g s in

In A h lg re n 's e x p erim en ts,

t r e a t m e n t was g i v e n t o t h e s o i l
te d seed,

no h e a t

sam ples or s o i l e x t r a c ­

y e t th e geranium seed g e rm in a te d .

experim ents w ith b u rie d seed ,
only in f l a t s

I n my

geranium seed s g e rm in a te d

exposed to h eat tre a tm e n t.

T h i s was f u r ­

t h e r s u b s t a n t i a t e d when i n 3 o f 6 c o n t r o l f l a t s

from

t h e 3 - y e a r and 3 5 - y e a r s i t e s , w h i c h showed no g e ra n iu m
g e r m i n a t i o n f o r 18 w e e k s , g e r a n i u m g e r m i n a t e d a f t e r
t h e s o i l was h e a t e d .

A lso,

seed s e x t r a c t e d from b u lk

s o i l s a m p le s fro m t h e 3 - y e a r s i t e when h e a t e d t o
90°C-30 m in .

sh o w e d 20% g e r m i n a t i o n ( 3 7 s e e d s t o t a l ) ,

w h e r e a s u n h e a t e d s e e d s sh o w ed 0% g e r m i n a t i o n

(30 s e e d s

156
to ta l)

(Abr ams a n d D i c k m a n n , u n p u b l . ) .
Geranium i n M in n e s o ta ,

in the f i e l d a t l e a s t ,

does n o t r e q u i r e h e a t from f i r e

fo r germ ination

(A hl­

g re n , p e rs o n a l co m m unication).

The a p p a r e n t h e a t

r e q u i r e m e n t f o r g e ra n iu m g e r m i n a t i o n i n n o r t h e r n low er
M ichigan s u g g e s ts t h a t e c o t y p ic d i f f e r e n c e s e x i s t f o r
th is

sp ecies.

CHAPTER V I I I
SUMMARY AND CONCLUSION
1.

B u rn in g prom oted th e e s t a b l is h m e n t o f a l a r g e
v a rie ty

o f s p e c i e s n o t fo u n d on unburned c l e a r c u t s .

The l o n g e v i t y o f m o s t o f t h e s e s p e c i e s was s h o r t
(tem porary f i r e - f o l l o w e r s ) , r e s u l t i n g in la r g e
d r o p s i n s p e c i e s r i c h n e s s b e t w e e n o n e a n d tw o
years a f te r
2.

fire.

Through y e a r s i x a f t e r
r e m a in e d more d i v e r s e

fire,

m ost burned s i t e s

th an unburned c l e a r c u t

s i t e s o f analogous a g es.

S ite

d iv ersity ,

however,

on b o t h b u r n e d and u n b u r n e d s i t e s was d i r e c t l y
a f f e c t e d by th e dom ination o f th e sedge,

Carex

p ensylvanica.
3.

The p r o l i f e r a t i o n o f C. p e n s y l v a n i c a
re la tiv e
th at i t

cover)

(up t o 86%

fo llo w in g d istu rb a n c e in d ic a te s

is an o p p o r tu n is tic sp e c ie s

m o nopolizing re s o u rc e s

lib erated

cap ab le o f

fo llo w in g d i s ­

tu r b a n c e and s u p p r e s s in g o r e x c l u d in g o t h e r s p e c i e s .
4.

U nburned c l e a r c u t s i t e s

co n sisten tly

r a p i d l y t o C a r e x meadows.
sh o w e d m u l t i p l e

converted

B urned s i t e s ,

su ccessio n al p a tte r n s ,

d o m i n a t i o n b y s h r u b a nd e a r l y

157

how ever,

in cluding

s u c c e s s i o n a l hardw ood

sp ecies,

c o n v e r s i o n t o C a re x meadows, and t h e

e stab lish m en t of ja c k pine rep ro d u c tio n .
C o m p a r i s o n s made b y t h e i n d e x o f s i m i l a r i t y
showed t h a t s u c c e s s i o n on e a c h s i t e
unique.

T herefore,

each s i t e ,

(S)

is h ig h ly

th e in d iv id u a lis tic n atu re of

r a t h e r th a n age f o l lo w in g d i s t u r b a n c e ,

becomes t h e d o m in a n t a s p e c t i n d i s c e r n i n g
s u c c e s s i o n a l r e l a t i o n s h i p s i n th e s e com m unities.
Through th e u se o f p erm anent p l o t s e s t a b l i s h e d
p r io r to b u rn in g in a ja c k p in e c le a r c u t a re a ,
i t w a s shown t h a t t h e i n t e n s i t y o f t h e same f i r e
g o i n g t h r o u g h two a d j a c e n t b l o c k s w i t h s i m i l a r
flo ristic

c o m p o s i t i o n a n d e d a p h i c f e a t u r e s was

m arkedly d i f f e r e n t .

F i r e i n t e n s i t y , m e a s u re d by

org an ic m atter re d u c tio n ,

proved to a f f e c t

a d d i t i o n and lo s s o f s p e c i e s ,
d iv ersity ,

the

s p e c i e s dom inance,

and p l a n t c o v e r on t h e b l o c k s .

F e r t i l i z a t i o n e x p e r i m e n ts on m a tu r e and c l e a r c u t
ja c k pine s i t e s

sh o w e d t h a t b i o m a s s o f u n d e r s t o r y

v eg etatio n in creased a fte r
pensylvanica c o n s is te n tly
resp o n se to f e r t i l i z e r ,
a s an o p p o r t u n i s t i c

fertiliz atio n .

C arex

sh o w ed a n a b o v e a v e r a g e

su p p o rtin g i t s

rep u tatio n

species.

E xperim ents conducted w ith s u rfa c e s o i l
from d i f f e r e n t - a g e d j a c k p i n e s i t e s

sam ples

in d icated th at

t h e o v erw h elm in g dom inance o f G eranium b i k n e l l i i

159
o n many o n e - y e a r - o l d b u r n e d s i t e s w a s d u e t o t h e
h e a t-s tim u la te d g erm ination of b u rie d seed.

APPENDIX A
RELATIVE COVER AND RELATIVE FREQUENCY DATA
FOR SPECIES SURVEYED ON UNBURNED
CLEARCUT SITES

TABLE A 1
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 1B80-81 SURVEYED AS A 0-YEAR AND 1-YEAR-OLD
CLEARCUT.1
1981

1 98 0

T r e e s and s h r u b s
Awef aiic/u c i ip^
A t c f c i t n p/ n y f c i l a’f l - u i i t
Cempfetii'a p e i e g i t n a
C in fa e g u i ip.
Epigen .lepcm
P l n n u l pumciln
Piunui i c t i i f t u n
Pi iom i iu’I j u H iI nn
O i i i /i c m i p p . ( r e d oak)
flu b U i
P l' <1 i t I V A II U ’ l l i
Re in b l amda
S a tri * g i n u c c p h ij t! t' c t d c 1
I'ac c t il ( urn i p p .

rlndicates

a vaLue l e s s

I

l_n

1.5

3]

16. 6

4!

_2

1.6
l.l

.9

7.5

4.

.5

1.

1
.1
.1
1.4
.

1.1
26.0

13.5
.4
.3
2.9

2.2
1.8
2.7

.6

5.4

31.0

chan . 0 5 .

160

1

2.

4. 3
.5
14.5
.5

.1

4!

33. 7

12.

5.9

3.6

1.1
2.7
.5
3.8
5.9
2.7
.5

2 .1

1. 3

.

4. 3
.5
.5
15.6

O

1.9

13.9

O

.6
9.7
.5
.5
6.4

\_r

.8
9.8
.1

O

P e r e n n i a l herbs
Ape c tj uum a i i d \ u l a cm< fje£Iurri
A i c l c p t a a i t j f ii a c a
Ai t c i i a e i ’-i j
Flag A lia v i i g i n i a u a
Ga ii l f /ic 1 <a pludumtie n i
Htc-iac ium a u ' i a n - t tncum
Ma iai: tficmum c Aiinde m e
P / j i / i j I I i u i l g i i U ana
P-f e l tdium a q ui C-inum
Sc I’ t dage i p p .
1/ ( I' f a a dune a

.5

.3
.

Freq

U 't d '

G r a s s e s and s e d g e s
Eu-rnui k a i m i i
Cnicx pc ii i i/i'i'iitu' cti
P.ui t / i e »] t ti i p t e a f.i
P-i c/if.nT/tcf i urn dcpn ape liilitm
P t i / c c p j . i i a i p e 11 j e C La
Q \ i j z c p h i i pungcui
Sc/i <m e / m e p u i p a i n i ecu 1

C o ve r

\o

A n n u a l s and b i e n n i a l s
L a c t uc n c a n a d v i: i T i
Mi't' amp s j i uin N i u e a n ;

F req

i>

Cover

L /1 Ln

Species

2.8

6.9
1.5
3.0
3.4
.5
7.9
5.0

2.8

7. 4

24.8

14.8

.6
.1
.4
3.5

TABLE A 2
RELATIVE COVER AND RELATIVE FREQUENCY OF
EACH SPECIES ON SITE 2 SURVEYED AS A
1-YEAR-OLD CLEARCUT
1 97 9
S pecies

Cover

Freq

4. 4
19.2

9.7
1 4.6
1.6

G r a s s e s and s e d g e s
Andnopogon gznandZZ
Ca n z x . p z n A y Z v a n Z c a
17 e . 6 c h a m p 6 l a f i Z z x u o A a
OnyzopAZA a A p z n Z i o Z Z a
OnyzopAZA pungznA

1.1
1 .6

P e re n n ia l herbs
An z m o n z q u n n q u z f i o Z Z a

.2

AntznnanZa n zg Z zcta
A A tz n ZazvZA
EpZgza nzpznA
Gau.Zthzn.Za p n o c u m b z n A
HzZZanthuA occZ dzntaZ ZA
HZzn.acZu.rn v z n o A u m
LZatnZA n o v a z - a n g Z Z a z

.7
6.2

MaZanthzmum c a n a d z n A Z
PtznZdZ um aquZZZnum
SoZZdago App.
S p Z n a n t h z A gnacZZZA

T re e s and sh ru b s
11.2
2.5
1.1
6.2
.4

AnctoAtaphyZoA uva-unAZ
C o m p to n Z a pzn.zgn.Zna
PZnuA b a n k A Z a n a
PnunuA pum ZZ a
PnunuA A z n o t Z n a
Q, uzncuA A p p . ( r e d o a k )
V a c c Z n Z u m App.

46.8

161

3.2
.5
5.4
1.1
6.0
1.6
1.6
.5
1 .1
.5
1.1
.5
13.5
10 .3
1.6
9.2
.5
14.1

TABLE A 3
RELATIVE COVER AND RELATIVE FREQUENCY OF
EACH SPECIES ON SITE 3A SURVEYED AS A
2 -YEAR-OLD CLEARCUT
1980
S pecies

Cover

Freq

G rasses and sedges
~AgnoAt ZA k y z m a Z Z A
B-'iomtti k a Z m Z Z
Canzx pznAyZvanZca
VanthonZa ApZcata
VZchantkzZZum dzpaupznatum
Ko.zZznZa m a c n a n t k a
OnyzopAZA aApznZ&oZZa
OnyzopAZA p u n g e n t
P a n Z c a m coZu.mbZa.num

..6
44 .3
.4
2.5
.1
1.7

.6
4.9
17.9
4.9
6.2
.6
1.2
5.6
1.8

P e re n n ia l herbs
Apocynum andnoAazmZ&oZZum
A A tzn ZazvZA
CampanuZa n o tu n d Z & o Z Z a
HZzn.acZu.rn a u n a n t Z a c u m
HZznacZum vznoAum
LZatnZA novae- a n g Z Z a z
P tzn Zd Z u m aquZZZnum

.1
.5
*
.6
19.6

So£^.dago 4 p p .

1.2
2.5
1.2
1.8
1.8
.6
10.5
.6

T re e s and sh ru b s
1.3
1 .1
1.7
.2

AmzZanckZzn. Ap.
AnctoAtaphyZn.A uva~unAZ
C o m p to n Z a pzn.zgn.Zna
C n a ta zg u * Ap.
PZnuA b a n k A Z a n a
PnunuA pumZZa
PnunuA A z n o t Z n a
QuzncuA A p p .
R u b iu p z n A Z Z v a n Z c u A
VaccZnZum App.

.6
1.6
2 .5
20 .6

162

3.1
1.8
4.9
.6
.6
6.8
1.2
1.2
15.4

TABLE A 4
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 3B79-80-81 SURVEYED AS A 2-YEAR-,
3-YEAR-, AND 4-YEAR-OLD CLEARCUT
1979
Cover

A n n u a l s and b i e n n i a l s
Me C a mp y t urn f t iic a '1C

3.8

5.7
28.8

15.3

.6

51.4

P t e ltd-turn n t j u t C o u i m
ScCtrfayc ip p .
Vn'Ca a d u n c a
Vt e £ a p e d a t t litrfa

T r e e s and s h r u b s
Ami; Cane (i t e * i p .
Al c f 0 i t apJ: </£ e l u i M - u t i t
CampCenta p c i e j - m i i i
P (' .!(!

