Montana Chrysothamnus
by Gail Lynn Winkler
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in
Range Science
Montana State University
© Copyright by Gail Lynn Winkler (1987)
Abstract:
During the summers of 1985 and 1986, a study was conducted to determine what Chrysothamnus taxa
occur in Montana and to record their respective distributions. Taxa found included C. linifolius. C.
nauseosus ssp. nauseosus, C. n. ssp. albicaulis. C. n. ssp. graveolens. C. n. ssp. consimilis. C.
viscidiflorus ssp. lanceolatus. C. v. ssp. viscidiflorus var. stenophyllus. C. v. ssp. viscidiflorus var.
viscidiflorus, and the previously uncharacterized, C. parryi ssp. glandulosus. The morphology, habitat,
and relationships of these taxa were discussed and their distributions described.
In 1985, a previously uncharacterized taxon of C. parryi was located in southwestern Montana. C. p.
ssp. glandulosus derived its name from conspicuous stalked glands. Two varieties of this taxon were
observed at 2 of 7 study sites. The typical form is named C. ssp. glandulosus var. glandulosus. The
other variety is covered with tomentum, which obscures the stalked glands and is named C. ssp.
glandulosus var. tomentosus.
Additional populations were located during the summer of 1986 and sites were sampled to characterize
the taxon’s habitat. All 7 sites sampled were found to have near-neutral, sandy loam to loam soils, and
supported Artemisia tridentata/grassland communities. Aspects, slopes, elevations, and annual
precipitation amounts varied among sites. An adjacent paired plot, lacking the taxon, was sampled at
each site in an attempt to ascertain what factor(s) account for this taxon’s occurrence. Disturbance and
soil type were the only factors found that likely influence the occurrence and distribution of C. p. ssp.
glandulosus. Evidence indicated this taxon is severely browsed yearly, which may be a factor limiting
the expansion of this taxon.
MONTANA CHRYSOTHAMNUS
by
G a i l Lynn W i n k l e r
A t h e s i s su b m itte d i n p a r t i a l f u l f i l l m e n t
o f th e re q u ire m e n ts f o r th e degree
of
M a s te r o f S c i e n c e
in
Range S c i e n c e
MONTANA STATE UNIVERSITY
Bozem ani Montana
November 1 987
a/ 3
7S
V J y ^ lq I
APPROVAL
o f a t h e s i s s u b m i t t e d by
G a i l Lynn W in k l e r
T h i s t h e s i s h a s b e e n r e a d by e a c h member o f t h e t h e s i s c o m m itte e
an d h a s b e e n fo u n d t o be s a t i s f a c t o r y r e g a r d i n g c o n t e n t , E n g l i s h u s a g e ,
fo rm a t, c i t a t i o n s , b ib lio g r a p h ic s t y l e , and c o n s is te n c y , and i s rea d y
f o r su b m issio n t o t h e C o lleg e o f G rad u ate S t u d ie s .
Date
C h a i r p e r s o n , G r a d u a te Com m ittee
Approved f o r t h e M ajor D e p a rtm e n t
Head, M ajor D e p a r tm e n t
D ate
Approved f o r t h e C o l l e g e o f G r a d u a te S t u d i e s
/ - - / 2 . - Z / " _____________
D ate
G r a d u a te Dean
iii
STATEMENT OF PERMISSION TO USE
In
p re se n tin g
th is
th e sis
in
re q u ire m e n ts o f a m a s t e r 's degree a t
p a rtia l
fu lfillm e n t
Montana S t a t e
U n iv e rsity ,
of
th e
I ag ree
t h a t t h e L i b r a r y s h a l l make i t a v a i l a b l e t o b o r r o w e r s u n d e r r u l e s o f t h e
L ib ra ry .
B r i e f q u o t a t i o n s from t h e t h e s i s a r e a l l o w a b l e w i t h o u t s p e c i a l
p e r m i s s i o n , p r o v i d e d t h a t a c c u r a t e ac k n o w led g m e n t o f s o u r c e i s made.
P e rm issio n
for
e x te n siv e
q u o ta tio n
from
t h e s i s may be g r a n t e d by my m a jo r p r o f e s s o r ,
Dean o f L i b r a r i e s when,
th e
m ate ria l
m aterial
is
for
in
th e
sc h o la rly
purposes.
Any
re p ro d u c tio n
or in h is absence,
o p in io n o f e i t h e r ,
of
th is
by t h e
th e proposed use o f
c o p y in g
or
use
of
th e
i n t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t
my w r i t t e n p e r m i s s i o n .
S ig n a tu re
D a te
or
.
/ 9 3 /
ACKNOWLEDGEMENTS
I
w ould
several
to
E.
lik e
to
th ank
th e
p e o p le t h a t a s s i s t e d
D.
M cA rthur a t
chromosome
c o u n ts
a p p re c ia tio n
c o lle c t
to
th e
on
a ll
sp ecim en s
of
tim e ,
me i n t h i s e n d e a v o r .
p arry i
ssp.
g lan d u lo su s.
p e rso n s aro u n d
C h r y s o th a m n u s .
th e
I 'd
B ray
c o lle c tin g
a ll
th e ir
h e lp
w ith
S ta te
T hese
s ig n ific a n tly to th is p ro je c t.
for
and
en co u rag em en t
of
Thanks a r e o f f e r e d
USES S h ru b S c i e n c e Lab i n
Ci
th e
e ffo rts ,
Provo,
U tah f o r
I
e x te n d
th a t
to o k
c o lle c tio n s
sp e cia l
tim e
to
c o n trib u te d
l i k e t o t h a n k S h e l l y D oyle a n d Rob
and
sa m p lin g .
To
C arl
Wambolt , my a c a d e m ic a d v i s o r , t h a n k you f o r t h e many h o u r s o f w o rk , t h e
p a ti e n c e , and t h e e n c o u ra g in g w ords.
and Jack T a y lo r,
w ords
and
in creased
illu stra tio n
much!
b e lie f
th a n k s f o r your g u id an c e and s u g g e s t i o n s .
sto rie s
my
a lw a y s
m eant
u n d e rsta n d in g
of
To c o m m itte e m em bers, J a c k Rumely
Ci
p arry i
of
ssp.
so
th e
m uch.
You b o t h
E n g lish
lan g u ag e.
g lan d u lo su s
S p e c i a l t h a n k s go t o my p a r e n t s .
in
me s u p p o r t me e v e r y
f r i e n d , S te v e.
day.
is
Your k i n d
have
b e a u tifu l!
g re a tly
Jack,
th e
Thanks
so
Your l o v e , e n c o u r a g e m e n t , a n d
L a stly ,
warm t h a n k s
F r i e n d s s u c h a s you a r e o n e i n a m i l l i o n !
t o my b e s t
V
TABLE OF CONTENTS
v
•
LIST OF TABLES...................................................................................................................
LIST OF FIGURES.......... ..................................................................................
ABSTRACT..................................
Eage
v ii
v iii
ix
I
LITERATURE REVIEW...........................................
3
P h y lo g e n y a n d C l a s s i f i c a t i o n o f C hry so th a m n u s
M o rp h o lo g y ..................................................................................
Anatomy.........................................................................................
E m b ry o lo g y ..................................................................................
Chromosome R e l a t i o n s h i p s ................................................
R e p r o d u c t i v e H a b i t s . .......................
D i s t r i b u t i o n a n d S i t e . C h a r a c t e r i s t i c s ................................................
L i f e H i s t o r y ............ ........................................................ ............................
U ses a n d Management O p p o r t u n i t i e s . ......................................................
STUDY AREAS AND METHOD........................
VO OO —3 U l LU
INTRODUCTION.......................................................................
10
11
16
19
24
G e n e r a l S u r v e y M e th o d s ..............................................................................
Unknown Taxon S tu d y Area a n d M e th o d s ...................................................
24
25
RESULTS AND DISCUSSION........................ .......................................................................
29
P h y lo g e n y a n d C l a s s i f i c a t i o n o f C h ry so th a m n u s p a r r y i
s s p . g l a n d u l o s u s . ...............................................................................................
M orphology o f C h ry so th am n u s p a r r y i s s p . g l a n d u l o s u s ...............
The R e l a t i o n s h i p o f C h ry so th a m n u s p a r r y i s s p . p a r r y i
t o o t h e r C h ry so th a m n u s T a x a ........................................................................
M orphology o f O t h e r C h ry so th a m n u s Taxa i n M o n ta n a ....................
C h ry so th a m n u s n a u s e o s u s ...........................................................................
C h ry so th a m n u s v i s c i d i f l o r u s ....................................................................
C h ry so th a m n u s l i n i f o l i u s ........................................................................
D i s t r i b u t i o n o f Montana C h r y s o th a m n u s .................................................
H a b i t a t o f Montana C h r y s o th a m n u s ............................................................
C h ry so th a m n u s p a r r y i s s p . g l a n d u l o s u s .........................................
C h ry so th a m n u s n a u s e o s u s .........................................................
C h ry so th a m n u s v i s c i d i f l o r u s .......... ......................................................
C h ry so th a m n u s l i n i f o l i u s ...................................................................
BI
34
35
37
37
40
42
42
53
53
62
68
70
vi
TABLE OF CONTENTS - c o n t i n u e d
O v e rla p p in g t a x a
.......... .......................
B ro w sin g o f Montana C h r y s o th a m n u s ..................... .................. ..
A d d i t i o n a l U ses f o r C h ry so th a m n u s Taxa i n M o n t a n a . . . . . . . . .
70
73
74
SUMMARY......... ................................ .U ............... ........................................................
77
LITERATURE CITED...................... .................................................................................... ..
31
APPENDIX...................... ................................... .......................................................................
89
vii
. LIST OF TABLES
Page
1
S e c t i o n s , s p e c i e s , a n d number o f s u b s p e c i e s o f
C h r y s o t h a m n i i ................. .................. ...........................................................
4
2
O c c u r r e n c e o f C h ry so th am n u s t a x a i n M o n tan a........................
29
3
C o m p a riso n o f C h ry so th a m n u s p a r r y j s s p . g l a n d u l o s u s
w i t h 3 m o r p h o l o g i c a l l y s i m i l a r C^ p a r r y ! t a x a ....................
33
/
38
4 '
D i a g n o s t i c c h a r a c t e r i s t i c s o f Montana C h r y s o th a m n u s . . .
5
L e g a l d e s c r i p t i o n s a n d r a n g e s i t e s f o r 7 C h ry s o th a m n d s .
p a r r y i s s p . g l a n d u l o s u s a n d s a m p le p l o t s i z e ......................
54
H a b i t a t d e s c r i p t i o n f o r 7 C h rv s o th a m n u s p a r r y i s s p .
g l a n d u l o s u s s a m p l i n g s i t e s .......... .............
56
pH m eans ( r a n g e ) o f s o i l s from 7 C h ry s o th a m n u s p a r r y i
s s p . g l a n d u l o s u s c o l l e c t i o n s i t e s ........................
58
S o i l e l e c t r i c a l c o n d u c t i v i t y (mmhos) means ( r a n g e ) o f
s o i l s fro m 7 C h ry s o th a m n u s p a r r y i s s p . g l a n d u l o s u s
c o l l e c t i o n s i t e s . . . ..........................................................................
59
P e r c e n t o f C h ry so th a m n u s p a r r y i s s p . g l a n d u l o s u s
p l a n t s b ro w se d a t 7 s a m p l in g s i t e s ............ ; . . . .......................
61
10
Range S i t e s o f C h ry so th a m n u s t a x a i n M o n t a n a . . . . . ..........
63
11
Range s i t e s a t w h ic h m u l t i p l e C h ry so th a m n u s t a x a w e re
c o l l e c t e d ........................................
71
Summary o f v e g e t a t i o n c h a r a c t e r i s t i c s m e a s u re d a t 7
C. p a r r y i s s p . g l a n d u l o s u s s a m p l i n g s i t e s .............................
90
H a b ita t n o t e s and l o c a t i o n s o f specim ens o f
C h ry s o th a m n u s n o t e d i n t h e R e s u l t s a n d D i s c u s s i o n
s e c t i o n ................................................ ; . . . ....................................................
92
6
7
8
9
12
13
v iii
LIST OF FIGURES
F igure
1
Page
I l l u s t r a t i o n s o f Chrysothamnus p a r r y i s sp .
g la n d u lo s u s ................................................................................ ..........
36
2
R e la tio n s h ip among Montana Cjl nauseosus s u b s p e c ie s .. . .
3
L o c a tio n s o f Chrysothamnus p a r r y i s s p . g la n d u lo su s
observed in M ontana................................................... ..................
43
4
L o catio n o f c o lle c tio n s o f Cl l l n i f o l i u s ..............................
44
5
L o catio n o f c o lle c tio n s and herbarium specim ens o f
Cl Ql s s p . a l b i c a u l i s .....................................................................
45
L o catio n o f c o lle c tio n s and herbarium specim ens o f
Cl Ql s s p . c o n s im ilis ......... ........................
46
L o catio n o f c o lle c tio n s and herbarium specim ens o f
Cl Ql s s p . g ra v e o le n s ......... ............................................................
47
L o catio n o f c o lle c tio n s and herbarium specim ens o f
Cl Ql s s p . n a u se o su s........................................... ............................
48
L o catio n o f c o lle c tio n s and herbarium specim ens of
Cl Vl s sp . I a n c e o la tu s ................................... ................................
49
L o c a tio n o f c o lle c tio n s and herbarium specim ens o f
Cl Vl s s p . v i s c i d i f l o r u s ........................ .......................................
50
L o catio n o f c o lle c tio n s o f Cl Vl s s p . v i s c i d i f l o r u s
v a r. s te n o p h y llu s ..................
51
L o catio n o f c o lle c tio n s o f Cl Vl s s p . v i s c i d i f l o r u s
v a r. v i s c i d i f l o r u s . ......... .......................................
52
6
7
8
9
10
11
12
40
ix
ABSTRACT
D u r in g t h e summers o f 1985 a n d 1986, a s t u d y w a s c o n d u c t e d t o
d e t e r m i n e w hat C h ry so th a m n u s t a x a o c c u r i n Montana a n d t o r e c o r d t h e i r
re s p e c tiv e d is tr ib u tio n s .
T axa f o u n d i n c l u d e d C^ l i n i f o l i u s . Cjl
n a u s e o s u s s s p . n a u s e o s u s , Cjl Hjl s s p . a l b i c a u l i s . C^ ru_ s s p . g r a v e o l e n s .
Cjl ru s s p . c o n s i m i l i s . C. v i s c i d i f l o r u s s s p . I a n c e o l a t u s . Cjl Vi s s p .
v i s c i d i f l o r u s v a r . s t e n o p h y l l u s . Ci Vi s s p . v i s c i d i f l o r u s v a r .
v i s c i d i f l o r u s , a n d t h e p r e v i o u s l y u n c h a r a c t e r i z e d , Ci p a r r y i s s p .
g la n d u lo su s.
The m o rp h o lo g y , h a b i t a t , a n d r e l a t i o n s h i p s o f t h e s e t a x a
w ere d i s c u s s e d a n d t h e i r d i s t r i b u t i o n s d e s c r i b e d .
I n 1 9 8 5 , a p r e v i o u s l y u n c h a r a c t e r i z e d t a x o n o f Ci p a r r y i was l o c a t e d
i n s o u t h w e s t e r n M on ta n a .
Ci Ei s s p . g l a n d u l o s u s d e r i v e d i t s name from
co n sp ic u o u s s ta lk e d g la n d s .
Two v a r i e t i e s o f t h i s t a x o n w ere o b s e r v e d
a t 2 o f 7 stu d y s i t e s .
The t y p i c a l f o r m i s nam ed Ci ^
ssp.
g la n d u lo su s v a r. g la n d u lo s u s .
The o t h e r v a r i e t y i s c o v e r e d w i t h
tom entum , w hich o b s c u r e s t h e . s t a l k e d g l a n d s a n d i s named Ci
ssp.
g la n d u lo s u s v a r . to m e n to su s.
A d d i t i o n a l p o p u l a t i o n s w e re l o c a t e d d u r i n g t h e summer o f 1 986 a n d
s i t e s w e re sa m p le d t o c h a r a c t e r i z e t h e t a x o n ’ s h a b i t a t .
A ll 7 s i t e s
sa m p le d w e re fo u n d t o h a v e n e a r - n e u t r a l , s a n d y loam t o loam s o i l s , a n d
s u p p o r t e d A r t e m i s i a t r i d e n t a t a / g r a s s l a n d c o m m u n it i e s . A s p e c t s , s l o p e s ,
e l e v a t i o n s , an d a n n u a l p r e c i p i t a t i o n a m o u n ts v a r i e d among s i t e s .
An
a d j a c e n t p a i r e d p l o t , l a c k i n g t h e t a x o n , was sa m pled a t e a c h s i t e i n a n
a t t e m p t t o a s c e r t a i n w hat f a c t o r ( s ) a c c o u n t f o r t h i s ta x o n ’ s
occurrence.
D i s t u r b a n c e a n d s o i l t y p e w e re t h e o n l y f a c t o r s fo u n d t h a t
l i k e l y i n f l u e n c e t h e o c c u r r e n c e a n d d i s t r i b u t i o n o f Ci ^ i s s p .
g l a n d u l o s u s . E v id e n c e i n d i c a t e d t h i s t a x o n i s s e v e r e l y bro w se d y e a r l y ,
w hich may be a f a c t o r l i m i t i n g t h e e x p a n s i o n o f t h i s t a x o n .
I
I
INTRODUCTION
The g e n u s
Chrysofchamnus
(ra b b itb ru sh )
th e w e s te rn U nited S t a t e s ,
is
w id e ly
d istrib u te d acro ss
from t h e G r e a t P l a i n s t o t h e P a c i f i c C o a s t .
I t i s e n d e m ic t o N o r th America a n d o c c u p i e s a v a r i e t y o f h a b i t a t s fro m
sea
lev e l
to
3300 m ( 1 0 ,9 0 0
ft)
(M cA rthur e t a l .
1979).
T h is g r e a t
g e o g r a p h i c a l d i s t r i b u t i o n i s r e f l e c t e d i n num erous d i v e r s e t a x a .
H a l l an d C le m e n ts (1923) o r g a n i z e d C h ry s o th a m n u s i n t o 4 S e c t i o n s ,
12
s p e c i e s , a n d 40 s u b s p e c i e s .
S i n c e t h e n L. C. A n d e rs o n h a s c o n t r i b u t e d
c o n sid e ra b le
C h ry s o th a m n u s a n d
k n o w led g e
of
re c e n tly
5
reco g n ized
S e c t i o n s , 16 s p e c i e s , a n d 41 s u b s p e c i e s (A n d e rso n 1 9 8 6 a ) .
U n til
th e
w ith in
present
th e
C h ry so th a m n u s
G e n e rally ,
n a tu ra l
e c o lo g ica l
in
Montana
resource
i m p o r ta n c e
has
gone
of
t a x o n o m ic
e sse n tia lly
d iv e rsity
un reco g n ized .
m a n a g e rs h a v e r e c o g n i z e d o n l y
2 tax a ,
C.
docum ented
in
v i s c i d i f l o r u s a n d C^ n a u s e o s u s .
As a
fo rag e
M o n ta n a .
so u rce,
u se fu ln e ss
T ra d itio n a lly
u n d e sira b le p la n ts ,
is
not
C h ry so th a m n u s
th o ro u g h ly
tax a
t h o u g h t t o be r e l a t i v e l y
have
been
c o n sid e re d
u n p a la ta b le and o ccu p an ts
o f l a n d s t h a t c o u ld o t h e r w i s e s u p p o r t more d e s i r a b l e f o r a g e .
However,
a Montana s t u d y ( E u s t a c e 1971) a n d a v a r i e t y o f r e p o r t s fro m e l s e w h e r e
(Cook a n d H a r r i s
K u f e ld e t a l .
1950,
Cook e t a l .
19 5 4 ,
1973, M cA rthur e t a l .
Sampson a n d J e s p e r s o n
1 974, Hanks e t a l .
196 3 ,
19 7 5 , M cA rthur
e t a l . 1 979) r e f u t e t h i s p o p u l a r c o n c e p t .
C o n sid erab le
uses
has
been
know ledge
g a in e d
of
C h ry so th a m n u s
re c e n tly
in
o th e r
tax o n o m y ,
w e stern
m anagem ent,
s ta te s.
and
T h is
2
i n f o r m a t i o n s u g g e s t s t h a t some C h ry so th a m n u s t a x a h a v e many u s e s (Weber
et a l.
The
1985).
lac k
of
in fo rm a tio n
re g a rd in g
th e
d iv e rsity
an d
v a lu e
of
C h ry s o th a m n u s i n Montana a n d t h e know le d g e g a i n e d on t h e i m p o r t a n c e o f
C h ry so th a m n u s
stu d y .
th eir
in
th e
w est
in sp ired
th e
o rig in a l
o b je c tiv e
of
th is
T h i s was t o d e t e r m i n e t h e t a x a o f C h ry so th a m n u s i n Montana a n d
re sp e c tiv e
d istrib u tio n s.
p re v io u sly
u n d escrib ed
becam e
c h a ra c te riz e
to
g lan d u lo su s.
taxon
th is
D u r in g
was
th e
lo ca ted .
ta x o n ,
now
course
of
Thus, a
know n
as
fie ld
second
(X
w o rk ,
a
o b je c tiv e
p arry i
ssp.
3
LITERATURE REVIEW
Phylogeny and C l a s s i f i c a t i o n o f Chrvsothamnus
The g e n u s C h ry s o th a m n u s i s a member o f one o f t h e l a r g e s t a n d m ost
advanced o f t h e f lo w e r in g p l a n t f a m i l i e s , t h e A s te ra c e a e (= C o m p o sita e ).
C h ry so th a m n u s,
and
its
b r i l l i a n t g o l d e n - y e l l o w f l o w e r , was e s t a b l i s h e d by N u t t a l l i n 1 840.
(X
p u m ilu s
is
nam ed
th e
fo r
ty p e
its
sp e c ie s
re la tio n s h ip
(H a ll
an d
to
C le m e n ts
Chrysocom a
1923).
H all
and
C le m e n ts (1923) d e s c r i b e d t h e h i s t o r y o f t h e g e n e r i c d e s i g n a t i o n .
C h r y s o th a m n u s . m eaning "g o ld e n w o o d " o r " g o l d e n b u s h " , b e l o n g s t o t h e
A s te re a e , th e A ster t r i b e .
H elia n th o id
or
H e lia n th o id
c h a ra c te rs
T h is t r i b e
p ro to -H e lia n th o id
co n sid e red
was m ost l i k e l y d e r i v e d fro m a
a n c e sto r
to
(C ro n q u ist
be r e l a t i v e l y
1 9 55).
p rim itiv e ,
The
present
i n some A s t e r e a e , i n c l u d e
s h ru b b y o r s u f f r u t e s c e n t h a b i t , l a r g e h e a d s ,
y e llo w
a
c o ro lla s,
C h ry so th a m n u s i s
H ap lo p ap p u s
and
and
re la te d
m ore
p a le a c e o u s
to S e c tio n
d ista n tly
pappus
1 9 5 5 )..
(C ro n q u ist
( S e c . ) Macronema o r E r i c a m e r i a o f
to
S o lid a g o
(A n d e rs o n
1964).
The
num erous s p e c i e s a n d s u b s p e c i e s o f t h e g e n u s C h ry s o th a m n u s a r e g ro u p e d
in to sev eral se ctio n s
( T a b le I ) .
A n d e rs o n a n d F i s h e r (1 9 7 0 ) d e v e l o p e d a s p e c i a l i z a t i o n i n d e x ( 0 - 1 0 ) ,
based
on
flo ra l
tax a
in
C h ry so th a m n u s.
sp e c ia liz e d .
a n a to m y ,
to
d e lin e a te
T axa
in
th e
s p e c ia liz a tio n
Sec.
N auseosi
of
th e
are
v a rio u s
th e
le a st
C^ p a r r y I (G ra y ) G reene s u b s p e c i e s r a n g e from 1 .2 t o 8 . 3
a n d Cl n a u s e o s u s ( P a l l a s ) B r i t t ,
s u b s p e c i e s from 2 . 3 t o 6 . 7 .
Cl p a r r y i
s s p . p a r r y i was d e t e r m i n e d t o be t h e m ost p r i m i t i v e t a x o n i n t h e g e n u s .
4
C ron q u ist
( 1 9 5 5 ) , r e v i e w i n g p h y lo g e n y i n
Compos i t a e ,
f e a t u r e s u c h a s few h e a d s , e a c h w i t h many f l o w e r s ,
c h a r a c t e r i s e x p r e s s e d by C^ p a r r v i .
su b sp ec ie s
of
C^_ n a u s e o s u s
i s p rim itiv e .
th at a
T h is
S u b s p e c i e s o f C^ p a r r v i and a few
c o n siste n tly
b u n d l e s t h a n o t h e r C h ry s o th a m n u s t a x a
su g g ested
have
m ore
o v a ria n
v a scu la r
(A n d e rs o n 1 9 7 0 ) , i n d i c a t i n g l e s s
s p e c i a l i z a t i o n t h a n o t h e r C h ry so th a m n u s t a x a
(A n d e rs o n 1 9 7 0 ) .
T a b l e I . S e c t i o n s , s p e c i e s , a n d n u m bers o f s u b s p e c i e s o f
Chry s o t h a m n i i a .
S e c tio n
S p e c ie s
C h ry s o th a m n u s
C^
C.
C.
C.
C.
C.
Number o f s u b s p e c i e s
a l b i d u s ( J o n e s e x G ra y ) G re e n e .
h u m ilis G reene.
I i n i f o l i u s G reene.
s p a t h u l a t u s L . C. A n d e r s o n .
g r e e n e i (G ray ) G re e n e .
v i s c i d i f l o r u s (H o o k .) N u t t .
G ram ini
C^ g r a m i n e u s H a l l .
C. e r e m o b iu s L .C . A n d e r s o n .
N auseosi
C.
C.
P u lc h e lli
C.
Cjl
C.
C.
P u n ctati
5
p a r r v i (G ray ) G r e e n e .
nauseosus ( P a lla s ) B r i t t .
12
22
v a s e y i (G ray ) G re e n e .
m o l e s t u s ( B l a k e ) L .C . A n d e rs o n .
d ep ressu s N u tt.
p u l c h e l l u s (G ray ) G r e e n e .
2
Cj. p a n i c u l a t u s (G ray) H a l l .
t e r e t i f o l i u s (D ur. & H i l g . ) H a l l & Clem.
a A n d e rso n 1986a
K a r y o t y p e s a n d b r e e d i n g s t r u c t u r e f u r t h e r i n d i c a t e t h a t Cjl p a r r y i i s
one o f t h e m ost p r i m i t i v e s p e c i e s o f t h e g e n u s ( A n d e rs o n 1 9 6 6 ) .
ssp.
p arry i
has
th e
lo n g est
chrom osomes
in
th e
genus,
th e
C2.
g rea test
h o m o g e n e ity i n chromosome l e n g t h , an d h a s chrom osomes w i t h e s s e n t i a l l y
5
m edian c e n t r o m e r e s (A n d e rs o n 1 9 6 9 ) .
A c c o r d in g t o H uziw ara (1 9 6 2 ) t h e s e
a re p rim itiv e c h a ra c te rs.
H all
& C le m e n ts
(1923)
th e o riz e d
th a t
s u b s p e c ie s and o th e r tax a in S e c . T y p ici
A nderson
1986a)
w ere
th e
le a st
m ost
Ci
(renam ed S e c .
e v o lu tio n a rily
v isc id iflo ru s
C h ry s o th a m n u s ;
advanced.
A nd e rso n
(1 9 6 6 ) p r e s e n t l y p r o p o s e s t h a t t h i s S e c t i o n c o n t a i n s t h e m o st a d v a n c e d
ta x a .
In
th e
C h r y s o th a m n u s ) ,
t o Ci Vi
Ci
v isc id iflo ru s
(H o o k .)
N u tt,
s p e c i a l i z a t i o n i n c r e a s e s from Ci Vi
s s p . I a n c e o l a t u s ( N u t t . ) H a l l & Clem, t o
(D. C. E a t . )
H a l l & Clem.
( A n d e rs o n 1 9 8 6 b ) .
Ci
co m p lex
(S ec.
ssp. v isc id iflo ru s
Ci Vi
s s p . pu b eru lu s
s s p . p u b e r u l u s was
Vi
p r o b a b l y d e r i v e d p h y l o g e n e t i c a l l y from Ci Vi s s p . I a n c e o l a t u s (A n d e rso n
1986b).
n a rr o w
C. p a n i c u l a t u s (G ray) H a l l , w i t h i t s s p e c i a l i z e d k a r y o t y p e a n d
e c o lo g ica l
req u irem e n ts,
is
c o n s i d e r e d t o be t h e m ost a d v a n c e d
s p e c ie s i n th e C h ry so th a m n ii.
M o rp h o logy
A ll
th e
tax a
of
C h ry so th a m n u s a r e
There a r e s e v e r a l t r u n k s ,
sh ru b s o r
s u ffr u te scent
coming from a s i n g l e b a s e , w h ic h a r e c o v e r e d
w i t h l o o s e , f i b r o u s , brown b a r k w h ic h p e e l s o f f i n
a r e a lw a y s a l t e r n a t e l y
b lad e s a re
Ci
ty p ic a lly
te re tifo liu s
strip s.
a rra n g e d and t h e m arg in s a r e
& H ilg .)
