AN ABSTRACT CF THE t
Kenric 3. Walburger for the d'gre c
SERTATION OF
torc Philosophy in Animal Science
presented on October 28, 20Q5.
Title: The Effects of Timber larJeta c -irL
ry on Understory Vetation and
Con pos'tion of Beef Cattle
hstrac1: Approved:
Redacted for privacy
Timoth4 DelCurto
The objectives of the experiment were to: 1) determine the relatfrnship
between overstory canopy cover and unqulate herbivory on uner:ory production
composition and diversity and 2) deb-rmine the effects of timber harvest, ungulate
herbivory and season of use on botanica composition and nutritional quaty o
beef cattie diets. Three randomly sccted grand
fir (Ahies grand&
and
ponderosa pine (Pfnusponderosa sites were established in 1986. Timber arest
and herbivory treatments were arranged in a spilt-plot design.
Results from grand fir forests indicated that timber havct had a greater
magnitude effect on understory species and shrub responses t-'r herbivory
treatments. Understory production was sic
antly greater in treatments that
aUowed domestic and wild ungulate herbivory compared to total removal of
ungulate herbivory. Production of understory 'egetation increased with
decreasing canopy cover. Plant communiticswer affcted by timber harvest and
;::-g all timber harvest treatments,
herbivory. There appeared
whereas, among herbivory treathierits, tote! rmoval of ungulate herbivory
appeared to be different th
other herhvcr treatments.
Results from ponderosa pin
treatnints indicated that there were no
directional responses attributed to either timber harvest or herbivory on
understory plant species frequency. Understory production was significantly
greater in treatments with no ungulate herbivory compared to treatments with
domestic and wild ungulate herbivory. Production was significantiy greater in
commercially thinned stands compared to non- harvested stands with greater
canopy cover. Plant communities were affected by timber harvest but not by
herbivory.
Nutritional quality of diets was significantly greater in June than in August
for both grand fir and ponderosa pine sites. Timber harvest treatments only
affected ADF content in grand
sites, whereas, herbivory treatments had no
affect on nutritional quality of diets. Graniinoids were the major constituent in the
diet, forbs were intermediate and shrubs were the least. Season of use did not
influence the botanical composition of
diets.
This research indicates that the effects of timber harvest may have a
greater effect on understory vegetation composition, structure and production
than herbivory. In addition, timing of grazing had a greater influence on diet
quality than did herbivory and(or) timber harvest.
©Copyrght by Kenric 3. Walburger
October 28, 2005
All Rights Reserved
The Effects of Timber Hr.:-: nd Herb yy n Understory Vegetation and
Composition of 3eef Cattle Det on Forested Rangelands
by
Kenric J. Waiburger
A Dissertation
submitted to
Oregon State Univeristy
in partial fulfillment of
the requirements of the
degree of
Doctor of Philosophy
Presented October 28, 2005
Commencement June 2006
Doctor of Philosoihy dissertton of
1cLaJurger presented on October 28,
2005.
APPROVED:
Redacted for privacy
Major Professor, representing Anim
Redacted for privacy
Hef the Depament of Animal Sciences
Redacted for privacy
Dean of the-Graduate School
I understand that my dissertation will become part of the permanent collection of
Oregon State University libraries. My signature below authorizes release of my
dissertation to any reader upon request.
Redacted for privacy
Kenric 3. Wa!burger, Author
CKNuWLEDGENTS
This research and graduate program were uported by the Eastern Oregon
Agriculture Center. I am tbrkfu for thc for;icht of Dr. Martin Vavra for
beginning this study in 198 and f011owing it though his tenure as superintendent
and also for Dr. Timothy DelCurto for helping maintain this project. Dr. Timothy
DelCurto, my major professor, mentor, and now a good friend. Tim, without your
assistance and support I would not have been able to complete this dissertation. I
would like to thank all the committee members Drs. Martin Vavra, Patricia
Kennedy and James Males for their time and constructive criticism throughout this
process. You all contributed your expertise to make this dissertation and program
better. Appreciation is also exo: to m t;aduate school representatives Dr.
Steven Strauss, for sitting on the other end of a phone for my preliminay exam
and Dr. Vasconcelos, for the final exam.
i would like to extend appreciation to tne summer students that were hired
to help with rumen evacuations and forage clipping. I would like to thank fellow
graduate students for their help and input aionq the way. I would especially like
to thank Micah "Weasel" Wells for the comic relief and friendship throughout the
time in Union.
Finally, I would like to express the sincerest thanks to my wife, Kimberly
for her moral support and understanding throughout our time in graduate school.
f not for her tremendous support I would not have been able to make it. This
degree is as much hers as nne. My kid 3n and Mackenzie, Jam finafly going
to be done and have a re& :ob
Thank you everyone'
CONTh UT1CN
Dr. Timothy DelCurto assisted
ir
pahnina
F tUTHORS
data
manuscript preparation. Dr. Martin Vavri
initiated and maintaining the xperment vr
toHection, data analysis and
in manuscript preparation and
years.