J

i i ' p L'>:i
f; i
pa" i

(iiT i,

I’1 in; u i

t
ia

1.5
.3

Re j a

.

1

2.7
.1
9.7
.3

.6

6.0

1.

.2

13.4

2.0

.6
12. 7

.6

9. 3

1.0

13 .

.4

3.

1.3
.7

2.3

10.0
2.7

l'.
.7

1.2

1
.1

11.5

.3

.4
6.7

.

1

6.0

1.3
5.7

1.3

9. 3

1.9

.3

.7
.7
1. 3
19.3

.6

j p p . ( r e d oak)

1

.6

36. 9

1 9. 1

.

1.3
3. 3
1. 3

1.4
4 .3

1 'iar.L i

Rn H u i p e n i l t i a i i c u i
P a c e t i l t urn 1 p p .

1 .3
1.3

3.2

f’lUllUi l l 1J I II Ulllil
(i ui.11 l a i

21.7

.6

.

i; a

44.8

2.0

.7

P I C 1 V l f l .1 t .; U' V la
Cp i g e a

2.9

"J ^ \Q "j

1. 3
1.9

1.5
.5

Lr t Cl ’ t i

e t t g <» t a n a
G a u i ! t h c i i a p.iocuiiibct: j
ft t e a a c t u m auAanCtaeurn
Hi 'c iac tum I'tiiL'ium
L i a f t f 4 ii l*!■a c - a i i g f t a e

.2
3. 3
17.3

1.9

P e r e n n ia l herbs
ApLieyuum aiid-toia c mt £et' tuin
Fiatjaita

Freq
1.4

1.9

S ; Ii i ; a c (me p u ' i p u i a i c c u i

AlteA

Cover

ro

puntjt' iii

Freq

o

Piy^Lipi t i

Cover

^'4 0'

G r a s s e s and s e d g e s
AiiiMi'jJi'Qlih ■)c i a d < t
Blii mu 4 Crifmtt
C a t e * pcuiyf.'ra»it ca
Paiif/ieiua i p t c a f n
On i c/inmpj t a ■Jfexuoia
P t c Ii a n f (ic f i um d c p i i u p c i a t u m
0 1 tj : u p 1 t i a i p e i i ii o f <a

19 81

1980
F r eq

163

40.4

.4
.7

1.6
4.2

<hD -'-j ro n.v•“j

S p ec i e s

8.0
1.5

.

1

45.8

') r)
2 1 ‘. 7

TABLE A 5

RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 4 A 7 9 - 8 Q - 8 1 SURVEYED AS A 3 - YEAR- ,
4 -YEAR- , AND 5 -YEAR-OLD CLEARCUT
1979
Species
A n n u a l s and b i e n n i a l s
Ai a 6 i i tj i a i *ta
G r a s s e s and s e d g e s
B to mu 1 tiU’mti
Caii!X pe m y I’vniit ca
Pan f fii’ii <a J p i c a f a
P m r fm inp i <a f ' l i ' i u i ' U
V <c fin n t /ic t turn licpaupc * a f urn
O i y r e p l i i a i p e i-i ^«.■t ta
O t y r v p i i i pa it 3 e >ii
P e r e n n ia l herbs
Aj t c 1 £ a c v i i
Cnmp.tiiufa l a t u h d-i jfc 1 i a
Hi i! 1 a c i u m it a a a ti 1 1 a c urn
Hi i? in c turn vc it a 1 am
I. y e a pad turn a b i c a l u i n
P f C l t d i u m ntfu i f ilium
S e ltd a g e ipp.
Pi a f a a d uu c a
T r e e s and s h r u b s
AniC f a DC ill C1 i p .
A i c t a jfapfi yl' u i u v a - u - . i i
Cernpteiun p e i C y l t i t a
Ptuui banhiitum
P r p u f u i f 1 emiHe t d e i
PltuiuJ i i i i e t i n i i
( • u c i e u i i p p . ( r e d oak)
Piii:c ituuin i p p .

Cover

.

1

.
77.
.
3.
1.

8
5
4
4
1

1 90 0
Cover

Freq

Cove r

Freq

6.4
27 . 3
1.8
15.5

64.1

78.1
.8
.5

.9
.9

.6
.7

30. 9
2. 1
6.4
1.1
2.1
7.4

31.6
2. 1
8.4
1.1
1.1
7.4

.1

1.1

*

1.1
1.1
7.4

.3
.3

.9
. 9
.3
4.5

1981

Freq

14. 5

.7

1.1
.7

3.2
2. 1
.

1

1.1

.2
2.1

.9

1.1

10. 3

1.8
3.6
2.7

1.0

.9
.9
20.0

31.6

164

1.0

.2

2. 1
1.1
1.1
4.3
2. 1
.4
1. 1
27.6

.

1

1.9
.7
15.4

1. 1
2.1
2.1
3.2
1. 1
1. 1
28.4

TA3LE A 6
R E L A T I V E COVER AND R E L A T I VE FREQUENCY OF EACH S P E C I E S
OH S I T E 5 A 7 9 - 8 0 - S 1 SURVEYED AS A 4 - Y E A R - ,
5-YEAR , AND 6 - Y E A R - O L D CLEAR

1979
Cover

Species
b r a s s e s a n d sedRc-s
"6 a. in a J f; .1T in t i
C,v i c x

C <• J y t v . a i: t u

P. ac* licuiia
Dt C

f iu’ I

Oa ij z v (J i t i

.2

1.2

76.7

34.5

Cover

66.8

iptccf.i

i

Uin

d L’ p . l u p C

t

It 17!

.4

puiitji'iri

P eren n ial herbs
Ai f 0 t ii Jti'iT
G . i u i ' f / i e a i n p H ’ c u i f i c <;i
P f i* 1 c urn a q u i (J < i . ufij

Se i <Jii ge
Vtvi'n

Freq

.6
■*
2.2

14. 3

.aduei: a

I
F re q
1.8
26.9
1.8
3.7
2.8

2. 2
" .1

1.8
2.8
13.9
.9

1981
Cover

Freq

64.6

2.0
30.0

.3
.7

1.0
2.0

.1
9.0

3.0
4.0
5 6
2.0

T r e e s and s h r u b s
Al C f i ' i f d / a / i r / f i 1 i
Ame i iiiic/uc a J p .

fptytit
P o :« i

u in

’j

nu j

3.6
2.4

1.9
.2

3.7
3.7

1.4
.2

3.6
5.9
1.2
1.2
32 . 1

5. 4
. 1

2.9
8.3

22.3

25.0

"CpL'tll
banfc j ua»in

P a u c u i punni'a
Pa

7.0
.2

i c ac 1

1 »: n

Qucacus i p p . (r e d oak)
Vac c m i um i p p .

12.0

165

.3
.2
3.2
.8
24 . 3

6.0
2.0
2.0
5.0
2.0
28. 0

TABLE A 7
RELATIVE COVER AND RELATIVE FREQUENCY OF
EACH SPECIES ON SITE 6 SURVEYED AS A
5 -YEAR-OLD CLEARCUT
19 79
S p ecies

Cover

Freq

66.0
6.0
5 .3

26.2
9.7
9,7

G ra ss e s and se d g es
Ca s i z x p e . n 6 y Z v a n x . c a
V z 6 c k a m p 6 Z a faZcxu 0 6 a
0siyzop6 Z6 a6pcsiZ^oZZa

P e re n n ia l herbs
1.9

Ga. uZt he. si Za p s i o c u m b z n 6
Pte.siZd.Zum a q u Z Z Z n u m
SoZZdago 6pp.

.1
1.0

T re e s and sh ru b s
As i c t o 6 t a p h y Z o 6 u v a ~ u s i 6 Z
Co mpto nZ a pcsizgsiZna
VZ&sivZZZa ZonZc,&sia
PZnu6 b a n k i Z a n a
PopuZuA t/L&muZoZde.6
PsiunuA pumZZa
PsiunuA i z s i o t Z n a
Ru.bu-4 p e m Z Z o a n Z c u i
SaZZx gZaucophyZZoZde.6
Vac c Zn Zu m 6 p p .

1 .5
.3
6.6
1.5
12.6

166

1.0
1.0
1.0
4 .8
1.9
2 .9
5.8
8.7
1.0
23.3

APPENDIX B
RELATIVE COVER AND RELATIVE FREQUENCY DATA
FOR SPECIES SURVEYED ON SITES BURNED BY
PRESCRIPTION OR WILDFIRE

TABLE B 1
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 7 A 8 0 - 8 1 SURVEYED AS A 1-YEAR- AND 2-YEAR-OLD
PRESCRIBED BURN
1980
A n n u a l s a nd b i e n n i a l s
C im ium ip .
C o n t j d a l i i i empc>iu.t4e.H4
E n i g e n o n canadeii4-t.4
Gc-tan-ium b i c k n e l l i i
Knigia o in g in ic a
Lactuiia c a n a d e m i i
lielampynum l i n e a n e
G r a s s e s a nd s e d g e s
Agfioptjxort i-’tac./ti/cauilum
AndJiopogon g e n a n d i i
Andnopogo n i c o p a n i u i
Canex p e m y l v a n i c a
D anthonia i p i c a t a
O eicham piia ^ le x u o ia
Vic.hantke.lium depaupenatum
Koe.ten.ia. m a c n a n t h a
t i u. h t e nl i e n . g i a m e x i c a n a
Onyzop&ii pung&m
P a n i c am c a p i t l a n e
SoJtg/ia4i<tam nu£an4
P eren n ia l herbs
Anem'one n i p a n i a
Antennania n e g le e ta
Aiten. j u n c i ^ o n m i i
A iten la e v ii
Campanula n o t u n d i & o l i a
P n a g a n i a vi^igiH-cana
H eXianthemum c a n a d e m e
U i . e n. a d u m wetioium
L ia tn ii novae-angliae
Mtt-tan-tfidmum c anai f £H4 £
P t e n i d i u m a q u i l i num
Snnec.t.0 t o m e n t o i u i
S o t i d a g o 4pp.
V i o l a adunca
V iola pedati& ida
T r e e s and s h r u b s
Ame I ancV-i&Ji 4 p .
A n d o ita p h y lo i uo.a-um i
C eanothui o v a tu i
Comptania pcneg n in a
C nataegui ip .
PnunuA p u m i t a
PnunuA ui n g i n i a n a
P o i a bt a n d a
Rubui pen4i£uanicu4
S{/mpbp-tica/ipo4 a £ b u i
Vaccinium ip p .

Cover
.1

1.2

1981
F r eq

Cover

Freq

.5
4.1

2.1
13.0

6.7

1.
1.1

.1

2.9

.2
44. 7
.3

17.0
3.5

.1

2.1
14.4
3.6

.6

3.6
.5

.2

.1

1. 2
1.2

1. 0
3 .6
1.5
.5

.2

.1

2.9
2 .3

2.0

5.9

.3

.2

.6

3.6

27.8

.5
.1

WO'O'O'

Species

.5
.5
2.5
.5

1.0
.1

1.5
1.5

1.8

1.0
2.1

,6
.6
.1

.1
.1

2.9

1. 2
1.2

1.2
1.2

1.2
.5

.6
.4
.1

1.7

4. 6

5.6

2.1

8.2

1.4

2. 1

1.0

1.2

.1
.5

.5
5.1

.5

1. 2
4. 1

5.0

10.3

17.4
2.7

12.9

27.7

17.5

1. 0

.2
40.9

167

.6

.5
.5

.6

1. 0
12. 3

.6
1. 2

TABLE B 2
SPECIES COVER AND FREQUENCY DATA FROM SITE 7 B ( 1 - , 2 - , AND 3-YEAR-OLD BURN),
SITE 1 2 A ( 3 5 -YEAR-OLD JACK PINE STAND), AND SITE 13C(55-YEAR-OLD JACK PINE STAND)
USED IN THE BURIED SEED EXPERIMENT
site
Species
A nn u a l s and b i e n n i a l s
CL* i i u m i p .
C o n y d a l i i itmpin.vine.ni
E'lige-’ion c a n a d e i u i i
Geranium t i c f e n e l l i i
Knigi a, v i n g i n i c a
Mcfampy’ium l i n z a n t
S ile n e aniirrfiina
G r a s s e s and s e d g e s
Agropyron i r a c / i y c a u l u m
Andropogou g t n .a n .d ii
A n d n o p o go n i c o p a n i u i
Bromui k a l m i i
Cantx i p p .
Vanthonia i p i c a t a
V acham piia ^texuoia
V i c h a n t h t l i u m depaupenatum
K o t l t n i a macnantha
O nyzopiii aiptni& olia
O nyzopiii pungtm
Panicum c o l u m b i a n a m
U uhltnbtngia mtxicana
S c h i z a c k n t p u^ p u ' i a i c e n i
S o t g b a J i l u m im-tani
P e r e n n ia l herbs
Anemone (jtmtque(Solid
Apoctjnum and no i a t m i { o H u m
Ai t t n j u n c i & o n m i i
A ittn iaeoij
Campanula n o t u n d i & o l i a
Cypniptdium acaule
E qu i i c t u m liyemale