H all
& Clem.
The
The l e a v e s
e n tire .
f l a t ; how ever, t e r e t e l e a v e s o c c u r i n
(D u r.
shrubs.
The l e a f
I sp e cie s,
num ber
of
v e in s
d e p e n d s l a r g e l y upon t h e w i d t h o f l e a f ( H a l l an d C le m e n ts 1 9 2 3 ) .
F lo ra l
heads
of
p a n i c l e s , a n d cym es.
C h ry so th am n u s a r e
T his la te r: ty p e i s
arran g ed
in
racem es,
t h e most common i n
sp ik e s,
the genus
6
o v e ra ll.
The ra c e m o s e t y p e i s
c h a ra c te ristic s,
a rra n g e m e n t,
such a s
and
c h a r a c t e r i s t i c o f Cl p a r r y i .
w h eth er
p h y lla ry
tip
th e
p h y lla rie s are
shapes,
In v o lu c re
k e e le d ,
c a n be h e l p f u l
in
p h y lla ry
id e n tify in g
tax a .
The h e a d s o f C h ry so th a m n u s a r e
present
in
o nly
2 p o p u la tio n s
s p a t h u l a t u s L . C. A n d e r s o n .
but
th e
to ta l
c h a ra c te rs
len g th
of
ty p ic a lly
(so u th e rn
d isc o id .
and
Ray f l o w e r s a r e
n o rth e rn
ranges)
of
(X
The s h a p e o f t h e c o r o l l a i s n o t d i a g n o s t i c
co ro lla
and c o r o l l a
( H a l l a n d C le m e n ts 1 9 2 3 ) .
lo b es a r e
d istin g u ish in g
The number o f f l o w e r s p e r h e a d
i s a l m o s t c o n s t a n t l y 5 e x c e p t i n Cl p a r r y i and (X p y r a m i d a t u s R o b in s o n
a n d G reenm an, w hich h a v e a g r e a t e r nu m b er.
D i f f e r e n t en v iro n m en ts and
seasonal
of
v a ria tio n s
v isc id iflo ru s
resu lt
su b sp ec ie s
season fre q u e n tly
in
v a ria b ility
(A n d e rs o n 1 9 6 4 ) .
flo ra l
fea tu re s
S am ples t a k e n e a r l y
a v e r a g e more f l o w e r s ' p e r h e a d t h a n
of
Cl
in th e
s a m p le s o b t a i n e d
fro m same p l a n t l a t e r i n t h e s e a s o n .
The p a p p u s
C le m e n ts 1 9 2 3 )•
d isp e rsa l,
The r a t i o
and
is
n e a rly
u n ifo rm
The p a p p u s i s
p o ssib ly
b e tw e e n l e n g t h
in a l l
C h ry so th a m n u s t a x a
and
i m p o r t a n t f o r a c h e n e o r i e n t a t i o n , w ind
m o istu re a c c u m u la tio n
of
(H all
(S te v e n s
s t y l e appendage and
et a l.
s ti g m a t i c
1986).
p o rtio n
n o t c o n s ta n t i n any I s p e c ie s o r p e rh a p s even i n any I s u b s p e c ie s .
is
The
o v e r a l l r a t i o i s h i g h e r f o r some s p e c i e s t h a n f o r o t h e r s .
Stamens a r e d ia g n o s tic only in (X a lb id u s (Jones ex Gray) G reene, in
which they a r e o b tu se and s h o r te r th an in o th e r ta x a .
In o th e r s , th e
t i p s a r e a c u te (H all and Clem ents 1923) .
The achene i s o f 2 form s, e i t h e r : I) o b s c u re ly 5-a n g le d or t e r e t e , 5
n e rv e d , and u s u a lly covered w ith dense pubescence; o r 2) n e a r ly t e r e t e
7
or s lig h tly
C le m e n ts
f l a t t e n e d , 10 s t r i a t e ,
1923).
fo u n d i n
The f o r m e r i s
and e s s e n t i a l l y
th e
g lab ro u s
common t y p e ;
th e
( H a l l an d
la tte r
ty p e
is
v a s e y i (G ra y ) G re e n e , a n d CL g r a m i n e u s H a l l .
Pubescence
nauseosus,
can
m ost
a
u sefu l
w id e
c h a ra c te ristic .
ran g in g
shows much i n t r a s p e c i f i c v a r i a t i o n w i t h r e s p e c t t o t h i s f e a t u r e
(Hanks
I n CL n a u s e o s u s . 2 g r o u p s ,
sp ecies
th e
in
CL
genus,
1975).
com plex a n d
d ia g n o stic
th e
et a l.
th e
be
g ray and t h e g re e n , a r e
d i s t i n g u i s h e d by t h e p r e s e n c e o r a b s e n c e o f v e s t i t u r e on t h e i n v o l u c r e
and th e ty p e o f v e s t i t u r e .
T h i s c h a r a c t e r c a n v a r y a n d s h o u ld be u s e d
in
c o m b i n a t i o n w i t h t h e c o l o r o f tomentum on t h e s te m .
on
th e
in v o lu c re s
gen eral
p lan t
ten d en cy
th an
th a t
in
th e
gray
t o w a rd a
group
is
o nly
g r e a t e r am ount
w h ic h o c c u r s
in
th e
of
green
an
The v e s t i t u r e
e x p re ssio n
pubescence
group
(H a ll
in
of
th e
and
th e
w h o le
C le m e n ts
1923).
)
Anatomy
A n d e rso n
(1 9 7 0 )
d e v e lo p m e n ta l
p ro v id e s a
a n a to m y
of
thorough
C h ry so th a m n u s
C h ry so th a m n u s p h y l l a r i e s a n d f l o w e r s .
c h a ra c te riz e d
by
p resen ce
t r i c h o m e s on t h e c o r o l l a
P u c h e lli lac k
th e
c o ro lla
sh o rter
and
se c re to ry
tu b e s.
m ore
(A n d e rs o n 1 9 7 0 ) .
account
of
tu b e.
and
of
of
th e
v ascu lar
th e
h isto lo g y
and
of
Taxa i n t h e S e c . N a u s e o s i a r e
se c re to ry
c a n als
Those t a x a i n S e c s .
and
u su a lly
have
C h ry s o th a m n u s a n d
c a n a l s a n d o n l y a few t a x a h a v e t r i c h o m e s on
In Sec.
c la v a te
P u n c ta ti, g la n d u la r h a ir s a re
th an
th e
g lan d u la r
h a irs
of
common b u t
Sec.
N auseosi
8
The v e n a t i o n i n C h ry a o th a m n u s f l o w e r s i s r e l a t i v e l y a d v a n c e d f o r t h e
C o m p o s ita e (A n d e rs o n 1 9 7 0 ) .
co ro lla ,
2
in
th e
D is k f l o w e r s t y p i c a l l y h a v e 5 v e i n s i n t h e
sty le ,
and
I
in
each
sta m e n .
C h ry so th a m n u s d o e s n o t c o n t a i n v a s c u l a t u r e .
The
V a sc u la r t r a c e
i n th e b r a c t s v a ry a t th e s p e c ie s and s u b s p e c ie s l e v e l s .
G reene,
ssp.
Cjl n a u s e o s u s
g rav e o len s
v isc id iflo ru s
ssp.
(N u tt.)
ssp.
a lb ic a u lis
P ip e r
in
(N u tt.)
have a
v isc id iflo ru s.
p u b e ru lus have 3 t r a c e s
b ra c ts.
b u t,
ssp.
C.
th e
H all
ssp.
& Clem,
p a tte rn s
and Cju ru_
p a tte rn .
I a n c e o l a t u s . and
o u te r b r a c t s and
in
C^ l i n i f o l j u s
3 tra c e/b rac t
v.
pappus
I in
th e
Cu
Cu
Vll
in n er
I n m ost t a x a o f C l p a r r y i 3 v a s c u l a r t r a c e s e n t e r e a c h b r a c t ,
i n Cl Pl
ssp.
an d Cl El s s p .
fre q u e n tly
have
p a r r y i . Cl ^ l s s p . n e v a d e n s i s
asper
5
( G reen e) H all
tra c e s/b ra c t.
(G ray ) H a l l & C l e m . ,
& C le m ., th e l a r g e r
In
Cl p a r r y i
th ese
(o u te r) b ra c ts
la te ra l
tra c e s
(A n d e rs o n
1971).
u s u a lly e x te n d h a l f th e le n g t h o f th e p h y l l a r i e s .
Em bryology
W ith in a h e a d ,
F lo w e rs
flo ra l
d e v e lo p
m a tu ra tio n
a c ro p e ta lly
is
on
sy n ch ro n ized
th e
re c e p ta c le
but
o fte n
m ic r o s p o r o g e n e s is and m e g a sp o ro g en e sis o c c u r s im u lta n e o u s ly r e g a r d l e s s
of
d iffe re n c e s
sp e cie s
is
in
Cl l i n i f o l i u s ,
embryo s a c s w h i l e
th e
in d iv id u a l
o v u le .
C h ry so th a m n u s
in
flo w er
w h ic h
th e youngest
E m bryo
sac
(A nderson
o ld er
flo w er i s
d e v e lo p m e n t
1983)
siz e .
and
The
flo w ers
still
is
embryo
o fte n
synchronous
have
m atu re
u n d erg o in g m e io s is i n
th e
sac
le a st
P o ly g o n u m - t y p e
siz e
v a rie s
in
w id e ly .
9
T y p i c a l l y m a t u r e s a c s a r e l o n g a n d n a r r o w (A n d e rs o n 1 9 7 1 ) .
P o ly p lo id y
h a s no s i g n i f i c a n t b e a r i n g on embryo s a c s i z e .
Embryogeny i s t h e A s t e r a d t y p e (A n d e rs o n 1 9 7 1 ) .
is
n u c le a r a t
endosperm .
first
M e io sis
m egaspores
u su a lly
a n a tro p o u s,
re s u lts
produce
u n ite g m ic ,
C h ry so th a m n u s a r e
flo ra l
th en w all
p rim o rd ia
Endosperm f o r m a t i o n
fo rm a tio n p r o g re s s e s to
in
4 c e llu la r
th e
embryo s a c .
and
m egaspores.
d iffe re n tia te
The
c h a la z a l
C h ry s o th a m n u s o v u l e s a r e
te n u in u c e lla r.
te tra sp o ra n g ia te .
fo rm a c e l l u l a r
The
a n th e rs
in
I n m ost s p e c i e s o f C h r y s o th a m n u s .
th ro u g h
sequence
of
c o ro lla ,
sta m e n s,
p a p p u s , an d p i s t i l .
Chromosome R e l a t i o n s h i p s
The b a s i c chromosome number o f C h ry so th a m n u s i s n = 9 .
modal
num ber
(S o lb rig
in
et
th e
a l.
a u to p o ly p lo id s
trib e
at
1 9 6 4 ).
both
th e
sp e c ific
C h ry so th a m n u s
(A nderson
1986b).
ssp.
p u b eru lu s
and
v isc id iflo ru s
com m only
have
g e n e ra lly
flo w er
e x te rn a l
1950).
3x ,
(A nderson
a d a p t e d t o w a rm e r,
A v a ila b ility
2x ,
fa c to r
and
le a v e s,
fru it
th e
and
is
site s
th an
le v e l
m o stly
to
C.
s s p . I a n c e o l a t u s a n d Cjl y \
6x o c c u r r i n g
th an d i p l o i d s .
larg e r
are
re stric te d
CLl v i s c i d i f l o r u s
la te r
u n e x p lo ite d
fav o rin g
5x ,
1986b).
u su a lly d r ie r
th ic k e r
of
4x,
g e n e ric
p o ly p lo id s
P o ly p lo id y
v i s c i d i f l o r u s , w i t h 2 x a n d 4x fo u n d i n (Ll
and
T his i s th e
in
C^ Vjl s s p .
p o ly p lo id s
A u to p o ly p lo id s
flo w e rs ,
larg e r
d ip lo id s
(S te b b in s
e n v iro n m e n ta l
e sta b lish m e n t
resources
of
are
is
p o ly p lo id y
fru its
th e
and
1947).
c h ie f
(S te b b in s
The f r e q u e n c y o f p o l y p l o i d s p e c i e s o f a n g i o s p e r m s a p p e a r s t o
10
in cre ase
from
s o u th
to
n o r th and
fro m
low
to
hig h
a ltitu d e s
(G rant
1971).
R ep ro d u c tiv e H a b its
P re d o m in a n tly ,
th e
sp e cie s
w ith
C h ry s o th a m n u s
cro ssb reed
own p o l l e n )
(A n d e rso n
a
p red o m in an c e
S e le c tio n
pressure
e n v iro n m e n ts
1980 b ) .
occurs
in
for
w h ich
of
ev ery
work
sp e cie s of
by
M cA rthur
se lf-p o llin a tio n
in b re e d in g
p ro b ab ly
C h ry so th a m n u s
but
on o c c a s i o n
Autogamy ( f l o w e r p o l l i n a t e d
in
ty p ic a lly
S e l f - f e r t i l i z a t i o n en co urages th e
co n tin u e d
C hry so th a m n u s
et
(X
re su lts
tax a
o f sympat r i e
a
se lf-fe rtiliz in g ,
C h r o m a to g r a p h ic
su g g e sts
1973).
(A n d e rso n
fre q u e n tly
1980b).
is
a l.
(1978a)
v isc id iflo ru s.
fro m
d ro u g h ty
in h ab it
(A n d e rso n
g e n e tic
se p ara tio n
s u b s p e c i e s a n d t h e f o r m a t i o n o f num erous l o c a l b i o t y p e s i n
su b sp ec ie s
(S te b b in s
1957).
The
success
of
C h r y s o th a m n u s
at
p r o d u c i n g new f o rm s a n d i n v a d i n g new e n v i r o n m e n t s i s a t t r i b u t e d t o t h e
a b ility
to
o ccasio n ally
gro u p , p rim itiv e
tax a
o u tcro ss
are
(A n d e rs o n 1 9 6 6 ) .
o fte n
o u tb reed ers
G e n e ra lly w ith in a
w hereas
sp e c ia liz e d
taxa
a r e i n b r e e d e r s (S te b b in s 1957).
Achene
fo rm a tio n
A pom ixis i s
in
known h y b r i d s
1980c).
Ci
in te rg e n e ric
a lb ic a u lis
C h ry so th a m n u s
n o t known t o o c c u r i n
H y b rid iz a tio n
A ll
in
occurs
in
have
b o la n d e ri,
h y b rid
an
genus
u n certain
b e tw e e n
fro m
C h ry so th a m n u s t a x a
C h ry so th a m n u s
th e
re su lts
in fre q u e n tly
in v o lv e d
tax o n ,
H ap lo p ap p u s
(A n d e rso n a n d R e v e a l 19 6 6 ) .
Ci
fe rtiliz a tio n .
(A n d e rs o n 1 9 7 1 ) .
1970 ) .
(A n d e rs o n
nauseosus
(A n d e rso n
has been r e d e f in e d a s an
macronema
and
C.
n.
ssp.
A d d itio n a l ty p e s o f h y b rid s a r e
11
known.
1966 ,
F o r e x a m p le , i n t e r s p e c i f i c h y b r i d s h a v e b e e n r e p o r t e d
A nderson
1983,
Nevada r e p r e s e n t
fo und
A n d e rs o n
th e f i r s t
in
th e
genus
fo u n d a t
th e
g eo g rap h ical
1984)
n a tu ra lly
(A n d e rs o n
or
Cjl Vi
ssp.
from
Ash
in te rse c tio n a l
The known h y b r i d s
e c o lo g ic a l
edge
Meadows,
o f one
h y b rid s
stu d ie d
or
b o th
of
(L e tte r
a lb ic a u lis
( G reene)
ssp.
H all
in te rg ra d e s
from
in te rg ra d e s
& C le m .,
h o lo le u c u s
Ci
(G ray)
( A n d e rs o n 1 9 8 6 c ) .
n a u s e o s u s a n d Ci
th e
I n t e r g r a d a t i o n o c c u r s b e tw e e n some o f
L.
C.
e x te n siv e ly
A nderson
w ith
5 /2 0 /8 6 ).
Cjl y\
ssp.
Ci
ssp.
ru
n a u s e o s u s s e e m i n g ly i n t e r g r a d e s o n l y w i t h Ci ru s s p . a l b i c a u l i s .
ssp.
w e re
v i s c i d i f l o r u s a n d some s u b s p e c i e s o f Cjl n a u s e o s u s .
Ia n c e o la tu s
v isc id iflo ru s
h y b rid s
o c c u rrin g
1973).
p a r e n t s ' r a n g e s (A n d e rs o n 1 9 7 3 ) .
th e su b sp ecies o f
and
(A n d e rso n
L ittle
o c c asio n ally
Qi
ssp.
H all
w ith
Ci
m
ssp.
Ci Iii
c o n sim ilis
n a u s e o s u s . an d p a r t i c u l a r l y
& C le m ,
w here
th e ir
ranges
Ci
Qi
o v e rla p
o r no i n t e g r a t i o n o c c u r s b e tw e e n Ci n .
ssp.
s s p . g r a v e o l e n s e v e n t h o u g h t h e y a r e sympat r i e o v e r
Qi
a b o u t h a l f t h e i r r e s p e c t i v e r a n g e s (A n d e rs o n 1986 c ) .
D i s t r i b u t i o n and S i t e C h a r a c t e r i s ti c s
C h ry so th a m n u s
o ccu rs
m o u n ta in s from B r i t i s h
a n d B a ja C a l i f o r n i a .
in
e le v a tio n
open
p la in s,
v a lle y s,
Colum bia an d S a s k a tc h e w a n ,
I t r a n g e s from s e a l e v e l
( H a l l a n d C le m e n ts
a n d W elch 1986 ) .
R e g io n
on
fo o th ills,
s o u th
e c o lo g ic a l a m p litu d e ;
1979a).
1 9 2 3 , M cA rthur e t ' a l .
Most
to w est Texas
t o 3 ,3 0 0 m ( 1 0 ,9 0 0 f t )
C h ry so th a m n u s i s m ost a b u n d a n t i n t h e
(M cA rthur e t a l .
1 9 7 4 , M cA rthur
In te rm o u n tain
C h ry so th a m n u s t a x a
th ey e x p re s s th e a b i l i t y
an d
have
great
t o i n h a b i t a w id e r a n g e
12
of e d a p h ic c o n d it io n s , l a t i t u d e s , and e le v a ti o n s
The
m any
su b sp e c ie s
and
e c o ty p e s
(A n d e rs o n 1 9 8 1 ) .
of
(%
nauseosus
r a b b i t b r u s h ) i n h a b i t a w id e v a r i e t y o f s o i l s a n d s i t e s .
heavy c l a y s and a l k a l i s o i l s i n s a l t y lo w lan d s
(ru b b er
S o ils in clu d e
(Hanks e t a l . 1975) a n d
sandy, g r a v e l l y , and c la y e y , n e u tr a l t o a c id s o i l s a t h ig h e r e le v a tio n s
(Hanks
et
a l.
E le v a tio n s
1975,
range
P in u s e d u l i s a l.
1978b).
fche
M cA rthur
from
150
to
et
a l.
1 978b,
M cA rthur
2750 m (5 0 0 -9 0 0 0
ft)
et a l.
1 979a).
i n A^ t r i d e n t a t a ,
J u n i p e r u s s p p . , an d P i n u s p o n d e r o s a z o n e s
(M cA rthur e t
Some t y p e s o f (X n a u s e o s u s a r e t h o u g h t t o be s u b c li m a x i n
trid e n ta ta
is
d i s t u r b e d by f i r e , t r a m p l i n g , o r w a s h o u t ( H a l l a n d G oodspeed 1 9 1 9 ) .
C.
—
trid e n ta ta
s s P* g t b i c a u l i s ,
clim a x .
Ci
th ey
re su lt
s s p . h o l o l e u c u s . Ci
Hi
s s p . c o n s i m i l i s , a n d Ci
O ften
!I l
ssp.
a lb ic a u lis
Ai
s s p . n a u s e o s u s . C. n .
Iii
ru s s p . g r a v e o l e n s a r e more w i d e s p r e a d t h a n t h e
o t h e r s u b s p e c i e s o f C. n a u s e o s u s (Weber e t a l .
—
if
(w h ite
rubber
1985).
rab b itb ru sh ),
a
fo o th ill
o c c u r s a t e l e v a t i o n s b e tw e e n 610 and 2135 m ( 2 0 0 0 -7 0 0 0 f t )
a l.
1974, A n d e rs o n 1986c) fro m B r i t i s h
1 979a).
stric ta ,
I n U ta h ,
Ai
p o n d ero sa
Ci
trid e n ta ta ,
c o m m u n ities,
Qi
ssp. a lb ic a u lis is
P in u s
1310
to
e d u lis
2290
(M cA rthur e t
Colum bia a n d M o n ta n a , s o u t h t o
New M e x ic o , C o l o r a d o , U ta h , N evada, an d e a s t e r n C a l i f o r n i a
a l.
ty p e ,
-
(M cA rthur e t
fo u n d i n D i s t i c h l i s
Ju n ip e ru s
m (4 3 0 0 -7 5 0 0
s p p .,
ft)
in
and
P jn u s
e le v a tio n
(W elsh 1 9 8 3 ) .
Ci
Qi
ssp.
g rav eo len s,
(g reen ru b b er r a b b itb ru s h )
e a s te rn h a l f of th e s p e c ie s ra n g e , i s
915 t o
a l.
1830 m ( 3 0 0 0 -6 0 0 0 f t )
1979a).
It
o c c u rrin g in th e
fo u n d i n f o o t h i l l s a n d v a l l e y s ,
from N o r th D akota
to
Idaho
(M cA rthur e t
i s a com ponent o f t h e w e s t e r n f o o t h i l l s o f t h e B la c k
13
H ills
P in e F o r e s t p l a n t a s s o c i a t i o n ,
p la n t
a sso c ia tio n
o ccu rs
in
d efin ed
th e
B lack
by K u c h l e r ( 1 9 6 4 ) .
H ills
of
S o u th
T h is
D akota
and
Wyoming.
In
U tah ,
s s P-
in h a b its
sa lin e
str ic ta
-
P in u 3
meadows,
S p o ro b o lu s
trid e n ta ta ,
rip a ria n
-
(Welsh
1983)»
In
and
A trip le x
Q uercus.
c o m m u n itie s
(th re a d le a f
zones,
a iro id e s.
C ercocarpus
p onderosa
e le v a tio n
c o n sim ilis
P in u s
from
1280
Nevada,
ra b b itb ru sh )
terrac e s
in
D istic h lis
c o n fe rtifo lia .
e d u lis
-
Ju n ip e r
A rtem isia
s p p . , and
3000 m ( 4200-9800
to
it
rubber
o c c u r s on s i t e s
ft)
in
re c e iv in g a s
l i t t l e a s 250 mm (10 i n ) p r e c i p i t a t i o n a n n u a l l y ( R o s e n t r e t e r 1 9 8 6 ) .
v isc id iflo ru s
d istrib u te d
(lo w
has g re a t e c o lo g ica l
not v ery s a l t t o le r a n t
N o rth
is
one
of
th e
m ost
w id e ly
s h r u b s on r a n g e l a n d s t h r o u g h o u t w e s t e r n N o r th A m e ric a .
v isc id iflo ru s
p lain s
ra b b itb ru sh )
and
m o u n tain s
D ak o ta,
so u th
ft)
v isc id iflo ru s
(M cA rthur e t a l .
on
poor
to
so ils
(A n d e rs o n 1986 b ) ,
1 978a).
fro m
New Mexico a n d
I t o c c u p ie s o p en , dry
B ritish
e a ste rn
but i s
C o lu m b ia ,
C a lifo rn ia
M o n ta n a ,
(H a ll
an d
v i s c i d i f l o r u s g ro w s a t e l e v a t i o n s o f 790 t o 3350 m
C le m e n ts 1 9 2 3 ) •
( 2 6 0 0 - 1 1 ,0 0 0
a m p litu d e
C.
(M cA rthur e t a l .
in c lu d e
1978a).
y e llo w b ru sh ,
O t h e r common names o f
y e llo w
rab b itb ru sh ,
C.
y e llo w sa g e ,
r a b b i t s a g e , s t i c k y - l e a v e d r a b b i t b r u s h , D o u g la s r a b b i t b r u s h , a n d s t i c k y flo w ere d I ris h m a n .
The
ty p es,
su b sp ecies o f
Cl v i s c i d i f l o r u s a r e
g la b ro u s and p u b e sc e n t.
ra b b itb ru sh ),
C o lu m b ia
and
a
pubescent
n o rth e rn
Cl Vl s s p .
ty p e,
New
ranges
M e x ic o
sp lit
in to
2 m o rp h o lo g ic a l
Ian c e o la tu s
b e tw e e n
(A nderson
( m o u n ta in low
so u th e rn
1 986b).
B ritish
In
th e
I n t e r m o u n t a i n R e g io n i t o c c u r s a t mid a n d h i g h e l e v a t i o n s , fro m 1525 t o
14
3200 m ( 5 0 0 0 - 1 0 ,5 0 0 f t )
(M cA rthur e t a l .
Ia n c e o la tu s
in h a b its
A rtem isia
tre m u lo id es,
Pseudots u s a
trld e n ta ta .
p u b e ru lu s
(h a iry
C o lo rad o ,
east
e le v a tio n s.
It
to
is
In U tah,
P in u s
low
abundant
on
and
dry
P icea -
Pop u l u s
A b i e s , an d
A n o th e r p u b e s c e n t t y p e , C. v .
ra b b itb ru sh )
C a lifo rn ia ,
ssp.
po n d ero sa.
m e n z e s ii. P in u s c o n to r t a ,
a l p i n e , meadow c o m m u n itie s (W elsh 1 9 8 3 ) .
ssp.
1 978a).
o ccu rs
in
h ills
b e tw e e n
B ritish
and
Montana
C o lu m b ia
in
dry
and
at
lo w
s t r e a m ways o f
w e s t e r n Nevada ( H a l l a n d C le m e n ts 1923)♦
A g la b ro u s
ra b b itb ru sh )
is
19 8 6 b ) a n d i s
ssp.
ty p e ,
g row s a t
Vi
ssp.
of
1981).
Ci v .
N evada,
so il
A rtem isia
Q uercus,
tax a
o ccu rs
in
n o rth e rn
so u th e rn
te x tu re s
A b ies
and
P in u s
c o n c o lo r.
C. I i n i f o l i u s
C a lifo rn ia
has
c o lle cte d
sin c e
and
Nevada.
M ontana,
on a
(M cA rthur
n o rth e rn
Vi
244 t o 3965 m ( 8 0 0 - 1 3 , 0 0 0 f t )
pHs
an d
w estern
Ci
It
e d u lis
P in u s
-
et
a l.
Ju n ip e ru s
ponderosa.
Ci ^ l i n i f o l i u s
1978a,
A nderson
and
s p p .,
C ercocarpus-
Pop u l u s
tre m u lo id es
(W elsh 1 9 8 3 ) .
is a ta ll,
w as
r o b u s t s h ru b up t o
o rig in a lly
v i s c i d i f l o r u s by H a l l a n d C le m e n ts
e sta b lish e d
by M cA rthur e t a l .
lin ifo liu s.
(A n d e rs o n
A rizo n a.
(sp re a d in g ra b b itb ru s h )
h e ig h t.
s u b s p e c i e s o f Ci
(1 9 6 4 )
genus
lo w
ssp. v is c id if lo r u s i s p resen t in A trip le x c o n f e r ti f o l i a .
trld e n ta ta ,
m in
th e
W ashington,
c o m m u n itie s b e tw e e n 1460-2900 m ( 4 8 0 0 -9 5 0 0 f t )
2 .4
in
(stic k y le a f
t h e m ost common s u b s p e c i e s o f Ci v i s c i d i f l o r u s .
low and h i g h e l e v a t i o n s ,
v a rie ty
v isc id iflo ru s
I o f t h e m o st w i d e s p r e a d
v isc id iflo ru s
n o rth w e ste rn
C^
it
as a
sp e cie s.
d e scrib ed
(1923).
as
a
A n d e rso n
C hrom atographic
d a ta
( 1 9 7 8 a ) s u p p o r t t h e s p e c i e s s t a t u s o f Ci
I t o c c u r s i n Wyoming, C o l o r a d o , M b n ta n a , New M exico, U ta h ,
In
U tah
it
is. found
on
stre a m
banks
and
terrac e s,
15
irrig a tio n
c a n a ls,
seeps
e l e v a t i o n s r a n g i n g fro m
and
sp rin g s
in
1130 t o 2535 m
rip a ria n
c o m m u n ities
( 3 7 0 0 -8 3 0 0 f t )
at
(W elsh 1 9 8 3 ) .
A n d e rs o n (1964) r e p o r t s t h a t CL l i n i f o l i u s o f t e n o c c u r s i n s a n d y s o i l s .
CL. g r e e n e i
( G ray)
v isc id iflo ru s
ssp.
v is c id if lo ru s
rab b itb ru sh )
and
in
h illsid e s,
(5 5 7 0 -8 5 2 5
ft)
in
p arry i
and
U tah,
ra b b itb ru sh )
G ray
1979a).