TABLE OF: CCNTTS
Page
Chapter 1: Introduction
LiteratureCited
1
.................................................................................
Chapter 2: The Effects of Herbivory and limber Harvest on
Understory Vegetation in a Ponderosa Pine (Pinuspondcrosa
Habitat in Northeastern Oregon ......................................................
7
11
Abstract.............................................................................................
12
Introduction......................................................................................
14
Materials and Methods
.....................................................................
15
Vegetation Sampflng ..................................................................
Statistica' AnaIysi5 ........................................................................
20
Results..............................................................................................
22
CanopyCover ............................................................................
L!nderstory Production ................................................................
Species Composition ..................................................................
Shrub Density and Cover .............................................................
Plant Community Responses ......................................................
22
22
25
29
33
Discussion................................................................................
36
Implications......................................................................................
40
Literature Clited ................................................................................
41
Chapter 3: The Effects of Herbivory and Timber Harvest on
Understory Vegetation in a Granc Fir (Ah'ësgrandis Habitt in
Northeastern Oregon ...................................................................
18
45
Abstract.........................................................................................
46
Introduction......................................................................................
48
Materials and Methods
......................................................................
49
..................................................................
52
Vegetation Sampling
TAE F CONTEIvTS Continued)
Statistical Analysis
...................................................................
Page
54
Results..............................................................................................
56
CanopyCover ............................................................................
Species Composition ...................................................................
Shrub Density and Cover .............................................................
Understory Production ................................................................
Plant Communfty Responses .......................................................
56
57
62
63
65
Discussion.........................................................................................
67
Implications......................................................................................
72
LiteratureCited
.................................................................................
73
Chapter 4: Effects of Timber Harvest and Herbivory on Subsequent
Cattle Diets on Forested Rangelands ..............................................
78
Abstract
79
............................
Introduction.....................................................................................
81
Materials and Methods
.....................................................................
82
Results..............................................................................................
88
GrandFir Sites ............................................................................
PonderosaPine Sites ..................................................................
88
92
Discussion.........................................................................................
94
Management implications
.............................................................
98
................................................................................
100
Chapter 5: Conclusions .........................................................................
103
Bibliography..........................................................................................
108
Appendices.............................................................................................
115
Literature Cited
.i.
£
Figure
2.1
Layout of timber h::ri:
cattle and
big game grazing
cntrd aid hnned) and herbivory
ne grr; CExc
(Graze
;
Lxc;
cattle exclosure,
x::sure, exclusion of
cattle and big game grng) tretn3 for each ponderosa
....................................................................................
17
The effects of herbivory treatments on the understory
production (kg/ha) within a ponderosa pine forest in eastern
Oregon......................................................................................
24
Changes in the production of elk sedge over time within a
ponderosa pine forest in northeastern Oregon ........................
25
The effects of herbivory treatments on snowberry production
(kg/ha) within a ponderosa pine forest in eastern Oregon
26
The changes in rose (Rosa gymnocarpa Nutt.) density over
time between 1988, 1991, 1994 and 2003 with 1985
34
pinesite
2.2
2.3
2.4
2.5
2.6
NMS Ordination of undertory species composition in 2003 of
rv-.t effects, (C) herbivory
(A) site differences,
effects in a ponderus p'c oret n northeastern Oregon
3.1
3.2
3.3
4d
....
35
Layout of the timber harvest (control, thinned and clearcut)
and herbivory (Graze cattle and big game grazing; CExc cattle exclosure, big game grazing only; TExc total
exclosure, exclusion of cattle and big game grazing)
treatments for each grand fir site ............................................
51
Overstory canopy cover (%) of grand fir sites in 1985, prior to
timber harvest, and subsequently in 2003 ..............................
56
NMS Ordination of the changes in understory species
composition from 1985 to 2003 of (A) site thfferences, (B)
timber harvest effects, and C) herhivorv effects in a grand fir
forest in northeastern Orcg.n ..................................................
66
Precipitation data (cm) from weather station located at
Eastern Oregon Atriculture Research Center's Hall Ranch
83
L1
G(JRE.
Li:rinued)
Figure
4.2
4.3
Layout of the timber hrvst (controL
nned and clearcut)
and herbivory (Graze catd and hie grazing; CExc
total
cattle exclosure, big qame azing o'; rEx
me grazing)
exciosure, exclusion of cattk and bg
treatments for each grand fIr site ............................................
84
Layout of timber harvest (control and thinned) and herbivory
(Graze cattle and big game grazing; CExc cattle exclosure,
big game grazing only; TExc total exciosure, exclusion of
cattle and big game grazing) treatments for each ponderosa
pinesite ....................................................................................
84
LIST QF TA2LES
Table
2.1
2.2
2.3
2.4
2.5
2.6
2.7
2.8
2.9
2.10
3.1
Page
Precipitation data (cm), for vegetation srnpUng years, from
weather station located äi Eastern Oregbn Agriculture
Research Center's HaU Ranch ..................................................