1-year
Cover
Free

2.8
22.lt
.5

7B

site

2-year

3-year
Cover
Freq

1.4
1.4

.6

8.8

.6

12A

site

35-year
Cover
Freq

13C

55-year
Cover
Freq

.9

.8

9.8

3.1
2.7
.4
12.4
5.3
2.6
.9

13.7

.5
15.5

.4
.7
1.1
.5
1.2
17.2

1.4
14.5

.9
1.6

.7
4.9
1.8

3.7
.6
.9
.5

3.5
3.2
1.1
1.8

.6

.8

1.2
.5

2.1
1.0

39.8
.1

17.0
2.8

41.7
1.5

16.1
3.2

6.1
5.1
3.4

3.6
.5
4.7
5.1

7.9
2.8
1.7
6.8
1.1

5.4
.3
1.6
2.5
.8

6.8

3.4

4.1

10.7
1.1
1.1
6.4
3.8
.5
3.2

.1
2.7

.5
4.4
.4

1. 0

.1

.3
2.1
3.2

1. 1

.9
.4
2.3
.5

.3

■k

1.6
.5

2.6

.7

1.5
2.6

mooi

CTi

cm

m

o\

CM

CM

CM

i—
t m m<r

<T i

OJ
M
**H
<n

O CD
m

MD

»r m
m

sO

sO so

sO

so oo so in

in

CM

OS

15.5

CM

CM

o

- r

23.9

m

15.8

Dl
<u
M M
(0 P k
01
>7
1
CM M
0>
>

A7.3

*sO

21.0

PQ

3.1

m -r

.2

c
•H
4M

.3

'o i

.9

mm

O' m O' -T

<3*

11.9

m

^

o s

A8 .A

C M H N

cm

m i—(i—i

OJ I
>
0
u

*T3
0)

to

c n cm

sO
<T

1A.5

so co t- i <r <r

<r m

O
in i—I

2.7

OS rH

M M
m C^l
0>
S>>
CO M

CM

m

cm

s|

i—iin

»-+
--T

.2

<n

r^
est

M M
nj £n|
01
>s
I
m m|
m oj
>
o|
u |

SI

N tO
00 rH

3.8

M ti4
CO
CJ

01 f
U
m m
♦H m iu
w
>
o
o

c
O
a
w

\0
rH

3. A

u
cn
rH

m

2.2

in o
so

2.3

O <T
H CN

o

a
C

u

i—I

sO<r

<r

H

* T f-H r-H

M

<r -3- p- -jI—I so

i

-j

.7

<r

sO

11.7

SI
u m
ta u*|

3. 9

H

1.1

pa

>\

0 > CM

^

sO

-o* m

O

P s h -

18.6

a|

9.0

Ml

169

<7
»3

ipp.

3

g£aucop/i;/ffo>idcJ

3 *tJ
"H
*
= OJ
: 3 u
-1 CLV Ck>
:
'M
s-*
Q -0 3 ^ >•7 3 3 ‘M
M
cj d, S *M<WV
V
3
X!'
3s*e 3 Q. -O *y ‘M 3 31 Q.
•ST <J
C L * tf £
J- ^ a
•O Q.’M -7 3 o O 3 3 CJ M-»
3-3 3 W 3
3 Q. -i
c
V
U-2 3 31 • a n -7 -7 o3
3
■0 3 V 3 a 3
in 3 3 3 V 3 w o 3 3 3 U
V *M 3 a v 31 3 5 3 “
o
<v U U 3 E 3 M 5: 3 3 3 CJ
M p* E 5J 3 -e CL’V ^ ^ 3
H -C < CJ CJ CM4 J G - G - G . ex cv

10
•§

Vacciimim

tO
0

S<i£ix

■n
*
3
I
3
3

/Uipidui
pens tfi'an^cui

in
OJ
•H
O
a*
O
CO

-7
-7
3
-i»
V)
cJ O
O
3
E
3
3 3 3
3 ■o
V
-o
3 €
V 2
£ t j CJ V U
■y
CU V 3 --I
3 : *o 3 3 W V «■>
-s; 3 E
3
V
W V E 0113 3 TS = 3 -7
>W
v W 3 *3 V 3 3 SC 3 E Q «S
o
*v. _ U W 3 O ■0 3 3 3 S *3 <v O
I 3 a i v w ■V V
3
3 2
SXW
O
O
4 je
< k CJ1
< J «* O
eJ ci
h 3 2
3
^ Q. Q 3
V ‘■MV CJ* CJ CL O) 3
CJ
-o ■V
3 onk 3 > E <1 o
3 E a
3 0 3 -7
3 3 EX 3
o -7 ■n
CjJ 3
V 3 E E E s; E
E V E
V
CJ 3 3 tw 3
o -=: "a
3 3 3
3 <1
. ' M O V V *v W *o ur i^j 2 ' Mw o 31 V 3
<t* -2 3 u o o ■v V 3 O V “3 V 3 2
o a w a 3 3 3 fj 2 31 Ol 2 W U 13 3 3
v cnw -V *- < r V 3 3k 3 \ V r* CJ t* V
2 V “V O
3 a 3 v ej rj
3
o CL
o ^ a 4) ‘M*M■v w 3 3 0 0 V
cj u. CDtc
5;
C X d .C i.P ^ c /> o ^ to ^

Rabuj
Rubui

o
o |

Roiti tfaiida

i

.2

oj

TABLE B 3
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES, BY SPECIES GROUP,
SURVEYED IN 1 9 7 9 ( SI TE l A , 0-YEAR CLEARCUT),
AND I 9 6 0 AND 1 9 8 1 ( SI TE 7 C 8 0 - 8 1 ,
1-AND 2-YEAR PRESCRIBED BURN)
Year o f S u rv ey
Species
A n n u a l s a nd b i e n n i a l s
iieTtampyaum t l n z a a z
Ge.tian4.um c a a o l i n + a n u m
Coaydat.ii izmpza.via.zni
C4.t1 i i .u m i p .
L a d uca c a nade.nii.i
G r a s s e s and s e d g e s
Cn>tex p z m t j l v a n i c a
V anihonia i p i c a i a
O a t j z o p i i i puagena
K o z l z a i a mactian-tha
Pa.n4.cum coluDib-catium
S-iomua k a l m i i
F e d u c a ip,
Poa p a a t z m i i
Schizachnz puapuaaiczni
O ayzopiii a ip za ifio lia
D zich a m p iia falzxuoia

1979
)over(2)
Freq(I)
_

_

.7
-

-

-

21.7
5.0
4 .6
.8
.4

14.5
.7
2.2
2.2
5. 1
1.5

28.7

2.9
3,6
2.9
7.2
5.8
-

*6
1.2

_

_

29.6
13.8
4.2
1.2
3.5
_

.2
-

13.5
4.8

-

1.8
2.2
.2
.8
2.1
•

-

.3
~
.4
-

1.5

.1

-

.1
.2

1.4
5.8
13.8
8.0
8.0
5.1
2.9
1.4
*7
■/
1.4
.7

_

3.8
2.2
.5
.5
14.1
2.2
5.4
3.8
1.6
2.7
.5
1.1
.5
-

-

.7
-

-

-

36.8
-

8.1
.2
.3
-

.2
. 1
.2
.7
-

2. 7
2.Q
.7
1.3
2.0
-

1.1
1.6
.5

_

..
1.3

.5
•J
.5

.

1

_
-

33.1
3.5
4.2
2.1
-

19.9
2.0
11.9
1.3
.7
.7
2.0

7.1
4.4
.5
3.3
.5
1.6
.5
3.3

15.2
4.9
7.1
1.1
4.4
-

.7
.7
1.3

40.2
2.2
6.9
1.8
1.7
-

19.9
6.0
11. 9
6.0
3.3
-

_

_

.5

-

-

-

.5
1.6

-

.7
.7
-*

-

-

-

-

-

.

170

1981
Cover(Z)
Freq

_

-

P eren n ial herbs
.4
Anemone q u i n q u z & o l i a
1.2
A iiza Z azvii
.2
S o l i d a g o ipV i o l a adunca
7.1
P tza id iu m aquiZinum
L ia ta ii novaz-angliaz
1.3
M a i a n t h emun c a n a d z m z
H zlianihui o c z id z n ia lii
A
A p o c y n u m a n d a o i a z m i & o t i u 1 *H
C onvotvului ip ith a m a zu i
HiC'tac-ium a u a a n t i a c u m
H i z a a c i u m u e n oi u m
“
Hizaacium c a n a d z m z
Campanula a o l u n d i & o l i a
P o l y g a l a po£ygama
1.3
Faagaaia v i n g i n i a n a
3.1
G a u lth za ia paocumbzm
T r e e s and S h r u b s
t/acci n. ium i p p .
A m ztanchiza ip .
Paunui pum ila
A a d o ita p h ylo i uva-uaii
ComptonZa p z a z g a i n a
Paunui v i a g i n i a n a
Pauhua i z a o t i n a
R e i a faiandn
Rubua p z m i t v a n i c u i
Pinui b a n k iia n a
Symphoaicaapoi a tb u i

1980
C o v e r (%)
Freq{/[)

1

TABLE B 4
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 7D79 - 8 0 - 8 1 SURVEYED AS A
1 - Y E A R - , 2 - Y E A R - , AND 3-YEAR-OLD
PRESCRIBED BURN

1980

L979
Species
A n n u a l s and b i e n n i a l s
Genanium c a n u t i n i a n u m
L a c t u c a canadenAxA
Melampynum l i n e a n e

Cover

.6
.3

G r a s s e s and s e d g e s
A g n o p y n o n t 4.ac.nyca.uium
AgAo-ttxA h y a m a l i i
. 3
Andnopogon g e n a n d i i
4.6
Canex. p e m y t v a n i c a
66.5
V anthonia ip ic a ta
.8
V e i c h a m p i i a fJtcxuoAa
.3
Di c h a n t h e l i u m d z p a u p e n a t u m
.2
Koe.te.nia m a c n a n t h a
O n y z o p i i i pungenA
Panxcum cotumbxanum
.3
Poa p n a t z m i i
.5
P e r e n n ia l herbs
Apocynum a n d n o i a z m i ^ o t i u m
A iten Zazvii
Campanula n o t u n d i ^ o t i a
Fnagania v i n g i n i a n a
Hiznacium aunantiacum
Hi e n a c i u m c u Ha d en At
H i & n a c i a m veno-Sum
He.lianthe.mu.rn c a n a d e n s e
He.£xantliuA o c c x d e n t a t x A
Lechea minon
L i a t n i i ito v a e - a n g l i a z
P h y ia tii vinginiana
P t e n i d i u m aqu i l i n u m
Rumex a c e t o i & l l a
Solidago ip .
V i o l a adunca
T r e e s a nd s h r u b s
An.c.to Ataplt i/ to a uva-uAAx
Comptonxa p e n e g n i n a
G a y tu a a c ia baccata
P i n u i b an f e i x a n a
Pnunui p u m ila
PnunuA i e n o t i n a
Pauijua v x A g t n t a H a
Q. uencui i p p . ( r e d o a k )
RoAa bt at tda
RubuA pe nAxt va ti xc uA
Vaccinium ip p .