It
( V asey
v a lle y s
Wyoming,
N evada,
th e w est
U tah
g ro w s
of
low
v a lle y s,
(H a ll
on
1700
New M e x ic o ,
m o u n ta in a n d
and
p lain s,
to
and
2600
m
A rizo n a
(M cA rthur
et a l.
in v ad es a l k a l i n e a r e a s .
of th e
c e n tra l C a lifo rn ia
1979a).
ty p e ,
is
C^ a l b i d u s
It
(a lk a li
i s a h a lo p h y te
I t i s m ost commonly fo u n d on
G r e a t S a l t Lake D e s e r t ,
(M cA rthur e t a l .
fo o th ill
C a l i f o r n i a and so u th to
i s th e o n ly taxon w ith w h ite flo w e rs .
sid e
(n arro w leaf
and
e le v a tio n s
C o lo rad o ,
ra b b itb ru sh ), a
A r iz o n a
and s u c c e s s f u l l y
v*_ s s p .
o c c u r s on p l a i n s ,
fo u n d fro m Wyoming t o w e s t e r n N ev a d a , w e s t t o
rab b itb ru sh )
(X
Cjl
1979a).
(P arry
New Mexico a n d
1980a)
ra b b itb ru sh )
at
re se m b le s
(c o n sid e re d
A nderson
New M ex ico ,
Cjl v a s e y i i
m o u n ta in s,
(M cA rthur e t a l .
C.
ste n o p h y llu s;
C o lo rad o ,
1923) •
(g reen
ste n o p h y llu s
(M cA rthur e t a l .
fo o th ills
C le m e n ts
v ar.
G reene
19 7 9 a ) .
th ro u g h
Nevada
to
<L d e p r e s s u s N u t t ,
east
(d w arf
r a b b i t b r u s h ) grow s on d r y p l a i n s , h i l l s , a n d r o c k y m o u n ta in s l o p e s f ro m
1000 t o
210 0 m ( 3 3 0 0 -6 9 0 0
ft)
in w e s te r n C o lo rad o ,
N evada, a n d s o u t h e a s t e r n C a l i f o r n i a
New M exico,
( H a l l a n d C le m e n ts 1 9 2 3 ) .
U ta h ,
16
L ife H isto ry
C h ry s o th a m n u s
1974 b ,
is
D a ubenm ire
a
re la tiv e ly
1975).
It
sh o rt
liv e d
reproduces
shrub
p rim a rily
(Young a n d
by
seed.
E vans
In itia l
e s t a b l i s h m e n t o f (X n a u s e o s u s by s e e d i s f a i r t o g o o d .
S tev en s e t a l .
(1986)
a lb ic a u lis
rep o rted
a
s ta rtlin g
in cre ase
of
(X
m
n a t u r a l s p r e a d fro m 10 t o 622 p l a n t s i n 9 y e a r s .
th e
effect
of
th e
pappus
on
g e rm in a tio n
e s t a b l i s h m e n t was more s u c c e s s f u l w i t h
ssp.
I n a s t u d y e x a m i n in g
and
u n c le a n e d
and h a l f b u rie d th a n o t h e r p o s i t i o n s and d e p th s
by
su rv iv a l,
seed
seed lin g
p lan ted
(S te v e n s e t a l .
u p rig h t
1986).
C. n . s s p . a l b i c a u l i s s e e d l i n g s u c c e s s was a u g m e n te d by s e e d p l a c e m e n t
and p o s i t i o n i n g .
In t h e f i e l d t h e pappus a c t s a s a p a ra c h u te , th e re b y
h e l p i n g t o p o s i t i o n t h e s e e d ( s c a r down) a n d a c t s a s a n a n c h o r .
Once C h ry so th a m n u s i s e s t a b l i s h e d , g r o w th i s r a p i d i f c o n d i t i o n s a r e
fav o ra b le
(M cA rthur
grow s v i g o r o u s l y
et
due
to
S e e d l i n g s a s young a s
e x tr e m e
drought
a l.
1974).
basal
When t r a n s p l a n t e d ,
sp ro u tin g
(H a ll
and
C hry so th a m n u s
C le m e n ts
3 m onths h a v e b e e n fo u n d t o r e s p r o u t
(McKell
and
C h ilco te
1957).
S c h la tte re r
1923).
fo llo w in g
and
T isd ale
(1 9 6 9 ) s u g g e s t e d t h a t (X n a u s e o s u s . c o n t a i n s f e w e r i n h i b i t o r y s u b s t a n c e s
a n d l e a c h a t e s t h a n o t h e r common a r i d z o n e s p e c i e s .
g e rm in a tio n
and
in c re a se d
P se u d o ro e g n e ria s p ic a ta
1 985)
under
th e
g ro w th
of
They s u g g e s t e d t h a t
E ly m u s
e ly m o id e s
and
( n o m e n c l a t u r e a c c o r d i n g t o B a r k w o r th an d Dewey
in flu e n c e
of
C h ry s o th a m n u s m ig h t
be
due
to
g ro w th
s t i m u l a t o r y s u b s t a n c e s i n C h ry s o th a m n u s .
F lo w e rin g tim e i n
nauseosus
b loom s a
C h ry so th a m n u s v a r i e s w i t h t a x o n a n d h a b i t a t .
few
to
several
w eeks l a t e r
th an
C.
(X_
v isc id iflo ru s
17
w hen
th ey
occur
to g e th e r
(Winward a n d
1986 ) .
A n d e rs o n
Cl
h u m ilis
G re e n e i s t h e e a r l i e s t b lo o m in g s p e c i e s o f t h e g e n u s ( A n d e rs o n 1 9 8 6 a ) .
P la n ts a t
h igh
e l e v a t i o n s bloom e a r l i e r
(M cA rthur e t a l .
(M cA rthur e t a l .
th e
to
c o m p a tib ility ,
c o n d itio n s ,
p h e n o ty p ic
in h a b itin g
is
to
w ould
d istu rb ed
and
c o lo n iz in g
for
and
rap id
very
Evans
(1974a)
sh o rt
d isp e rsal,
v a ria tio n ,
v ig o ro u s
fe rtility
and
w ith
have
them
th a t
severe
in
p lan ts
Lowest
d isp e rsa l
(Young e t
if
C^ V l
a l.
ssp.
c a n o p i e s w e re s h a p e d i n a p e r f e c t h e m i s p h e r e t h e a v e r a g e
produce
for
v isc id iflo ru s
S e lf-
1957).
3 0 ,0 0 0
achenes
(6 a c h e n e s /c m 2 o f
w i t h a mean s h r u b canopy h e i g h t a n d d i a m e t e r o f 55 cm ).
im p o rta n t
se lf
e n v iro n m e n ta l
(S te b b in s
e stim a te d
as
fa c u lta tiv e
e sta b lish m e n t.
seeds
areas
lists
d istan ce
co n tin u e d
drought
su rro u n d in g
(1965)
fa v o ra b le
se ed lin g
sm all
C h rv s o th a m n u s t a x a
Baker
e d a p h ic
in su rin g
p e rio d ic
many,
in
and
p l a n t c o m m u n itie s
sp e cie s:
lo n g and
c lim a tic
show er
th a t
p ro d u ctio n
m echanism
produce
Y oung
of
e q u ip p e d
a
w h ich
v isc id iflo ru s
p lan t
seed
e n v iro n m en ts o f
m ec h a n ism s
1 972).
h ig h
re p ro d u c tio n ,
sp e cie s
c o lo n iz e
c h a ra c te ris tic s
success
p la stic ity
fe rtiliz a tio n
serai
th e
seeds
v e g e ta tiv e
o fte n
1 978b, M cA rthur an d W elch 1 9 8 6 ) .
fo llo w in g
c o n trib u tin g
low e l e v a t i o n s
19 7 9 a).
Some C h ry so th a m n u s t a x a
ex p ress
th an th o se a t
w ind
p lan ts
drought
d isp e rsa l
failed
(Y oung
p h e n o ty p ic p l a s t i c i t y
and
to
(S te v e n s
et
flo w e r a t
a
Evans
1 9 7 4 a).
i n C h ry s o th a m n u s .
a l.
canopy
The p a p p u s i s
1986).
site
T h is
th a t
is
su rface
Ci
Vl
ssp.
e x p erien ced
an
exam ple
a
of
C h ry so th a m n u s s e e d e s t a b l i s h e s
r a p i d l y ; w i t h c o o l n i g h t s a n d warm d a y s i t
can g e r m i n a t e w i t h i n 2 d a y s
18
(Weber e t a l .
1985).
a r e fav o ra b le
(M cA rthur e t a l .
P lan t
Once e s t a b l i s h e d ,
c o m p e titio n i s
re p ro d u c tio n
of
m e a s u r in g
e ffe c ts
flo w er
th e
c o m p e titio n .
v ery im p o rta n t in
of
g ro w th
C h ry so th a m n u s,
p a rtic u la rly
w here
on
1 9 5 3 )•
if
c o n d itio n s
c o n t r o l l i n g t h e g r o w th
open
on
to
be
w hen
u n d e rsto ry
(F risch k n ech t e t a l .
th e
c o m p e titio n
ste m
rap id
1974).
C h ry so th a m n u s
p ro d u c tio n and
g r o w th i s
McKell
C h ry s o th a m n u s
c o n tro lle d ,
fo und
reduced
rein v a d es
v e g e ta tio n
th e v i a b i l i t y
(1956),
g ro w th ,
sig n ific a n tly
h erb aceo u s
S till,
range.
and
is
by
s lo w ly ,
abundant
o f s e e d s fro m p l a n t s
w i t h a n d w i t h o u t c o m p e t i t i o n i s t h e same (McKell a n d C h i l c o t e 1 9 5 7 ) .
C h ry s o th a m n u s
(1 9 7 4 b )
fo u n d t h a t
community
They
re sp ro u ts
a fte r
fo u n d
fire
th at
d o m in a te d t h e
Ci
in
response
v isc id iflo ru s
by
fire
to p k ill.
Young a n d
d o m in a te d a n A r t e m i s i a
re sp ro u tin g
enhanced
to
and
achene
by
seed lin g
p ro d u ctio n .
E v ans
trid e n ta ta
e sta b lish m e n t.
Ci
s i t e and p e r i o d i c a l ly r e e s t a b l is h e d
its e lf
C h ry s o th a m n u s h a s a C3 p h o t o s y n t h e t i c p a th w a y .
I n Ci
v isc id iflo ru s
for
15 y e a r s
a t th is s ite .
ra te
of
p h o to sy n th e sis
is
become l i g h t - s a t u r a t e d a t
t h i s m ig h t be due t o
th e
is
c h lo ro p la sts.
an a d a p tiv e
tem p e ra tu re s.
h ig h
fu ll
for
sun.
a
woody C3 p l a n t .
Weber e t a l .
to m e n to se v e s t i t u r e
They c i t e
fea tu re ,
nauseosus th e
It
does
not
(1 9 8 5 ) s u g g e s t t h a t
on t h e l e a f
su rface
sh a d in g
o th e r a u th o r s s u g g e s tin g t h a t pubescence
red u c in g
heat
lo a d s
by
lo w e rin g
in te rn a l
19
U ses a n d Management O p p o r t u n i t i e s
The t a x a
of
C h ry so th a m n u s t e n d
to
form
r e s i n s and o i l s .
a s s o c ia te d w ith th e x e ro p h y tic h a b i t a t of th e p la n ts
1923) .
(X
nauseosus
is
ric h
(M cA rthur a n d W elch 1 9 8 6 ) .
in
th e
secondary
H a l l & G oodspeed
T h is i s
( H a l l a n d C le m e n ts
m e ta b o lite ,
rubber
(1919) c o n c l u d e d t h a t i f
e x t r a c t a b l e o v e r 1 5 0 ,0 0 0 t o n s o f r u b b e r w e re a v a i l a b l e i n n a t i v e s t a n d s
of
C h ry so th a m n u s.
T he
rubber
in
C h ry so th a m n u s
in d iv id u a l c e ll s ; i t i s not a la te x ru b b er.
is
to o
sm all
to
w arrant
h arv est
(H a ll
is
present
n o ted t h a t
enhanced
because
d istrib u te d
are
p o te n tia l
of
th e
for
an d
G oodspeed
1919),
but
no
O stle r e t a l .
r u b b e r p r o d u c t i o n by (% n a u s e o s u s i s
fo llo w in g
c h a ra c te ristic s:
I)
it
is
w id e ly
t h r o u g h o u t w e s t e r n N o r th America w here w i n t e r t e m p e r a t u r e s
o fte n
b e lo w - 2 0 F ;
th u s i t s
p o te n tia l
o t h e r known r u b b e r - p r o d u c i n g p l a n t ;
a lk a lin e
th e
The p e r c e n t i n e a c h p l a n t
g e n e t i c s e l e c t i o n s h a v e b e e n made ( O s t l e r e t a l . 1986 ) .
(1986)
in
s o i l s ; and
3)
its
a b ility
2)
to
range
it
exceeds
th a t
grow s on m a r g i n a l
r e s p r o u t m akes i t
of any
lan d and
re sista n t
to
mowing a n d h a r v e s t i n g .
Many C h ry s o th a m n u s t a x a h a v e b e e n n o t e d a s u s e f u l s o i l
C.
Iin ifo liu s
is
v a lu a b le
for
s ta b iliz in g
s tr o n g un d erg ro u n d s p re a d in g c h a r a c t e r i s t i c
nauseosus
he a v y
is
u sefu l
litte r,
a b ility
for
to
ero sio n
c o n tro l
e sta b lish
fo rag e
s o il
(M cA rthur e t a l .
because
on s e v e r e
1978b), easy e s ta b lis h m e n t, and r a p id sp read
T h e re i s
d istu rb ed
sta b iliz e rs.
of
site s
its
due
to
1974).
a
C.
deep
ro o ts,
(M cA rthur
et a l.
(M cA rthur e t a l . 1 9 7 4 ).
some c o n t r o v e r s y o v e r t h e v a l u e o f C h ry s o th a m n u s t a x a a s a
reso u rce.
A c c o rd in g
to
M c K e ll
and
C h ilco te
(1 9 5 7 ),
C^
20
v lsc id iflo ru s
S ev eral
to
b o th
rep o rted
ste n o p h y I l u s
d ie t
of
nauseosus
situ a tio n s
c o n trib u te d
(1 9 5 0 )
and
have
liv e sto c k
th at
re sp e c tiv e ly .
re la tiv e ly
re p o rte d
and
on w i n t e r
range
S everson
in
C h ry so th a m n u s
Cook a n d
ssp.
v isc id iflo ru s
U tah
managed a s
h ig h ly
cover.
a
n e e d ed
p referred
In
was fo u n d t o
v ar.
d u rin g
1946-47
and
1947-48,
C h ry s o th a m n u s t a x a
L e a c h (1 9 5 6 )
d u rin g se v ere
Yoakum (1 9 8 6 ) s u g g e s t s C h ry s o th a m n u s s h o u l d be
com ponent
fo rag e
M o n ta n a ,
H a rris
(1966) e s t i m a t e d t h a t C h ry s o th a m n u s c o m p r is e d
G reat B asin d e e r u t i l i z e d
w in ters in C a lifo rn ia .
has
11.31% a n d 8.2% by w e i g h t o f.
30% o f t h e p r o n g h o r n d i e t i n t h e Red D e s e r t o f Wyoming.
fo u n d t h a t
shrubs.
d iets.
(C^
1980a) c o n t r i b u t e d
u n p a la ta b le
w here
w ild life
C^ s t e n o p h y l l u s
A n d e rs o n
sheep
been
are
on p r o n g h o r n
sp e cie s
for
ran g elan d s
p ro nghorn and
C h r y s o th a m n u s . a l o n g w i t h
c o m p r i s e 60% o f mule d e e r d i e t s
sin c e
p ro v id e s
A rtem isia
in
it
is a
needed
and J u n i p e r ,
G a r f i e l d a n d Rosebud
c o u n t i e s , MT ( E u s t a c e 1 9 7 1 ) .
The u s e
of
C h ry so th a m n u s t a x a
may be p a r t i a l l y
and abundance o f a s s o c i a te d f o r a g e .
d e te rm in e d
I n a s t u d y on t h e n u t r i t i v e v a l u e
o f w i n t e r r a n g e p l a n t s i n t h e G r e a t B a s i n (Cook e t a l .
s te n o p h y llu s
co m p rised
8% o f
th e
d ie t
s a l t b u s h r a n g e , a n d 2% on g r a s s r a n g e .
on d e p l e t e d
w in ter
game r a n g e s
when more
(M cA rthur e t
d e sira b le
by t y p e
on
1 9 5 4 ) , u s e o f Ci
sagebrush
range,
6% on
C h ry so th a m n u s s p e c i e s a r e u s e d
a l.
fo rag es a re
1974) a n d
u n a v a ilab le
in
th e
fa ll
(M cA rthur
and
et a l.
1978b, L e a c h 1 9 5 6 ) .
M cA rthur e t a l .
(1 9 7 4 ) s u g g e s t t h a t t h e f o r a g e v a l u e o f Ci n a u s e o s u s
v a rie s
w ith
su b sp ec ie s,
lik e ly
tru e
of a l l
th e
e c o ty p e ,
and
season
C h ry so th a m h u s t a x a .
of
th e
I n Ci
y ear.
T h is
is
n au seo su s t h e gray
21
group
th an
su b sp ecies
th o se
g e n e ra lly
su b sp ec ie s
c o n s im ilis and
p a la ta b le
report
gam e a n d
and
ra b b itb ru sh )
in
th e
p a la tab le
green
liv e sto c k
ru
ssp.
(M cA rthur e t
p a la ta b ility
to
group.
th an
gray
s a lic ifo liu s
a l.
ratin g s
s t a t e s by s u b s p e c i e s .
nauseosus a r e
more
game a n d
In
U tah,
liv e sto c k
Cjl ru
ssp.
n ^ s s p . g r a v e o le n s (g re e n group s u b s p e c ie s ) a r e l e s s
to
a lb ic a u lis
are
1974).
of
group
tax a ,
R ydberg
D ittb e rn e r
( m o u n ta in
and
C h ry so th a m n u s t a x a
Cjl ru
in
ssp.
rubber
O lso n
(1983)
several
w estern
F o r e x a m p le , i n Montana s e v e r a l s u b s p e c i e s o f C.
r a t e d good i n
p a la ta b ility
for w h ite - ta il
deer as is
C2.
v i s c i d i f l o r u s f o r m ule d e e r .
Sampson a n d
Jesperson
(1963)
n o ted
th at
th e
n a u s e o s u s w as 9% d u r i n g d o rm a n t m o n th s a n d
lea v e s a re
found t o
form ed.
low
11.8% i n
p ro te in
of
C2^
s p r i n g a f t e r new
I n a n a n a l y s i s o f C2^ s t e n o p h y l l u s . c a r o t e n e was
be r e l a t i v e l y
re la tiv e ly
crude
h ig h
(d ig e stib le
( 2 .1
m g /lb ),
p ro te in
but
v a lu e
p h o sp h o ru s and
= 4% O c t o b e r
31
p ro te in
an d
2.2%
December 11) (Cook e t a l . 1 9 5 4 ) .
F risc h k n e c h t
A g ro p y ro n
and
(1 9 6 3 )
c rista tu rn
av erag e
c u lm
p ro d u c tio n and
h e ig h t
C h ry s o th a m n u s had h o t
th e
g ro w th
of
d e te rm in e d
of
e ffe c ts
of
fo u n d
herbage
y ie ld
Ajl c r i s t a t u m
b e e n rem o v e d .
A2. c r i s t a t u m
th e
by
w ere
Cjl n a u s e o s u s
a ll
of
seedheads,
g rea ter
C h ry so th a m n u s seem ed t o
p ro v id in g
e v a p o r a t i o n , a n d by n o t c o m p e tin g f o r w a t e r .
shade,
hence
on
w h e re
enhance
d e c re a sin g
C o m p e t i t i o n f o r w a t e r was
m in im iz e d b e c a u s e A2. c r i s t a t u m a n d Cjl n a u s e o s u s do n o t a c t i v e l y grow a t
t h e same t i m e s a n d s e c o n d a r y l a t e r a l s o f Cl n a u s e o s u s s e e m i n g ly o f f e r
little
c o m p e titio n t o g r a s s .
Thus Cl n a u s e o s u s c a n i n c r e a s e t h e v a l u e
22
Al C f i s t a t u m r a n g e f o r f a l l g r a z i n g by a l l o w i n g u n d e r s t o r y p l a n t s t o
r e m a in s u c c u l e n t .
C h ry s o th a m n u s t a x a h a v e s e v e r a l o t h e r p o s s i b l e u s e s .
(1 9 8 5 )
such
suggest
as
th at
Weber e t a l .
n a u s e o s u s h a s many c u r r e n t a n d p o t e n t i a l
la n d sc a p in g ,
n u tritiv e
enhance
v a lu e
of
th e
in cre ase
im prove
reduce th e
chances of d e s tr u c tiv e
O ften
lan d sc a p es
th e
d istu rb ed .
1931).
dye
C ontrol
and
R o b e rtso n
and
y ie ld s,
a v a ila b le
of
th e
in fe s ta tio n s
C h ry so th a m n u s
as h a b ita t
im prove
w in ter
seed in g ,
spp.
and
th e
fo rag e,
have
in d ic a to r
and
1986 ) .
(S te v e n s
of
P e e b le s
in
M ille r
C h ry so th a m n u s
g reat
and
2 ,4 - D
c o n tro lle d
and
at
(M a rio n e t a l .
is
et
a l.
d iffic u lt
re p ro d u c tiv e
C ords
w ith
(1 9 6 8 )
season
M cA rthur
1979a),
and
1979 a ) recommended C h ry s o th a m n u s f o r l a t e
C^ v i s c i d i f l o r u s
w ith
by
c a n be u s e f u l
(1957)
c o n t r o l l i n g C h r y s o th a m n u s .
g r o w in g
in se c t
in se c tic id e s
been
(D a y to n
F l o w e r s a n d i n n e r b a r k o f C h ry so th a m n u s make y e l l o w and g r e e n
M cA rthur e t a l .
and
grass
th e a e s t h e ti c s
d o m in a te d
C h ry so th a m n u s
(K e a rn e y
h a b it
enhance
se ed in g ,
snow e n t r a p m e n t ,
n a tu ra l
p o te n tia l
When C h ry so th a m n u s i s i n c l u d e d i n a h e r b a c e o u s s e e d i n g , i t
p ro d u c tio n ,
of
ru b b er,
fu n g ic id e s.
to ta l
source
n a tu ra l
crop,
in cre ase
p o te n tia l
of
hydrocarbon
can
and
p ro d u c tio n
uses
c a p a c ity
fo u n d
d e te rm in e d
rate
1986).
2 ,4 -D
2 ,4 - D
T o rdon
e ffe c tiv e
low a p p l i c a t i o n s
any
( McKelI
of
and
su p e rio r
its
season honey.
resp ro u tin g
C h ilco te
to
(in
1957).
2 ,4 ,5 -T
in
E vans a n d Young (1973) g o t 97% c o n t r o l o f
2 .4 k g /h a
was m ost
because
B ooth
b e tw e e n
in
of
1 .2 8
+ 0 .6 k g / h a p i c l o r a m .
22-K a t 2 . 4
e ra d ic a tin g
T o rd o n .
to
1 .6 8
k g /h a
la te r
P au lsen
in
th e
C^ p a r r y i
com pared
C^ n a u s e o s u s
was n o t
k g /h a
of
T e b u th iu ro n
The optimum t i m e t o a p p l y 2 ,4 - D t o C h ry so th a m n u s
23
is
when t h e
1973).
a d v a n ta g e
c u rre n t annual
Bange
im p ro v e m e n t
because
its
sh o o ts a r e
p ra c tic e s
g r o w th a n d
c o m p e tin g v e g e t a t i o n i s r e m o v e d „
7 »6 cm l o n g
may g i v e
seed
(E v a n s an d
C h ry so th a m n u s
p ro d u ctio n
are
Young
tax a
enhanced
an
when
24
STUDY AREAS AND METHODS
G e n e r a l S u rv e y M ethods
C h ry s o th a m n u s s p e c i m e n s w e re c o l l e c t e d t h r o u g h o u t Montana d u r i n g t h e
summers o f
th e ir
1985 an d
1986
d istrib u tio n s.
A ll
su b sp ecies le v e l
a n n o ta te d
by
L.
to
in v en to ry
c o lle c te d
u s i n g A n d e r s o n 's
C.
A nderson,
Montana
tax a and to
s p e c i m e n s w e re
d e te rm in e d
(1 9 8 6 a ) d i a g n o s t i c k e y .
o b tain ed
on
lo an
d e te rm in e
from
th e
to
th e
S p e c im e n s
fo llo w in g
h e r b a r i a ; U n i v e r s i t y o f M ontana, M i s s o u l a (MONTU); U n i v e r s i t y o f I d a h o ,
Moscow ( I D ) ; W a sh in g to n S t a t e U n i v e r s i t y , P u llm a n (WS); a n d U n i v e r s i t y
of
W yom ing,
L a ram ie
(RM)
(H o lm g ren
and
Keuken
1974),
a id e d
in
v e rific a tio n .
Each p l a n t
c o lle c tio n
site
was n o t e d on a
g e n e r a l h a b i t a t i n f o r m a t i o n was r e c o r d e d .
by t h e hand e s t i m a t i o n m ethod
S o i l t e x t u r e was d e t e r m i n e d
(USDA 1 9 7 5 ) .
I o f 3 c a t e g o r i e s ; v e ry sh a llo w (0-2 i n ) ,
(6+ i n ) .
S o i l d e p t h was a s s i g n e d t o
s h a l l o w ( 2 - 6 i n ) , a n d n o rm al
A com pass was em ployed t o d e t e r m i n e a s p e c t a n d p e r c e n t s l o p e
was e s t i m a t e d .
B ro w sin g on
(<15?> l e a d e r s
(40+%
Montana h ighw ay map a n d
brow sed),
le a d e rs
C h ry so th a m n u s
m o d e r a te
bro w sed ) .
p lan ts
was n o t e d a s
lig h t
b row sed),
heavy
( 15-40% l e a d e r s
D o m in an t
a sso c ia te s
and
or
d istu rb an c e
i n d i c a t o r s , i f p r e s e n t , w ere n o t e d .
In a d d itio n ,
fro m Montana
w e re
v e rifie d
h erb ariu m
S ta te
to
s p e c im e n s
U n iv e rsity
su b sp ec ie s.
p rev io u sly
(MONT) a n d
th e
Any a v a i l a b l e
c o lle c te d
in
M o n ta n a ,
U n iv e rsity
of
M ontana,
h a b ita t
i n f o r m a t i o n was
n o t e d a n d l o c a t i o n s w e re r e c o r d e d s e p a r a t e l y on d i s t r i b u t i o n m aps.
25
Unknown Taxon S t u d y Area a n d M ethods
D u r in g
1 985,
d iscovered
in
a
p re v io u sly
so u th w e ste rn
a d d itio n a l l o c a l it ie s
c h a ra c te ristic s
w e re
u n c h a ra c te riz e d
M on ta n a .
Thus,
tax o n
in
th e
o f t h e new t a x o n w e re s o u g h t .
know n,
sim ila r a re a s
w ere
of
p arry i
sum m er
was
of
1986
Once l i k e l y
site
surveyed
for
the
new
tax o n .
At 2 l o c a l i t i e s ,
s i t e s lo c a te d
u n d escrib ed
Jerry
C reek a n d Q u a r tz H i l l
in th e R e s u lts and D isc u ssio n
taxon apppeared
to
have
l e a v e s a n d I w i t h o u t to m en tu m .
w e re
c o lle c te d
p l a n t s w ere
siz e
at
soaked
(A n d e rs o n
th ese
in
1964).
2
sectio n ,
fo rm s,
site s.
Leaves and heads
50% e t h y l
a lc o h o l
to
the
tomentum on t h e
fe w e r due t o
fro m
re sto re
V e g e tativ e and f l o r a l
(o c c a sio n a lly
I w ith
T a b les 5 ,6 ) ,
P l a n t s w i t h m a t u r e f l o w e r s o f e a c h form
f o r com parative and d e s c r i p t i v e in f o r m a tio n .
10 m a tu r e h e a d s
( d e s c r ip tio n o f th ese
th ese
them t o
pressed
o rig in a l
m e a s u r e m e n ts w e re t a k e n
F o l l o w i n g A nd e rso n (1 9 6 4 )
t h e l a c k o f m a tu r e h e a d s )
from e a c h s a m p le w e re m e a s u re d f o r i n v o l u c r e f e a t u r e s a n d I f l o w e r fro m
e a c h h e a d was m e a s u re d
tom entum form an d
for
flo ra l
fe a tu re s.
S ev en teen p l a n t s
7 o f fo rm w i t h o u t tom entum ) w e re m e a s u r e d
(10 o f
from t h e
J e r r y C re e k s i t e a n d 20 (1 0 o f e a c h fo rm ) w ere m e a s u re d from t h e Q u a r tz
H ill
site .
A ll
to g e th e r
298
heads
and
flo w e rs
w ere
m easured.
M e asu re m e n ts w e re made on 10 l e a v e s fro m e a c h p l a n t .