16
The effects of timber harvest on the understory production
(kg/ha) within a ponderosa pine forest in eastern Oregon
23
Species frequency means in 1985, prior to experiment
initiation, in a ponderosa pine forest in northeastern Oregon
..
27
The effects of timber harvest and year on the difference in
species frequency from 1985 in a ponderosa pine forest in
northeastern Oregon ................................................................
28
The effects of timber harvest and herbivory on the change in
species frequency from 1985 in a ponderosa pine forest in
northeastern Oregon ................................................................
29
The effects of timber harvest and herbvory on the change in
understory species frequency from 1985 in a ponderoa pine
forest in northeastern Or:!"n ..................................................
30
Plant species richness, diversity, and evenness responses to
timber harvest and herhivory in a ponderosa pine forest in
northeastern Oregon ...............................................................
31
Understory species turnover rates from 1985 to 1988, 1991,
1994, 1997 and 2003 in a ponderosa pine forest in
northeastern Oregon ................................................................
32
Shrub species density (#/ha) and cover in 1985, prior to
experiment initiation, in a ponderosa pine forest in
northeastern Oregon .................................................................
32
The effects of timber harvest and herbivory on the changes
from 1985 of understory shr.ib density (#/ha) and cover in a
ponderosa pine forest in northeastern Oregon .......................
33
Precipitation data (cm), for vegetation sampling years, from
weather station located at Eastern Oregon Agriculture
Research Center's Hall Ranch ..................................................
50
15t3FT1ABLES :.niied)
Table
3.2
3.3
3.4
3.5
3.6
3.7
3.8
3.9
3.10
4.1
4.2
Page
Species frequency means in 1985, pror to experiment
initiation, in a grand fir ret in nor:heastern Oregon
57
The average shrub spedes densfty (#fh a) and cover (%) in
1985, prior to experiment initiation, in a grand fir forest in
northeastern Oregon .................................................................
58
The effects of timber harvest and herbivory treatmentson the
changes in species composition, from 1985 through 2003, in a
grand fir forest in northeastern Oregon ..................................
59
Changes in species composition, since 1985, in a grand fir
forest in northeastern Oregon .................................................
60
Changes in species composition, from 1985, as affected by
time and timber harvest treatments in a grand fir forest in
northeastern Oregon ..................................................................
60
Changes in species composition, from 1985 to 2003, as
affected by timber harvest and herbivory treatments in a
grand fir forest in northeastern Oregon ..................................
61
Plant species richness, diversty, and evenness responses to
timber harvest and herbivory in a grand fir forest in
northeastern Oregon ..............................................................
62
The effects of timber harvest and herbivory on understory
species turnover rates from 1985 to 2003 in a grand fir forest
in northeastern Oregon ............................................................
63
The effects of timber harvest and herbivoiy on understory
production in a grand fir forest in northeastern Oregon
64
The effect of season of use and timber harvest on the
subsequent quality of steer diets in a grand fir forested
habitat.......................................................................................
89
The effects of timber harvest and herbivory treatments on the
botanical composition (%) of steer diets in a grand fir forested
habitat......................................................................................
90
LsT CF TABLES cConnued)
Table
4.3
4.4
4.5
4.6
4.7
The effects of timber harvest and herbivo!y treatments on the
relative preference index RPI) of uriiimoids, forbs, and
shrubs for steer diets in grand Ill forest:ed habitat
91
The effects of timber harvest on understory production (kg.
ha-'), 18 years post-harvest, in a grand flr forest in
northeastern Oregon .................................................................
92
The effects of season of use on the diet quality of steers
grazing a ponderosa pine forested habitat ..............................
92
The effects of herbivory on the botanical composition (%) and
relative preference index (RPI)* of steer diets within a
ponderosa pine forest type .......................................................
93
The effects of timber harvest on understory production (kg.
ha-'), 18 years post-harvest, in a ponderosa pine forest in
northeastern Oregon ................................................................
95
LJ T OF APN DICES
Appendix A: List of Species Identied in Grrd Fir and Ponderosa Pine
Sites...................................................................................
166
Appendix B: Ponderosa Pine Understory Vegetation Changes
123
Appendix C: Ponderosa Pine Shrub Density and Cover Changes
154
Appendix D: Grand Fir and Ponderosa Pine Understory Production
161
......................
166
Appendix F: Grand Fir Shrub Density and Cover Changes ..................
191
.....................................
200
Appendix E: Grand Fir Understory Vegetation Changes
Appendix G: Nutritional Quality of Steer Diets
LIST OF APPNflx TABLES
Table
A.1
A.2
6.1
B.2
B.3
B.4
B.5
B.6
B.7
B.8
B.9
B.10
Cd
C.2
Page
List of all species identifle by frquercy counts for ponderosa
pinesites ..................................................................................
117
List of all species identified by frequency counts in grand fir
sizes..........................................................................................
120
The changes in species composition in a ponderosa pine
forest from 1985 to 1988 using a 30 cm x 30 cm plot frame
...
124
The changes in species composition in a ponderosa pine
forest from 1985 to 1988 using a 30 cm x 60 cm plot frame
...