1.0
1.0
.6

F r eq

1981

Cover

F r eq

2.7
50.0
3.4
.4
11.2
1.5
.3
2.1

1.4
4.2
20.4
5.6
.7
12.7
1.4
1.4
8.4

.7

3.5

Cover

Fr eq

.3
50.7
4.0

3.3
23.8
3.3

8.6

9.8
2. 5
3. 3
6.6
.4

.6
.6
1.7
.6
1.1
7.9
19.8
7.9
.6

6.2
.6
1.1
3.4
1.7
.6
.6
.6
4.5

.1
2.5

.1

4. 9

.7
3. 3
2.5
.3

1.1

1.6

4.2
.6
1.6
3.6
.6

8.5
.6
1.7

11.3
.3
.1

2.1
7.8
2.1

.2

9.8
1.6
1.6

14.1

.4
13.2

14.7

.7
. 7

.3
.5

12.3

.7

17.0

.6
13.0
.6
1. 7
.6
.6
3.4
.6
1.3
5.1

171

14.7

.8

.1
.6

1. 4
.7 .
2.8
2.1

2.9

1.6

2.8

2.5
2.5

TABLE B 5

RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 8 A 7 9 - 8 0 - 8 1 SURVEYED AS A
2-YEAR-, 3-YEAR-, AND 4 -YEAR-OLD WILDFIRE
1979

Aipic t i i n c l i t c i
ip.
A c e To i t r i p l y f ip l l U M ' i n i i

Ccauulftui o o a t u i
Compt oni a p e a c g A l n a
Caalaegui ip.
P l n u i jJanfcil ana
PtunuA p u m l l a
P a un u i o l a g l u l a n a
Queacui ip p . ( r e d oak)
Roia b la n da
Rubui p e r i i l l i 1a t i l c u i
S a il* glaucopfigCtotdei
Sgmpl t oAl c a apui a l b u i
1/acclnlum i p p .

.4
.4
.9

Fr eq

Co v e r

Freq

*

1.7
.4

.2

.5
.4
.9

1.3

1.3
.4

1.5

27.4

11.8
1.3
.4
2.6
.9
3.5
2.2
7.0

34. 4

2.5

1.7

.4

.1

.4
3. 5
4,8

.3
.6

1 ., 3]

,4
.4

.9

.2

. 7

29.3
.3
1.1
1.5
1.7

12.3
1.7
.4
3.1
.4
4.9

1.6

.9
1. 1
.2

.4

7.0
.4
.4
.4

6.5
.1

.4
.6
•
1fc
*

.8
4.8
.4
1.3
it
.4

.3

.4

.2

.4
.4
.9
.4

1.4
.2
.1

.2
.3
13.6
.2
9.8
.4

3.5
.9

.4
.9

1.3

172

I.
5.
.9

6.0
.1

.4
.9
2.6
2.6
.4

.1

l".

.1

,

5.4
.3

2.
1.
1.

.7
1.0
.8
10.4

2.
2.

1.4
4.8

4.
9.

2.0

,

2.6
.4
1.6
.2
1.1

5.7
1.8
3.9

2.4
4.4
2.6
.8

5.7
9.6
1.3
.9

.3
40.6

.4
11.3

.

4.

1.

.9
.9
11.5

.9

l\

.2
28.2

5.0

.1

.4
2.2

5.7
.4
.9
4.8
.9
3.5
10.6

.5
.7
.4

.5
3.1
.9
1.8
13.5
1.3
1.8
2.2
.9
2.2
1.4
6.7
.9

r* ^ C iJ s o i* u iL n

.4
4.0
3.1

u e -v j

4.3
4.0

27.2

13!

u i f ’ NjhJ

T r e e s end sh r u b s

.1

1981

Co v e r

o yi

P e re n n ia l herbs
Anemone a t p a ' i c a
Aitlennaala n e g l e c t s
Apocynum a n d a o i d e m l j o l l u m
Aitca laevla
BtaAilca ip.
C a mp a n u l a \ o t u n d i & o t i a
Cuituofuufui ip-tlhamaeui
E q u l i c t u m Ayemale
F'taga'ila v t s g l i t l a n a
C a u t t f i c a l a paocumbetiA
H ellanlfmi o c c i d t n t a t i i
Mlc-taclum au.fi an-t in cun
H le a a e l u m ucn o i u m
Houitonla l o n g l / o l l a
L ia tn ii npvac-augtlae
Matautfiemum c a n a d c n i e
P/iyialli visglnlana
P o t y g a l a puli/gama
P f e A l d l um a q u l l l r t u m
S c n e c l e t omei i t a i uA
Solidago ipp.
y i i i d c n i l i S l c d mo n a c a l
V m t a adunca
Viola p c d a tc l id a

Freq

f-

G r a s s e s a n d s ed R c s
A g t u p g a o n c a a c Ay c a u l u m
Andaopogon g e a a a d t l
Andaupogon i c o p a a l u i
Baumui h a l m l l
Cuat u p e n i y l v a n l c a
Pdntfioma i p t e a l a
Ocicbampila jl e n u u i a
0 ccltanthelium depaupeaalum
Fcituca ip.
K u e t e a l a ma c a an l / t a
MuhltnbeAgla mexlcana
O a g c o p ili pungeni
Paitlcum c i p f u m t u a i i u m
Pad p A a t c i i i l i
Suaghai - t aum nuldpt i

1980

Co v e r

O si) Ln Oo

Species
A n n u a l s and b i e n n i a l s
C o i y d a l i i icmpcav-r Aeni
Ealgenon c a n a d e n i li
ll tJioi peAmum a a u e n i e
PclampyAum l i n e a - t e

TABLE B 6

RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SITE 9 A 7 9 - 8 0 - 8 1 SURVEYED AS A 3 - YEAR- ,
4 - YE A R- , AND 5-YEAR-OLD PRESCRIBED BURN
1979
Species

Cover

A n n u a l s a nd b i e n n i a l s
Ci ai Lum i p .
E n i g e n o n cartadeni-ii
K nigia v in g in ic a
Lac- t uca caii adetti' t. i
Sie.ta.mpynum t i n e a n e
G r a s s e s and s e d g e s
Agaopi / aon t n a c h y c a u t u m
A ndnopogon ge/iaA.dtt
Andnopogon ic o p a n i u i
Bn omu i k a l m i i
Canex p e m y t v a n i c a
V anthonia i p i c a t a
V e &c h am p& ia rft e x u o i a
D i c h a n t h e t i a m depaupe.fia.tam
F a tu c a ip.
K o e te n ia macnantha
O nyzopiii pungem
P a n i c u m co i u m b i a n u m
S c h iza c h n e puapuaaiceni
P e r e n n ia l herbs
Anemcne q u i n q u e & o t i a
Aht e n n a n i a n e g l e c t a
Kpocynum a n d n o i a e m i ^ o t i a m
Ait e n ju n c i& o n m ii
Ait e n l a e v i i
Campantiia n o t a n d i l o l i a
Convotvului ipitkatnaeui
Pnagania v i n g i n i a n a
helianthai o c c id e n ta tii
Hienacium aunantiacum
H ienacium vcnoium
Lechea minon
Li a t n i i ho v a e - a n g ^ La e .
M aianthemum c a n a d c n i c
P o te n tilla tn id en ta ta
P t e n i d - cam aq u-ii-aium
Scnecio tom ento i u i
Solidago ip p .
V i o l a adunca
T r e e s a nd s h r u b s
AwicCancA-ta^i i p .
A n c t o i t a p h y t e i uva-uai.L
Ccanothui o v a tu i
Comptonia p e n e g n in a
P i n u i ba nki - c ana
Pa unui p u mi £ a
Pa unui i e n o t i n a
Pa unui v i n g i n i a n a
Q u c a c u i i p p . ( r e d o a k)
Rei d b l a n d a
Rubui p e p i i i £ van i c u i
S a lix gtaucoplujiEoidei
Stjmphonicanpu J a£bui
Vaccinium ipp.

1980
Fr e q

Cover

198 1
Fr e q

Cover

.5

.1
.2

Fr e q

.4
.3
.5
★
2.7
.6

1.8
.2

.9
5.1
3.3
.5
13.5
.9
5.1
1. 9

.4
k
.1
.2

2.3
4.6
.9
1,9

57.6

2.1
.7
54.1
1.8
1.4
.9
1.7

4.1
1.8
2.3
13.9
1.8
5.1
1.8
2.3
6.0
.5
.9

3.2

3.6

1.0
39.3
1.4
.2
.3
.2
1.6
2.5

2.0
15.3
3.1
4.6
1.0
6. 6
.5

1.5
.5
*
.6

5.6

.1
.2
.3

.9
1.4
.5
1.4

*
.7
*

.1
.2

.9
6.5
.5
.5
.5
1.8
2.3

1.5
1.4

1.5
1.5
.5
.1

1.5
1.0
1.0

1.5

.5

.2

1.0
1.5

.5
.1

*
.9
.2
2.5
2.6
.3
4.2

.5
.9
.9
1.9

.9
.5
.5
.9
.5
1.8

.2
.6
1.9
1.3
1. 7
11.7

4.1
1.5
1.5
5.1

3.1

.5

3.3
1.9
2.8
6.5

.6
.4
.5
.8

5.1
2.3
1.8
6.0

10.7
.5
1.9

9.1

9.3
.5
.5

.5

1.6

.5

3.5
3.6

3.3
.5

1.4

2.3

.4
.2
7.2
1.8

16.7

12. 6

21.7

12.0

18.2

173

8.2

6.6

.5
4.6
1.0
1.5
14.3

TABLE B 7
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH S P E C IE S
ON S I T E 9 B 7 9 - 8 0 - 8 1 SURVEYED AS A 3 - Y E A R - ,
4 - Y E A R - , AND 5 -Y E A R -O L D W ILDFIRE

1979
Species
G r a s s e s a nd s e d g e s
A n d A a p o g a n ge.ftaA.d-ii
perny tva n ica
Vanthonia i p i c a t a
V ic h a n th e liu m depaupeaatum
K o e te A ia macAantha
OAyzop&ii pungcni
Pan.ic.um co i u m b i a n u m
P e r e n n i a l herbs
ne.gle.cta
Apocynum a n d A o i a e m i & o l i u m
A iteA l a e v i i
Campanula A o t u n d i f o l i a
H-te-’i a c i u m v e n o i a m
Maianthemum c a n a d e m e
P o te n tilla tA identata
Solidago ipp.
U nidentified d icot
1! i o t a a d u n c a
\)iola p e d a t i f i d a
T r e e s and s h r u b s
Am elanckieA i p .
A a c to ita p h tjlo i uua-uA ii
C v m p t o n i a p e A CQ A in a
P-inui banfei-taria
Pa unui p u m i l a
Q u e a c u i i p p . ( r e d oak)
Vaccinium ip p .

I960

1981

Co v e r

Freq

Cover

Fr e q

Cover

Fr e q

3.3
51. 3
.4
4.4
.1
.7

2.6
25.2
2.6
11.3
.8
3.5
3.5

.3
67.3
.3
2.1

2.8
21.1
.7
12.0
2.1
8.4
2.8

.7
60. 6

3.3
24,8
.8
9.9

1.8

.8
.7

2.5
1.7

.9

.1

.9

.2
it

1.4
2.1
.8

.2
.3

.9

.7
3.5

1.6

1.1
.9

1.3

2.6
6.1

3.5
.2
33.5

2.6
10.4
.9
24.4

174

.7
1.4
'

1.3
.6
.7
.2
2.7

2.8
4.9
1.4
1.4
10.6

1.4
.4
.1
.3
7.5

4.1
4.1
1.7
2.5
18.2

22.2

19.0

27.6

24.0

TABLE B 8

RELATIVE COVER AND RELATIVE' FREQUENCY OF EACH SPECIES
ON SITE 1 0 A 7 9 - 8 0 - 8 L SURVEYED AS A 4 - YEAR- ,
5 - YE A R- , AND 6-YEAR-OLD PRESCRIBED BURN
1979
Species

Cover

A n n u a l s and b i e n n i a l s
A a a b Z i g t 1a 6 / i n
C Z t i i Z am Ap.
Lac.-Cana ca.nade.m-Li
G r a s s e s and s e d g e s
Ag / to p t/ n on t a a c n y c a u Z u m
A g a a itZ i kyemaZZi
SA.omuA k a Z m L i
Ca-tex p & m t j Z v a n Z c a
VanthonZa i p Z c a ta
V acham piZa fttexuoia
VZchanthe.lZu.rn d e p a u p e a a t u m
Feituca ip .
MuhZenbeagZa mexZcana
QayzopiZi aipeaZioZZa
OnyzopiZi p u n g e n A
Pan-icum c<j£ambifl(ium
PanZcum x a n t h o p h y i u m
Pea p n a t e m Z i
Sc/t-t z a c / m e p u a p u n a i c e m
Se-tghaA-CAum

Cover

T r e e s a nd s h r u b s
A c e * •'lub'ium
Ame Za nc hZe n. i p .
C aataegui ip .
VZeavZZZa ZonZceaa
PZnui b a u k A - i a n o .
Popa^aA t a e m u Z o Z d e i
P a u n u i pumZZa
Paunui ie a o ttn a .
App.

RubuA p e m Z Z v a n Z c u i
VaccZnZum i p p .