F lo w e r b u d s , a t t h e s t a g e o f a n t h e r s - e n l a r g e d ,: b u t n o t y e l l o w , w ere
c o lle c te d ,
p r e s e r v e d i n a c e t i c a c i d a n d l a t e r t r a n s f e r r e d t o 50% e t h y l
a l c o h o l f o r m e i o t i c chromosome c o u n t s .
Chromosome c o u n t s w e re made by
26
D r.
E. D. M cA rthur a t
th e
U n ited S t a t e s
F orest
S e rv ic e
S h ru b S c i e n c e
L a b o r a t o r y i n P r o v o , U ta h .
Seven
site s
w ith
fea tu re s.
A ll
1976)
th e y
but
P re c ip ita tio n
e le v a tio n s
su p p o rte d
v a ria b le
c o m p le te
th e
taxon
7 lo c a tio n s
w ere
re c e iv e d
v a rin g
a m o u n ts
ranged
ranged
from
A rtem isia
and
new
a ll
d e sc rip tio n s
silty
to
356
-
except
th ese
range
2227
trid e n ta ta
of
stu d ie d
am o u n ts
fro m
1708
a sp ec ts
w e re
d e scrib e
h a b ita t
(R oss and
H unter
annual
588
n o rth
ft).
c o m m u n it i e s .
w ere
see
p re c ip ita tio n .
mm ( 1 4 - 2 3 . i n )
m (5 6 0 0 -7 3 0 0
grass
s ite s
site s
of
to
to
A ll
site s
S lo p es
w e re
For
more
e n c o u n te re d .
th e
R e su lts
and
and
D iscu ssio n
s e c t i o n (T ab le 5 , 6 ) .
The f o l l o w i n g p r o c e d u r e was. em ployed a t e a c h s i t e
a re a o f th e
s a m p l in g p l o t s .
F irst,
th e reg io n
t o d e te rm in e th e
c o n ta in in g th e h ig h est
d e n s i t y o f Cl p a r r y i was o c u l a r l y e s t i m a t e d a n d t h e a r e a w as m e a s u r e d .
T hen,
w ith in
th is
area,
w e re m e a s u r e d from a
th e
"clo sest
n e ig h b o r"
in te r-p la n t
sam ple o f 10 ra n d o m ly c h o s e n
d istan ces
p lan ts.
Twice t h e
mean " c l o s e s t n e i g h b o r " d i s t a n c e w as a d d e d t o e a c h s i d e o f t h e o c u l a r l y
e stim a ted
area,
th e re b y
e sta b lish in g
a
p lo t
w h ich
in clu d e d
some
o u tly in g p la n ts .
The f o l l o w i n g m e a s u r e m e n ts w e re t a k e n o f
d e n sity ;
average
in to
2)
crown
lea d e r
su b p lo ts.
coverage;
le n g th
In
(cm ).
each
3)
average
p la n t
To s i m p l i f y
su b p lo t
a ll
th e
h e ig h t
c o u n ts,
Cl p a r r y i
T his in c lu d e d o n ly i n d iv i d u a l s w ith a l l
Cl p a r r y i
basal
p lo ts
p lan ts
p lan ts;
(cm );
w e re
w e re
s te m s w i t h i n
and
I)
4)
d iv id e d
c o u n te d .
th e
p lo t.
F o r t h e r e m a i n i n g m e a s u r e m e n ts , a p r o p o r t i o n o f p l a n t s w e r e sam pled so
t h a t 300 l e a d e r s w e re m e a s u r e d p e r p l o t .
The number o f p l a n t s n e e d ed
27
to
in su re
th is
num ber
of
lea d e rs
was
d e te rm in e d
by
a v e r a g e number o f l e a d e r s p e r p l a n t a t e a c h p l o t .
e stim a tin g
th e
T h i s was n e c e s s a r y
b e c a u s e t h e num ber o f l e a d e r s p e r p l a n t v a r i e d b e tw e e n p l o t s a n d w i t h i n
p lo ts.
The number o f
e s tim a tin g th e
The f i r s t
p lan ts
m easured
p ro p o rtio n of th e t o t a l
p lan t
per
s u b p l o t was d e t e r m i n e d
by
count p r e s e n t i n each s u b p lo t.
m e a s u r e d was a r b i t r a r i l y
chosen and fo llo w in g p l a n t s
w e re m e a s u r e d a t random i n t e r v a l s by c h o o s i n g a number b e tw e e n I a n d 10
f o r each s u b p lo t.
Use was n o t e d on e a c h p l a n t m e a s u r e d .
Ten p e r c e n t
( b u t n o t <15) o f t h e l e a d e r s w ere c o u n te d p e r i n d i v i d u a l .
The p e r c e n t
ca n o p y c o v e r a g e o f
p a r r y i was d e t e r m i n e d by m e a s u r in g t h e
sh o rte st
and th e l o n g e s t d im e n sio n s o f each p l a n t .
To d e t e r m i n e w h a t e n v i r o n m e n t a l
an a d ja c e n t
p a ire d
m a t c h in g s l o p e ,
p lo t
p lo t
a sp ec t,
c o n ta in in g
w ith o u t
f a c t o r s m ig h t f a v o r t h e new t a x o n ,
th e
tax o n
and v e g e ta ti o n a s
p a rry i.
p resen t
w as
c lo se ly a s
T h is p l o t , a s i n t h e
chosen
p o ssib le
by
to
th e
CL p a r r y i p l o t ,
was
d iv id e d i n to s u b p lo ts .
A lin e
t r a n s e c t was p l a c e d i n e a c h s u b p l o t i n
both th e p a ir e d p l o t s .
To m e a s u r e b a s a l c o v e r a g e o f g r a s s e s a n d f o r b s ,
by s p e c i e s , a 2 x 5 dm r e c t a n g u l a r q u a d r a t was p l a c e d e v e r y o t h e r m e t e r
a lo n g a tap e s tr e tc h e d th e le n g th of th e s u b p lo t.
o f t h e q u a d r a t was p l a c e d on t h e m e t e r m a rk .
bob,
canopy
cover
of
shrubs
o th er
re c o rd in g th e d is ta n c e each p la n t
than
The t o p l e f t c o r n e r
By t h e
Cl p a r r y i
spanned a lo n g
th e
use
of a
plumb
was m ea su red
tap e.
by
Only l i v e
crow n was r e c o r d e d a s c o v e r a g e .
D e n sity
of
sh ru b s
o th er
th an
Cl p a r r y i
was m e a s u re d
by
i n d iv i d u a l s of each s p e c ie s w ith in I m of each l i n e t r a n s e c t .
c o u n tin g
28
S o i l s a m p l e s w ere t a k e n i n e a c h q u a r t e r o f t h e p a i r e d p l o t s .
taxon
p lo t,
s a m p le s
w ere
tak en
d e p t h s : 0 t o 5 cm ( 0 - 2 i n ) , 5 t o
c lo se
to
e le c tric a l
c o n d u c tiv ity
Cl p a r r v i
p lan t.
T h re e
15 cm ( 2 - 6 i n ) , a n d 15 t o 3 0 . 5 cm ( 6 -
12 i n ) w ere s e p a r a t e d a n d s t o r e d i n b a g s .
1936),
a
In th e
T e x t u r e a n a l y s i s ( Bouyoucous
(USDA 1 9 5 4 ) ,
and
pH (USDA 1954)
w ere
a n a ly z e d .
A d d itio n a l
d istu rb an c e ,
annual
in fo rm a tio n
a sso c ia te d
p re c ip ita tio n s a t
w as
sp e cie s,
each
recorded
a sp ec t,
stu d y
site
and
at
each
s lo p e .
s ite
such
as
E le v a tio n s and
w e re e s t i m a t e d
from Montana
c l i m a t o l o g i c a l r e c o r d s ( N a t i o n a l O c e a n i c a n d A tm o s p h e r ic A d m i n i s t r a t i o n
1984).
29
RESULTS AND DISCUSSION
D u rin g
th e
sum m ers
C h ry so th a m n u s
A d d itio n a lly ,
tax o n
of
U n til
now,
1986a).
tax a
of
w ere
in A ugust,
and
1986,
8
to
occur
in
found
p arry i
su b sp ec ie s
sta lk e d
P^-1T y i
w ith
of
was l o c a t e d
C^ p a r r y i
had
^a x a
g lan d s
th e
d iffe re n tia te
in
(T a b le
so u th w e ste rn
been
e x c ep tio n
of
th e
Ct
2 ).
d e scrib ed
M ontana.
(A n d e rso n
it
is
new t a x o n fro m o t h e r Ct
ssp.
C le m ., known o n l y from C a l i f o r n i a a n d N evada.
asper
( G reen e) H all
B a se d on t h e s e
p r o p o s e d t h a t t h i s new t a x o n be known a s
g l a n d u l o s u s W i n k l e r a n d Wambolt .
s s P-
M o n ta n a
d e scrib ed
None o f t h i s s u b s p e c i e s h a v e b e e n n o t e d t o o c c u r i n M ontana.
C o n s p ic u o u s
g lan d s
p re v io u sly
1985 a p r e v i o u s l y u n c h a r a c t e r i z e d s u b s p e c i f i c
C h ry s o th a m n u s
12
1985
g la n d u lo su s
v ar.
g lan d u lo su s
Two v a r i e t i e s
and
Ct
are
ssp.
&
sta lk e d
C. p a r r y i s s p .
proposed,
g lan d u lo su s
C.
v ar.
to m e n to su s.
T ab le 2 .
O c c u r r e n c e o f C h ry so th a m n u s t a x a i n M ontana.
L o c a te d d u r i n g
t h i s stu d y
C. I i n i f o l j u s
R e p o r te d i n t h e l i t e r a t u r e
A n d e rs o n 19 8 6 a . Ct v i s c i d i f l o r u s
ssp.
I i n i f o l i u s by H a l l and
C le m e n ts 1923.
Ct n a u s e o s u s
ssp. a lb ic a u lis
x
ssp . c o n sim ilis
x
ssp . g rav e o len s
x
A n d e rs o n 19 8 6 a , H a l l a n d C le m e n ts
1 923.
A n d e rso n 1986a.
A n d e rs o n 19 8 6 a , H a l l a n d C le m e n ts
19 23.
30
Table 2 .
co n tin u ed .
Taxon
L o c a te d d u r i n g
t h i s stu d y
R e p o r te d i n t h e l i t e r a t u r e
C. n a u s e o s u s
s s p . na u s e o s u s
x
ssp . sp ecio su s
A n d e rs o n 1 9 8 6 a .
(X
ssp.
t y p i c u s by H a l l a n d C le m e n ts
1 923.
H a l l a n d C le m e n ts 1 9 2 3 . I n c l u d e d
i n C. n . s s p . a l b i c a u l i s i n
A n d e rso n 1 9 8 6 a . , .
C. p a r r y i
s s p . g lan d u lo su s
x
C. v i s c i d i f l o r u s
ssp . Ia n c e o la tu s
x
A n d e rs o n 19 8 6 a , H a l l a n d C le m e n ts
1923.
ss p . p u b eru lu s
H a l l an d C le m e n ts 1 923.
ssp.
p u m ilis
H a l l an d C le m e n ts 1 9 2 3 . P a r t o f
C. v . s s p . v i s c i d i f l o r u s bv
A n d e rso n 198 6 a .
ssp.
ste n o p h y llu s
H a l l a n d C le m e n ts 1 9 2 3 . V a r i e t y
o f C. n . s s p . v i s c i d i f l o r u s bv
A n d e rso n 1980b.
ssp. v is c id iflo ru s
v a r. ste n o p h y llu s
x
A n d e rs o n 1 9 8 6 a .
L a b e le d Ci
s s p . t y p i c u s by H a l l an d C le m e n ts
1923.
x
v ar. v is c id if lo r u s x
A n d e rs o n 1980b.
31
PhylOKeny and c l a s s i f i c a t i o n o f Chryaothamnus p a rrv i
sa p . K landulosus
The r e p r o d u c t i v e s t r a t e g y o f f a c u l t a t i v e s e l f - f e r t i l i z a t i o n w i t h t h e
c a p ac ity
to
o c c asio n ally
o u tcro ss,
as
o ccu rs
p r o m o te s t h e d e v e lo p m e n t o f new v a r i e t i e s .
e c o ty p e s,
su b sp ec ie s,
and
sp e c ie s.
in
C h r v s o th a m n u s . o f t e n
T h e se v a r i e t i e s i n c l u d e new
H ow ever,
th is
c o n d u c iv e t o e v o l u t i o n o f new g e n e r a a n d f a m i l i e s
sp e cie s
is
a
re stric tio n
b a rrie rs
p la n ts
p o p u la tio n
of
gene
(S te b b in s
co n fo rm
c o n se q u e n tly
e n tity
exchange
1950).
w e ll
m ain ta in e d
by
S e v e n ty
to
th is
(S te b b in s
1965).
d e te rm in e
if
ssp.
Ci
80) o f
b io lo g ic a l
show m o r p h o l o g i c a l
iso la tio n
to
No r e p r o d u c t i v e
g lan d u lo su s
is
in a b ility
or
g e n e tic
th e
not
great
of
h ig h er
d e fin itio n
based
te sts
or
on
w ere
and
re p ro d u c tiv e
perform ed
re p ro d u c tiv e ly
A
iso la tio n
sp ecies
sp e c ie s
d isc o n tin u ity
is
( S te b b in s 1957).
by t h e
p h y sio lo g ic a l
tra it
iso la te d
to
from
o t h e r Ci p a r r y i t a x a .
S u b sp e c ie s
c e rta in
are
se rie s
m o rp h o lo g ic a l
and
of
p o p u latio n s
p h y sio lo g ic a l
rese m b lin g
each
o th er
c h a ra c te ristic s,
in
each
in h a b itin g a g eo g rap h ic s u b d iv is io n o f th e s p e c ie s range o r a s e r i e s o f
sim ila r h a b ita ts .
ty p ic a l
of
them
d iffe re n t
members o f o t h e r s u b s p e c i e s ,
by a
su b sp ecies
se rie s
can
le a s t p a rtly
1950).
They a r e
c o e x ist
iso la te d
M u lle r
evo l u ti o n a r i ly
of
in
yet are
in te rg ra d in g
over
th e
several
fo rm s
same a r e a ,
c h a ra c te ristic s
co n n ected w ith
(S te b b in s
but a r e
sig n ific a n t
sta te s
in
th a t
th a t
it
su b sp ec ie s
I o r more
1 9 50).
lik e ly
from e a c h o t h e r by h a b i t a t p r e f e r e n c e s
(1940)
fro m
to
Two
be a t
(S te b b in s
e sta b lish m e n t
p r o m o te s a l l o p a t r i c
is
sp e c ia tio n .
32
He s u g g e s t s
p o p u la tio n
ten d s
to
th at
th e
s y s te m
in to
re stric t
th e
se g re g a tio n
a
p re v io u sly
2 o r more r e p r o d u c t i v e l y
su p p ly
p o p u la tio n s
and
E co ty p es,
e c o lo g ica l
an
of
c h a n n e ls
of
them
an d
genes
in to
iso la te d
a v a ila b le
c e rta in
a d a p ta tio n a l
in te rb re e d in g
to
each
p a th s
co n cep t,
p o p u la tio n s
of
are
of
th e se
a d a p ta tio n .
d istin g u ish e d
p r i m a r i l y by t h e i r r e a c t i o n t o t h e e n v ir o n m e n t a n d may p o s s e s s d e f i n e d
m o rp h o lo g ic a l
d iffe re n c es
m o rp h o lo g ic a lly
and
(S te b b in s 1950).
a p p a re n tly
C.
g e o g ra p h ic a lly
ssp.
g lan d u lo su s i s
se p a ra te d
from
o th e r
s u b s p e c ie s and th u s i s proposed a s a s u b s p e c ie s r a t h e r th a n a n e c o ty p e .
A n d e rs o n (19 7 8 ) i n f e r r e d t h a t Ci Pi
s s p . m o n ta n u s , a l o c a l
e n d e m ic ,
i s r e l a t e d t o Ci jx s s p . p a r r y i b e c a u s e o f i t s m a n y - f lo w e r e d h e a d s .
If
t h i s i s s o , t h e n Ci . Pi s s p . g l a n d u l o s u s i s l i k e l y d e r i v e d from t h e same
lin e a g e , sin c e i t
s im ila r
l i k e w i s e h a s many f l o w e r s p e r h e a d .
m o rp h o lo g ic a lly ,
yet
d iffe r
p rim a rily
in
T hese t a x a a r e
th at
Ci
Pi
ssp.
S-L&ndulo s u s h a s s t a l k e d g l a n d s a n d a g r e a t e r number o f f l o w e r s p e r h e a d
than
Ci
Pi
m o n ta n u s
ssp.
is
m on ta n u s
C la rk
C o .,
( T a b le 3 ) ;
Idaho
The t y p e l o c a t i o n o f Ci
on
exposed
ro c k y
slo p e s
of
Pi
th e
ssp.
Red
C o n g lo m e r a te P e a k s , l o c a t e d i n t h e I r v i n g C re e k d r a i n a g e , 28 a i r m i l e s
n o rth w est
o f D u b o is,
so u th w e st
border
of
Idaho
(A n d e rso n
Montana
1978).
w h e re a
few
T his l i e s
p o p u la tio n s
c lo se
of
Ci
to
Pi
th e
ssp.
g l a n d u l o s u s a r e known t o o c c u r .
It
is
from Ci
s p e c u l a t e d t h a t Ci
Pi
ssp.
Pi
ssp.
g lan d u lo su s v a r.
g lan d u lo su s v a r.
g la n d u lo su s.
Ci
Pi
to m en to su s a r o s e
s s p . g lan d u lo su s
v a r * g l a n d u l o s u s h a s a w i d e r r a n g e , o c c u r s more f r e q u e n t l y , and o c c u r s
i n d e p e n d e n t l y "of Ci
g lan d u lo u s v a r.
Pi
ssp.
g lan d u lo su s v a r.
to m e n to su s.
Ci
Pi
ssp.
t o m e n to s u s was fo u n d a t o n l y 2 o f t h e 7 s a m p l i n g s i t e s
T able 3 .
C om parison o f Chr y s o th a m n u s
y i vflxo. •
Taxon
Cl L l s s p .
g la n d u lo su s
L eaf
.len g th
(cm)
Leaf
w id th
(mm)
- 1-5
1 -4
p a rry !
s s p . g l a n d u l o s u s w i t h 3 m o r p h o l o g i c a l l y s i m i l a r C.
Leaf
su rface
Leaf
shape
T ip
sh ap e
F l ower
len g th
(mid)
Flow er
lo b e
len gth
(mm)
sta lk ed
glan d s
lin e a r lin ea r
o b la n ce o la te
acu te
6-9
O•8—1 .6
-[-S h
short
. sta lk ed
g la n d sb
o b la n ce o la te a
up t o
Sa
9 - 10ad
9 - 10b
.
C. Ll s s p .
asper
2-5b
C. p. s s p .
montanus
2 - 3 . 5ad
1-2d
v isc id u lo u s
Iin e a r d
C. p. s s p .
im u lu s
1-1 . 5 ab
2-3b
gray
w ith
d e n se
tcmentumb
sp atu Iatelin ea r
sp atu I a t e bi
^Anderson
bHall and
‘'McArthur
“Anderson
1986a.
Clements 1923.
et a l . 1979a.
1978.
-
"""
O b tu se b
In flo r e sc e n c e
arr a n g e m e n t
' cyme
No.
flo w ers
per
head
10-14
short
racem eb
5 - 1 Obc
1 . 4 - 1 .7 d
few -h ead ed ,
Cymosed
5-1 Id
1 - 1 .5 b
reduced.
racem eb
11-15°
34
a n d was a l w a y s i n t e r m i x e d w i t h C2, ^
ssp . g lan d u lo su s v a r . g lan d u lo su s.
M orphology o f C h ry so th a m n u s p a r r y j
C h ry so th a m n u s p a r r y i s s p .
much b r a n c h e d
tax a
underground s h o o ts .
t omentum.
M ille r
g lan d u lo su s
.
g l a n d u l o s u s i s a low s h r u b ,
from a woody c a u d e x .
(P a u lse n and
ssd
1968,
M cA rthur e t
The s te m s a r e
C h a ra c te ristic a lly ,
T his ta x o n ,
Ci
le a fy
a l.
lik e
1-3 dm t a l l ,
o t h e r Ci
1979b),
p arry i
sp read s
by
and covered w ith lo o s e w h ite
p a r r y i tax a have f e l t - l i k e
w h ite to
g r e e n t omentum w h ic h i s n o t a s d e n s e o r r e s i n o u s a s t h a t on s te m s o f C.
n a u s e o s u s (M cA rthur e t a l .
are
m o stly
e re c t,
a lth o u g h
19793-) •
Stem s o f Ci p a r r y i s s p . g l a n d u l o s u s
some a r e
fo u n d
sp re a d in g
B r a n c h in g o c c u r s o n l y i n t h e i n f l o r e s c e n c e .
se ssile ,
on
th e
ground.
The l e a v e s a r e a l t e r n a t e ,
n a rr o w t o b r o a d l y I i n e a r - o b l a n c e o l a t e , w i t h a c u t e t i p s .
l e a v e s m easure
(I)
2 t o 4 ( 5 ) cm l o n g a n d 1 .5 t o 4 mm w i d e .
The
They have
I p r o m in e n t n e r v e , y e t o c c a s i o n a l l y t h e r e a r e 2 o r 4 a d d i t i o n a l n e r v e s .
H a ll
and
dependent
C le m e n ts
on
le a f
(1 9 2 3 )
w id th .
n o te
th a t
The l e a v e s
n erve
are
number
is
roughened,
v a ria b le
w ith
and
num erous
I
sta lk e d
re sin
g lan d s.
The m a r g i n s a r e
e n t i r e and
th e
upper le a v e s
o c c a sio n a lly b arely su rp a ss th e in flo re s c e n c e .
The f l o w e r h e a d s o f Ci p a r r y i s s p . g l a n d u l o s u s a r e a r r a n g e d i n l e a f y
t e r m i n a l cymes ( F i g .
I ) , d i f f e r i n g fro m o t h e r Ci p a r r y i i n f l o r e s c e n c e s ,
w h ic h a r e t y p i c a l l y
ra c e m e s
( H a l l a n d C le m e n ts 1 9 2 3 ) .
The i n v o l u c r e s
o f Ci p a r r y i s s p . g l a n d u l o s u s a v e r a g e 1 .1 3 cm h i g h ; b r a c t number r a n g e s
from
(9)
k e e le d ,
12 t o
have
16.
T hese i n v o l u c r a l
n arro w ly acu m in ate
tip s
b rac ts a re
somewhat r a n k e d ,
and
nerved,
are
I
not
o ccasio n ally
35
w ith 2 a d d itio n a l
nerv es.
T here a r e
(9)
The d i s k f l o w e r s , t u b u l a r s h a p e d , a r e
I to
1 .3 mm l o n g .
stig m a tic
57%).
The
lin e s are
sty le
u su a lly
branch
sh o rte r
2 n = l8 .
14 ( 1 6 )
flo w e rs/h e a d .
(6 ) 7 t o 8 ( 9 ) mm l o n g ; l o b e s a r e
ranges
fro m 2 t o
th an th e
The p a p p u s l e n g t h i s 7 t o 8 mm.
The chromosome number i s
10 t o
3 .5
mm.
s ty l e appendages
The
(35 t o
The a c h e n e i s 2 t o 5 mm l o n g .
T h is i s
t h e modal num ber t h r o u g h o u t
*
th e
genus
from J u l y
(S o lb rig
et a l.
t o A u g u s t.
1969).
Cl p a r r y i
(% p a r r y i
ssp.
ssp.
g lan d u lo su s v a r .
l e a v e s a n d i n v o l u c r a l b r a c t s t h a t h a v e no o r l i t t l e
Cl p a r r y i
b rac ts
ssp.
g lan d u lo su s v a r.
co v ered
w ith
dense
g lan d u lo su s
g lan d u lo su s h as
to m e n to s ity w hereas
to m en to su s h a s l e a v e s and
tom entum w h ic h
makes
bloom s
th e
in v o lu c ral
sta lk e d
g lan d s
in co n sp ic u o u s (F ig . I ) .
T he
R e la tio n sh ip
of
C h ry so th a m n u s
p arry i
ssp.
g lan d u lo su s
to
o th er
C h ry so th a m n u s Taxa
C. p . s s p . g l a n d u l o s u s i s m o r p h o l o g i c a l l y c l o s e t o 3 o t h e r C^ p a r r y i
t a x a : Cl
ssp.
m o n ta n u s
L.
g la n d u lo su s
T y p ic a lly ,
m on ta n u s
Leaf
im u lu s H a l l & C l e m ., Ol
C.
and
A nderson.
C^
Cl ^ l s s p .
h a v e more
le n g th ,
ssp.
le a f
In tra s p e c if ic a I l y ,
asper
g la n d u lo su s.
than
7 flo w ers
w id th ,
s s p . a s p e r . a n d Cl p^_ s s p .
have
co n sp ic u o u s
Cl ^ l s s p .
flo w e r
num ber
Cl
ELl
C a lifo rn ia
Cl
sta lk e d
ssp.
g lan d s.
i m u l u s . a n d Cl ^ l s s p .
p e r h e a d an d a r e
m o r p h o l o g i c a l c h a r a c t e r i s t i c s d i s t i n g u i s h Cl
3 ).
o n ly
per
I
to
head,
2 dm t a l l .
and
o th e r
s s p . g l a n d u l o s u s ( T a b le
s s p . a s p e r o c c u r s on s l o p e s b o r d e r i n g t h e d e s e r t o f e a s t e r n
an d w e s t e r n
Nevada
from 2100
to
2600 m (6 8 8 5 -8 5 2 5 f t )
in
36
F ig u re I .
I l l u s t r a t i o n s o f C h ry so th a m n u s p a r r y ! s s p . g l a n d u l o s u s .
A.
H ab it x 2 / 3 .
B.
I n v o l u c r a l b r a c t o f Ci
ssp.
g l a n d u l o s u s v a r . g l a n d u l o s u s . show ing s t a l k e d g l a n d s
x 4 . C. I n v o l u c r a l b r a c t o f Ci
ssp. g lan d u lo su s
v a r . t o m e n t o s u s . show ing t omentum x 4 .
37
e le v a tio n
( H a l l a n d C le m e n ts 1 9 2 3 ); (% ^
ssp.
im u lu s i s
t h e San B e r n a r d i n o M o u n ta in s o f s o u t h e r n C a l i f o r n i a
1923);
and
C_^
ssp.
m o n ta n u s
has
been
fo u n d o n l y i n
( H a l l a n d C le m e n ts
lo ca ted
o n ly
in
th e
Red
C o n g lo m e ra te p e a k s a t t h e Montana — I d a h o b o r d e r ( A n d e rs o n 19 7 8 ) .
As a
H all and
sp e cie s
Cl p a r r y i
C le m e n ts
(1 9 2 3 )
is
m ost
c lo se ly
sta te
th a t
th e re
common o r i g i n o f t h e s e 2 s p e c i e s .
t omentum on
th e
ste m s
and
re la te d
to
Cl n a u s e o s u s .
can be no d o u b t a s t o
th e
They s h a r e c h a r a c t e r i s t i c s s u c h a s
sim ila r
shapes
of
c o ro lla s
and
sty le
branches.
In
th e
p a st,
n a tu ra l
resource
m a n a g e rs
have
reco g n ized
C h ry so th a m n u s t a x a i n M ontana, Cl n a u s e o s u s a n d Cl v i s c i d i f l o r u s
and
W rig h t
1966) .
R e c e n tly
4
D o rn
(1 9 8 4 )
illu s tra te s
c ite d
som e
of
a
th ird
th e
only
2
(B ooth
taxon,
Cl
lin ifo liu s .
T a b le
m o rp h o lo g ic a l
c h a ra c te ristic s
d i f f e r e n t i a t i n g Cl ^ l s s p . g l a n d u l o s u s fro m o t h e r t a x a
in th e S ta te .
M orphology o f o t h e r C h ry s o th a m n u s Taxa i n Montana
C h ry so th a m n u s n a u s e o s u s
M o r p h o l o g i c a l l y Cl n a u s e o s u s s u b s p e c i e s f a l l i n t o 2 g r o u p s , g r a y and
green,
based
in v o lu c re
on
ste m
( A n d e rs o n
c o lo r
and p re se n c e
1986 c ) .
The g r a y
or absence
group t y p i c a l l y
of
h a irs
h as gray
on t h e
s te m s
a n d p u b e s c e n c e t o t omentum on t h e i n v o l u c r e s a n d i n c l u d e s Montana t a x a
Cl Iil s s p .
n a u s e o s u s a n d (X m
ssp. a lb ic a u lis .
The p u b e s c e n c e on t h e
Table 4 .
D i a g n o s t i c c h a r a c t e r i s t i c s o f Montana C h r y s o t h a m n u s . a
Stems Tomentose
Stems Kot Tomentose
glabrous to puberulent
P h y llarie s a tte n u a te , ra th e r membranous
P h y llarie s obtuse to a c u te , chartaceous
Inflorescence racemose
5-20 flowers/head
Inflorescence mostly cycose
Usually 5 flowers/heed
Chrvsothaanus parrv l
Chrvsothamnus nauseosus
Leaves with
stalked
glands
Involucres pubescent to
tomentose (rarely nearly
nearly glabrous)
Stems w hitish
ssp.