127
The changes in species composition in a ponderosa pine
forest from 1985 to 1991 using a 30 cm x 30 cm plot frame
...
130
The changes in species composition in a ponderosa pine
forest from 1985 to 1991 using a 30 cm x 60 cm plot frame
...
133
The changes in species composition in a ponderosa pine
forest from 1985 to 1994 using a 30 cm x 30 cm plot frame
..
136
The changes in species composon in a ponderosa pine
forest from 1985 to 1994 using a 30 cm x 60 cm plot frame
...
139
The changes in species composition in a ponderosa pine
forest from 1985 to 1997 using a 30 cm x 30 cm plot frame
...
142
a ponderosa pine
The changes in species composition
forest from 1985 to 1997 using a 30 cm x 60 cm plot frame
...
145
The changes in species composition in a ponderosa pine
forest from 1985 to 2005 using a 30 cm x 30 cm plot frame
...
148
The changes in species composition jr a ponderosa pine
forest from 1985 to 2005 using a 30 cm x 60 cm plot frame
...
151
Changes in shrub density from 1995 to 1988, 1991, 1994, and
2003 .........................................................................................
155
Changes in shrub cover from 1995 to 19S8, 1991, 1994, and
2003 .........................................................................................
158
ifl
LIST C-
f ?NDIX
(Continued)
Page
Table
E.1
E.2
E.3
E.4
E.5
E.6
F.1
F.2
G.1
G.2
G.3
G.4
The changes in species c npositon r1 a grand fir forest from
1985 to 1994 using a 30 cm x 30 cm plot frame ....................
167
The changes in species composition in a grand fir forest from
1985 to 1994 using a 30 cm x 60 cm plot frame ....................
171
The changes in species composition in a grand fir forest from
1985 to 1997 using a 30 cm x 30 cm plot frame ....................
175
The changes in species composition in a grand fir forest from
1985 to 1997 using a 30 cm x 60 cm plot frame ....................
179
The changes in species composition in a grand fir forest from
1985 to 2003 using a 30 cm x 30 cm plot frame .....................
183
The changes in species composition in a grand fir forest from
1985 to 2003 using a 30 cm x 60 cm plot frame ....................
187
The changes in shrub density from 1995 to 1991, 1994, and
2003 ..........................................................................................
192
The changes in shrub densft\' from 1995 to 1991, 1994, and
2003 ..........................................................................................
196
Nutritional quality of diets collected In June and August of
2001 and 2002 for ponderosa pine sites .................................
201
Botanical composition and relative preference index for steer
diets collected in June and August of 2001 and 2002 for the
ponderosa pine sites ................................................................
203
Nutritional quality of diets co'lected in June and August of
2001 and 2002 for grand fir sites ............................................
205
Botanical composition and relative preference index for steer
diets collected in June and August of 2001 and 2002 for the
grandfir sites ...........................................................................
208
ri: Mit*IJ
INTRODUCTION
2
Forested rangelands comp;-e a sghThant portion of the public land in
the western US. Within the northwestern US. (Idaho, Montana, Oregon, and
Washington) forested rangends crnprite
land area, and public ownership comprises
proximately 35 percent of the total
:er:ent of these forested
rangelands (Smith et al. 2002). Porests of the interior Northwest, on average,
receive greater precipitation, have deeper soil profiles, and have greater soil
moisture holding capacity than adjacent grasslands (Riggs et al. 2004). Ponderosa
pine (Pinusponderosa and grand fir (Abies grandis are two of the forest types
that are commonly found within the region. These productive forests provide
valuable habitat for many wildlife species and summer grazing allotments for beef
cattle.
However, over the last 100 years, mary of these forests have developed
into relative homogeneous stanc;
pcsn and structure of fire-sensitive
and disease-susceptible species (eisky and Blumenthal 1997; Hessburg et al.
2005), thereby reducing the overa! understosy productivity of the area. Many
areas with high potential for timber harvest and forage production have the lowest
output due to dense canopy cover (Hedrck et al. 1969).
To restore productivity to these dense stands, in terms of forage and
timber production, it may be necessary to open the canopy. Forage production
response to overstory canopy cover is we documented (Mcconnell and Smith
1965 and 1970; Jameson 1967; Young et aL 1967; Thompson and Gartner 1971;
Miller et al. 1986) and results suggest that as overstory canopy cover decreases
understory production (kg/ha) increases. Clary et al. (1975) also documented this
3
nd f- prductipn, but noted there was
relationship between canopy c':
an economic optimum between rrasir:g orace production for cattle grazing and
Hovr, this optimum may not be
increased timber growth for
ecologically sustainable.
Ur
tOi
considered if grazing and loggn
plartspci
responses/trajectories must be
ar to- be sustnable.