1981
Freq

Cover

£res

.4
1.6

.1

.2
.6

1.3
.5

.1
.3
17. A
9.6
.2

.8
9.6
4.0
.8
2.8

.9
21.8
3.0

10.9
5.3

30 . 9

13 . 7
.5
1.6

1.3
.4

.2
32.5

.A
10. B
.8
.8
.4

25.5
.5

A. 4

6.0

1.8

6.4

.2
.2
1.0

.1

1.6
2.8

ie

6.4
.4

.7

.4
5.3
.4
.9
1.8

.5
.2
10.8

iiu-tanA

P e r e n n i a l herbs
Anemone q u Z n q u e ri o Z Z a
Apo cyitum a n d a o i a e m Z i o Z Z u m
A itea ZaevZi
Ai^e^i i a g Z t t Z & o t Z u i Camp a nu Za a o t u n d Z & o Z Z a
Couvol uul uA i p Z t h a m a e u i
E p Z g c a aepenA
F a a g a a Z a s f Z a gZ nZ a n a
GauZtheaZa p/tocumbem
HcZZanthui occZ d en ta Z Z i
HZea.acZu.rn oenOAum
LZataZi n o va e-a n g tZ a e
Ma-iaa-t/temum c a n a d e n i e
Poi ygoi t am c Z Z Z n o d e
P o t e n t Z Z Z a A-cmp-tex
P t e a Z d Z u m a quZ ZZ n u m
Rumex a c e £ o A e l £ a
S e n e c Z o { omi ' i UoAui
S o Z Z d a g o App.
VZoZa a d u n c a

Q u e . 'i c u A

1980
Fr e q

1.0
.1

.4

10.5
2.2
.9
1.3

25.6
.2

14.7
1.1

.1

1.1

4.0
.9

9.3

6.6

.4
3.5
3.5
1.8
2. 6

.4

2.2

.6
.4
.1
.5

4.9
2.2
1.6
1.1

1.3
.1

2.7
1.1
1.1

.1

.2

4.0

*
.2

11.5

.4
10.8

15.9

.2
.3

.4
.4
5.2

.2

10.1
.4
.4
.4
.9

.2
1.7

1.3

.5

.4
.8
.8
.4
.4

.4
.5

4.9
.6

4.5

9.8

.1
.1

1.1
1.1

1. 1

.1
1. 9

1-9

2.0

.3

9.6
.6

9.6
8.8

2. 9
.8
11.0
7.5

.4
1.3
1.3
12.7
10.1

10.3
.7
10.8

1.6
.6
2.5
2.2
.6
9.3
9.8

TABLE B 9
RELATIVE COVER AND RELATIVE FREQUENCY OF EACH SPECIES
ON SI TE 1 0 B 7 9 - 8 0 - 8 1 SURVEYED AS A 4 - YEAR- ,
5 -YEAR- , , AND 6-■YEAR-OLD PRESCRIBED BURN

A n n u a l s and b i e n n i a l s
E n Z g e n o n c a na dc ud- c i
LacTuca. canadenA-i.4
Li-t/icMpedmum a n v z m e
G r a s s e s and s e d g e s
Ag>icpyA0 rt t Aacf t i / cauIum
Ag^ioi Z Z i h y e m a Z Z i
Bn o mu i kaZmZZ
Canex p e m y Z v a n Z c a
VanthonZa ipZcaZa
O ei c h a m p i Z a f r Z e x u d i a
Oi ch a . ni he .ZZu m d e p a u p e n a Z u m
KoeZenZa macnanZha
Mti/iicHbeAg.da mex.Zca.na
Q n y zo p iZ i pungeni
Q n y z o p iZ i aipenZ&oZZa
Pan^cum co£umb.i<muffl
Paa p n a Z e m Z i
SchZzachne punpunaiceni
PanZcum x a n Z h o p h y i u m
P e r e n n ia l herbs
Apocytutm a nd n oi ae mZ &o ZZ t im
A i Z e n Za&vZi
ConvoZvu.lu.& i p Z Z h a m a e a i
Fn.agan.Za v Z n g Z n Z a n a
GaaZZkenZa p n o c u m b e n i
HZenacium aunanZZacum
MaZanZhemum c a n a d e m e
P o Z e nZ ZZ Za a n g u Z a
PoZenZZZta ZnZdenZaZa
PZenZdZum a q u Z Z Z n u m
Se ne c - t d -tc>man.toiai
So Z id a g o ipp.
VZoZa a d u n c a
T r e e s a nd s h r u b s
AmeZanchZen i p .
AncZoiZaphyZm a v a -u m Z
Ceano-tfiu-i o v a t u i
CompZo n Za pen.egn.Zna
CnaZaegui i p .
VZenvZZZa Za n Z cen a
PZnui b a n k i Z a n a
Pop uZu a Z n e m u Z o Z d e i
P fi u nu i pumZZa
Pnunui ic n o Z Z n a
Que Ac ui i p p .
R j i a bZanda
Rabiii p e m Z Z v a n Z c u i
SaZZx gZaucophyZZoZdet,
SymphonZcanpu& a£6ui
V ac c Z n Z u m i p p .

19 81

1980

1979
Species

Co v e r

Freq

Cover

Freq

Cover

Fr e q

.2

3.8

.3

.8

2.3

.5

3.6
1.2
.6

.3
44.9

.6
18.1

38.8

1. 2
17.4

2.5
,3

4.2
.6

.9
.3

.6
2.9
1.2

4.8
4.1
.4

8.4
1.8
1.2
1.2

3.0
4.8

8.1
4.1
1.2

.6

2.5

1.7
.6

.1
.9
.3
.1

.7
1.3
43.5
.1

19.4
1.3

1.3

5.8
.7
.6
3.2

.6

.9

1.3
1.3
3.2
.6

7.7

1.3
.6
.7

1.6

.6

.6

.1

1.2

.7
.6
.6

*
.1

.2

22.4

14. a

8.3

1.8
.6
.6
13.2
2.4

2.3
1.7
1.2
2.3
.6
.6

9.8
.9

.6
7.0
1. 7

.4

1.3
1.9

.1

3.0

.3

3. 5

.6
1.3
3.2
.7

.2
.4
.2
.4
14.9
.7
.1

3. 9
2.6
.6
1.3
.7
3.9
.6
1. 3
11.6

.4
.2
.6
5.2

176

1.2
.6

.6
.7
1.5
.2
.6
10.1
2.3
2.8
1.8
.8
.2
.5
12.1

2.4
1.2
1.2
.6
3.6
3.6
.6
2.4

2.5
14.2
1. 2
2.2
2.0

4.8
2.4
.6
12.1

.4
12 . 1

4.7
1.7
1.7
.6
1. 7
4.6
1.7
1.7
1.7
1.7
14. 5

TABLE B 10
RELATIVE COVER AND RELATIVE FREQUENCY FOR
EACH SPECIES ON SITE 11 SURVEYED AS A
6-YEAR-OLD WILDFIRE
1 97 9
S pecies
G r a s s e s and s e d g e s
Ca/iex pen-6 y l v a n l c a
V a n t h c n l a 6pj.ca.tcL
V c 6 c h a m p 6 l a &lcxu.0 6 a
V tc k a n tk c llu m dcpaupc^atum
0 / i y z o p 6 t 6 pungcn.6
Panlcum co ta m b la n u m

Cover

Freq

86.0
3.5

28.2
10.3
2.6
14.1
2.6
1.3

2.0
.3

P e re n n ia l herbs
.5

PZc/Llti[lu.m a q u l l l n u m

10. 3

T re e s and sh ru b s
1 .3
2. 6

A n.cto.6tapkyZo6 a va -u A .6 l
C om ptonla pe/ieg/ilna
Ptnu6 b a n k 6 la n a
Q u c x. c u 6 a l b a
Q ue-tc iu 6 p p . ( r e d o a k )
V acclnlum 6pp.

2.8
5.0

177

1 .3
2.6
23.1

APPENDIX C
RELATIVE COVER AND RELATIVE FREQUENCY DATA
FOR UNDERSTORY SPECIES SURVEYED ON
INTERMEDIATE-AGE AND MATURE JACK PINE STANDS

TABLE C 1
RELATIVE COVER AND RELATIVE FREQUENCY FOR
EACH SPECIES ON SITE 12 B SURVEYED
AS AN INTERMEDIATE-AGE (35-YEAR-OLD)
JACK PINE STAND
1 98 1
S p ecies

Cover

Freq

A nnuals and b i e n n i a l s
McZam py ttu m Z Z n c a t t e .

*

2.6

G ra sses and sed g es
A n d tto p o g o n gcttattdZZ
B4.omcu k a Z m Z Z
Cax.e.x p c n A y Z v a n Z c a
V a n th o n Z a ip.i c a t a
VcAckam pAZa fiZcxuoAa
V Z c k a n t h e Z Z u m de.pau.pzfiatu.rn
MuhZcnbe.ttgZa me.xtc.ana
O tt y z o p A Z A pu.nge.nA
S c h Z z a c h n c pu.tipu.fta6 c c n A

P e re n n ia l herbs
Apocyuum a n d t t o A a c m Z ^ o Z Z u m
A6 t e . f i Z a c v Z A
Ft t a g a t i Z a v Z t t g Z n Z a n a
Ga u.Zth e.tiZ a ptLacambe,n6
HcZZanthcmum canad& nAc
He.ZZanthu.6 o c c Z d c n t a Z Z A
HZe.ttacZu.rn v c n o A u m
LZatftZA n o v a e - a n g Z Z a c
P ot& ntZZZ a t a Z d c n t a t a
SoZZdago ip .
VZoZa a d u n c a

.2
1.7
17.5
.5
4.7
.3
.4
.4

3.1
3.7
15.1
1.6
6.8
1.0
5.2
.8
.5

2.3
*

.1
.3
.5
.1

4.2
1.0
.5
1.0
2.6
6.3
1.0
2.6
.5
1.6

T r e e s and sh ru b s
.5
1.9
5.3
.8
9.3
1.5
.2
5 0.7

’K m c Z a n c K Z c t t ~ i ^ ~ .
A ficto A ta p h yZ o A uva-uttA Z
C om ptonZa petiegttZna
PZnuA b a n k A Z a n a
PttunuA p u m Z Z a
PftunuA A C f t o t Z n a
[lue.ticu.6 i p p . ( r e d o a k )
VaccZnZum i p p .

178

3.1
3.1
7.8
1.6
9.9
13.5

TABLE C 2
RELATIVE COVER AND RELATIVE FREQUENCY FOR
EACH SPECIES ON SITE 13B SURVEYED
AS A MATURE JACK PINE STAND
1981
Cover

Freq

Me.ZampyA.um Z Z n z a / t z

1. 2

14. 9

G ra ss e s and sed ges
An d n . o p o g o n g z n . a ^ d Z Z
An d n o p o g o n AC OpanZuA
Catizx p z n A y Z v a n Z c a

2.2
.7
12.9

5.

S p ecies
A nnuals and b i e n n i a l s

20.

M u k Z z n b z n . g Z a mzx.Ze.ana
0n.yzopAZA p u n g z n A

P e re n n ia l herbs
Aa Z z a Z a z v Z A
G auZ£hzn.Za p n o c u m b zn A
H zZZantkuA o c zZ d zn ta Z Z A
HZ zn .a zZ um v z n o A u m
P otzntZ Z Z a tn .Z d zn ta ta
Ptzn.ZdZum aquZZZnum
SoZZdago. Ap.
VZoZa a d u n z a

.3

2.

7.8
.1

9.
1 .4

T re e s and sh ru b s
Am z Z a n z h Z z h . A p .

1.0
.6
3.7
1.7
.8
3.7
6 3 .4

An.ztoAtaphyZQ.A u\/a-un.AZ
C o m p to n Z a pzn.zgn.Zna
PZnuA b a n k A Z a n a
PnunuA pumZZa
Qugaciu App. ( r e d oak)

Uae.cx.nlum 4 p p .

179

2.0
6.8
3.4
.7
5.4
4.0
19.6

TABLE C 3
RELATIVE COVER AND RELATIVE FREQUENCY FOR
EACH SPECIES ON SITE I 3D SURVEYED AS A
MATURE JACK PINE STAND
19 81
S pecies

Cover

A nnuals and b i e n n i a l s
MeZampyA.um ZZneaA.c

.6

Freq
7.4

G ra ss e s and sed ges
8 .6
.1

Ca f i z x p e n 6 y Z v a n X c a
V anthonZa 6 p Z ca ta
0A.yzop6Z6 p u n g e n t
SciiA.za.chne, p u A .p u A .a 6 c e n 6

1.0

1.6
1.6

.2
.3
.2

1.1
2.6

*

P e re n n ia l herbs
Ap o c y n u m and A. o6 ae mZ& oZ Zu m
A 6 t e A . Zae.v Z6
ChA.mapkA.ta u m b e Z Z a t a
pA.a gaA Za VA.KgA.nA.ana
G a u Z t h e A Z a pA.ocambe.n6
H eZZanthu6 o c c Z d e n ta Z Z 6
Ma Z a n t h e m a m c a n a d e n 6 c
Pt eA .Z dZ um aquA.ZA.nam
SoZZdago 6pp.