Elandulosus
Leaves without stalked
glands
Leaves overtop
inflorescence
Upper stems glabrous
Leaves > 2 ma
wide, h a iry to
glabrous
Leaves 1-10 nm wide,
c i l i a t e margins
otherw ise glabrous
Leaves I m
wide or l e s s ,
I nerved.
Involucre
7-8 nm long.
Corolla 7 9.5 mm long.
Corolla lobes
1-2.5 nm long
Often b rig h t
green stem.
ssp.
salmonensis
ssp.
a lb ic a u lls
ssp.
Kraveolens
ssp.
consim ilis
fo lia g e
without
t omentum
van.
tomentosus
var.
Klandulosus
a I n c lu d e s two
Chrvsothamnus lin if o liu s
Upper stems hairy
Leaves 1-3
mm wide,
1-5 nerved.
Involucre
6.7-9«5 QQ
long.
Corolla 7-9
mm long.
Corolla lobes
.6-1.5 nn
long.
Often t o l l ,
ro b u st.
fo lia g e with
tomentun
Chrysothamnus v lsc id iflo ru s
Stems greenish
Corolla (8)
9-10 mm long.
Involucre
9-11 m long.
Shrub 4-15
dm t o l l .
ssp.
nauseosus
Leaves never tw isted,
lanceolate
Soboliferous
Thll plant
> 7 dm
Involucres glabrous
5-7 flow ers/ 8-20 flowers Corolla 6 head.
/head.
8.5 mm long.
Corolla p i e Leaves 2-3-5 Involucre
yellow, 8-10 cm long.
7-9-5 mm
™ long.
Shrub 1-2
long.
Leaves 1-3
dm t o l l .
Shrub 2-7
mm wide.
dm t a l l .
ssp.
montonus
Leaves o ften tw isted , lin e a r to oblong
lan c eo la te
S a a lle r p lant
UU
OD
ssp.
Ianceolatus
ssp.
v lsc id iflo ru s
Leaves > I .5 on Leaves 1-1.5 mm
wide; up to I m wide; mostly
t o ll
< 3 dm t o ll
var.
v lsc id iflo ru s
B arryi taxa from Idaho.
var.
stenophvllus
These taxa were not found in Montana but might occur.
\
39
i n v o l u c r e i s a n e x p r e s s i o n o f t h e t e n d e n c y to w a r d a b u n d a n t p u b e s c e n c e
i n t h e w h o le p l a n t
( H a l l a n d C le m e n ts 1 9 2 3 ) .
The s p e c i m e n s o f
n^ ssp.
nauseosus
r a n g e d i n h e i g h t f ro m 1 . 6 - 1 0 dm.
th e
in tra su b sp e c ific
v a ria tio n
c o lle c te d
d u rin g t h i s
stu d y
The f o l l o w i n g c o l l e c t i o n s i l l u s t r a t e
in
s te m c o l o r o f
th is
tax o n :
8 / I 6A b ,
8 / l 6 / 5 e , a n d 8 / 2 7 / A a had b r i g h t g r e e n s te m s ;
8 / 2 7 / 9b h a d a d a r k g r e e n
s te m ; a n d
S p e c im e n s o f
a lb ic a u lis
8 /2 6 /2b
had a c o t t o n y w h i t e s te m .
c o lle c te d
ranged
in
h e ig h t
from
2 .5 -1 1
dm.
n^ ssp.
Stem
c o lo r
c h a r a c t e r i s t i c s o f t h i s taxon v a rie d c o n s id e ra b ly . C o lle c tio n s 8 / 2 9 / la ,
8 / 2 9 / Ib , 8 / 3 1/ 3 b , 8 / 3 1 / 3 c , a n d 8 / 3 1/ 3 d had b r i g h t g r e e n s t e m s , h o w e v e r ,
c o l l e c t i o n 8 / 3 1 / 2 c ha d w h i t e s t e m s .
T y p i c a l l y ste m p u b e s c e n c e i n C. n .
s a p . a l b i c a u l i s r a n g e s from l i g h t g r e e n t o w h i t e v a r y i n g w i t h e c o t y p e
(Hanks e t a l . 1 9 7 5 ) .
The g r e e n g r o u p , composed o f
g rav eo len s,
has
green
s te m s a n d
C^ ru
ssp.
g lab ro u s
c o n s i m i l i s a n d CL n ^ s s p .
in v o lu c re s.
The. h e i g h t
c o l l e c t e d s p e c i m e n s o f Cr n ^ s s p . g r a v e o l e n s r a n g e d fro m 3 .1
Most s p e c i m e n s o f t h i s t a x o n w e re f a i r l y
n arrow er l e a v e s .
S p e c im e n s o f Cr m
ty p ic a l, y e t,
of
t o 12 dm.
some p l a n t s h a d
s s p . c o n s i m i l i s r a n g e d from 2 . 4 t o
7 dm i n h e i g h t .
V a r i a t i o n i n t h e a b o v e t a x a I may be e x p l a i n e d by i n t e r g r a d a t i o n t h a t
o ccu rs
b e tw e e n
(F ig . 2 ) .
p a ra p a trie
and
sy m p atric
I n a r e a s w h e re 2 A c i u r i n a
su b sp ec ie s
of
Cr
nauseosus
s p e c i e s ( f r u i t f l y ) c o n c u r a n d Cr
n a u s e o s u s s u b s p e c i e s a r e s y m p a t r i c , I f l y s p e c i e s (A c i u r i r m b i g e l o v i a e )
i n d u c e s c o t t o n g a l l s on t h e g r e e n g r o u p o f Cr n a u s e o s u s w h e r e a s a n o t h e r
fly sp ecies
(Ar t r i x a )
i n d u c e s c a l l u s g a l l s on t h e w h i t e g r o u p .
G a lls
40
in
areas
of
s y m p a try
can a i d
in
id e n tific a tio n
of
su b sp ecies
of
C.
n a u s e o s u s (M cA rthur e t a l . 1979 b , Weber e t a l . 19 8 5 ) .
C. n . s s p . g r a v e o l e n s
C. n . s s p . a l b i c a u l i s
- 3 Ci
ssp.
%
-----------^
— / j —>
F ig u re 2 .
c o n sim ilis
Sympat r i e ; m o d e r a te g e n i c e x c h a n g e
S y m p a t r i c ; e x c h a n g e unknown
A llo p a tric
R e l a t i o n s h i p s among Montana C hry so th a m n u s n a u s e o s u s
su b sp ecies.
A dapted f o r Montana from A n d e rso n 1 9 8 6 c .
C hry so th a m n u s v i s c i d i f l o r u s
I n tr a s p e c ific
nauseosus
are
p rim a rily
d iffic u lt
d istin c tio n s
and
su b sp ec ie s
(1986a),
Ci
Ci
by t h e l a c k
c la ssify
(A n d e rs o n
ste m s,
in
if
ssp.
an d
v isc id iflo ru s,
of
due
to m e n to s e v e s t i t u r e
to
th e
lac k
p resen t,
a id s
A c c o r d in g
lea v e s
c o lle c tio n s
of
The am ount an d
in
from
Ci
on t h e
ste m s,
su ffic ie n t
flo ra l
to
are
ranged
in
type
of
d e te rm in in g
Ci
d ia g n o stic
key
a
Ian c e o la tu s has h isp id
th e
se p a ra te d
The f l o r a l f e a t u r e s v a r y w i t h d i f f e r e n t
19 6 4 ) .
M o n ta n a .
Vi
in flo re sc en c e
Ia n c e o la tu s
to
of
(A n d e rs o n 1 9 8 0 a ) .
en v iro n m en ts
le a v e s
v a ria n ts
h irsu te
h e ig h t
v e stitu re
on
v isc id iflo ru s
by
A nderson
h a i r on t h e s te m s n e a r t h e
to
g lab ro u s.
from
I .4
to
Ci
3 dm.
Vi
ssp.
A ll
c o l l e c t i o n s made d u r i n g
M cA rthur e t a l .
le a v e s
th is
Vl
fla t
( 1 9 7 8 a ) , who r e p o r t e d t h a t
2.5-6mm w i d e ,
1 .5-4cm
8 / 2 0 / Ib had t w i s t i n g l e a v e s .
Cl
s t u d y had
lo n g ,
A lso ,
and
l e a v e s w h ic h a g r e e s w i t h
Cl Vl s s p . I a n c e o l a t u s h a s
fla t.
H ow ever,
c o lle c tio n
s e v e r a l n a rro w l e a v e d s p e c i m e n s o f
s s p . I a n c e o l a t u s w e re f o u n d .
The l e a v e s o f c o l l e c t i o n s 8 / l 6 / 2 a ,
8 / 2 0 / 2 C , 8 / 2 0 / 3 d , a n d . 8 / 3 1 / 5 a a l l m e a s u re d l e s s t h a n 1 .7 mm.
e x a m p le s o f p h e n o t y p i c v a r i a t i o n , w h ic h may r e f l e c t
These a r e
d i f f e r e n t e c o ty p e s
and e n v iro n m en ts.
The
s te m s
of
Cl Vl s s p .
et
a l.
v isc id iflo ru s
a lth o u g h ,
M cA rthur
(1978 a )
n o te s
sp a rse ly
p u b e ru le n t.
th ey a r e
g l a b r o u s a n d r a n g e i n w i d t h fro m I t o
u su a lly
th e
ste m s
are
g lab ro u s,
are
so m e tim e s
The l e a v e s o f t e n h a v e c i l i a t e m a r g i n s , o t h e r w i s e
10 mm.
No Montana
s p e c i m e n s o f Cl Vl s s p . v i s c i d i f l o r u s w e re fo u n d w i t h l e a v e s w i d e r t h a n
3 -5 mm.
A n d e rso n
Cl
Vl
(1 9 8 0 a ) d e s c r i b e s 2 v a r i e t i e s o f Cl Vl s s p . v i s c i d i f l o r u s :
ssp.
v isc id iflo ru s
v isc id iflo ru s v ar.
and
p lan t
h e ig h t.
w id th .
Cl
v isc id iflo ru s
The p r i n c i p a l
ssp.
Vl
in
th e
c h a ra c te ristic
used
c o l l e c t i o n s made f o r
v isc id iflo ru s
v ar.
I
to
v isc id iflo ru s
v isc id iflo ru s
1 9 8 0 a).
1 .5
mm w i d e .
v ar.
v ar.
Cl
R e p o rte d
v isc id iflo ru s
ste n o p h y llu s
C o lle c te d
sp ecim en s
range
is
of
p la n t
up
u su a lly
Cl
Vl
to
Vl
ssp.
I
s tu d y was
has le a v e s
ste n o p h y llu s
h e ig h ts
to
d istin g u ish
th is
v isc id iflo ru s
g r e a t e r t h a n 1 .5 mm a n d Cl , Vl s s p . v i s c i d i f l o r u s v a r .
le a v e s
and
s t e n o p h y l l u s . w h ic h a r e d i s t i n g u i s h e d by l e a f w i d t h
b e tw e e n t h e s e 2 v a r i e t i e s
lea f
v ar.
has
for
Cl
Vl
ssp.
m w h ile
Cl
yV
ssp.
less
th an
3 dm (A n d e rso n
ssp.
v isc id iflo ru s
v ar.
42
v isc id iflo ru s
r a n g e d i n h e i g h t from 1 .5 t o 5 . 5 dm.
C. Vjl s s p . v i s c i d i f l o r u s v a r .
A l l s p e c im e n s o f
s t e n o p h y l l u s w e re l e s s t h a n 3 dm w i t h t h e
e x c e p t i o n o f 8 / 1 6 / Ib ( 6 . 6 dm) a n d 8 / 2 4 / Ia ( 5 . 6 dm ).
A lth o u g h ,
ste m s,
and
V jl
Vl s s p .
8 /1 7 /4 d
ha d
ssp.
v isc id iflo ru s
v is c id iflo ru s v ar.
some h a i r
on t h e
is
rep o rted
to
v isc id iflo ru s
ste m s.
have
g lab ro u s
specim ens 8 / l6 / 2 b
C o lle c tio n s
of
Cl Vl
ssp.
v i s c i d i f l o r u s v a r . s t e n o p h y l l u s ( 8 / l 6 / 4 d a n d B annack 8 /V 1 1 /8 5 ) had some
h i s p id h a i r s on th e
ste m s.
to
b e tw e en
in te rg ra d a tio n
v isc id iflo ru s
T hese s p e c i m e n s may e x p r e s s v e s t i t u r e due
Cjl
vv
ssp.
Ia n c e o la tu s
and
Cl
Vl
ssp.
( L e t t e r from L. C. A n d e rs o n 5 / 2 0 / 8 6 ) , w h ic h a r e sympat r i e
o v e r p a r t o f t h e i r r a n g e s i n Montana ( F i g s . 9 , 1 0 ) .
C h ry so th a m n u s l i n i f o l i u s
C.
Iin ifo liu s
A n d e rs o n 19 8 6 ) .
has
broad,
s m o o th ,
Some s p e c im e n s o f
dark
Montana
averaged
1 .8 3
m ta ll.
r o o t s t h a t fo rm a d v e n t i o u s s h o o t s
(Winward a n d
Cl l i n i f o l i u s grow 3 . 6 6 m t a l l
s o u t h e r n U ta h (Winward a n d A n d e rs o n 1 9 8 6 ) .
in
green le a v e s
in
The few s p e c i m e n s c o l l e c t e d
Cl l i n i f o l i u s
(M cA rthur e t a l .
sp read s
by
1979a).
la te ra l
It is
th is
c h a r a c t e r i s t i c t h a t g i v e s Cl l i n i f o l i u s i t s l a r g e r o u n d f o r m .
D i s t r i b u t i o n o f Montana C h ry so th a m n u s
T he
o n ly
know n
s o u t h w e s t e r n Montana
tax a
o c c u rrin g
in
p o p u la tio n s
(F ig .
3) •
M o n ta n a ;
of
Cl ^ l
ssp.
T able 2 l i s t s
F ig u re s
4 -1 1
g lan d u lo su s
occur
in
t h e o t h e r C hry so th a m n u s
show
th e ir
re sp e c tiv e
'------------
^sam pling s i t e s '
o f Ci 2^. s s p .
I
clandulosus
• sampling s i t e s
<
containing
Bi
ssp . clandulosus
v a r . tomentosus
and v a r . c l a n d u l o s u s
a
l o c a t i o n s wh er e
C. p . s s p . c l a n d u l o s u s
was o b s e r v e d .
F ig u re 3•
L o catio n s o f Chrysothamnus p a r r y i ssp . c la n d u lo su s found i n Montana.
1
* study
collections
Figure 4 .
Location o f c o l l e c t i o n s o f Ci l i n i f o l i u s .
• study
collection s
A Montana S t a t e
U niversity
herbarium
specim ens
Af- U n i v e r s it y of
Montana herbarium
specim ens
Figure 5 .
Location o f c o l l e c t i o n s and herbarium specimens o f (L_ ru s s p . a l b i c a u l i s .
Figure 6.
L o c a t i o n o f c o l l e c t i o n s and h e r b a r i u m s p e c i m e n s o f Ci n^ s s p .
con sim ilis.
•------------
1
-Cr
-a
• study
c o lle c t io n s
a Montana
S ta te
U n iversity
herbarium
specimens
J f U n iv e r s it y o f
Montana herbarium
specimens
Figure 7 .
Location o f c o l l e c t i o n s and herbarium specimens o f Ci n^ s sp . g r a v e o l e n s .
-Cr
OO
• study
c o lle c t io n s
a Montana
S ta te
U n iv ersity
herbarium
specimens
J fU n iversity of
Montana herbarium
specimens
F igu re 8 .
Location o f c o l l e c t i o n s and herbarium specimens o f
n^ s sp . n a u se o su s.
• study
collections
* Montana S t a t e
U niversity
herbarium
specimens
••
f
r
af-U n i v e r s i t y o f
Montana h e r b a r i u m
specimens
Figure 9 .
L ocation o f c o l l e c t i o n s and herbarium specimens o f Cl Vl s sp . I a n c e o l a t u s .
• study
c o lle c t io n s
a Montana
S ta te
U n iversity
herbarium
specimens
JfU n iversity of
Montana herbarium
specimens
F igu re 10.
Location o f c o l l e c t i o n s and herbarium specimens o f
v^ s s p . v i s c i d i f l o r u s .
• study
collections
a
Montana S t a t e
U n iv e r s it y
herbarium
specim ens
F igu re 11.
Location o f c o l l e c t i o n s o f
Vl s s p . v i s c i d i f l o r u s v a r . s t e n o p h y l l u s .
v
rv
• study
collection s
Figure 12.
Location o f c o l l e c t i o n s o f Ci Vi s s p . v i s c i d i f l o r u s var. v i s c i d i f l o r u s .
53
d istrib u tio n s.
H e rb a riu m
sp ecim en s w ith
in su ffic ie n t
in fo rm a tio n
on
l o c a t i o n h a v e b e e n n o t e d a t t h e c o u n ty s e a t .
£ l S i ’ s s P • nau seosus i s
8)•
t h e m ost w i d e s p r e a d t a x o n i n Montana
(F ig .
l i n i f o l i u s i s l i m i t e d t o a sm all a r e a i n t h e so u th c e n t r a l p a r t
of th e S ta te
(F ig 4 ) .
A lth o u g h H a l l a n d C le m e n ts (1 9 2 3 ) l i s t e d Ci Vi
s s p . p u b e r u l u s a s o c c u r r i n g i n Montana ( T a b le 5 ) , no s p e c im e n s o f Ci y \
ssp.
p u b eru lu s
w e re f o u n d .
A n d e rso n ( 1 9 8 6 a ) , Ci
M ontana.
s s P-
A n d e rs o n
Ci
c o lle c te d
Vi
m ore
ssp.
Vi
v ar.
ssp.
map o f
d istrib u tio n
by
p u b eru lu s does not o ccu r a s f a r n o rth a s
(1 9 8 0 b ) i l l u s t r a t e d
v isc id iflo ru s
H ow ever,
A c c o r d in g t o a
t h a t t h e d i s t r i b u t i o n o f Ci
s te n o p h y llu s
v isc id iflo ru s
fre q u e n tly
th an
did
v ar.
Ci
not
e x te n d
s te n o p h y llu s
Vi
ssp.
in to
was
Vi
M ontana.
fo u n d
v isc id iflo ru s
and
v ar.
v isc id iflo ru s.
The m a j o r i t y
o f C h ry so th a m n u s t a x a a r e
o f t h e i r r a n g e i n M o n ta n a , p a r t i c u l a r l y
11).
Ci
Iii
ssp.
n a u s e o s u s a n d Ci
g rav eo len s a p p e a rs
lin ifo liu s
sy m p atric o v er a l l
in th e w e stern h a l f (F ig s. 3-
sym p atric
(F ig . 4 , 7 , 8 ) .
i s s y m p a t r i c o v e r p a r t o f t h e r a n g e o f Ci
or p a rt
w ith
o n ly
Ci
Qi
ssp.
A d d i t i o n a l l y , Ci l i n i f o l i u s
Qi
ss p . n auseosus (F ig . 4 , 8 ) .
H a b i t a t o f Montana C h ry so th a m n u s
C h ry so th a m n u s p a r r y i s s p . g l a n d u l o s u s
C h a ra c te ristic a lly
p o p u la tio n s.
Ci
The 7 s i t e s
Qi
ssp.
g lan d u lo su s
sa m pled a v e r a g i n g
w as
85 m2 i n
S i t e s w ere g r a s s y o p e n i n g s among A r t e m i s i a t r i d e n t a t a
fo u n d
siz e
in
sm all
( T a b le 5 ) .
- g r a s s la n d ty p e
T ab le 5 .
L e g a l d e s c r i p t i o n s a n d r a n g e S i t e s a f o r 7 C h ry s o th a m n u s p a r r y ! s s p . g l a n d u l o s u s a n d
sam ple p l o t s i z e .
S ite
County
Legal
d e sc rip tio n
Range s i t e
P lo t s iz e
(m)
B u r e a u o f Land
Management
B e a v e rh e a d
NW 1 / 4 , S e c . I ,
T1S, RIIW
s i l t y r a n g e s i t e 1 5 -1 9 " P. Z.
4 .0 x 1 0 .5
Coal C reek
M adison
NE 1 / 4 , S e c . 2 9 ,
. Tl I S , R3W
s i l t y r a n g e s i t e 2 0 - 2 4 " P. Z.
5 .8 x 6 .8
C e n te n n ia l V a lle y
B e a v e rh e a d
SW 1 / 4 , S e c . 5 ,
T13S, R4W
s ilty
range s i t e
1 0 -14" P . Z.
5 .0 x 8 .5
H e le n L a r s o n
B e a v e rh e a d
SW 1 / 4 , S e c . 6
T6S, R14W
silty
range s i t e
1 5 -1 9 " P . Z.
6«i0x10
Jerry
S ilv e rb o w
NW 1 / 4 , S e c . 30
TIN, RIOW
s i l t y r a n g e s i t e 1 5 -1 9 " P . Z.
b o r d e r i n g on s u b a l p i n e f i r ,
Dou g l a s - f i r c li m a x f o r e s t s on
deep s o i l s w ith c r y i c tem per­
a t u r e r e g i m e s a n d p a l e brown d r
l i g h t b r o w n is h g r a y s u r f a c e s on
m o d era te ly s te e p t o v e ry s te e p
s l o p e s 2 0 - 4 5 " P. Z.
C reek
4 .0 x 1 3 .7
Table 5 .
continued
S ite
County
Q u a rtz H i l l
B e a v e rh e a d
R iv ersid e
B e a v e rh e a d
Legal
d e sc rip tio n
NE 1 / 4 , S e c . 8
T1S, RIOW
NW 1 / 4 , S e c . I
H S , RIlW
Range s i t e
P lo t s iz e
(m)
s i l t y r a n g e s i t e 1 5 -1 9 " P. Z.
b o r d e r i n g on s u b a l p i n e f i r ,
D o u g la s - f i r c li m a x f o r e s t s on
deep s o i l s w ith c r y i c tem p er­
a t u r e r e g i m e s a n d p a l e brown o r
l i g h t b r o w n is h g r a y s u r f a c e s on
m o d era te ly s te e p t o v e ry s te e p
s l o p e s 2 0 - 4 5 " P . Z.
8 .0 x 1 0
s i l t y r a n g e s i t e 1 5 - 1 9 " P. Z.
8 .0 x 1 6 .4
T ab le 6 .
H a b i t a t d e s c r i p t i o n f o r 7 C h rv s o th a m n u s p a r r y i s s p . K l a n d u l o s u s s a m p l in g s i t e s
S ite
R iv ersid e
taxon p lo t
E le v atio n
(m)
1708
A nnual
Ppt
1739
J e r r y C reek
taxon p l o t
p a ire d p lo t
1769
C e n te n n ia l
V a lle y
taxon p l o t
p a ire d p lo t
1830
S o il
D epth
406
p a ire d p lo t
Q u a rtz H i l l
taxon p l o t
p a ire d p lo t
S o il
T ex tu re
sandyIoam
sandyIoam
sh a llo w
loam
sandyIoam
n o rm a l
norm al
loam
loam
n o rm a l
norm al
sandyIoam
sandyIoam
n o rm a l
A spect
S lo p e
(%)
Type o f
d istu rb an ce3
east
6-10
r,f,d s
lev e l
0-2
r
lev el
0-2
r
lev el
0-2
r
sh a llo w
419
432
■ 356
n o rm a l
T able 6 .
con tinued
S ite
B u r e a u o f Land
Management
E le v a tio n
W
1983
A nnual
Ppt
2135
588
sandyIoam
Ioam s a n d y - lo a m
p a ire d p lo t
S lope
(%)
Type o f
d istu rb an ce3
so u th ­
w est
3-5
g
w est
10-20
wr
2227
w est
3-5
norm al
norm al
£
p a ire d p lo t
A sp e c t
sandyn o rm a l
Icam
Ioam n o rm a l
s a n d y -lo a m
. p a ire d p lo t
H e le n L a r s o n
taxon p lo t
S o il
D epth
406
taxon p lo t
Coal Creek
tax o n p lo t
S o il
T ex tu re
sandyloam
sandyIoam
sh a llo w
sh a llo w
g
58
c o m m u n it i e s .
E le v a tio n s' a t
th e
7 site s
ranged
fro m
1788 t o
2227
m
( 5862-7300 f t ) a n d a n n u a l p r e c i p i t a t i o n v a r i e d from 356 t o 588 mm ( 1 4 23 i n )
(T ab le 6) .
S l o p e s r a n g e d f ro m 0-20% a n d a s p e c t s v a r i e d
( T a b le
6).
S o il
pH a v e r a g e d
(T ab le 7 ) .
th e
b e tw e e n
T h e r e w as no d i f f e r e n c e i n
p a ire d
p lo ts.
E le c tric a l
i n d i c a t i n g low c o n t e n t o f s o l u b l e
in
so lu b le s a l t
2 e x c e p tio n s
w ith
d e p th
pH b e tw e e n t h e t a x o n p l o t s a n d
c o n d u c tiv itie s
sa lts.
w e re a l l
very
lo w ,
T h e re was no r e a l d i f f e r e n c e
( T a b le 8 ) .
sa ndy loam i n t e x t u r e t h r o u g h o u t t h e i r p r o f i l e
(T ab le 6 ) .
Land lfe n a g e m e n t, t e x t u r e s w ere
taxon p l o t s .
s lig h tly
c o n t e n t b e tw e e n t h e t a x o n a n d p a i r e d p l o t s
A ll s i t e s w e re loam y t o
w ith
8 , in cre asin g
7 and
At 2 s i t e s ,
s ilt
Jerry
loam a t
th e
C re e k a n d B u re a u o f
6-12"
d e p t h s on t h e
S o il c h a r a c t e r i s t i c s m easured p ro v id e d l i t t l e
in sig h t a s
t o w h at e d a p h i c f a c t o r ( s ) m ig h t be d e t e r m i n i n g o c c u r r e n c e o f Ci
ssp.
g lan d u lo su s.
T ab le 7 .
S o i l D epth
pH m eans ( r a n g e s ) o f s o i l s from 7 C h ry s o th a m n u s
p a r r y i s s p . g lan d u lo su s c o ll e c t i o n s i t e s .
C. p . s s p . g l a n d u l o s u s p l o t
p a ire d p lo t
0 - 2 in
7 . 1 3 ( 6.01 - 8 .1 7 )
7 .2 8 ( 6 . 0 6 - 8 . 3 4 )
2 - 6 in
7 .5 3 ( 6 .2 2 - 8 .8 7 )
6. - 12 i n
7-82 (6 .3 9 - 8 .8 7 )
;
;
7 . 7 3 ( 6 .3 0 - 8 . 7 8 )
7 . 8 3 ( 6 . 3 9 - 9 .2 0 )
59
T ab le 8 .
S o i l e l e c t r i c a l c o n d u c t i v i t y (mmhos) m eans ( r a n g e ) fro m 7
C h ry so th m n u s p a r r y ! s s p . g l a n d u l o s u s c o l l e c t i o n s i t e s .
S o i l D epth
C. p . s s p . g l a n d u l o s u s p l o t
p a ire d p lo t
...
0 -2 i n
0 .7 2 ( 0 .2 - 1 . 7 )
0 .51
2 - 6 in
0 .3 1 ( 0.1 - 0 . 7 )
0 . 3 0 ( 0 .1 - 0 . 5 )
6 - 12 i n
0 . 3 4 ( 0.1 - 0 . 7 )
0 . 3 0 ( 0 .1 - 0 . 5 )
C o m m unities
co v erag e,
common
w ith
Cr
ssp.
( 0 . 3 - 1 . 1)
13% g r a s s b a s a l
g lan d u lo su s averag ed
9% s h r u b
ca n o p y
co v erag e,
a sso c ia te d
w oody
sp e c ie s
and
17% b a r e
in c lu d e d
ground.
A rtem isia
The m o st
frig id a .
t r i d e n t a t a . Ar t r i p a r t i t a . Cr ^ n a u s e o s u s . a n d C. v i s c i d i f l o r u s .
c o lle c tio n
site s
w ere
silty
m o u n ta in s g e o g ra p h ic a l
(T ab le 5 ) .
co v erag e,
As w i t h
range
area,
so il
but
site s
lo c a te d
in
th e
A.
A ll 7
f o o t h i l l s and
had v a r y i n g a m o u n ts o f p r e c i p i t a t i o n
p a ra m ete rs,
p lan t
p aram eters o f g ra s s
s h r u b c a n o p y c o v e r a g e a n d b a r e g ro u n d a l s o
basal
f a i l e d to e x p la in
d iffe re n c es.
D istu rb a n c e
had
o ccu rred
at
a ll
7
s ite s
on
th e
Cr
g la n d u lo s u s p l o t s and t h e a s s o c i a t e d p a i r e d p l o t s (T a b le 6 ) .
except
for
th e
th e se
site s,
Coal
C reek s i t e
Q u a rtz
H ill,
w e r e fo u n d c l o s e
Jerry
C reek,
to a
road.
C e n te n n ia l
ssp.
A ll s i t e s
Four o f
V a lle y ,
and
R i v e r s i d e , a p p e a r e d d i s t u r b e d a s a r e s u l t o f m e c h a n ic a l m eans, p o s s i b l y
p a rk in g l o t s
f o r heavy
e q u ip m e n t.