Timber harvest on forested rangelands sets back succession, and generally
the overstory development proceeds with the following pattern: herb and shrub
to tall shrub to juvenile tree stage development and, finally, replacement of shadeintolerant for shade-tolerant species (Bainbridge and Strong 2005). However,
understory vegetation successional development can be influenced by the level of
disturbance (Griffis et al. 2001; Gibbs et al. 2004), residual species (Halpern 1988;
Selmants and Knight 2003), ur icrttrv
undisturbed stands (Selmants -ui
composition of adjacent
iiyht: 20)3), and possibly by soils (Bennett et
a. 1987). Schoonmaker and McKee (1988) reported that species richness
increased to peak at 15 to 20 years after togging and burning in a coniferous
forest in western Oregon. They also documented that species diversity is greatest
up to 20 years following disturbance.
Following disturbances, such as timber harvest, forested rangelands often
become focal points of ungulate herbivory for 2. reasons: 1) vegetation within
disturbed sites is often more palatable than undisturbed areas an
2) surrounding
undisturbed forest communities conLain Umited understory forage (Vavra et al.
2004). As a result, grazing or herbivory may influence plant community structure
and diversity within these environments. Hobbs (1996), Augustine and
'S
McNaughton (1998), Riggs
(Z000), ar.i
documented that ungulate herbivc'ry can
nd Lehmkuhl (2001) all
h1tce plant community structure and
composition. Ungulate herbivor directly affects vegetation through selective
feeding and the ability of a plant to recover rron herbivory (Augustine and
McNaughton 1998). However, plant species diversity is not consistently affected
by grazing (Stohlgren et al. 1999; Riggs et al. 2000; Brockway and Lewis 2003),
but individual species, at the local level, are affected by herbivory and logging and
their interaction. Herbivory has the greatest effect in altering plant community
structure within clearcuts (Riggs et al, 2000). Stohlgren et al. (1999) also
documents that grazing effects local species; however few plant species showed
consistent directional responses to grazing.
Timber harvest increases the opportunity for cattle/wildlife to forage and
obtain a higher quality diet and subsequeridy increase productivity. Several
researchers (Harris 1954; Young et al. 1967; Roath and Krueger 1982; Gillen et al.
1984) have documented that cattle se increased in areas with greater forage
production. Young et al. (1967) also noted that it was more difficult to get
moderate to heavy utilization under dense overstory canopy covers than more
open canopy covers. The understery vegetation under open canopies is typically
grass dominated with greater forage production than under more closed canopies
(McConnell and Smith 1965 and 1970). As a resuft, cattle grazing on forested
rangelands typically selected diets that were high in grass, moderate i
forbs and
minimal in shrubs (Holeckek et al. 1982; Quinton 1984; Mitchell and Rodgers
1985; Uresk and Paintner 1985). However, cattle diets can vary as a grazing
5
season progresses, increasu1i prcrtior;;'
c:s and shrubs, while
decreasing in grasses (Holechek et &. 1982; Mitthell and Rodgers 1985; Uresk
and Paintner 1985). Although cattle diets dJqe with progression of the grazing
season, nutritional quality of caWe diets ondis to decline as well (Walker et al.
1989).
Holechek et al. (1981), Kirby and Parman (1986), and Griggs et al. (1995,
2001) documented that nutritional quality of cattle diets declined with progression
of the grazing season. Holechek et al. (1981) further noted, in a mixed-conifer
forest, that CP content was not affected by season of use but IVOMD declined
with progression of the grazing season. However, Kirby and Parman (1986) and
Griggs et al. (1995, 2001) documented, on grassland sites, that CP and IVOMD
declined with progression of the grazing season. Therefore, the nutritionaF quaRty
of cattle diets may be affected by previous ungulate herbivory, canopy cover
and/or plant community changes.
Although, much is known about how quantity of vegetation responds to
canopy cover changes, less is known about how herbivory can affect plant
community structure. Even tess is known about how plant community responds to
the combined effects of herbivory and overstory canopy within forested
environments. Therefore the objectives of th
1.
study are:
Determine the relationship between overstory canopy cover and ungulate
herbivory on understory production, understory species composition and
diversity.
2.
Determine the effects ;f t'1ibr hr':, ugulate herbivory and season
of use on botanical cornpoition and nutrftional quality of beef cattle diets.
The following chapters wi precnt the cit coflected to answer the two
objectives. Chapter 2 conta1s the results of the changes in plant community
structure, production and cornpositon of the ponderosa pine sites. Chapter 3
contains the plant community structure, production, and compositional changes of
the grand fir sites. Finally, Chapter 4 contains the botanical composition and
nutritional quality of steer diets grazing both the grand fir and ponderosa pine
sites.
7
LITERATURE CITED
Augustine, Di. and Si. McNaughtor. 19)5. Ungulate Effects on the Functional
Species Composition of Plant Cornmunit!es Herbivore Selectivity and Plant
Tolerance. Journal of Wildlife ' 'iagrment 62: 1165-1183.
Bennett, D.L., G.D. Lemme, and P.D. Even;h, 1937. Understory herbage
production of major soils wfthin the E3ak HIHs of South Dakota. Journal
Range Management. 40:166-170,
of
Bainbridge, E.L. and W.L. Strong. 2005. P/pus contorta understory vegetation
dynamics following clearcutting in west-central Alberta, Canada. Forest Ecology
and Management 213:133-150.