2.7
38.1
*

T re e s and sh ru b s
AmcZanchZcA. 6 p .
A/lc t o 6 t a p k y Z o 6 uva-uA.6Z
C omptonZ a p&A.egAZna
CA.atae.gu6 6 p .
Ep-tgea A.epen6
PZnu6 bank.6Zana
PZnu6 6 t A o b u 6
PopuZu6 tA.emaZoZde6
PA.unu6 pumZZa
Pauhu 6 6CA.otZna
Qu &a.cu6 6 p p . ( r e d o a k )
Rubu6 p e n 6 Z Z v a n Z c u 6
VaccZnZum 6 p p .

.9
.5
.8
3.0
.4
1.9
.9
.6
39.0

180

15 .3

.5
9.5
.5
2.6
13.6
.5

3.7
2.1
4.2
1.1
5.8
.5
.5
5.3
2 .1
.5
2.1
15.8

Appendix D

National Cooperative S o il Survey
f or the Grayling S e r ie s

Nati onal Cooperat ive S o i l Survey f o r the Grayling S e r i e s .

c rattling

series

The C r a y l l t i g a e r i e s I s a mixed, f r i g i d Typic UdipsamBeot*. These s o i l s e r e s e n d t h r o u g h o u t*
T y p i c a l l y , t h e y have b l a c k en d g r e y i s h brown A h o r i z o n s , d a r k brown sod s t r o n g brown B h o r i z o n s *
and L ig h t brown C h o r i z o n s .
T y p i c a l Fed on i
sta te d ,)

G rayling send - fo re ste d *

(Colors e r e f o r n o l s i s o i l u n le s s o th e r w is e

Al & A2— 0 t o 3 i n c h e s ; b l a c k (H 2 / ) ( A l ) , en d g r a y i s h brown (10YR 5 / 2 ) s e n d , (A2); c o a t e d
and u n c o a t e d sa nd g r a i n s n i x e d t h r o u g h o u t t b s h o r i z o n , g i v i n g a s a l t and p e p p e r a p p e a r a n c e ;
D o d s r s t s o r g a n i c m a t t e r co o t t a t I n u pper p a r t ; week a e d l i a g r a n u l a r s t r u c t u r e ; v e r y f r i a b l e ;
v e r y s t r o n g l y a c i d ; a b r u p t smooth b o u n d a r y ,
(2 Co 4 inch** t h i c k )
B 2 i l r —<-3 co 9 I n c h e s ; d a r k brown (7.5YR 4 / 6 ) s a n d ; weak c o a r s e g r a n u l a r s t r u c t u r e ; v e r y
f r i a b l e ; s t r o n g l y a c i d ; c l e a r smooth bo u n d ary ,
(4 t o B I n c h e s t h i c k )
B 2 2 i r — 9 t o 15 I n c h e s ; s t r o n g brown ( 7 , SYR 5 / 6 ) se n d; v e r y week c o a r s e g r a n u l a r s t r u c t u r e ;
very f r i a b l e ; o e d l m a c i d ; c l e a r I r r e g u l a r boundary.
(4 t o 14 I n c h e s c h i c k )
B3— LS co 23 I n c h e s ; brown (7.5TR S /4) s a n d ; s i n g l e g r a i n e d ; l o o s e ; medium a c i d ; g r a d u a l
smooth b o u n d a r y ,
(3 t o 10 l a c h e s t h i c k )
C— 23 co 60 I n c h e s ; l i g h t brown ( 7 . STR 6 / 4 ) s a n d ; s i n g l e g r a i n e d ; l o o s e ; medium acid *
Type L o c a t i o n : D e l t a C ounty, M ic h ig a n ; S i * , S i * , SEk, S I* , Sec. 34, T. 41 H ., H. 21 W.;
0 . 2 a l l e s o u t h o f U. S. 2 on 1-21 t h e n e a s t 100 f e e t .
Range I n C h a r a c t e r i s t i c s : T h i c k n e s s o f t h e solum r a n g e s f r o a a b o u t 15 t o 30 I n c h e s , The
u p p e r e i g h t I n c h e s o f t h e solum i s sa nd o r l o a a y s a n d and t h e r e s t o f t h e solum l a sand;
medium sand l a d o m in a n t. The u p p e r p a r t o f t h e s o i l I s very s t r o n g l y a c i d o r s t r o n g l y a c i d
and t h e low er p a r t I s medium a c i d o r s l i g h t l y a c i d .
Kean a n n u al s o i l t e m p e r a t u r e J s e s t i m a t e d
t o r a n g e f r o a a b o u t 3& t o 43° P. Some pedons have an 01 h o r i z o n ,
t o lH I n c h e s t h i c k .
It is
d a r k g r a y i s h brown (10YR 4 / 2 ) t o b l a c k (10YR 2 /L ) and i s composed o f o ak l e a v e s o r Ja ck p i n e
n e e d l e s , and some t w i g s an d r o o t s In v a r i o u s s t a g e s
o f d e c o m p o s i ti o n . The Al an d A2 h o r i z o n s
a r e n o r m a l l y I n t e r m i x e d in a s i n g l e l a y e r , b ut some p e d a n t have a s e p a r a t e A2 h o r i z o n .
In t h e
l a t t e r pedons t h e Al h o r i z o n r a n g e s from 1 t o 3 I n c h e s i n t h i c k n e s s an d t h e A2 h o r i z o n from I
t o 3 I n c h e s In t h i c k n e s s , b u t t h e A2 h o r i z o n i s i n t e r m i t t e n t w i t h i n t h e pedon* The Al h o r i z o n
h a s h u e o f 7.5TR o r IOTA, v a l u e o f 2 t h r o u g h 4 and chroma o f 1 o r 2. The A2 h o r i z o n h a s hue of
7.5VR o r 10YR, v a l u e o f 5 o r 6 , and chroma o f 2. The B 2 l l r h o r i z o n h a s hue o f 5YR, 7.5YR o r
1GTR, v a l u e o f 4 o r S, and chroma o f 4; t h e B221r h o r i z o n has s i m i l a r h ue b u t v a l u e r a d g e s
from 4 th r o u g h 6, and chroma from 6 t h r o u g h B. The B3 h o r i z o n h a s hue o f SYR, 7.6YR or 10YR,
v a l u e o f 5 o r 6, sod chroma o f 6 th r o u g h B. The C h o r i z o n has hue o f 2.5YR, 6YR, 7.5YK o r
10YR, v a l u e o f 6 o r 7 , and chroma o f 2 t h r o u g h 4 . Some pedons have c o a r s e sa nd In t h e C
h o r i z o n . Dark m i n e r a l s a r e more co neo n In t h e n o r t h w e s t e r n p a r t o f t h e s o i l s r a n g e .
CcQpetlog S e r i e s an d T h e i r D i f f e r e n t i a e : These a r e t h e C l a i r e , F r i e n d s h i p , Hen ah ga, Kymore,
P l a l & b o , S a r t e l l , S c f d c a and Shawano s e r i e s i o t h e &aar f am ily and t h e C r c s w e l l , Deer P a r k ,
Hiuood, O a k v i l l e , Omega, P l a i n f i e l d , Rubicon, V i l a s an d Windsor s o i l s . C l a i r e and S erden s o i l s
are le s s acid.
F r i e n d s h i p s o i l s have m o t t l e s i n t h e c o n t r o l s e c t i o n * Menahga s o i l s ( s e e
Rem ark s). Kymore s o i l s have c o a r s e r s a n d s . P l a i n b o s o i l s have b ed r o c k a t d e p t h s of 20 t o 40
I n c h e s . S a r t e l l an d Shawano s o i l s have f i n e r s a n d s . C r o s w e l l , R ubicon, V i l a s and Windsor
s o i l s have s p o d l c h o r i z o n s . Deer P ark s o i l s have a l b l c h o r i z o n s o v e r l y i n g d a r k e r c o l o r e d B
h o r i z o n s and h av e l e s s a c i d s o l a .
Hlwopd s o i l s have d o m in a n tly f i n e sand t h r o u g h o u t t h e
co n tro l se ctio n .
O a k v i l l e , P l a i n f i e l d and Windsor s o i l s h^ve mean a n n u a l s o i l t e m p e r a t u r e
g r e a t e r t h a n 47° F. Omega s o i l s have loamy sand B h o rizo n s*
S e t c l n i t : G r a y l i n g s o i l s a r e on o u t v a s h p l a t s * and l a k e p l a i n s o f W isconsin a p e .
S lo pe
g r a d i e n t s ace d o m i n a n t l y l e s s t h a n 6 p e r c e n t b u t r a n g e f r o a 0 t o a b o u t 15 p e r c e n t * These s o i l s
formed in sandy g l a c i o f l u v l a l se dim e n ts * The c l i m a t e i s c o n t i n e n t a l . Aver ag e a n n u a l p r e c i p i ­
t a t i o n r a n g e s from 26 t o 33 i n c h e s ; mean a n n u a l t e m p e r a t u r e from a b o u t 61 t o 6 5 ° F . ; and mean
5_c=!er t e m p e r a t u r e from 60 t o 6fl° F.

181

BIBLIOGRAPHY

BIBLIOGRAPHY
A b r a m s , M. D. a n d D. I . D i c k m a n n .
1982.
E arly
r e v e g e t a t i o n o f c l e a r c u t and b u rn ed j a c k p in e
s i t e s i n n o r t h e r n low er M ichigan.
Can. J . B o t.
(in p ress) .
A hlgren,
C. E.
1959. V e g e t a t i o n a l d e v e lo p m e n t f o l l o w ­
in g b u rn in g in the n o rth e rn c o n ife ro u s f o r e s t in
M innesota.
P r o c . S o c . Amer. F o r . 1 9 6 0 : 2 1 - 2 2 .
A hlgren,
C. E.
1 9 6 0 . Some e f f e c t s o f f i r e on r e p r o ­
d u c tio n and gro w th o f v e g e t a t i o n i n n o r t h e a s t e r n
M innesota.
E cology 4 1 :431-445.
A hlgren,
C. E.
1974. E f f e c t s o f f i r e on t e m p e r a t e
fo rests:
N orth C e n tr a l U n ited S t a t e s , In
F i r e a n d E c o s y s t e m s , T. X. K o z l o w s k i a n d C. E. A h l g r e n ( E d . ) , A c a d e m i c P r e s s I n c . , N. Y.
pp. 195-219.
A hlgren,
C. E.
1976. R e g e n e r a t i o n o f r e d p i n e and
w h ite p in e fo llo w in g w i l d f i r e and lo g g in g in
n o r t h e a s t e r n M innesota,
J . F o r e s t r y 74:135140 .
A h l g r e n , C. E.
1979A.
B uried seed in the f o r e s t flo o r
o f t h e B o u n d a r y W a t e r s Ca n oe A r e a .
M in n . F o r .
R es. Note 271, 4 p p .
A h l g r e n , C. E.
1979B.
B uried seed in p re s c rib e d burned ja c k p in e f o r e s t s o i l s , n o r th e a s te r n
M innesota,
M inn. F o r . R e s . N o t e 2 7 2 , 3 p p .
A h l g r e n , C. E .
1 9 7 9C.
E m ergent s e e d l i n g s on s o i l from
b u rn e d and u n burned r e d p i n e f o r e s t .
M in n. F o r .
Res, N ote 273, 3 p p .
A h lg r e n , I . F.
effects

a n d C. E. A h l g r e n .
1960.
E co lo g ical
of fo rest fire s .
B o t. Rev. 2 6 : 4 8 3 - 5 3 3 .

B a k e l a a r , F . H. a n d E. P. Odum.
1978.
Community
and p o p u l a t i o n l e v e l r e s p o n s e s t o f e r t i l i s a ­
t i o n i n an o l d - f i e l d eco sy stem .
E c o l o g y 59 :
660-665.

182

183

B e a u f a i t , W. R.
1962.
Jack pine rep ro d u ctio n .
Mich.
C o l l . M i n i n g T e c h n o l . T e c h . B u l l . 9, 37 p p .
B ell,

D. T . a n d J . M. K o c h .
1980.
P o s t- f ir e su ccessio n
i n th e n o r t h e r n j a r r a h f o r e s t o f W estern Aus­
t r a l i a , A ust. J . E col, 5 :9 -1 4 .

B e n z i e , J . W.
1977.
M a n a g e r's handbook f o r j a c k p in e
in th e N orth C e n tra l S t a t e s .
USDA F o r . S e r .
Gen. T e c h . Rep. N C - 3 2 , 18 p p .
B o r m a n n , F . H . , G. E. L i k e n s , D. W. F i s h e r , a n d R. S.
P ierce.
1968.
N u t r i e n t l o s s a c c e l e r a t e d by
c l e a r c u t t i n g o f a f o r e s t ecosystem .
S cience
159:882-884.
B u e l l , M. F . a n d J . E. C a n t l o n .
1953.
E ffects of
p r e s c r i b e d b u r n i n g o n g r o u n d c o v e r i n t h e New
J e rse y p ine region.
E cology 3 4 :5 2 2 -5 2 8 .
C a i n , S . A.
M idi.