The 2 o t h e r
road
site
lo c a tio n s,
H e le n L a r s o n Ranch and B u re a u o f Land M anagem ent, w e re h e a v i l y u s e d by
c a ttle .
The r e m a i n i n g l o c a t i o n ,
t h e summit o f a r i d g e .
Coal
C reek,
was a
b lo w o u t
site
near
60
A c o lo n i z e r i s any ta x o n h a v in g a n a b i l i t y
as
a
re su lt
appears to
fa c to r
of
human o r
be a
stra te g y
in v o lv e d
C h ry so th a m n u s
c o m m u n itie s
m em b ers
n a tu ra l
in
tax a
are
th e
(S te b b in s
ssp . g lan d u lo su s.
o ccu rren ce
re c o g n ize d
(M cA rthur e t a l .
of
d istu rb an c e
o f Cl j x
its
to e s t a b l i s h and sp read
and
of
ex p ress
O th e r
d istu rb ed
197 8b , M cA rthur a n d W elch
genus
T his
D istu rb a n c e i s a
d istrib u tio n .
c o lo n iz e rs
C h ry so th a m n u s
1965).
p lan t
19 8 6 ) .
th e
Known
fo llo w in g
c h a r a c t e r i s t i c s t h a t B a k e r (19 6 5 ) l i s t s a s c o n t r i b u t i n g t o t h e s u c c e s s
of
c o lo n iz in g
p ro d u ctio n
lo n g and
and
in
sh o rt
e d a p h ic
seed lin g
sp e c ie s:
fa c u lta tiv e
fa v o ra b le
e n v iro n m e n ta l
d ista n c e
v a ria tio n ;
d isp e rsa l;
v ig o ro u s
e sta b lish m e n t.
p ro d u ctio n
stra te g ie s
Cl
c o m p a tib ility ;
c o n d itio n s;
p h e n o ty p ic
v e g e ta tiv e
The r a t e
of
se lf
of
ssp.
seeds
h ig h
seed
e q u ip p e d
p la stic ity
to
re p ro d u c tio n ;
for
c lim a tic
and
ra p id
se e d lin g e s ta b lis h m e n t and seed
g lan d u lo su s
are
not
yet
know n,
how ever, t h i s tax o n d o es e x p r e s s th e o t h e r c h a r a c t e r i s t i c s .
An e x p l a n a t i o n f o r t h e o c c u r r e n c e o f Cl ^ l s s p . g l a n d u l o s u s i n s m a l l
i s o l a t e d p o p u l a t i o n s , b u t i t s a b s e n c e on s i m i l a r a d j a c e n t s i t e s ,
be t h a t i t
g razin g .
has stro n g c o lo n iz in g a b i l i t y ,
Cl ^ l s s p .
s e n s i t i v e to e x tre m e
g l a n d u l o s u s a p p e a r s t o r e q u i r e medium t o l i g h t e r
te x tu re d , n e a r-n e u tra l s o ils ,
8) .
but i s
m ig h t
t h a t a r e low i n s o l u b l e s a l t s
P erhaps i t i s a b le to e x p lo i t th e s e
( T a b l e s 6-
s i t e s when t h e y a r e o p e n e d up
by d i s t u r b a n c e s s u c h a s c o m p a c tio n a n d v e g e t a t i o n r e m o v a l due t o h e a v y
m a c h in e r y
tra m p lin g ,
it
tra ffic ,
heavy
o r by s o i l
g ra z in g
and
b ro w sin g
l a y e r re m o v a l by w in d .
s p r e a d s v i g o r o u s l y by u n d e r g r o u n d r o o t s .
w ith
Once i t
a c co m p a n y in g
i s e sta b lish e d ,
E v i d e n c e w as a p p a r e n t a t
e a c h s t u d y s i t e t h a t t h e t a x o n i s b ro w s e d h e a v i l y ( n e a r 100 % o f c u r r e n t
61
y e a r s g r o w t h ) e a c h y e a r ( T a b le 9 ) • T h i s may e x p l a i n why t h i s t a x o n h a s
n o t been p r e v io u s ly r e c o g n iz e d .
lim itin g
fu rth e r
ex p a n sio n
S e v e r e b r o w s in g o f t h i s t a x o n m ig h t be
a n d /o r
e lim in a tin g
it
fro m
a d ja c e n t
d istu rb ed s i t e s .
It
is
p o ssib le
th at
th e
site
p aram eters
m easured
s a m p l in g t e c h n i q u e s em ployed d i d n o t i s o l a t e
of
fa c to rs
successfu l
d e te rm in in g
c o lo n iz e r
s u c c e ssio n a l
it
sp ecies.
th e
lo c a tio n
may l a c k
of
or
perhaps
th e
th e f a c t o r o r c o m b in a tio n
th is
th e a b i l i t y
tax o n .
to
S in c e
it
c o m p e te w i t h
The s a m p l in g em ployed d i d n o t
is
a
h ig h er
d e te c t advanced
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 a s s o c i a t e d p a i r e d p l o t s com pared t o t h e
new t a x o n p l o t s .
T ab le 9 .
P e r c e n t o f C h ry so th a m n u s p a r r y i s s p . g l a n d u l o s u s p l a n t s
b ro w se d a t 7 s a m p l in g s i t e s .
s ite
P l a n t s brow sed
(in c lu d es past
and t h i s season)
P l a n t s bro w se d by
raid-sum m er, 1986
Ia
2b
100
100
40
a B u re a u o f Land Management
^C oal C reek
° C e n te n n ia l V a lle y
^ H e le n L a r s o n Ranch
e J e r r y C reek
f Q u a r tz H i l l
^ R iv e rsid e
■■■
3°
4d
5e
6f
76
97
100
91
80
98
3
86
5
■we
7
62
C h ry s o th a m n u s n a u s e o s u s
In
Montana
nauseosus
su b sp ecies
w e re
( F i g s . 5 - 8 ) on a v a r i e t y o f h a b i t a t s .
nauseosus i s
in
Montana on t h e
C le m e n ts 1 9 2 3 ) •
s o ils
ran g in g
a sp e c ts.
S p e c im e n s o f
fro m
c la y
E stim a te d
to
annual
fo u n d
w id e ly
d istrib u te d
The t y p e s p e c im e n o f Ci n ^ s s p .
banks o f th e M isso u ri R iv er (H a ll and
Ci
Qi
sandy
ssp.
loam
n a u s e o s u s w e re c o l l e c t e d on
on a
p re c ip ita tio n
on
v a rie ty
of
c o lle c tio n
slo p e s
site s
and
ran g ed
fro m 280 t o 510 mm ( 1 1 - 2 0 i n ) a n d e l e v a t i o n s r a n g e d f r o m 700 t o 2300 m
(2 3 0 0 -7 5 0 0 f t ) .
-
Ci
P se u d o ro e g n e ria
d o m in a te d
sp ic a ta
c o m m u n itie s;
stric ta .
In
a sso c ia te d
w ith
S p e c im e n s w e re
a lo n g
s s p . n a u s e o s u s was fo u n d i n A r t e m i s i a t r i d e n t a t a
Qi
and
On
so u th e a ste rn
ro a d sid e s.
M on ta n a ,
c o lle c te d
It
trid e n ta ta
o ccasio n
S ch izach y riu m
o fte n
Ai
was
it
Ci
was
s c o pariurn an d
in
se v e re ly
found
on
B o u t e lo u a
fo u n d
ssp.
Qi
-
w ith
D istic h lis
nauseosus
C a la m o v ilfa
o v ergrazed
many
g ra c ilis
ran g e
was
found
lo n g ifo lia .
p a stu re s
s ite s
in
and
a ll
5
g e o g r a p h i c a l a r e a s o f Montana ( T a b le 1 0 ) .
Ci
Qi
ssp.
g r a v e o l e n s was fo u n d o n s i t e s w i t h a v a r i e t y
a n d a s p e c t s , a t e l e v a t i o n s fro m 640 t o
of slo p e s
1500 m ( 2 1 0 0 -5 0 0 0 f t ) .
Annual
p r e c i p i t a t i o n on t h e s e s i t e s r a n g e d fro m 280 t o 380 mm ( 1 1 - 1 5 i n ) .
was o f t e n
and i n
th e
a sso c ia te d
w ith
S arco b atu s
so u th e a ste rn p a rt of
v e rm ic u la tu s and A trio le x
th e S ta te w ith
A rtem isia
It
s p p .,
trid e n ta ta
■-
a n d B o u t e lo u a
g ra c ilis.
o v e r l a p s t h a t o f Ci
fo u n d
w ith
Qi
S arco b atu s
A lth o u g h
th e
range
of
Ci
s s p . g r a v e o l e n s . r a r e l y was Ci
v e rm ic u la tu s.
g r a v e o l e n s i s m ore s a l t t o l e r a n t t h a n Ci
T his
Qi
su g g e sts
Qi
Qi
ssp.
nauseosus
ssp . nauseosus
th at
ssp. nauseosus.
Ci
Qi
ssp.
T ab le 10.
Range s i t e s 3 o f C h ry s o th a m n u s t a x a i n M o n ta n a .
G e o g ra p h ic a l a r e a and ran g e s i t e
. E a s te rn G la c ia te d P la in s
r"
O c c u rrin g tax a
~
"
~
s i l t y - c l a y e y r a n g e s i t e c o m plex 1 0 -1 4 " P . Z.
C. n . s s p . g r a v e o l e n s
C. n . ssp.. n a u s e o s u s
W estern G la c ia te d P l a i n s
silty
range s i t e ,
1 0 -1 4 " P . Z.
C. n . s s p . n a u s e o s u s
s i l t y - c l a y e y r a n g e s i t e c o m p le x , 1 0 - 1 4 " P . Z.
c l a y e y an d s h a l l o w c l a y r a n g e s i t e a s s o c i a t i o n ,
C. h . s s p . n a u s e o s u s
1 0 -1 4 " P . Z.
C. n . s s p . n a u s e o s u s
E a s t e r n S e d i m e n ta r y P l a i n s
. silty
range s i t e ,
1 0 -1 4 " P . Z.
C. n . s s p . K r a v e o l e n s
C1. n j _ s s p . n a u s e o s u s
s i l t y - c l a y e y r a n g e s i t e c o m p le x , 1 5 -1 9 " P . Z.
C. n . s s p . n a u s e o s u s
c l a y e y an d s h a l l o w r a n g e s i t e a s s o c i a t i o n ,
C. n . s s p . K r a v e b l e n s
C. n . s s p . n a u s e o s u s
1 0 -1 4 " P . Z.
r i v e r b r e a k s , 1 0 -1 4 " P . Z.
C. n . s s p . g r a v e o l e n s
C. n . s s p . n a u s e o s u s
b a d l a n d s , 1 0 -1 4 " P . Z.
C. n . s s p . g r a v e o l e n s
T able 10.
con tin u ed
G eo g ra p h ic a l a r e a and ran g e s i t e
O ccu rrin g tax a
W e s te r n S e d i m e n ta r y P l a i n s
c l a y e y an d s h a l l o w c l a y r a n g e s i t e a s s o c i a t i o n 1 0 -1 4 " P . Z.
C. n . s s p . g r a v e o l e n s
C. n . s s p . n a u s e o s u s
c l a y e y a n d s h a l l o w c l a y r a n g e s i t e a s s o c i a t i o n 1 5 -1 9 " P . Z.
C. n . s s p . n a u s e o s u s
d e n s e c l a y - c l a y e y - s a l i n e u p l a n d r a n g e s i t e c o m p le x , 5 - 9 " P . Z.
C. I i n i f o l i u s
C. n . s s p . g r a v e o l e n s
r i v e r b r e a k s . 10 - 1 4 " P . Z.
Cl n ^ s s p . g r a v e o l e n s
F o o t h i l l s a n d M o u n ta in s
silty
range s i t e
1 0 -1 4 " P . Z.
C. n . s s p . a l b i c a u l i s
Cl IIl SSp. c o n s i m i l i s
Cl Ul s s p . n a u s e o s u s
Cl Pl s s p . g l a n d u l o s u s
Cl Vl s s p . l a n c e o l a t u s
Cl Vl ssp . v is c ld if lo r u s
var. sten o p h vllus
var. v i s c id f I orus
s ilty
range s i t e
1 5 -1 9 " P . Z.
Cl
Cl
Cl
Cl
Cl
Iil ssp .
Ql ssp .
Ql ssp .
Ql . ssp .
Vl ssp .
a lb ic a u lis
c o n s im ilis
nauseosus
glandulosus
la n c e o la tu s
Table 10.
con tin u ed
G e o g ra p h ic a l a r e a an d ran g e s i t e
O c c u rrin g taxa
F o o t h i l l s a n d M o u n ta in s
silty
range s i t e
1 5 -1 9 " P . Z.
C. y . s s p . v i s c i d i f l o r u s
v a r . s t e n o D h v llu s
v ar. v isc id flo ru s
silty
r a n g e s i t e 2 0 - 2 4 " P . Z.
C. n_. s s p . n a u s e o s'us
C. p . s s p . p a r r y i
C_!_ y_. s s p . v i s c i d i f l o r u s
v a r . s t e n o D h v llu s
' s i l t y - c l a y e y r a n g e s i t e c o m p le x , 1 0 -1 4 " P . Z.
Gi. n_. s s p . a l b i c a u l i s
C. n , s s p . n a u s e o s u s
C_!_ y , s s p . v i s c i d i f l o r u s
v a r . s t e n o D h v llu sv ar. v is c id iflo ru s
s i l t y - c l a y e y r a n g e s i t e c o m p le x , 1 5 -1 9 " P . Z.
C.
Ci
C.
C.
n. ssp . a lb ic a u lis
n. s s p . g r a v e d ens
n. s s p . nauseosus
v, ssp . v is c id if lo r u s
v ar. v isc id iflo ru s
l i m y - s h a l I o w -v e ry s h a l l o w r a n g e s i t e
Ci
Ci
Ci
Ci
n. ssp . a lb ic a u lis
n. s s p . nauseosus
v. s s p . Ia n c e o la tU s
v. ssp. v isc id iflo ru s
v a r. ste n o p h v llu s
1 0 -1 4 " P . Z.
cr,
Table 10.
c o n tin u ed .
G eo g ra p h ic a l a re a and ran g e s i t e
O c c u rrin g tax a
F o o t h i l l s a n d M o u n ta in s
s u b a lp in e f i r , d o u g l a s - f i r . a n d p o n d ero sa p in e clim a x f o r e s t s
on d e e p s o i l s w i t h c r y i c o r f r i g i d t e m p e r a t u r e r e g i m e s a n d
g r a y i s h bro w n , l i g h t b r o w n is h g r a y , o r l i g h t y e l l o w i s h brown
s u r f a c e s on m o d e r a t e l y s t e e p t o v e r y s t e e p m o u n ta in s l o p e s
w e s t o f t h e c o n t i n e n t i a l d i v i d e 1 6 -7 0 " P . Z.
£i_ O i s s p . a l b i c a u l i s
Ci Qi s s p . c o n s i m i l i s
67
The t y p e l o c a l i t y
M isso u ri
R iv er i n
of
m
sa p . g r a v e o le n s i s on t h e banks o f th e
denuded s o i l
( H a l l an d C le m e n ts 1 9 2 3 ) .
g r a v e o l e n s was m o st o f t e n fo u n d on s i t e s w i t h h i g h l y
d e v e lo p e d
in clu d e
so ils,
h ig h ly
How ever,
it
c la y
to
sa n d
e ro d ib le
was a l s o
in
b a d la n d
te x tu re .
s o ils,
n^ ssp.
eroded o r p o o rly
E xam ples
d ra in a g e s,
of
su ch
and
site s
re a d o u ts.
fo u n d i n d e e p e r s o i l s s u c h a s c l a y b r e a k s o f t h e
M i s s o u r i r i v e r a n d r e c e d e d s h o r e l i n e s o f F o r t P eck l a k e .
Cr Hr s s p .
g r a v e o l e n s w as f o u n d on s e v e r a l
range s i t e s i n t h e w e s te rn and e a s t e r n
se d im e n ta ry
in
p l a i n s and
lo c a lly
th e
so u th w e ste rn e a s t e r n
g lac iate d
p l a i n s (T a b le 1 0 ).
C. n . s s p . a l b i c a u l i s i s a f o o t h i l l
1975»
A nderson
19 8 6 c ).
Montana w e re fo u n d a t
in
an d m o u n ta in t y p e (Hanks e t a l .
H e rb a riu m
specim ens and
71.0 mm ( 1 1 - 2 8 i n ) a n n u a l
Most C h ry s o th a m n u s grow i n b a s i c s o i l s ,
so ils
from
s i t e s r a n g i n g fro m 732 t o 2440 m ( 2 4 0 0 -8 0 0 0 f t )
e l e v a t i o n a n d from 290 t o
o c c u r on a c i d
c o lle c tio n s
(Plummer 1 9 7 7 ) .
b u t Cr
Iir
p re c ip ita tio n .
s s p . a l b i c a u l i s may
A l l t h e s p e c i m e n s o f Cr Hr s s p .
a l b i c a u l i s c o l l e c t e d w e re fo u n d o n medium t o l i g h t t e x t u r e d s o i l s ( s i l t
to
san d ).
One h e r b a r i u m
s p e c im e n n o t e d a
c o ll e c t i o n s i t e w ith a c la y
loam s o i l .
C.
n.
ssp.
P se u d o ro e g n e ria
o ana, S tip a
was
a lb ic a u lis
sp ic a ta ,
occurred
A.
in
a sso c ia tio n
t r i d e n t a t a . F e stu c a
c o m a t a , an d O r y z o p s i s h y m e n o i d e s .
c o lle c te d
w ith
R hus
trilo b a te ,
P u rsh ia
w ith
su c h
tax a
as
i d a h o e n s i s . A rtem isia
On m ore m e s i c s i t e s i t
trid e n ta ta .
an d
Prunus
V irg in ia n a.
Cr Qr s s p . a l b i c a u l i s was fo u n d on s e v e r a l r a n g e s i t e s b u t
was l i m i t e d
to
10 ) .
th e
fo o th ills
and m o u n ta in s g e o g ra p h ic a l
area
( T a b le
68
Few
sp e cim en s
h erb ariu m
m o u n ts o f
c o n sim ilis
a lk a li
of
Cjl Hjl s s p .
th is
c h a ra c te ristic a lly
m ig h t
be
re stric tiv e
g r a v e o l e n s (Hanks e t a l .
a sso c ia tio n
a l.
taxon
w ith
1985).
c o n sim ilis
w e re
lo ca ted .
to
Ci
1975).
th is
c o lle c te d
In
U ta h ,
Ci
and
n.
no
ssp.
g ro w s .o n d r y l o w l a n d s w h e r e d r o u g h t a n d
Iii
ssp.
a l b i c a u l i s a n d Ci
O f t e n Ci Iii s s p .
S arco b atu s v e rm ic u la tu s
However,
w e re
was n o t
th e
in
ssp.
c o n s im ilis occurs in
on a l k a l i n e
case
Iii
fla ts
(Weber e t
M on ta n a .
Ci
Qi
ssp.
c o n s i m i l i s w as m ost commonly fo u n d a s s o c i a t e d w i t h A r t e m i s i a t r i d e n t a t a
and
P se u d o ro e g n e ria s p i c a t a .
Ci
Qi
annual
ssp.
p re c ip ita tio n
e le v a tio n .
a sp ec ts.
ha d
c o n sim ilis s i t e s
1373
to
1 912
m (4 500-6270
A n d e rso n (1 9 8 1 ) r e p o r t e d t h a t s o i l s o f Ci
to
fro m 5 .6
250 mmhos/cm.
fo u n d on a few r a n g e
area
fro m
ft)
in
I t w as f o u n d on s i l t t o loam s o i l s w i t h v a r i a b l e s l o p e s a n d
pHs r a n g i n g
from 0 . 2
and
r a n g e d from 280 t o 420 mm ( 1 1 - 1 6 . 5 i n )
site s
8 .6 a n d e l e c t r i c a l
to
( A n d e rs o n
1981).
Ci
ssp . c o n sim ilis
Qi
c o n d u c tiv ity
Qi
ssp.
ran g in g
c o n s i m i l i s was
in
th e
f o o t h i l l s and m o u n tain s g e o g ra p h ic
of
th e
m ost w i d e l y
(T ab le 1 0 ).
C h ry so th a m n u s v i s c i d i f l o r u s
Ci
v isc id iflo ru s
is
one
d istrib u te d
shrubs
on
r a n g e l a n d s t h r o u g h o u t N o r th America a n d h a s g r e a t e c o l o g i c a l a m p l i t u d e
(A n d e rs o n
h a b ita ts.
1986 b ) .
w estern
Montana
it
in h a b its
H e rb a riu m m ounts a n d c o l l e c t i o n s o f Ci
fro m M ontana h a v e
e le v a tio n .
In
been found
fro m
1190 t o
Vi
a
w id e
ssp.
range
of
Ian c e o la tu s
2380 m ( 3 9 0 0 -7 8 0 0
ft)
in
O f t e n t h i s t y p e was f o u n d i n h e a v i l y g r a z e d p a s t u r e s w i t h
69
A rtem isia
trid e n ta ta
-
Poa_ s p p . ,
A rtem isia
frig id a
-
A rtem isia
f i l i f o lia -
P s e u d o ro e g n e ria s p i c a t a . and K o e le ria p y ram id ata - F e stu o a
id ah o en sis -
S tip a
spp.
t o 510 mm ( 1 2 - 2 0 i n ) .
P r e c i p i t a t i o n on th e s e
s i t e s r a n g e d fro m 300
C o l l e c t i o n s d u r i n g t h i s s t u d y w ere made o n l y on
l i g h t t e x t u r e d s o i l s r a n g i n g from s i l t t o s a n d .
M ounted s p e c i m e n s a c c .
# 14391 (MONTU) a n d a c c . # 45706 (MONT) r e p o r t e d c l a y loam s o i l s .
t a x o n was l i m i t e d t o a few r a n g e
site s
in
T h is
th e f o o t h i l l s and m o u n tain s
g e o g ra p h ic a l r e g io n (T ab le 1 0 ).
Vl s s p . v l s c i d i f l o r u s i s one o f t h e m ost w i d e s p r e a d t a x a i n t h e
genus
arid
( A n d e rs o n 1 9 8 6 b ) .
d istu rb ed
rocks
(H a ll
s ite s
and
(M cA rthur e t a l .
v.
ssp.
1986b).
It
is
r e p o r t e d t o be w e l l s u i t e d on m o i s t t o
(A n d e rs o n 1 9 8 6 b ) , d r y p l a i n s a n d h i l l s i d e s
C le m e n ts
1978a).
v isc id iflo ru s
1923) ,
and
from
te ste d
ranged
in
w ere k e y e d o n l y t o t h e s u b s p e c i e s l e v e l .
re p o rte d
silty
to
sandy
e x c e p t i o n s , a c c . # 20053 (MONT), a c c .
broader le a v e s lik e
to
h ig h
e le v a tio n s
A n d e rso n (1 9 8 6 b ) r e p o r t s t h e few s o i l s o f Cl
pH from
H e rb a riu m s p e c im e n s o f Cl Vl s s p .
in fo rm a tio n
low
among
to
v isc id iflo ru s
8 .4
(A n d e rs o n
from Montana
Most s p e c i m e n s t h a t h a d s o i l
te x tu re d
s o ils
# 44715 (MONT)
Cl Vl s s p . v i s c i d i f l o r u s v a r .
# 45107 (MONT), a n d a c c . # 44726
6
w ith
a
fe w
(n o te d a s h a v in g
v isc id iflo ru s) . acc.
(MONT), w h ic h w e re n o t e d t o
be fo u n d
on c l a y e y s o i l s .
C o llec ted
fu rth e r
sp ecim en s o f
keyed
to
th e
Cl Vl s s p .
v a rie ty .
Cl
v isc id iflo ru s
Vl s s p .
fro m Montana w e re
v isc id iflo ru s
v ar.
v i s c i d i f l o r u s w as c o l l e c t e d on s i t e s w i t h s i l t t o sa n d y loam s o i l s w i t h
a v a rie ty
of s lo p e s and a s p e c t s .
P r e c i p i t a t i o n r a n g e d on t h e s e s i t e s
fro m 330 t o 460 mm ( 1 3 - 1 8 i n ) a n d e l e v a t i o n r a n g e d f r o m . 1403 t o 2104 m
70
( 4 6 0 0 -6 9 0 0 f t ) .
Cl V l s s p . v l s c i d i f l o r u s v a r .
a sso c ia te d
w ith
c in e re u s♦
I t w as f o u n d t o o c c u r i n f o o t h i l l an d m o u n t a i n s g e o g r a p h i c a l
area
Rosa s p p . , A r t e m i s i a
v i s c i d i f l o r u s was f o u n d
t r i d e n t a t a . Poa s p p . , and Levmus
Cl Vl s s p . v i s c i d i f l o r u s v a r .
(T a b le 1 0 ) .
s te n o p h y llu s specim ens
w ere c o l l e c t e d on v a r i a b l e s l o p e s a n d a s p e c t s o f l i g h t t e x t u r e d s o i l s .
T h is
ta x o n
w as
P se u d o ro e g n e ria
found
o c c u rrin g
sp ic a ta .
e l e v a t i o n s fro m 1220 t o
S tip a
w ith
co m a ta ,
A rte m isia
and
trid e n ta ta -
P a scopyrum
sm ith ii
2196 m (4 0 0 0 -7 2 0 0 f t ) a n d p r e c i p i t a t i o n
from 310 t o 510 mm ( 1 3 - 2 0 i n ) y e a r l y .
at
zones
I t was l o c a t e d o n s e v e r a l r a n g e
s i t e s i n t h e f o o t h i l l s a n d m o u n ta in s g e o g r a p h i c a l r e g i o n
(T ab le 1 0 ).
C h ry so th a m n u s l i n i f o l i u s
In
M on ta n a ,
lo c a tio n .
specim ens
of
Cl l i n i f o l i u s
w ere
c o lle c te d
T h i s l o c a t i o n was d i s t u r b e d a n d l i e s
County i n h a b i t e d by s a l t d e s e r t s h r u b s p e c i e s .
o th er
s ta te s as
favor
sim ila r
g r o w in g i n a l k a l i n e a n d
site s
in
M ontana.
th e w e s te rn s e d im e n ta ry p l a i n s
in
a
p art
at
o n ly
of
I
C arbon
T h i s t a x o n i s known i n
sandy s o i l s .
It
I t was fo u n d i n o n l y
appears
to
I range s i t e
in
(T a b le 1 0 ).
O v e rla p p in g ta x a
S ev eral
h a b ita t
C h ry s o th a m n u s
req u irem e n ts
ex p ress th e a b i l i t y
T h is
a b ility
to
e n a b le s
tax a
( T a b le
in
Montana
have
o v e rla p p in g
e v id e n c e d
above,
i n h a b i t and a d a p t t o
a v a rie ty
of c o n d itio n s.
tax a
And,
to
as
th e se
11).
appear
to
d isp e rse
to
new
several
site s
w hen
71
c o m p e t i t i o n from h i g h e r s u c c e s s i o n s ! s p e c i e s o c c u r s .
I t r e in f o rc e s th e
n o t i o n t h a t t h e e c o l o g i c a l r o l e o f s e v e r a l C h ry s o th a m n u s t a x a i s on e o f
c o lo n iz a tio n .
T ab le 11.
Range s i t e s 3 a t w hich m u l t i p l e C h ry s o th a m n u s t a x a w ere
c o lle c te d .
G eo g ra p h ic a l a r e a an d ran g e s i t e
F o o t h i l l s a n d M o u n ta in s
s i l t y r a n g e s i t e 1 0 -1 4 " P . Z.
silty
range s i t e
1 5 -1 9 " P . Z.
Taxa c o l l e c t e d a t
site
same
C. n . s s p . n a u s e o s u s
C. v . s s p . I a n c e o l a t u s
Cl Ul s s p . a l b i c a u l i s
C. n . s s p . n a u s e o s u s
C. v . s s p . v i s c i d i f l o r u s
v a r. ste n o p h v llu s
C. n . s s p . a l b i c a u l i s
C. n . s s p . n a u s e o s u s
C. n . s s p . n a u s e o s u s
C. Vl s s p . l a n c e o l a t u s
silty
r a n g e s i t e 2 0 - 2 4 " P . Z.
s i l t y - c l a y e y r a n g e s i t e com plex
1 0 -1 4 " P . Z.
Cl n^ s s p . a l b i c a u l i s
Cl P l s s p . n a u s e o s u s
Cjl Vl s s p . l a n c e o l a t u s
C. n. ssp . a lb ic a u lis
Cl Hl s sp . nauseosus
Cl Vl s sp . v is c id if lo r u s
var. v is c id if lo r u s
Cl Vl s sp . v is c id if lo r u s
var. sten op h vllu s
var. v is c id if lo r u s
Cl P l s s p . a l b i c a u l i s
C. v . s s p . v i s c i d i f l o r u s
v a r. ste n o p h v llu s
G l Ii l s s p .
nauseosus
Cl
la n c e o la tu s
Vl s s p .
72
T a b l e 11.
c o n tin u e d
G eo g ra p h ic a l a r e a and ran g e s i t e
s i l t y - c l a y e y r a n g e s i t e com plex
1 0 -1 4 " P . Z.