Belsky, J.A. and D.M. Blumenthal. 1997. Effects of Livestock Grazing on Stand
Dynamics and Soils in Upland Forests of the Interior West. Conservation
Biology. 11: 315-327.
Brockway, D.G. and C.L. Lewis. 2003. influence of Deer, Cattle Grazing and
Timber Harvest on Plant Species Diversity in a Longleaf Pine Bluestem
Ecosystem. Forest Ecology and 1''anagement 175: 49-69.
Gillen, R.L., W.C. Krueger, and R.F.. MiUer. t9&4. Cattle Distribution on Mountain
Rangeland in Northeastern Orçon. o-rlof Range Management. 37: 549553.
Griffis, K.L., J.A. Crawford, M.R. Waqner, W.H. Moir. 2001. Understory response to
management treatments n northern Arizona ponderosa pine forests.
Forest Ecology and Management. 146:239-245.
Grings, E.E., R.E. Short, M.R. Haferkamp, and R.K. Heitschrnidt. 2001. Animal age
and sex effects on diets or grazing cattle. Journal of Range Management
54:77-81.
Grings, E.E., D.C. Adams, and R.E. Short. 1995. Diet quality of suckling calves and
mature steers in Northern Great Plains rangelands. Jouiial of Range
Management 48:438-441.
Halpern, C.B. 1989. Early successional patterns of forest species: interactions of
life history traits and disturbance. Ecology. 70:704-720.
Harris, R.W. 1954. Fluctuations in Fo'age Utihzation on Ponderosa Pine Ranges in
Eastern Oregon. Journal of Range Management 7: 250-255.
Hedrick, D.W., B.R. Eller, J... L. !'l:.Arthur, .......... 9. Pettit. 1969. Steer Grazing
on Mixed Coniferous Forest Rargs
Management. 22: 322-325.
Nerhastern Oregon. Journal of Range
Hobbs, N.T. 1996. Modiflcaton of Ecosysten
Management. 60: 695-713.
by iJngulates. Journal of Wildlife
Holechek, J.L., M. Vavra, and JSkov!n. 1981 D quality and performance of
cattle on forest and grassland rarge.Jcir,iaiofAnimalScience. 53:291-198.
Holechek, J.L., M. Vavra, J. Skovlin, and W.C. Krueger. 1982. Cattle Diets in the
Blue Mountains of Oregon II. Forests. Journal of Range Management. 239-242.
Holechek, J.L., R.D. Pieper, and C.H. Herbel. 1995. Range Management: Practices
and Principles (2 Edition). Prentice Hall. Englewood Cliffs, New Jersey. p.
195-916.
Jameson, D.A. 1967. The Relationship of Tree Overstory and Herbaceous
Understory Vegetation. Journal of Range Management. 20: 247-249.
Kie, J.G. and J.F. Lehmkuhl. 2001. Herbivory by Wild and Domestic Ungulates in
the Intermountain West. Northwest Science. 75: 55-61.
Kirby, D.R. and M. Parman. 1986. Botanical composition and diet quality of cattle
under a short duration grazing system. Journal of Range Management. 39:509512.
McConnell, B.R. and J.G. Smith. 1965. Understory Response Three Years After
Thinning Pine. Journal of Range Management. 18: 129-132.
McConnell, B.R. and J.G. Smith. 1970. Response of Understory Vegetation to
Ponderosa Pine Thinning in Eastern Washington. Journal of Range
Management. 23: 208-212.
Miller, R.F., W.C. Krueger, and M. Vavra. 1986. Twelve Years of Plant Succession
on a Seeded Clearcut Under Grazing and Protection from Cattle. Special Report
773. Corvallis, OR: Oregon State University, Agricultural Experiment Station. p
4-10.
Mitchell, i.E. and R.T. Rodgers. 1985. Food Habits and Distribution of Cattle on a
Forest and Pasture Range in Northern Idaho. Journal of Range Management.
38: 214-220.
Quinton, D.A. 1984. Cattle Diets on Seeded Cearcut Areas in Central Interior
British Columbia. Journal of Range Management. 37: 349-352.
Riggs, R.A., A.R. Tiedemanf', G. Coak, T,N.
rd, P.1 Edgerton, M. Vavra,
W.C. Krueger, F.C. Hall; L.D. Bryant, L.L. rvin, and 1. DelCurto. 2000.
Modification of Mixed-Conifer' Forests by Rumiiiant Herbivores in the Blue
Mountains Ecological Province, Res. Pj: PNW-RP-527. Portland, OR: U.S.
Department of Agriculture, Forest: Service. Padfic Northwest Research Station.
77 p.
Roath, L.R. and W.C. Krueger. 1982. Cattle Grazing Behavior on a Forested Range.
Journal of Range Management. 35: 332-338.
Schoonmaker, P. and A. McKee. 1988. Species composition and diversity during
secondary succession in coniferous forests in the western Cascade Mountains
of Oregon. Forest Science. 34:960-979.