1938.
The s p e c i e s - a r e a c u r v e .
N at. 19:573-581.

Amer.

C a t t e l i n o , P. J . , I . R. N o b l e , R. 0 . S l a t y e r , a n d
S . R. K e s s e l l .
1979.
P re d ic tin g th e m u ltip le
pathw ays o f p l a n t s u c c e s s i o n .
E n v i r o n . Manage.
3:41-50.
C a y f o r d , J . H.
1970.
The r o l e o f f i r e i n t h e e c o l o g y
and s i l v i c u l t u r e o f j a c k p i n e .
P r o c . 1 0 t h Ann.
T a l l T im bers F i r e E c o l. C onf.
pp. 221-244.
C h r i s t e n s e n , N. L,
1973.
F i r e and th e n i t r o g e n c y c le
in C a lifo rn ia c h ap a rra l.
S cien ce 181:66-68.
C h r i s t e n s e n , N. L. a n d C. H. M u l l e r .
1975A.
E ffects
o f f i r e on f a c t o r s c o n t r o l l i n g p l a n t gro w th i n
Adenostom a c h a p a r r a l .
E c o l . Monogr. 4 5 : 2 9 - 5 5 .
C h r i s t e n s e n , N. L. a n d C. H. M u l l e r .
1975B.
R elativ e
im portance o f f a c t o r s c o n t r o l l i n g g e rm in a tio n
and s e e d l i n g s u r v i v a l i n A denostom a c h a p a r r a l .
Amer. M i d i . N a t . 9 3 : 7 1 - 7 8 .
C h r i s t e n s e n , P . E. a n d P . C. K i m b e r .
1975.
E ffect of
p r e s c r i b e d b u r n i n g on t h e f l o r a a n d f a u n a o f
south-w est A u s tra lia n f o r e s ts .
P roc. E col.
Soc. A u s t.
9:85-106.

184
C l e m e n t s , F. E.
1916.
P la n t S uccession:
o f th e developm ent o f v e g e t a t i o n .
I n s t . W ash. P u b l . 2 4 2 , 512 p p .

An a n a l y s i s
C arnegie

C o n n e l l , J . H. a n d R. 0 . S l a t y e r .
1977.
Mechanisms
o f s u c c e s s i o n i n n a t u r a l com m unities and t h e i r
r o l e i n c o m m u n i ty s t a b i l i t y a n d o r g a n i z a t i o n .
Am er. N a t u r . 1 1 1 : 1 1 1 9 - 1 1 4 4 .
C o o p e r , W. S .
1913.
The c l i m a x f o r e s t o f I s l e R o y a l e ,
Lake S u p e r i o r , and i t s d e v e lo p m e n t.
B ot. G azet.
55:189-235.
C u s h w a , C. T . , R. E. M a r t i n , a n d R. L. M i l l e r .
The e f f e c t s o f f i r e on s e e d g e r m i n a t i o n .
Range Manage. 2 1 : 2 5 0 - 2 5 4 .

1968.
J.

D e b a n o , L . F. a n d C. E. C o n r a d .
1978.
The e f f e c t s
o f f i r e on n u t r i e n t s i n a c h a p a r r a l e c o s y s t e m .
E c o lo g y 59: 4 8 9 -4 9 7 .
Denslow , J . S.
1980.
P a tte rn s of p la n t sp ecies d iv e r­
s i t y d u rin g su c ce ssio n under d i f f e r e n t d i s t u r ­
bance regim es.
O ecologia (B e rl.) 46:18-21.
D i x , R. L .
1960.
The e f f e c t s o f b u r n i n g on t h e m u lch
s t r u c t u r e and s p e c ie s c o m p o sitio n of g r a s s la n d s
i n w e s te r n N o rth D akota.
E cology 4 1 :4 9 -5 6 .
D y r n e s s , C. T.
1973.
E arly sta g es of p la n t su ccessio n
f o l l o w i n g lo g g in g and b u r n i n g i n w e s t e r n C ascades
i n O regon.
Ecology 5 4 :5 7 -6 9 .
E g ler,

F . E.
1954.
V e g e t a t i o n s c i e n c e c o n c e p t s 1.
I n i t i a l F l o r i s t i c C om position, a f a c t o r in o ld f i e l d v e g e t a t i o n developm ent.
V e g e t a t i o 4:
412-417.

E y r e , F . H. a n d R. K. L e B a r r o n .
1944.
Management o f
j a c k p i n e i n th e Lake S t a t e s .
U.S.D .A . Tech.
B u l l . No. 8 6 3 .
66 p p .
F e r n a l d , M. L.
1950.
G ra y 's m anual o f b o ta n y .
N o s t r a n d C o . , N.Y .
1632 p p .

D. Van

F l i n n , M. A. a n d R. W. W ein .
1977.
D epth o f u n d e r­
g round p l a n t o r g a n s and t h e o r e t i c a l s u r v i v a l
during f i r e .
Can. J . B o t. 5 5 :2 5 5 0 - 2 5 5 4 .

185
F l o y d , A. G.
1966.
E f f e c t s o f f i r e upon weed s e e d s
in th e wet s c l e r o p h y l l f o r e s t s o f n o r t h e r n
New S o u t h W a l e s .
A u st. J . B ot. 1 4 :243-256.
F l o y d , A. G.
1976.
E f f e c t o f b u r n i n g on r e g e n e r a t i o n
from se e d s i n w et s c l e r o p h y l l f o r e s t s .
A ust.
F or. 39:210-220.
G a s h w i l e r , J . S.
1970.
P l a n t a n d mammal c h a n g e s o n a
c l e a r c u t i n w e s t - c e n t r a l Oregon.
E co lo g y 51:
1018-1026.
G i l l , A. M.
1977.
P la n t t r a i t s a d a p tiv e to f i r e s in
m e d ite rra n e a n land ecosystem s.
I n Sym posiu m on
the en v iro n m en tal consequences
aF f i r e and f u e l
management i n m e d i t e r r a n e a n e c o s y s t e m s .
(ed .)
H.A . Mooney a n d C. E. C o n r a d .
U .S .D .A . F o r.
S e r v . Gen. T e c h , R e p . WO- 3 , p p . 1 7 - 2 6 .
G l e a s o n , H. A.
1926.
The i n d i v i d u a l i s t i c
the p la n t a s s o c ia tio n .
B u ll. T orr.
53:7-26.

concept of
B ot. C lub.

G o d w in , H.
1929.
The s u b - c l i m a x and d e f l e c t e d
sion.
J . E cology 1 7 :1 4 4 -1 4 7 .

succes­

G r a h a m , S . A.
1941.
C lim ax f o r e s t s o f t h e u p p e r p e n i n ­
s u l a o f M ichigan.
E cology 2 2 :3 5 5 -3 6 2 .
H a l l , J . B. a n d M. D. S w a i n e .
G hanaian f o r e s t s o i l s .

1980.
Seed s t o c k s i n
B io tro p . 12:256-263.

H a n e s , T. L.
1971.
S uccession a f t e r f i r e in th e chapar­
r a l of Southern C a lif o r n ia .
E c o l . Monogr. 41:
27-52.
H a r c o m b e , P . A.
1977.
The i n f l u e n c e o f f e r t i l i z a t i o n
o n some a s p e c t s o f s u c c e s s i o n i n a h u m i d t r o p i ­
cal fo rest.
E cology 58:1375-1383.
H a r p e r , J . L.
1977.
P o p u latio n b io lo g y o f p la n ts .
A c a d e m i c P r e s s , New Y o r k , N.Y.
H e i n s e l m a n , M. L.
1973.
F ire in the v i r g in f o r e s ts of
t h e B o undary W a te rs Canoe A re a M i n n e s o t a .
Q uat.
Res. 3:329 -38 2.
H o r t o n , J . S. a n d C. J . K r a e b e l .
1955.
D evelopm ent
o f v e g e t a t i o n a f t e r f i r e i n th e cham ise
chaparral
of sou th ern C a lifo rn ia .
E cology
36:244-262.

186

J e f f e r y , D, W. a n d C. D. P o g o t t .
1973.
The r e s p o n s e
o f g r a s s l a n d s on s u g a r - l i m e s t o n e i n T e e s d a l e
to a p p l i c a t i o n o f p h o sp h o ru s and n i t r o g e n .
J . E cology 6 1 :8 5 -9 2 .
J o r g e n s e n , J . R. a n d C. G. W e l l s .
1971.
Apparent
n i t r o g e n f i x a t i o n i n s o i l i n f l u e n c e d by p r e ­
s c rib e d burning.
S o i l S c i . S o c . Amer. P r o c .
35:806-810.
K e l l m a n , M.
1980.
G eographic p a t t e r n i n g i n t r o p i c a l
weed c o m m u n itie s and e a r l y s e c o n d a r y s u c c e s s i o n .
T r o p ic a l s u c c e s s io n , pp. 34-39.
K e s s e l l , S. R. a n d W. C. F i s c h e r .
1981.
P red ictin g
p o s t f i r e p l a n t s u c c e s s i o n f o r f i r e management
planning.
U . S . D . A . F o r , S e r v . Gen. T e c h . R e p.
I N T - 9 4 . 19 p p .
K e s s e l l , S . R. a n d M. W. P o t t e r .
1980.
A q u an titativ e
s u c c e s s i o n m o d e l f o r n i n e M o n t a n a f o r e s t com m uni­
tie s.
E n v i r o n . Manage. 4 : 2 2 7 - 2 4 0 .
K r e f t i n g , L. W. a n d G. E. A h l g r e n .
1974.
Sm all
mammals a n d v e g e t a t i o n c h a n g e s a f t e r f i r e i n
a m ixed c o n if e r - h a r d w o o d f o r e s t .
Ecology
55:1391-1398.
L ik en s,

G, E . , F . H. B o r m a n n , R. S. P i e r c e , a n d W. A.
R ein ers.
1978.
R ecovery o f a d e f o r e s te d e co ­
system .
S cien ce 199:492-496.

L ittle ,

S . a n d E. B. M o o r e .
1949.
The e c o l o g i c a l
r o l e o f p r e s c r i b e d b urns in the p in e -o a k f o r e s t s
o f s o u t h e r n New J e r s e y .
E cology 3 0 :22 3-2 33.

L i v i n g s t o n , R. B. a n d M. L. A l l e s s i o .
1968.
B uried
v i a b l e s e e d i n s u c c e s s i o n a l f i e l d s and f o r e s t
stan d s,
H arvard F o r e s t,
M ass. B u l l . T o r r .
B o t. Club 9 5 :5 8 -6 9 .
L o u c k s , 0 . L.
1970.
E v o lu tio n of d iv e r s ity , e f f i ­
c i e n c y , and comm unity s t a b i l i t y .
Amer. Z o o l .
10:17-25.
L u g o , A. E.
1980.
Mangrove e c o s y s t e m s :
su ccessio n al
or stead y s ta te ?
T r o p ic a l S u c c e s s io n , pp. 65-72.

187

L y o n , J . L. a n d P . F. S t i c k n e y .
1974.
E arly v e g e ta l
s u c c e s s i o n f o l l o w i n g l a r g e n o r t h e r n R oc ky
M ountain w i l d f i r e s .
P r o c . 1 4 t h Ann. T a l l
T im bers F i r e E c o l. Conf, pp. 355-375.
M a i s s . u r o w , D. K.
1941.
The r o l e o f f i r e i n t h e
p erp etu atio n of v irg in fo re s ts of no rth ern
W iscon sin .
J . F o r e s t. 39:201-207.
M a j o r , J . a n d W. T. P y o t t .
1966.
B uried, v ia b le
s e e d i n two C a l i f o r n i a b u n c h g r a s s s i t e s a n d
t h e i r b e a r i n g on t h e d e f i n i t i o n o f f l o r a .
V eg etatio
13:253-282.
M a r k s , P . L.
1974.
The r o l e o f p i n c h e r r y (P r u n u s
p e n s y lv a n ic a L .) in th e m aintenance o f s t a ­
b i l i t y i n n o r t h e r n hardw ood e c o s y s te m s .
E col.
M onogr. 4 4 : 7 3 - 8 8 .
M a r g a l e f , R.
1963.
On c e r t a i n u n i f y i n g p r i n c i p l e s
in ecology.
Amer. N a t u r . 9 7 : 3 5 7 - 3 7 4 .
M a r s h a l l , J . D.
1980.
F acto rs a ff e c tin g jac k pine
r e g e n e r a t i o n on th e sand p l a i n s o f n o r t h e r n
low er M ichigan.
M. S . t h e s i s , M i c h i g a n S t a t e
U n iv ersity .
M a r t i n , R. E . , R. L. M i l l e r , a n d C. T. Cushwa.
1975.
G e r m i n a t i o n r e s p o n s e o f legum e s e e d s s u b j e c t e d
to m o is t and d ry h e a t .
E cology 56:1441-1445.
McDonough, W. T.
1977.
Serv. 4:155-184.