Taxa c o l l e c t e d a t
site
same
C. n . s s p . n a u s e o s u s
Ci . v . s s p . v i s c i d i f l o r u s
v a r. ste n o p h v llu s
v ar. v isc id iflo ru s
s i l t y - c l a y e y r a n g e s i t e com plex
1 5 -1 9 P. Z.
C. v . s s p . v i s c i d i f l o r u s
v a r. ste n o p h v llu s
v ar. v is c id iflo ru s
C. n . s s p . n a u s e o s u s
Ci v_. s s p . v i s c i d i f l o r u s
C. n . s s p . n a u s e o s u s
C. v , s s p . v i s c i d i f l o r u s
v ar. v is c id iflo ru s
Ci h . s s p . a l b i c a u l i s
C. n . s s p . n a u s e o s u s
su b a lp in e f i r ,
d o u g l a s - f i r c li m a x
Ci n . s s p . n a u s e o s u s
f o r e s t s on d e e p s o i l s w i t h c r y i c
t e m p e r a t u r e r e g i m e s a n d p a l e brown
o r I i g h b r o w n i s h g r a y s u r f a c e s on
m o d era te ly s te e p t o v e ry s te e p
m o u n ta in s l o p e s 2 0 - 4 5 " P . Z.
Ci v . s s p . I a n c e o l a t u s
s u b a lp in e f i r , d o u g l a s - f i r , ponderosa
p i n e c l i m a x f o r e s t s on d e e p s o i l s w i t h
c r y i c o r f r i g i d t e m p e r a t u r e r e g i m e s on
a n d g r a y i s h brow n, l i g h t b r o w n is h g r a y ,
o r l i g h t y e l l o w i s h brown s u r f a c e s on
m o d e r a t e l y s t e e p t o v e r y s t e e p m o u n ta in
s lo p e s w est o f th e c o n tin e n t i a l d iv id e
1 6 -7 0 " P . Z.
Ci n . s s p . a l b i c a u l i s
C. n . s s p . c o n s i m i l i s
E a ste rn G la c ia te d P la in s
s i l t y - c l a y e y r a n g e s i t e com plex 1 0 -14"
P . Z.
Ci n . s s p . g r a v e d e n s
C. n . s s p . n a u s e o s u s
73
T a b l e 11.
c o n tin u e d
G eo g ra p h ic a l a r e a and ran g e s i t e
Taxa c o l l e c t e d a t same
site
W e s te r n S e d i m e n t a r y P l a i n s
c la y e y and s h a llo w c la y range s i t e
a s s o c i a t i o n 1 0 -1 4 " P . Z.
Ci n ^ s s p . g r a v e o l e n s
Ci Iii s s p . n a u s e o s u s
r i v e r b r e a k s 1 0 -1 4 " P . Z.
Ci Iii s s p . g ra v e o le n s
Ci Hi s s p . nauseosus
B ro w s in g o f Montana C h ry so th a m n u s Taxa
Ci
Ei
ssp.
liv e sto c k .
g r o w th i s
g lan d u lo su s
appears
to
be
p a la tab le
to
w ild life
and
P a s t e v i d e n c e o f b r o w s in g i n d i c a t e s t h a t t h e c u r r e n t y e a r ' s
g e n e ra lly
bro w se d t o
th e
g ro u n d a n n u a l l y
(T a b le
9).
On 2
s i t e s , w hich w e re a c t i v e l y s t o c k e d w i t h c a t t l e a t t h e t i m e o f s a m p l i n g ,
86% and 40% u s e ha d b e e n r e a c h e d by midsummer . ( T a b l e 9 ) .
Ci
Ri
ssp.
g l a n d u l o s u s o c c u r s so i n f r e q u e n t l y a n d i n s u c h s m a l l q u a n t i t i e s t h a t i t
i s u n lik e ly
t o be a m a jo r component o f w i l d l i f e d i e t s i n s o u t h w e s t e r n
M ontana.
Ci
Hi
ssp.
ty p es
of
C h ry so th a m n u s
M o n ta n a .
O ften
P l a n t s o f Ci
Qi
nauseosus and
tax a
p lan ts of
Ci
Ci
Qi
ssp.
observed
Qi
ssp.
to
g r a v e o le n s w ere
be
brow sed
little
o f t h e b a s e r e m a i n e d , m aking i t a p p e a r d e f o r m e d .
M cA rthur
et
a l.
in
s s p . n a u s e o s u s c o l l e c t e d a t s i t e 8 / 2 6 / 2 w ere brow sed t o
One
of
2
n a u s e o s u s w e re s e v e r e l y g r a z e d .
form s.
use
o th er
s ig n ific a n tly
dw arf
The
th e
Ci
c o lle c tio n
Qi
ssp.
(1 9 7 9 b )
(8 /2 6 /2 a )
was he d g e d
nauseosus a g re e s w ith
th at
th e
gray
group,
th e
so
se v e re ly
th a t
o b se rv a tio n s
c o n ta in in g
Ci
Qi
of
ssp.
74
na u s e o s u s , i s m ore p a l a t a b l e t h a n t h e g r e e n g r o u p .
How ever,
n^ s s p .
g r a v e d e n s ( b e l o n g i n g t o t h e g r e e n g r o u p ) was f o u n d t o be brow sed o f t e n
a lso ,
o ccasio n ally
q u ite
h e a v ily
(site
8 /2 7 /2 ).
C urrent
use o f
th is
t a x o n w as o b s e r v e d i n May o f 1986 o n t h e b a n k s o f t h e M i s s o u r i r i v e r .
B r o w s in g was m ost
o fte n
observed
T his i s t h e e x te n t o f th e range o f
in
th e
e a ste rn
h a lf
of
Ito n tan a.
n^ ss p . g ra v e o le n s (F ig . 7 ) , y e t,
— — s s P- n a u s e o s u s e x t e n d s o v e r t o t h e w e s t e r n h a l f o f Ito n ta n a ( F i g .
8 ).
N o ta b le
use
on
p la n ts
of
n^
ssp.
c o n sim ilis
and
C.
v i s c i d i f l o r u s was o b s e r v e d i n a r e a s n o r t h o f Y e l l o w s t o n e N a t i o n a l P h rk
n e a r G a r d i n e r , MT.
P r o n g h o rn a n d m ule d e e r a r e l i k e l y c o n s u m e rs o f C h rv s o th a m n u s t a x a i n
M ontana.
I n Rosebud a n d G a r f i e l d c o u n t i e s , M on ta n a , Chrvsotham nn= was
fo u n d
be a n
to
im p o rta n t
w in ter
brow se
of
mule
d eer.
A lo n g
w ith
A r t e m i s i a s p e c i e s a n d J u n i p e r u s s p e c i e s , i t made up g r e a t e r t h a n 60? o f
th e d i e t a t b oth s i t e s
(E u sta c e 1 9 7 1 ).
I n Rosebud c o u n t y C h rv so th a m n n s
a c c o u n t e d f o r 3 6 . 8 ? o f t h e w i n t e r d i e t , t h e l a r g e s t s i n g l e c o m p o n e n t.
H a l l a n d C le m e n ts (1 9 2 3 ) s u g g e s t e d t h e p a l a t a b l i t y o f C h r y s o t h a m n n =
t a x a i s d e p e n d e n t on t h e s p e c i e s i n w h ic h t h e y a r e a s s o c i a t e d .
th is
is
th e
case
in
M ontana.
L ittle
lite ra tu re
Perhaps
w as fo u n d r e g a r d i n g
C h ry s o th a m n u s t a x a c o n t r i b u t i o n s t o w i l d l i f e i n M on ta n a .
A d d i t i o n a l U ses f o r C h ry s o th a m n u s Taxa i n Montana
to
T here a r e
several a d d itio n a l
c o n trib u tio n s
M o n ta n a .
C h ry so th a m n u s t a x a
w e re o f t e n
w ith
so ils
th at
w e re
se v e re ly
eroded.
C h ry s o th a m n u s t a x a make
fo u n d on s i t e s
S ev eral
tax a
i n Montana
c o u ld
be
v ery
75
u s e f u l i n r e c l a i m i n g m ine s p o i l s , r o a d s i d e s , a n d o t h e r d i s t u r b e d
in
th e ir
ta x a
re sp e c tiv e
h ig h
fo r
ran g es.
Plummer
re c la m a tio n
e sta b lish m e n t,
(1 9 7 7 )
purp o ses
s p r e a d , g r o w th r a t e ,
rate d
based
s o il
a ll
on
so ils
C h ry so th a m n u s
fa c to rs
such
as
s t a b i l i z a t i o n , and a d a p ta tio n
to d istu rb an c e .
M cA rthur (1 9 8 3 ) recommended s e v e r a l C h ry so th a m n u s t a x a
for
d istu rb ed
sta b iliz in g
th ese
taxa
occur
in
so ils
in
th e
In te rm o u n tain
Montana a n d m ig h t be
sim ila rly
A rea.
Some o f
e m p lo y e d .
C^ n ^
s s p . c o n s i m i l i s i s recommended f o r s t a b i l i z i n g d i s t u r b e d s o i l s t h a t a r e
p o o rly
d rain e d
or
heavy
te x tu re d .
i n te r m i tt e n t d rain ag e c a n a ls .
so il
a l.
Cr
lin ifo liu s
can
I t i s v a lu a b le fo r s ta b i l i z i n g d istu rb e d
due t o a s t r o n g u n d e r g r o u n d s p r e a d i n g c h a r a c t e r i s t i c
1974) • Cr
sim ila r
Ur
ssp.
sta b iliz e
a lb ic a u lis
sta b iliz a tio n
on
w ell
C h ry s o th a m n u s t a x a
in
Montana
and
d rain e d ,
Cr
v isc id iflo ru s
dry,
ro ck y
(M cA rthur e t
w ill
so ils
p ro v id e
(M c A rth u r
1983).
a fte r
o th e r
b rig h t
flo w er l a t e
flo w e re d
C h ry so th a m n u s t a x a , p a r t i c u l a r l y
p la n ts
Cr
in
have
th e
summer g e n e r a l l y
fin ish e d
nauseosus su b sp e c ie s,
b lo o m in g .
c a n be u s e d
in la n d s c a p in g t o add c o lo r to p a rk s , y a rd s , and r o a d s i d e s .
a lb ic a u lis,
w ith
y e llo w ,
o r n a m e n t a l (M cA rthur e t a l .
In a d d itio n
pronghorn.
to
v ery
show y
flo w e rs,
is
Cr Qr s s p .
su ita b le
as
an
1979a).
p ro v id in g brow se,
C h ry so th a m n u s
p ro v id e s
cover f o r
Yoakum (1 9 8 6 ) s u g g e s t s t h a t C h ry so th a m n u s s h o u l d be managed
a s a n e e d e d com ponent on p r o n g h o r n r a n g e l a n d s .
Laycock
and
Shoop
n a u s e o s u s make e x c e l l e n t
(1 9 8 2 )
liv in g
re p o rte d
th a t
Cr
lin ifo liu s
snow f e n c e s i n n o r t h e a s t e r n
a n d -Cr
C olo rad o .
S i m i l a r l y C h ry s o th a m n u s t a x a c o u ld be em ployed a s n a t u r a l snow f e n c e i n
76
Montana e s p e c i a l l y
Cjl
iu
ssp.
g rav e o len s.
I t h a s a w id e r a n g e i n t h e
e a s t e r n h a l f o f t h e S t a t e a n d i t g ro w s t o 12 dm.
■
• v .
;;
,
Y: : ^ -
.
'
77
SUMMARY
D u r in g t h e summers 1985 a n d 1 9 8 6 , 8 p r e v i o u s l y d e s c r i b e d t a x a a n d
u n c h a ra c te riz e d
t a x o n o f C h ry so th a m n u s w e re l o c a t e d
i n M ontana.
I
T his
new t a x o n , C h ry so th a m n u s p a r r y i s s p . g l a n d u l o s u s i s d i f f e r e n t i a t e d fro m
o th e r
C\
p arry i
c o n sp ic u o u s
tax a,
sta lk e d
w ith
th e
g lan d s.
i n f l o r e s c e n c e a rra n g e m e n t,
The i n f l o r e s c e n c e o f Ci
e x c ep tio n
of
d iffe rs
from
It
num ber o f
flo w ers
ssp.
C^ jx
ssp.
per head,
10
flo w e rs
per
C a lifo rn ia .
Ci Ei
Ci
m o n ta n u s,
Ez. s s p .
asper
by
s s p . g l a n d u l o s u s i s a r r a n g e d i n a cyme w h i l e
jk
14 f l o w e r s p e r h e a d .
head.
by
and l o c a t i o n .
Ci Ei s s p . a s p e r h a s a n i n f l o r e s c e n c e a r r a n g e d i n a r a c e m e .
g l a n d u l o s u s h a s 10 t o
asper.
Ci
^i
ssp.
Ci ^ i
asper
is
Ci ^ i s s p .
ssp . a sp e r has 5 to .
lo c a te d
in
Nevada
and
s s p . g l a n d u l o s u s i s s i m i l a r t o Ci Ei s s p . im u lu s a n d
but
d iffe rs
by
c h a ra c te ristic s
such
as
le a f
Ci
Ez.
ssp.
l e n g t h , l e a f w i d t h , a n d t h e number o f f l o w e r s p e r h e a d .
Ci
Ez.
s s P-
g lan d u lo su s
ex p resses
2
v a rie tie s.
g l a n d u l o s u s v a r . t o m e n to s u s i s c o v e r e d w i t h tomentum w h ic h o b s c u r e s t h e
sta lk e d
g lan d s.
The
ty p ic a l
fo rm ,
Ci
ssp.
g lan d u lo su s
ssp.
g l a n d u l o s u s l a c k s tom entum .
Four
su b sp ecies
of
Ci
nauseosus
Two, Ci Oi s s p . n a u s e o s u s a n d Ci Qi
g r o u p , w h ic h
ty p ic a lly
th e in v o lu c re s.
c o tto n y w h ite ,
Oi
ssp.
have g ray
V a ria n ts in
w ere
and g la b ro u s i n v o lu c r e s .
occur
s te m s a n d p u b e s c e n c e t o
ste m c o l o r ,
th e
to
in
M o n ta n a .
s s p . a l b i c a u l i s . f a l l in to th e gray
w ere fo u n d i n b o t h t a x a .
g r a v e o le n s b e lo n g t o
fo u n d
tomentum on
r a n g i n g fro m b r i g h t g r e e n t o
Ci
Qi
g reen group,
s s p . c o n s i m i l i s a n d Ci
w hich h a v e g r e e n s te m s
78
T hree ta x a
ssp.
th e
of
Ci
v iso id iflo ru s
Ci Vi s s p .
Ia n c e o la tu s and
presence
v a rie tie s
and
of
ty p e
Vi
Ci
of
ssp.
w e re f o u n d .
v iso id iflo ru s . are
v e stitu re
on
v iso id iflo ru s
th e
V'
in
yv
d iffe re n tia te d
s te m s a n d
occur
lea v e s.
M o n ta n a .
by
Two
T hese w e re
Ci Vi s s p . v i s o i d i f l o r u s v a r .
d i f f e r e n t i a t e d p r i m a r i l y by l e a f w i d t h .
v iso id iflo ru s
Two s u b s p e c i e s ,
h a s l e a v e s g r e a t e r t h a n I .5 mm w ide a n d t h e l e a v e s o f Ci
ssp.
v i s o i d i f l o r u s v a r . s t e n o p h y l l u s r a n g e b e tw e e n I a n d 1 .5 mm
'
■
Many v a r i a n t s w e re f o u n d o f t h e s e 3 t a x a .
T h i s v a r i a t i o n m ig h t
w id e.
be
e x p la in e d
by
in te rg ra d a tio n ,
w hich
fre q u e n tly
happens
in
Ci
v iso id iflo ru s ta x a .
A lso Ci
su b sp e c ie s
l i n i f o l i u s , o rig in a lly
of
v iso id iflo ru s.
Ci
it
d e s c r i b e d by H a l l a n d C le m e n ts a s a
v iso id iflo ru s.
la c k s
o ccu rs
in
t omentum c h a r a c t e r i s t i c
d i f f e r s fro m Ci v i s o i d i f l o r u s t a x a
by i t s
M o n ta n a .
of
Ci
L ik e
nauseosus.
Ci
It
t a l l n e s s a n d by i t s a b i l i t y
t o s p r e a d by l a t e r a l r o o t s .
Ci
Ejl s s p .
p o p u la tio n s in
type
g la n d u lo su s
grassy
c o m m u n it i e s .
h a b ita t
ranged
c h a ra c te ris tic a lly
o p e n in g s among A r t e m i s i a
The
in
w as
s ite s
e le v a tio n
sa m p le d
from
1788
to
to
fo u n d
trid e n ta ta
c h a ra c te riz e
2227
m ( 5 8 6 2 -7 3 0 0
w ere
canopy
s ilty
coverage,
range
ft)
and
S lo p es and
s ite s
and
The c o m m u n itie s a v e r a g e d 13% c o v e r a g e ,
17% b a r e g r o u n d .
lo c a te d
in
the.
A ll
fo o th ills
7 site s
and
g e o g r a p h i c a r e a , b u t h a d v a r y i n g a m o u n ts o f p r e c i p i t a t i o n .
h a d o c c u r r e d on a l l
tax o n * s
A l l s i t e s h a d loam t o sandy loam t e x t u r e d s o i l s
w i t h pH v a l u e s b e tw e e n 7 a n d 8 .
9% s h r u b
sm all
- g rasslan d
th is
a n n u a l p r e c i p i t a t i o n v a r i e d fro m 356 t o 588 mm ( 1 4 - 2 3 i n ) .
a s p e c t s w e re v a r i a b l e .
in
site s.
sa m pled
m o u n ta in s
D istu rb a n c e
79
A d jacen t
p l o ts w ith o u t
th e
Cjl
ssp.
g l a n d u l o s u s w e re sam pled t o
d e t e r m i n e w h a t f a c t o r ( s ) w e re r e s p o n s i b l e f o r t h i s
A lth o u g h
fac to r,
d istu rb an ce
a lo n g
w ith
ha d a l s o
so il
d i s t r i b u t i o n o f Ci
n e u tra l,
loam t o
ty p e ,
occurred
seem ed
to
ssp . g la n d u lo su s.
san d y loam s o i l s ,
on
th e
ta x o n 's o c c u rre n c e .
a d ja c e n t
in flu e n c e
ex trem e
a d ja c e n t
g razin g
P a st e v id e n c e
C.
p.
ssp.
fo u n d t o
s m a ll
low i n
so lu b le
sa lts,
of
(X
ssp.
h o w e v e r,
it
S e n sitiv ity
g l a n d u l o s u s on
o f p o p u l a t i o n s i n w h ic h i t
o ccu rs.
o f b r o w s in g i n d i c a t e s t h a t t h e c u r r e n t y e a r ' s g ro w th o f
g lan d u lo su s
be
siz e
and
I t seem s a b l e t o c o l o n i z e n e a r
may e x p l a i n a b s e n c e
s i t e s and the
th is
o ccu rren ce
was a b s e n t on a d j a c e n t s i t e s w i t h s i m i l a r c h a r a c t e r i s t i c s .
to
p lo ts,
40 an d
is
g e n e ra lly
86? by
bro w se d t o
m id-summer on 2
th e
site s
ground.
Use was
t h a t w e re a c t i v e l y
s t o c k e d a t t h e tim e o f s a m p l i n g .
C h ry so th a m n u s t a x a
ssp.
w e re
d istrib u te d
g l a n d u l o s u s was fo u n d o n l y
in
th ro u g h o u t
th e
S ta te .
s o u t h w e s t e r n M o ntana.
C^
C^ n ^ s s p .
n a u s e o s u s , t h e m ost w i d e s p r e a d t a x o n , was fo u n d o v e r t h e e n t i r e S t a t e .
C. n . s s p . a l b i c a u l i s was l i m i t e d t o t h e w e s t e r n h a l f o f Jtontana a n d C^
n^ ssp.
g r a v e o l e n s was d i s t r i b u t e d
in
th e
e a ste rn
h a lf.
Cjl Hjl s s p .
c o n s im il is o c c u rre d o n ly in th e s o u th e rn p o r tio n s o f th e S t a t e .
ssp.
Ia n c e o la tu s
J to n t a n a .
Cl
ranged
in
th e low er h a l f of c e n t r a l
to
s o u th w e ste rn
s s p . v i s c i d i f l o r u s s h a r e d t h e same r a n g e a s Cl
Vl
C l Vl
Vl
ssp.
I a n c e o l a t u s b u t e x t e n d e d i n t o n o r t h w e s t e r n J to n t a n a .
Cl l i n i f o l i u s was
fo u n d
h a lf
o nly
in
Carbon
County
lo ca ted
in
th e. lo w er
of
c e n tra l
J to n t a n a .
C.
nauseosus
h a b ita ts.
su b sp ec ie s
Cl Hl s s p .
w ere
d istrib u te d
n a u s e o s u s w as c o l l e c t e d
on a
w id e
on c l a y
to
v a rie ty
of
sa ndy loam
80
s o ils.
C o llec tio n
280 t o
site s
v aried
510 mm ( 1 1 - 2 0 i n )
( 2 3 0 0 -7 5 0 0 f t ) .
in annual
p re c ip ita tio n
w ith e l e v a t i o n s
As w i t h
Ci ru
was o f t e n fo u n d on d i s t u r b e d
It
fro m
r a n g i n g fro m 700 t o 2300 m
ssp. nauseosus,
site s.
a m o u n ts
Ci m
ssp . g r a v e d ens
was commonly a s s o c i a t e d
w ith
S a rco b a tu s v e rm ic u la tu s and A tr ip le x spp.
—
v isc id iflo ru s
d istu rb an ce
tax a
was e v i d e n t .
w ere
a lso
o fte n
C o llec tio n s o f
made on s o i l s r a n g i n g fro m s i l t t o s a n d .
y isc id iflo ru s
w e re a l s o
c o lle c te d
—
was fo u n d i n
Ci
found
Vi
on
ssp.
areas
I a n c e o l a t u s w e re
The 2 v a r i e t i e s o f Ci
p rim a rily
w here
on l i g h t
tex tu red
c o m m u n itie s d o m in a te d by s a l t
ssp.
Vi
so ils.
d e s e r t shrub
sp e cie s.
C h ry so th a m n u s
M o n ta n a .
Ci . ^ i
c o n trib u te s l i t t l e
q u a n titie s .
a n d Ci
ru
tax a
seem
ssp.
g lan d u lo su s
to
However,
ssp.
to
w ild life
be
d iets
im p o rta n t
ap p ears
to
because i t
C h ry so th a m n u s t a x a
g r a v e d ens,
lo c a lly
to
be
p a la ta b le ,
o ccu rs
su c h a s Ci Pi
in
ssp.
a r e m ore a b u n d a n t a n d w i d e l y
T hese t a x a w e re fo u n d s e v e r e l y
grazed in
w ild life
such
in
but
sm all
nauseosus
d istrib u te d .
s e v e ra l l o c a t i o n s and a p p e a r
t o c o n t r i b u t e t o w i l d l i f e d i e t s , p a r t i c u l a r l y i n e a s t e r n M o n ta n a .
Chirysothamnus t a x a
re a d ily
e sta b lish
make a d d i t i o n a l
flo w e re d
a lb ic a u lis.
They
on d i s t u r b e d a n d d e p l e t e d a r e a s m ak in g them u s e f u l
f o r re c la m a tio n p u rp o se s.
b rig h t
c o n t r i b u t i o n s t o M on ta n a .
p la n ts
In a d d it io n ,
They o f f e r
have
d rie d
c o lo r to lan d scap es a f t e r o th e r
up,
p a rtic u la rly
C h ry so th a m n u s t a x a
w i l d l i f e a n d can be u s e d f o r l i v e snow f e n c e s .
p ro v id e
Ci . n .
ssp.
needed co v er f o r
LITERATURE CITED
82
A n d e r s o n , L . C. 1 9 6 4 .
T a x o n o m ic n o t e s o n t h e C h r y s o t h a m n u s
v i s c i d i f l o r u s c o m plex ( A s t e r e a e , C o m p o s it a e ) . Madrono 1 7 : 2 2 2 - 2 2 7 .
A n d e r s o n , L. C. 1 9 6 6 .
C y to tax o n o m ic s t u d i e s i n
( A s t e r e a e , Coinpos i t a e ) ♦ Amer. J . B o t . 5 3 : 2 0 4 - 2 1 2 .
A n d e r s o n , L. C. 1969«
and i t s im p lic a tio n s .
A nderson,
503.
L.
C.
1970.
C h ry so th a m n u s
The k a r y o t e o f C h ry so th a m n u s p a r r y i s s p . p a r r y i
T r a n s . K a n s. A cad. S c i . 72 ( 3 ) : 3 9 9 - 4 0 1 .
F lo ral
an a to m y o f
A n d e r s o n , L. C. 1 9 7 1 .
E m b ry o lo g y
Compos i t a e ) . Madrono 2 0 ( 7 ) : 3 3 7 - 3 8 4 .
of
C h ry s o th a m n u s .
S id a 3 : 4 6 6 -
C h ry so th a m n u s
(A s te r e a e .
A n d e r s o n , L . C. 1 9 7 3 .
Unique C h ry so th a m n u s h y b r i d i z a t i o n s
Meadows, N ev a d a . B u l l . T o r r e y B o t . Club 1 0 0 : 1 7 1 - 1 7 7 .
A n d e r s o n , L. C. 1 9 7 8 .
P h y to lo g ia 38: 309-320.
New t a x a . i n
C h ry s o th a m n u s ,
se ctio n
in
Ash
N au seo si.
A n d e r s o n , L . C. 1 9 8 0 a . I d e n t i t y o f n a r r o w - l e a v e d C h r y s o t h a m n u s
v i s c i d i f l o r u s (A s t e r a c e a e ) . G r e a t B a s i n N a t. 4 0 : 1 1 7 - 1 2 0 .
A n d e r s o n , L. C. 1980b. M orphology a n d b i o g e o g r a p h y o f C h ry so th a m n u s
n a u s e o s u s s s p . t e x e n s i s ( A s t e r a c e a e ) : a new G u a d a l u p e M o u n ta in s
e n d e m ic . S outhw . N a t . 2 5 : 1 9 7 - 2 0 6 .
A n d e rs o n , L . C. 1 9 8 1 .
C h ry so th a m n u s n a u se o su s s s p . i r i d i s
(A s t e r a c e a e ) : a new endem ic fro m U ta h .
G reat B asin N at. 41:3 1 1 -3 1 3 .
A n d e r s o n , L. C. 1 9 8 3 .
C h ry so th a m n u s e r e m o b iu s
s p e c i e s fro m N evada. B r i t t o n i a 3 5 : 2 3 - 2 7 .
(A s t e r e a c e a e ) : a
A n d e r s o n , L. C. 19 8 4 .
C h ry so th a m n u s n a u s e o s u s s s p .
s t a b i l i z e d h y b rid .
G reat B asin N a t. 4 4 ( 3 ):4 l6 - 4 2 0 .
new
u in ta h e n sis: a
A n d e r s o n , L . C. 1 9 8 6 a .
An o v e r v i e w o f t h e g e n u s C h ry s o th a m n u s
( A s t e r a c e a e ) . p . 2 9 - 4 5 . I n : E. D. M cA rthur a n d B. L. Welch c o m p i l e r s .
P r o c e e d in g s -S y m p o s iu m o f t h e b i o l o g y o f A r t e m i s i a a n d C h ry s o th a m n u s .
USDA FS Gen. T e c h . Rep. IN T -200. O g d e n , U ta h .
A nderson,
p .9 3 -9 7 .
Symposium
T e c h . R ep.
L. C.
1 9 86b. C y to g e o g ra p h y o f C h ry s o th a m n u s V i s c i d i f l o r u s .
I n : E. D. M cA rthur a n d B. L. Welch c o m p i l e r s . P r o c e e d i n g s o f t h e b i o l o g y o f A r t e m i s i a a n d C h r y s o th a m n u s .
USDA FS Gen.
IN T -200. O g d e n , U t a h . ’
83
A n d e r s o n , L. C. 1 9 8 6 c .
S y m p a tr ic s u b s p e c i e s i n C h ry so th a m n u s
nau se osus.
p .9 3 -1 0 3 .
I n : E. D. M c A rth u r, E. D. a n d B. L. Welch
c o m p i l e r s . P r o c e e d in g s -S y m p o s iu m o f t h e b io lo g y of A rtem isia and
C h r y s o th a m n u s . USDA FS Gen. T e c h . Rep. I N t - 2 0 0 . O gden, U ta h .
A n d e r s o n , L. C. a n d P . S . F i s h e r .
1970.
P h y l o g e n e t i c i n d i c a t o r s fro m
f l o r a l a natom y i n C h ry so th a m n u s . ( A s t e r e a e . Compos i t a e ) .
P h y to m o rp h .
20 : 112- 118 .
A n d e r s o n , L. C. a n d J . L. R e v e a l .
.1966.
C h ry s o th a m n u s b o l a n d e r i . a n
i n t e r g e n e r i c h y b r i d . Madrono 1 8 ( 8 ) : 2 2 5 - 2 3 2 .
B a k e r H. G. 1 9 6 5 .