Selmants, P.C. and D.H. Knight. 2003. Understory plant species composition 30-50
years after clearcutting in southeastern Wyoming coniferous forests. Forest
Ecology and Management. 185:275-289.
Smith, W.B., J.S. Vissage, D.R. Darr, and R.M, Sheffield. 2002. Forest Resources of
the United States, 1997, Metric Units. GTR-NC-222. St. Paul, MN: U.S.
Department of Agriculture, Forest Service, North Central Research Station. 127
p.
Strickler, Gerald S. 1959. Use of the Densiometer to Estimate Density of Forest
Canopy on Permanent Sample Plots. Research Note No. 180. Portland, OR:
U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and
Range Experiment Station. S p.
Thompson, W.W. and F.R. Gartner. 1971. Native Forage Response to Clearing Low
Quality Ponderosa Pine. Journal of Range Management. 24: 272-277.
Uresk, D.W. and W.W. Paintner. 1985. Cattle Diets in a Ponderosa Pine Forest in
the Northern Black Hills. Journal of Range Management. 38: 440-442.
Vavra, M., M.3. Wisdom, 1G. Kie, 1G. Cook, and R.A. Riggs. 2004. The role of
ungulate herbivory and management on ecosystem patterns and processes:
future direction of the Starkey project. In: 1 Rahm (ED.), Transactions of the
Sixty-ninth North American Wildhfe and Natural Resources Conference. 16-20
March 2004; Spokane, WA: Wildlife Management Institute. p. 785-797.
Walker, J.W., R.K. Heitschmidt, E.A. e Moraes, M.M. Kothmanri and S.L.
Dowhower. 1989. Quality and Botanical Composition of Cattle Diets Under
Rotational and Continuous Grazing Treatments. Journal of Range
Management. 72: 239-242.
10
Young, IA., A.B. McArthur, ziid 1W. Hedrkk.
Forage Utilization in a
Mixed-Coniferous forest of NOrtheastern Oregon. Journal of Forestiy. 65:
393.
391-
11
CHAPTEfl 2
The Effects of Herbivory and Timber Harvest on Understory Vegetation
in a Ponderosa Pine (Pinusponderosa) Habitat in Northeastern Oregon.
Kenric J. Wa/burger,' Timothy De/Curto,2 and Martin Vavr
Authors are: 1 Graduate Research Assfstar,t. 2 Associate Professor, Department of
Animal Sciences, Eastern Oregon Agriculture Research Center, Oregon State
University, Union, OR 97883,' Superv/soiy Range/and Scientist, Pacific Northwest
('PNW) Research Station, USDA Forest Scricc, Porestiy and Range Sciences
Laboratory, La Grande, OR 97850
12
ABSTRACT
The objectives of this study were t document the effects of logging and
herbivory, cattle and native ungulates, oh understory vegetation within a
ponderosa pine (Pinusponderosa) habitat. Three randomly selected sites were
established in 1986. Ponderosa pine sites were arranged as a split-plot design and
timber harvest treatments [1) commercial thinning (TH), 2) control (CON)] were
whole plots and herbivory treatments [1) cattle and native ungulate grazing (GR),
2) native ungulate grazing (BG), and 3) exclusion of cattle and native ungulates
grazing (EX)] were the sub-plots.
Understory production was affected by timber
harvest and herbivory. Totai production was greater in (P < 0.05) TH than in CON
for all years sampled and production was greater in the BG and EX when
compared to GR. A total of 149 species were identified on these sites, but only 35
species occurred in every treatment combination which were used for analysis. In
general, there were no directiona! responses attributed to either timber harvest or
herbivory on understory plant species frequency. Herbivory and timber harvest
did not affect (P> 0.10) understory plant species richness, Shannon diversity, or
evenness.
Differences occurred among timber harvest and herbivory treatments
on individual shrub species density and cover. Non-metric multidimensional
scaling (NMS) ordination and multi-response permutation procedures (MMRP)
analyses revealed that plant communities were different in 1985 prior to study
initiation and these differences were maintained through 2003. Plant communities
13
in TH were different than i
plant community structure.
howev:r, it-:ory did not appear to affect
his earc ihdicates that the effects of timber
harvest may have a greater effet oh uncerstOry vegetation composition,
structure, and diversity than
herbiJory.
Keywords: Beef cattle, Native ungulates, Mule ceer, Elk, Succession
14
INTRODUCTION
Forested rangelands comprise
western US. Ponderosa pine (P.
siqriiflcant p!tiüfl of the public land in the
1:nc-as P& C. Lawson) forests provide the
most extensive and important forested grazing
nds !n North America (Skoviin
1976). These forests also provide valuable habitat for many wildlife species.
However, over the last 100 years, many of these forests have developed into
relative homogeneous stands in composition and structure of fire-sensitive and
disease-susceptible species (Belsky and Blumenthal 1997; Hessburg et al. 2005),
thereby reducing the overall productivity of the area. Traditionally! many of these
areas with high potential for timber harvest and forage production have the lowest
output due to dense canopy cover (Hedrick et al. 1969).