Seed p h y s io lo g y .

R a n ge S c i .

McGraw, J . B.
1980.
Seed bank s i z e and d i s t r i b u t i o n
of seeds i n c o tto n g r a s s tu s s o c k tu n d r a , E agle
C reek, A laska.
Can. J . B o t . 5 8 : 1 6 0 7 - 1 6 1 1 .
M cL ean , A.
1968.
F ire r e s is ta n c e of f o r e s t species
as i n f lu e n c e d by r o o t sy ste m s.
J . Range
M a n ag e . 2 2 : 1 2 0 - 1 2 2 .
M c P h e r s o n , J . K. a n d C. H. M u l l e r .
1969.
A llelo p ath ic
e f f e c t s o f A denostom a f a s c i c u l a t u m " c h a m is e " , i n
the C a lifo rn ia c h a p a rra l.
E c o l . M o n og r. 3 9 : 1 7 7 198.
M o r r i s o n , I . K. a n d N. W. F o s t e r .
1977.
F ate of u rea
f e r t i l i z e r added to a b o r e a l f o r e s t P inus
b a n k s i a n a Lamb, s t a n d .
S o i l S c i . Amer. J . 44:
441-448.

188

M u l l e r , C. H . , R. B. H a n a w a l t , a n d J . K. M c P h e r s o n .
1968.
A l l e l o p a t h i c c o n t r o l o f h e rb grow th
in the f i r e c y c le o f C a li f o r n ia c h a p a rr a l.
B u ll. T o rr. B ot. C lub. 95:225-231.
N i e r i n g , J . A. a n d R. H. Goo d w in .
1974.
C reatio n
of r e l a t i v e l y s ta b le shrublands w ith h e rb ic id e s :
a r r e s t i n g s u c c e s s i o n on r i g h t - o f - w a y and p a s t u r e
land.
E cology 5 5 :784-795.
N oble,

I . R.
1980.
I n t e r a c t i o n s betw een tu s s o c k
g r a s s (Poa s p p .) and E u c a l y p t u s p a u c i f l o r a
seedlings near tr e e lin e in so u th eastern
A u stralia.
O eco lo g ia ( B e rl.) 45:350-353.

N oble,

I . R. a n d R. 0 . S l a t y e r .
1980.
The u s e o f
v i t a l a t t r i b u t e s to p r e d i c t s u c c e s s io n a l changes
i n p l a n t com m unities s u b j e c t t o r e c u r r e n t d i s ­
turbances.
V e g e ta tio 43:5-21.

N o b l e , J . C . , A. D. B e l l , a n d J . L. H a r p e r .
1979.
The p o p u l a t i o n b i o l o g y o f p l a n t s w i t h c l o n a l
grow th.
J . Ecology 6 7 :983-1009.
Odum, E. P .
1971.
Fundam entals o f E cology.
W. B.
S a u n d e r s C o . , P h i l a d e l p h i a , PA. 574 p p .
Ohmann, L. F . a n d D. F. G r i g a l .
1979.
E arly rev eg e­
t a t i o n and n u t r i e n t dynam ics f o l l o w i n g th e
19 71 L i t t l e S i o u x f o r e s t f i r e i n n o r t h e a s t e r n
M innesota.
F o r . S c i . M o no g r. 2 1 : 1 - 8 0 .
O u t c a l t , K. W. a n d E.
c a l changes in
tim b er h a r v e s t
J . F o r. Res.

H. W h i t e .
1981.
P h y to so cio lo g iu n d e rsto ry v e g e ta tio n fo llo w in g
i n n o r th e r n M innesota.
Can.
11:175-183.

P a r s o n s , D. J .
1976.
The r o l e o f f i r e i n n a t u r a l
c o m m u n i t i e s : An e x a m p l e f r o m t h e S o u t h e r n
S i e r r a Nevada, C a l i f o r n i a . E n v iro n . C onserv.
3:91-99.
P u r d i e , R. W.
1977.
E arly stag es of re g e n e ra tio n
a f t e r b u rn in g in dry s c le r o p h y ll v e g e ta tio n ,
I I R e g e n e r a t i o n by s e e d g e r m i n a t i o n .
A ust. J .
B ot. 2 5 :3 5 -4 6 .
P u r d i e , R. W.
a n d R. 0 . S l a t y e r .
1976.
V eg etatio n
s u c c e s s i o n a f t e r f i r e i n s c l e r o p h y l l w oodland
com m unities i n s o u t h - e a s t e r n A u s t r a l i a .
A ust.
J . E cology 1 :2 2 3 -2 3 6 .

189

R i c h a r d s , P . W.
1955.
The s e c o n d a r y s u c c e s s i o n i n
the t r o p ic a l r a i n f o r e s t.
S cience P ro g re ss ,
Lond. 4 3 : 4 5 - 5 7 .
S c h e i n e r , S. M. a n d J . A. T e e r i .
1981.
A 53-year
re c o rd of f o r e s t su ccessio n fo llo w in g f i r e in
n o r t h e r n low er M ichigan.
The M ich. B o t. 2 0 : 3 - 1 4 .
S h a f i , M. X. a n d G. A. Y a r r a n t o n .
1973.
D iv ersity ,
f l o r i s t i c r i c h n e s s , and s p e c ie s evenness d u rin g
a secondary ( p o s t - f i r e ) su ccessio n .
E co lo g y 54:
897-902.
S i m a r d , A. J . a n d R. W. B l a n k .
1981.
F ire h is to ry of
a M ichigan j a c k p in e f o r e s t .
Mich. Acad. S c i . ,
A r t s , and L e t t e r s ( s u b m itte d ) .
S o m m e r s , L. M.
1977.
A t l a s o f M ichigan.
S ta te U n iv ersity P ress.

M ichigan

S p e c h t , R. L . , D. L. C o n n e r , a n d H. T. C l i f f o r d .
1977.
The h e a t h - s a v a n n a p r o b l e m :
the e f f e c t o f f e r t i l i ­
z e r on s a n d - h e a t h v e g e t a t i o n o f N o rth S t r a d b r o k e
I s l a n d , Q u eensland.
A u st. J . Ecology 2 :1 7 9 -1 8 6 .
S t.

J o h n , T . V. a n d P . W. R u n d e l .
1976.
The r o l e o f
f i r e as a m in e ra liz in g agent in a S ie rra n c o n if ­
erous f o r e s t .
O ecologia (B e rl.) 25:35-45.

S t a f f o r d , L. 0 . a n d S. M. F i l i p .
1974.
Biom ass and
n u t r i e n t c o n te n t o f 4 - y e a r - o ld f e r t i l i z e d and
u n f e r t i l i z e d n o r t h e r n hardw ood s t a n d s .
Can. J .
F o r. Res. 4 :5 4 9 -5 5 4 .
S t e a r n s , F . W.
1949.
N in e ty y e a r s change i n a n o r t h e r n
hardw ood f o r e s t i n W is c o n s in .
Ecology 3 0 :3 5 0 358.
S t e p h e n s o n , S. N.
1973.
A com parison o f p r o d u c t i v i t y
and d i v e r s i t y i n e a r l y and l a t e s e a s o n o l d - f i e l d
p l a n t com m unities.
M i c h . Academ. 5 : 3 2 5 - 3 3 4 .
S t o n e , E. C. a n d G. J u h r e n .
1951.
The e f f e c t o f f i r e
o n t h e g e r m i n a t i o n o f t h e s e e d o f Rhus o v a t a .
Amer. J . B o t . 3 8 : 3 6 8 - 3 7 2 .
S w a i n , A. M.
1973.
A h i s t o r y o f f i r e and v e g e t a t i o n
i n n o r t h e a s t e r n M in n e so ta a s r e c o r d e d by l a k e
sedim ents.
Q uat. Res. 3 :3 8 3 -3 9 6 .

190

S w e e n e y , J . R.
1956.
Response o f v e g e t a t i o n to f i r e .
U niv. C a l i f . P u b l. B o t. 2 8 :1 4 3 -2 5 0
T h o m s o n , J . W.
1943.
P l a n t s u c c e s s i o n on abandoned
f i e l d s i n th e c e n t r a l W iscon sin sand p l a i n
area.
B u ll. T o r r. B ot. C lub. 7 0 :3 4 -4 1 .
T r a b a u d , L. and J . L e p a r t .
1980.
D i v e r s i t y and
s t a b i l i t y in G a rrig u e ecosystem s a f t e r f i r e .
V e g e ta tio 43:49-57.
T r e d i c i , P . D.
1971.
T he b u r i e d s e e d o f C o m p t o n i a
p e r e g r i n a , th e sw eet f e r n .
B u l l. T o r r . B ot.
Club 1 0 4 :2 7 0 -2 7 5 .
U h l , C . , K. C l a r k , H. C l a r k , a n d P . M u r p h y .
1981.
E a r l y p l a n t s u c c e s s i o n a f t e r c u t t i n g and b u r n in g
i n t h e U p p e r R i o N e g r o R e g i o n o f t h e Amazon
B asin.
J . E cology 6 9 :6 3 1 -6 4 9 .
U .S .D .A .
1971.
M ic h ig a n W eath er S e r v i c e f o r Houghton
L ake, G r a y l i n g , Mio, and H i g g i n s Lake S t a t i o n .
V eatch,

J . W . , L. R. S c h o e m a n n , a n d J . W, Moon.
1924.
S o i l S u r v e y o f Roscommon C o u n t y , M i c h i g a n .
U .S.D .A .

V o g l , R. J .
1964.
V e g e t a t i o n a l h i s t o r y o f Crex
Meadows, a p r a i r i e s a v a n n a i n N o r t h w e s t e r n
W isconsin.
Amer. M i d i . N a t . 7 2 : 1 5 7 - 1 7 5 .
V o g l , R.
J . 1970.
F i r e and t h e n o r t h e r n W is c o n s in
pine b a rre n s.
P r o c . 1 0 t h Ann, T a l l T i m b e r s
F i r e E c o l. Conf. pp. 175-209.
V o s s , E.
G. 1 9 7 2 .
M i c h i g a n F l o r a , P a r t X Gymnospe rms
and M onocots.
C r a n b r o o k I n s t . S c i . B u l l . 55,
4 88 p p .
W e i n , A. J . a n d L . C. B l i s s .
1973.
Changes i n a r t i c
E riophorum t u s s o c k com m unities f o llo w in g ' f i r e .
E cology 5 4 :8 4 5 -8 5 2 .
W e l l s , C. G.
1979.
E f f e c t s o f f i r e on s o i l .
F o r . S e r v . Gen. T e c h . R e p . WO-7 , 34 p p .

U.S.D .A .

W e n t , F . W . ,G. J u h r e n , a n d M. C. J u h r e n .
1952.
and b i o t i c f a c t o r s a f f e c t i n g g e r m in a tio n .
E cology 33:351-364.

F ire

191

W e s t , N. E. a n d W. W. C h i l c o t e .
1968.
S enecio
s y lv a tic u s in r e l a t i o n to D o u g la s-F ir c l e a r c u t s u c c e s s i o n i n th e Oregon c o a s t r a n g e .
E cology 49:1101-1107.
W i l l i a m s , W. T . , G. N. L a n c e , L. J . Webb, J . G.
T r a c e y , a n d M. B. D a l e .
1969.
S tu d ie s in the
n u m e r ic a l a n a l y s i s of com plex r a i n - f o r e s t
com m unities.
J . E cology 5 7 :5 1 5 -5 3 5 .
W i l l i s , A. J .
1963.
B ra u n to n Burrows:
The e f f e c t s
on th e v e g e t a t i o n o f t h e a d d i t i o n o f m i n e r a l
n u t r i e n t s to t h e dune s o i l .
J . E c o l o g y 51:
353-374.
W o o d w e l l , G, H.
1967.
R a d ia tio n and p a t t e r n s
n atu re.
S c i e n c e 156: 4 6 1 -4 7 0 .

in

Z av itk o v sk i, J .
1976.
Ground v e g e t a t i o n b i o m a s s ,
p r o d u c t i o n , and e f f i c i e n c y o f e n e rg y u t i l i z a t i o n
i n some n o r t h e r n W i s c o n s i n f o r e s t e c o s y s t e m s .
E cology 5 7 :6 9 4 -7 0 6 .