C h a r a c t e r i s t i c a n d modes o f t h e o r i g i n o f w e e d s , p .
1 4 7 -1 6 8 .
I n : B a k e r , H. G. a n d G. L. S t e b b i n s ( e d s . ) The g e n e t i c s o f
c o lo n iz in g s p e c ie s ; p ro c e e d in g s.
Academic P r e s s , New Y ork.
B a r k w o r t h , M. E. a n d D. R. Dewey.
1985.
G e n o m ic a lly b a s e d g e n e r a i n
t h e p e r e n n i a l T r i t i c e a e o f N o rth A m e ric a: i d e n t i f i c a t i o n a n d
m e m b e rs h ip .
Amer. J . B o t . 7 2 ( 5 ) : 7 6 7 - 7 7 6 .
B o o th , W. E. a n d J . C. W r i g h t .
1966.
F l o r a o f M o n ta n a , d i c o t y l e d o n s
part I I .
D e p t , o f B o ta n y a n d M i c r o b i o l o g y . MSU. Bozem an, MT.
B o u y o u c o s, C. J .
1936.
D i r e c t i o n s f o r m aking m e c h a n i c a l a n a l y s e s o f
s o i l s by t h e h y d r o m e t e r m e th o d . S o i l S c i e n c e 4 2 ( 3 ) : 2 2 5 - 2 2 8 .
Cook, C. W. a n d L. E. H a r r i s .
195 0 .
The n u t r i t i v e c o n t e n t o f t h e
g r a z i n g s h e e p ' s d i e t on summer a n d w i n t e r r a n g e s i n U ta h .
U tah S t .
A g r i c . C o l l . A g r i c . Exp. S t a . B u l l . 3 4 2 .
Cook, C. W., L. A. S t o d d a r t , a n d L. E. H a r r i s .
1954.
The n u t r i t i v e
v a lu e o f w in te r range p l a n ts in th e G reat B a sin .
Utah
S t. A g ric.
C o l l . A g r i c . E xp. S t a . B u l l . 3 7 2 .
C r o n q u i s t , A.
1955.
P h y lo g e n y an d taxonom y o f t h e Compos i t a e .
M id i. N a t . 5 3 : 4 7 8 - 5 1 1 .
Amer.
Daubenmi r e , R. F .
1975.
P l a n t s u c c e s s i o n on a b a n d o n e d f i e l d s , a n d
f i r e i n f l u e n c e s , i n a s te p p e a r e a i n s o u t h e a s t e r n W a sh in g to n .
N o r th w e s t S c i e n c e . 4 9 ( 1 ) : 3 6 - 4 8 .
D a y to n , W. A.
1 9 3 1 . I m p o r t a n t w e s t e r n bro w se p l a n t s .
USDA M is c . P ub.
n o . 101.
U . . S. G overnm ent P r i n t i n g O f f i c e , W a s h in g to n D. C.
D i t t b e r n e r , P . L. an d M. R. O l s o n . 1 9 8 3 . The p l a n t i n f o r m a t i o n n e tw o r k
(PIN ) d a t a b a s e : C o l o r a d o , M ontana, N o r th D a k o t a , U t a h , a n d Wyoming.
FW S/OBS-83/36. W a s h in g to n , D. C .: U. S. D e p a r tm e n t o f t h e I n t e r i o r ,
F is h and W il d li f e S e r v ic e . 786p.
D o r n , R. D.
1984.
V a sc u la r
P u b l i s h i n g . C he y e n n e , WY.
p la n ts
of
M ontana,
M o u n ta in West
84
E u s t a c e , C. D. 1971.
Mule d e e r f o o d h a b i t s a n d brow se u s e s t u d y .
Montana F i s h a n d Game F e d . Aid F i n a l Rep. P r o j . W-13 O-R-I a n d 2 , J ^ l -
E v a n s , R. A. a n d J . A. Young.
1973. C u rren t a p p ro a c h e s t o r a b b i t b r u s h
c o n t r o l w i t h h e r b i c i d e s . Down t o E a r t h 2 9 ( 2 ) : 1 - 4 .
F r i s c h k n e c h t , N. C. 1 963.
C o n tr a s tin g e f f e c t s o f b ig sag eb ru sh and
r u b b e r r a b b i t b r u s h on p r o d u c t i o n o f c r e s t e d w h e a t g r a s s .
J . Range
Manage. 1 6 : 7 0 - 7 4 .
F r i s c h k n e c h t , N. C . , L. E. H a r r i s , a n d H. K. Woodward.
1 953.
C a ttle
g a i n s a n d v e g e t a l c h a n g e s a s i n f l u e n c e d by g r a z i n g t r e a t m e n t s o n
c re s te d w h e a tg ra ss.
J . Range R h n a g e . 6 : 1 5 1 - 1 5 8 .
G ra n t,
Y ork.
V.
1971.
P lan t
sp e c ia tio n .
Colum bia
U n iv e rsity
P ress,
New
H a l l , H. M. a n d F . E. C l e m e n t s .
1923.
The p h y l o g e n e t i c m ethod o f
tax o n o m y .
The N o r th A m erican s u b s p e c i e s o f A r t e m i s i a . C h r y s o th a m n u s .
a n d A t r i p l e x . C a r n e g i e I n s t , o f W a s h . , W a s h in g to n .
H a l l , H. M. an d T. H. G o o d sp e e d . 1 919.
C h r y s i l , a new r u b b e r from
C h ry so th a m n u s n a u s e o s u s . U n iv . o f C a l i f . P ub. i n B o t . 7 : 1 8 3 - 2 6 4 .
H anks, D. L . , E. D. M c a r t h u r , A. P . Plummer, B. C. G i u n t a , a n d A. C.
B la u e r.
1975. C h r o m a to g r a p h ic r e c o g n i t i o n o f some p a l a t a b l e a n d
u n p a l a t a b l e s u b s p e c i e s o f r u b b e r r a b b i t b r u s h i n a n d a r o u n d U ta h .
J.
Range tfe n a g e . 2 8 ( 2 ) : 1 4 4 - 1 4 8 .
Holmgrem, P. K. a n d W. K euken.
1974.
I n d e x H e r b a r i o r u m . P a r t I . The
H e r b a r i a o f t h e W o rld . E d . 6 . I n t e r n a t i o n a l B u re a u f o r P l a n t Taxonomy
a n d N o m e n c la tu r e . U t r e c h t , N e t h e r l a n d s .
H u z iw a r a , Y.
1962.
K a r y o ty p e a n a l y s i s i n some g e n e r a o f C o m p o s ita e
V III.
F u r t h e r s t u d i e s o n t h e chrom osom es o f A s t e r .
Amer. J . o f B o t .
49:1 1 6 -1 1 9 .
K u c h l e r , A. W. 1 9 6 4 .
M a n u al t o a c c o m p a n y t h e map o f p o t e n t i a l
v e g e t a t i o n o f t h e c o n t e r m i n o u s U n i t e d S t a t e s . A m erican G e o g r a p h i c a l
S o c i e t y . S p e c i a l P u b l i c a t i o n No. 3 6 .
K u f e l d , R. C . , 0 . C. W allm o, a n d C. F e d d e m a . 1973« F o o d s o f t h e r o c k y
m o u n ta in m u le d e e r . USDA FS R e s. P a p e r RM-11 1 . F t . C o l l i n s , CO.
L a y c o c k , W. A. a n d M. C. S h o o p .
1982.
S u r v i v a l a n d g ro w th o f s h r u b
s p e c i e s p l a n t e d b e h in d s n o w f e n c e s on t h e c e n t r a l G r e a t P l a i n s ,
p. 17.
I n : A b s t r a c t s o f P a p e r s , 3 5 t h A nnual M e e t i n g .
S o c i e t y f o r Range
M anagement; C a l g a r y , A l b e r t a . L e a c h , H. R.
1956.
Food h a b i t s o f t h e
G re a t B asin d e e r h e rd s o f C a l i f o r n i a .
C a l i f . F i s h a n d Game. 4 2 : 2 4 3 93.
85
M a rio n , J . H . , E. I . H acketfc, a n d J . W. B u r k h a r d t o
19 86 o Range b r u s h
c o n tr o l w ith G ra sla n p e l l e t s .
p . 1 2 2 -1 2 5 I n : E. D. M cA rthur a n d B.
L. Welch c o m p i l e r s . P r o c e e d in g s -S y m p o s iu m o f t h e b i o l o g y o f A r t e m i s i a
a n d C h r y s o th a m n u s . USDA FS Gen. T e c h . Rep. IN T -2 0 0 . O gden, U ta h .
M c A r t h u r , E. D.
1983.
Im p o rta n t sh ru b s f o r w ild la n d p la n tin g s ,
C o m p o sita e (A s t e r a c e a e ) .
p . 15 0 -1 5 7 •
I n : M anaging I n t e r m o u n t a i n
ran g e lan d s-im p ro v em e n t o f range and w i l d l i f e h a b i t a t s .
USDA FS Gen.
T e c h . Rep. I N T - I 5 7 . O gden, U ta h .
M c A rth u r, E. D . , A. C. B l a u e r , A. P. Plummer, a n d R. S t e v e n s .
197 9 a .
C h a r a c t e r i s ti c s and h y b r id iz a tio n o f im p o rta n t In te rm o u n tain sh ru b s.
I I I . S u n f lo w e r f a m i l y .
USDA FS R e s. P a p . IN T -2 0 0 . O gden, U ta h .
M c A rth u r, E. D . , B. C. G u i n t a , a n d A. P . Plumm er.
r e s t o r a t i o n o f d e p le te d ran g es and d is tu rb e d a r e a s .
33-
1974.
Shrubs f o r
U tah S c i . 3 5 : 2 8 -
M c A rth u r, E. D . , D. L. H anks, A. P . Plummer, a n d A. C. B l a u e r . 19 7 8 a .
C o n t r i b u t i o n s t o t h e taxonom y o f C h ry so th a m n u s v i s c i d i f l o r u s a n d o t h e r
C h ry s o th a m n u s s p e c i e s u s i n g p a p e r c h r o m a to g r a p h y .
J . Range tfe n a g e.
3 1 :216-223.
M c A r t h u r , E . D . , A. P . P l u m m e r , a n d J . N. D a v i s .
1978b.
R e h a b i l i t a t i o n o f game r a n g e i n t h e S a l t D e s e r t .
Wyoming s h r u b l a n d s ,
P r o c . o f 7 t h Wy S hrub E c o l . W orkshop.
U. o f WY., L a r a m ie .
M c A rth u r, E. D . , C. F . T i e r n a n , a n d B. L. W e lc h .
1979b.
S u b sp e c ie s
s p e c i f i c i t y o f g a l l fo rm s on C h ry so th a m n u s n a u s e o s u s .
G reat B asin
N a tu r. 3 9 (1 ):8 1 -8 7 .
M c A rth u r, E. D. a n d B. L. W elch.
1986.
In tro d u c tio n :
A rte m isia and
C h r y s o th a m n u s .
p. 2 -4 .
I n : E. D. M cA rthur a n d B. L. Welch c o m p i l e r s .
P r o c e e d in g s -S y m p o s iu m o f t h e b i o l o g y o f A r t e m i s i a a n d C h r y s o th a m n u s .
USDA FS Gen. T e c h . Rep. IN T -200. O g d e n , U ta h .
M c K e ll, C. M. a n d W. W. C h i l c o t e .
1957.
f o l l o w i n g re m o v a l o f c o m p e tin g v e g e t a t i o n .
230.
R e sponse o f r a b b i t b r u s h
J . Range tfe n a g e . 1 0 : 2 2 8 -
M u l l e r , H. J .
1 9 4 0 . B e a r i n g s o f t h e " D r o s o p h i l a " work on s y s t e m s t i c s ,
p . 18 5 - 2 6 8 . I n J . H u x le y , e d . The new s y s te m s t i c s . C la r e n d o n P r e s s ,
O xford.
N a t i o n a l O c e a n ic a n d A tm o s p h e r ic A d m i n i s t r a t i o n .
1984. C lim a to lo g ic a l
D ata A nnual Summary M ontana. V p l . 87. No. 13.
D e p a r tm e n t o f Commerce,
A s h e v i l l e , N. C.
86
O s t l e r , W. K . , C. M. McKelI , an d S . W h it e .
1986.
C hry so th a m n u s
n a u s e o s u s ; A p o t e n t i a l source o f n a tu r a l ru b b e r,
p. 389-394.
I n : E.
D. M cA rthur a n d B. L. Welch c o m p i l e r s . P r o c e e d in g s -S y m p o s iu m of. t h e
b i o l o g y o f A r t e m i s i a a n d C h r y a o th a m n u s .
USDA FS Gen. T e c h . Rep. INT2 0 0 . O gden, U t a h .
P a u l s e n , H. A. a n d J . C. M i l l e r .
19 6 8 .
C ontrol o f p a rry r a b b itb ru s h
on m o u n ta in g r a s s l a n d s o f w e s t e r n C o l o r a d o .
J . Range M hnage. 2 1 : 1 7 5 177.
-;■
Plummer, A. P .
1977
R e v e g e ta tio n o f d is tu rb e d In te rm o u n ta in a re a
site s,
p . 302-339•
I n : J . L. Thames, J . L. e d .
R eclam atio n and u se
o f d is tu r b e d la n d i n t h e s o u th w e st.
U n iv . A riz o n a P r e s s .
T u c so n .
R o b e r t s o n , J . H. a n d H. P. C o r d s .
1957.
C h ry so th a m n u s s p p . , f o ll o w i n g c h e m i c a l ,
tre a tm e n ts.
J . I h n g e Ife n a g e . 1 0 : 8 3 - 8 9 .
R o s e n t r e t e r , R.
1986.
( C h r y s o th a m n u s ) com m unity
I n : E. D. M cA rthur an d B.
th e b io lo g y o f A rte m isia
IN T -200. O g d e n , U t a h .
S u rv iv a l
b u rn in g ,
of rab b itb ru sh ,
a n d m ec h a n ic a l
C o m p o s it io n a l p a t t e r n s w i t h i n a r a b b i t b r u s h
o f t h e I d a h o S n a k e 'R i v e r P l a i n ,
p. 273-277.
L. Welch c o m p i l e r s . P r o c e e d in g s -S y m p o s iu m o f
a n d C h r y s o th a m n u s .
USDA FS Gen. T e c h . Rep.
R o s s , R. L. a n d H. E. H u n t e r . 1 9 7 6 . C lim ax v e g e t a t i o n i n Montana b a s e d
on s o i l s a n d c l i m a t e .
USDA SCS Bozeman, MT.
Sam pson, A. W. a n d B. S . J e s p e r s o n .
1963. C a l i f o r n i a ran g e b r u s h la n d s
a n d brow se p l a n t s .
U n i v . C a l i f . D i v . A g r. S c i . Mfenual 3 3 , B e r k e l e y .
S c h l a t t e r e r , E. F . a n d E. W. T i s d a l e .
1 969.
E ffe c ts of l i t t e r
A r t e m i s i a , C h r y s o th a m n u s , an d T o r t u l a on g e r m i n a t i o n a n d g ro w th
th re e p eren n ial g ra s s e s .
E c o lo g y 5 0 ( 5 ) : 86 9 - 8 7 3 .
of
of
S e v e r s o n , K. E.
1966.
G r a z in g c a p a c i t i e s a n d c o m p e t i t i o n o f p r o n g h o r n
a n t e l o p e a n d d o m e s t i c s h e e p i n W yom ing's Red D e s e r t .
PhD D i s s e r t a t i o n ,
U n i v e r s i t y o f Wyoming, L a r a m ie , WY.
S o l b r i g , 0 . T . , L. C. A n d e rs o n , D. W. K yhos, a n d P . H. R aven.
19 6 9 .
Chromosome num bers i n C o m p o s ita e V I I : A s t e r e a e I I I .
Amer. J . B o t .
5 6 (3)5348-353• 1
S o l b r i g , 0 . T . , L. C. A n d e r s o n , D. W. K yhos, P . H. R aven, a n d L.
R udenberg.
1964.
Chromosome num bers i n C o m p o s ita e V: A s t e r e a e I I .
Amer. J . B o t . 5 1 ( 5 ) : 5 1 3 - 5 1 9 S t e b b i n s , G. L.
1947.
T ypes o f p o l y p l o i d s : t h e i r c l a s s i f i c a t i o n a n d
sig n ific a n c e .
Advances i n G e n e tic s I :4 0 3 -4 2 9 .
S t e b b i n s , Gi L. . 1 9 5 0 .
V a r ia tio n and e v o lu tio n i n p l a n t s .
U n i v e r s i t y P r e s s , New Y ork.
Colum bia
87
S t e b b i n s , G. L.
1957.
S e l f f e r t i l i z a t i o n and p o p u la tio n v a r i a b i l i t y
in th e h ig h er p la n ts .
Am. N a t. 9 1 : 3 3 7 - 3 5 3 .
S t e b b i n s , G. L. 1965.
C o lo n izin g s p e c ie s o f th e n a tiv e C a lif o r n ia
flo ra .
p. 173-191.
I n : H. G. B a k e r a n d G. L. S t e b b i n s ( e d s . ) The
g e n e t i c s o f c o l o n i z i n g s p e c i e s ; p r o c e e d i n g s . A cadem ic P r e s s , New Y o rk .
S t e v e n s , R.
1986.
P o p u l a t i o n d y n a m ic s o f two s a g e b r u s h s p e c i e s a n d
r u b b e r r a b b i t b r u s h o v e r 22 y e a r s o f g r a z i n g u s e by t h r e e a n i m a l
c la sse s.
p. 278-285.
I n : E. D. M cA rthur a n d B. L. Welch c o m p i l e r s .
P r o c e e d in g s -S y m p o s iu m o f t h e b i o l o g y o f A r t e m i s i a a n d C h ry s o th a m n u s ♦
USDA FS Gen. T e c h . Rep. IN T -200. O g d e n , U ta h .
S t e v e n s , R . , K. R. J o r g e n s e n , J . N. D a v i s , a n d S . B. Monsen.
198 6 .
Seed p a p p u s a n d p l a c e m e n t i n f l u e n c e s o n w h i t e r u b b e r r a b b i t b r u s h
e sta b lish m e n t.
p. 353-357.
I n : E. D. M cA rthur a n d B. L. Welch
c o m p i l e r s . P r o c e e d in g s -S y m p o s iu m o f t h e b io lo g y o f A rtem isia and
C h r y s o th a m n u s . USDA ES Gen. T e c h . Rep. IN T -2 0 0 . O gd e n , U ta h .
U n ite d S t a t e s D e p a rtm e n t o f A g r i c u l t u r e .
1954.
D ia g n o s is and
im p ro v e m e n t o f s a l i n e a n d a l k a l i s o i l s .
S o i l s A g r i c u l t u r a l Handbook
No. 6 0 .
U. S . G ovt. P r i n t i n g O f f i c e , W a s h in g to n , D. C.
U n i t e d S t a t e s D e p a r tm e n t o f A g r i c u l t u r e .
1975.
S o i l Taxonom y.
A g r i c u l t u r a l H a n d b o o k No. 4 3 6 . U. S . G o v t . P r i n t i n g O f f i c e ,
W a s h i n g t o n , D. C.
W eber, D. J . , J . D. D a v i s , E. D. M c A rth u r,
a n d N. S a n k h l a .1 9 8 5 .
C h ry so th a m n u s n a u s e o s u s ( r u b b e r r a b b i t b r u s h ) : m u l t i p l e - u s e s h ru b o f t h e
d e se rt.
D esert P la n ts 7(4 )1 7 2 -2 1 0 .
W e ls h , S.
N a t. 43(2)
L. 1983•
179-357.
Utah f l o r a :
Compos i t a e
(A s t e r a c e a e ) .
G. B a s i n
W inw ard, A. H. a n d L. C. A n d e r s o n .
1986.
F ie ld t o u r . . th e b io lo g y
o f A r t e m i s i a a n d C h r y s o th a m n u s .
p. 6-11.
I n : E. D. M cA rthur a n d B.
L. Welch c o m p i l e r s . P r o c e e d in g s -S y m p o s iu m o f t h e b i o l o g y o f A r t e m i s i a
a n d C h r y s o th a m n u s . USDA FS Gen. T e c h . Rep. IN T -2 0 0 . O gden, U ta h .
Yoakum, J .
1986.
Use o f A r t e m i s i a a n d C h ry s o th a m n u s by p r o n g h o r n s .
P . 176-180.
I n : E. D. M c A r t h u r a n d B. L. W e lc h c o m p i l e r s .
P ro c e e d in g s -S y m p o s iu m o f t h e b i o l o g y o f A r t e m i s i a a n d C h ry s o th a m n u s .
USDA FS Gen. T e c h . Rep. IN T -200. O g d e n , U t a h .
Young, J . A. a n d R. A. E v a n s .
1974a.
P h e n o lo g y o f C h ry so th a m n u s
v i s c i d i f l o r u s s u b s p e c i e s v i s c i d i f l o r u s (H o o k .) N u t t . Weed S c i . 2 2 : 4 6 9 475.
Young, J . A. a n d R. A.
ra b b itb ru sh in d istu rb ed
2 7 ( 2 ): 127- 132 .
E vans 1974 b .
b ig sagebrush
P o p u l a t i o n d y n a m ic s o f g r e e n
c o m m u n it i e s . J . Range lfe n a g e .
88
Young, J . A . , R. A. E v a n s , a n d J . M a j o r s .
G reat B a sin .
J . Range Mhnage. 2 5 : 1 9 4 - 2 0 1 .
1972.
A lien p l a n t s i n
th e
APPENDIX
T ab le 12.
Summary o f v e g e t a t i o n p a r a m e t e r s m e a s u r e d a t 7
R iv ersid e
taxon
. p lo t
$ to ta l
g r a s s cover
% shrub canopy
cover
to ta l
C. p a r r v i
o t h e r C h ry so th a m n u s
ta x a .
' A rte m is ia tax a
8 .5
6 .0 4
J e r r y C re e k
p a ire d
p lo t
9 .0
4 .3
3 .5 4
taxon
p lo t
1 3 .2
9 .8 9
3 .5
——---
1 .5
0 .5
1 .6
0 .9 8
% B a re g ro u n d
5 .4
2 1 .2
9 .6
8 .2 9
1 .0
C. p a r r v i s s p .
g lan d u lo su s
d e n sity /m ^
p a ire d
p lo t
7 .5
Q u a r tz H i l l
taxon
p lo t
1 0 .5
5 .2
p a ire d
p lo t
8.1
2 4 .7
2 .6 2
1 .5
1 .5
0 .5
3 .2 6
1 .8
3 .7
H e le n L a r s o n
Ranch
taxon
p lo t
1 6 .3
6 .1 9
p a ire d
p lo t
9 .7
5.1
2 .5 9
— — — —
3 .2 8
2 .2
p a r r v i s s p . g la n d u lo s u s sam p lin g s i t e s .
2 .2
3 .7
2 .9
1 .4
1 .5 8
5 .7
3 .2
1 1 .2
Table 12 .
con tin u ed
Coal C reek
BLM
C e n te n n ia l
taxon
p lo t
p a ire d
p lo t
taxon
p lo t
p a ire d
p lo t
taxon
p lo t
% to ta l
g ra s s cover
1 2 .5
1 1 .0
1 1 .3
1 4 .0
1 1 .7
7 .2
% s h r u b canopycover
to ta l
1 6 .9 4
5 .8
1 0 .4 6
9.1
7 .3
18.2
C. p a r r v i
o t h e r C h ry so th a m n u s
tax a .
A rte m isia tax a
C. p a r r v i s s d .
g lan d u lo su s
d e 'n s ity /m ^
% B a re g ro u n d
1 1 .6 4
7 .7 6
2 .6
- 1.1
1 .5
0 .9
0 .5
1.1
4 .2
0 .2
1 .8
5 .9
3 .6
3-04
3 3 .4
6 .0 5
3 8 .7
3 6 .3
p a ire d
p lo t
18.1
1 .11
2 2 .8
3 0 .7
3 4 .3
Table 13
H a b i t a t n o t e s a n d l o c a t i o n s o f s p e c i m e n s o f C h ry so th a m n u s a n d s i t e s n o t e d i n t h e R e s u l t s
and D isc u ssio n s e c t i o n .
Taxon and specimen
S o il te x tu r e
Aspect
Slope
%
C. nauseosus
ss p . nauseosus
8 /l6 /4 b
s ilty
ea st . 1 0 - 2 0
Dominant a sso c ia te d
p la n ts
Grazing
at s ite
P seudoreeneria sp ica ta
A rtem isia arbuscula
y es
Location d e sc r ip tio n
38.1 m iles from Alder on
Ruby River R eservoir
road , Hbdison County
8 / l 6 / 5e
s ilty ,
gra v el
0-2
A rtem isia tr id e n ta ta imoderate
A. fr ie id a
P seudoreeneria sp ica ta
34.1 m iles from Alder on
Ruby River R eservoir
road, hfedison County
8 / 2 7 / 9b
loam
0-2
A rtem isia tr id e n ta ta
Bouteloua g r a c i l i s
yes
West sid e o f highway 19,
7.45 m iles south of
ju n ctio n 191 n o rth ,
Fergus County
west
6-10
Pseudoresneria sp ica ta • y e s
G rindelia. souarrosa
South sid e o f 2 0 0 s, 15.6
m ile s so u th ea st o f
C ir c le , Dawson County
ea st
3-5
A rtem isia tr id e n ta ta
West sid e o f 8 9 , 9 .1 5
m ile s north o f ju n ctio n
. south
3-5
Elymus can ad en sis
Poa spp.
Stip a comata
8/26/2b
.
cla y ey
s s p . a lb i c a u l i s
8 / 2 9 / 1a and Ib
8 /3 1 /3 b and 3c .
■ loam
loam
heavy
moderate
East sid e o f Bear Trap
R ecreation road, 2 . 1
m ile s south o f 84,
G a lla tin County
12 E ast, Meagher County
Table 13.
con tin u ed
Taxon and specimen
S o il te x tu r e
Aspect
S lop e
%
8 /3 1 /2 C
■8/26/2a
C. v i s c i d i f l o r u s
s s p . Ia n c e o la tu s
8 / l6 /2 a
• 8 / 2 0 / Ib
8 /2 0 /2 a
loam
cla y ey
west
Dominant a s s o c ia te d
p la n ts
Grazing
a t s ite
L ocation d e s c r ip tio n
0-2
Pseudoregneria sp ic a ta heavy
North sid e o f .84 , 7 . 5 5
m ile s where 86 m eets th e
Madison River on th e
e a s t sid e o f 84, Pkdison
County
6-10
Pseudoregneria sp ic a ta
G rin d elia squarrosa
South sid e o f 2 0 0 s, 15.6
m ile s so u th ea st o f
C ir c le , Dawson County
y es
VO
LO
south
6-10
A rtem isia tr id e n ta ta
yes
west
0-2
A rtem isia tr id e n ta ta
Juniperus seopulorum
heavy
south
6-10
K o eleria c r i s t a ta
, moderate
F estuca id a h o e n sis
Stip a spp.
south
6-10
1 .6 m iles e a s t o f
V irg in ia City
southw est o f 287 , Pkdison
County
I .05 m iles north of
H elm v ille on 141, Powell
County
P r i e s t ' s Pass road 6.4
m ile s northw est o f 1 2 ,
8 . 3 m iles w est o f H elena,
Lewis and Clark County
s s p . Ia n c e o la tu s
8 /2 0 /3 d
K o eleria c r i s t a ta
Festuca id a h o e n sis
S tip a spp.
moderate
P r i e s t ' s Pass road 6 . 2
m ile s northw est o f 1 2 ,
H elena, Lewis and Cla>k
County
Table 13.
con tin u ed
Taxon and specimen
S o il te x tu r e
Aspect
Slope
%
ss p . Ia n c e o la tu s
8 / 3 1/5a
C. v. ssp .
v is c id iflo r u s
v a r. ste n o n h v llu s
8 / l6 A d
8/2it/1a
sandyIoam
s ilty
sandyIoam
C. v . ss p .
v is c id iflo r u s
v a r. v i s c i d i f l o r u s
8 /l6 /2 b
var. v is c id iflo r u s
SZITZ1Id
.
Grazing
a t s ite
L ocation d e s c r ip tio n
west
20+
P seudorenneria sp ic a ta
A rtem isia cana
K o eleria c r i s t a ta
Poa spp.
yes
West sid e o f 287 , 4.1
m ile s south o f M orris,
Madison County
ea st
'10+20
P seudorenneria sp ica ta
A rtem isia arbuscula
yes
38.1 m iles w est from
Alder on Ruby River
R eservoir road, Madison
County
north
20+
Pseudorenneria sp ic a ta
A rtem isia cana
A rtem isia fr in id a
A rtem isia tr id e n ta ta
no
On 1 -9 0 , 0 .6 m ile s west
o f Springdale e x i t , Park
County
6-10
A rtem isia tr id e n ta ta
Juniperus son .
y es
1 .6 m iles e a s t o f
V irgin ia City southw est
o f 28?, Ifedison County
0-2
A rtem isia tr id e n ta ta
heavy
■ 4 ,6 m iles e a s t o f Twin
B rid ges - M elrose ro a d ,
Madison County
sou th ­
west
s ilty
Dominant a s s o c ia te d
p la n ts
libraries
3 1762