Two uses common to western ponderosa pine forests are timber and
forage production. To enhance these, thinning of overstory canopy maybe a
useful tool, but the effects of this manipulation on forage production, and
understory community structure and diversity is not well known. Timber harvest
on forested rangelands sets back succession and, in most cases, increases
understory forage production. Forage production response to overstory canopy
cover is well documented (McConnell and Smith 1965 and 1?70; Jameson 1967;
Young et al. 1967; Thompson and Gartner 1971) in that as overstory canopy cover
decreases understory production (kq/h) increases.
Herbivory may also influence plant community structure, diversity, and
production within forested environments. Hobbs (1996), Augustine and
McNaughton (1998), and R;
&. (20Cij.
documented that herbivory can ihi!ucnce
nd Lehmkuhl (2001) all
t
composition. Ungulate herbvory drctly äft
feeding and the ability of a pmt
mmunity structure and
vegetation through selective
covr ftom nerbivor (Augustine and
McNaughton 1998). However, pknt species thvrsity was not consistently affected
by ungulate herbivory (Stohlgren et at, 1999, Riggs èt al. 2000 and Brockway and
Lewis 2003), but individual species, at the local level, were affected by herbivory
and timber harvest. Herbivory had the greatest effect in altering plant community
structure within clearcuts (Riggs et at. 2000). Stohlgren et at. (1999) also
documented grazing effects on local species, however few plant species showed
consistent directional responses to grazing.
As stated earlier, much is known about how quantity of vegetation
responds to canopy cover. However, little is known about how plant community
structure and composition responds when combining the effects of herbivory and
overstory canopy within a ponderosa pine forest. Therefore the objective of this
study was to document the effects of timber harvest and herbivory on understory
vegetation composition, structure and diversity.
MATERIALS AND METHODS
The study area is located at the Eastern Qregon Agriculture Research Center's Hall
Ranch that is approximately 16 km east Of the city' of Union in the Waltowa
Mountains of northeastern Oregon with an elevation range from 1050 to 1250 ni.
I
16
Summers tend to be dry and :3rm
ith t:;mp-ures rarely exceeding 38°C,
though freezing or near-freezing temperture are possible all year. Winters are
cold and wet, with the majority of th predpftton coming in the form of snow
between November and May. Avraqe annua credpitation for the Hall Ranch was
56 cm (Table 1). Elk (Cèivus elaphus
L.)
and mule deer (Odoco/leus hem/onus
Raf.) are indigenous to the area and can be found throughout the year; however,
heaviest use occurs in the spring and the fall.
Table 1. Precipitation data (cm), for vegetation sampling years, from weather
station located at Eastern Ore9cuiture Research Center's Hall Ranch.
Precipitation (cm)
Year
Jul AuQçp Oct Nov Dec Total
Jan Feb Mar Apr
y)n
1985
1988
1989
1991
1992
1994
1995
1997
2003
1.3
6.1
8.9
4.6
2.0
6.1
9.7
5.8
6.1
4.3
4.1
4.1
3.0
4.6
7.6
3.0
5.3
6.4
4.9
3.8
6.4
9.7
6.6
4.1
4.3
1.3
2.3
4.6
7.4
9.7
10.4
7.4
6.1
9.1
1.5
9.9
5.5
5d
6.1
5.8
5.6
6.1
13.0
1.5
7.4
5.9
4.3
7.1
3.3
8.1
3.6
7.9
9.1
3.3
7.6
3.6
1.0
6.64.7
1.0
0.0
0.5
0.5
2.8
LC
2.0
5.3
0.0
1.6
2.5
1.0
9.1
0.3
0.5
0.0
4.1
1.3
3.6
1.8
5.3
0.8
2.5
0.0
3.0
0.8
1.3
3.3
2.5
2.3
5.6
0.5
3.8
3.8
4.3
3.6
4.1
3.8
2.0
3.6
8.9
11.9
4,3
1L4
6.4
10.4
12.4
4.1
3.6
6.6
2.5
7.4
2.0
2.0
8.1
3.6
9.1
7.6
8.9
5.9
48.5
56.1
59.7
59.2
48.3
53.3
79.0
56.4
58.4
55.6
Three replicated sites of a Piiusponderosa P.& C. Lawson /
Symphoricarpos a/bus (L.) Blake (ponderosa pine) habitat type, 15 ha in size, were
selected to analyze the effects of herbivory and timber harvest on understory plant
communities. Potential areas, with relatively homogeneous stand structure and of
the necessary size, were initially identified and then research sites were selected
from these areas. The ponderosa pine community had two timber harvest
17
treatments applied to them (Fiy.e 1); 1) c':
''''
e:'cial thinning (thinned) and 2)
uncut (Control). Commercial thiniing was don by removing merchantable timber
to achieve homogeneous reduction in stands to a basal area of less than 24 m2/ha
(tree spacing of approx. 8 m). Timber h'rst began in 1985 and was completed
by 1986. In order to protect herbaceous vegetation and minimize soil disturbance
from the impact of skidding, spacing between skid trails were at least 36.5 m if
soils were not frozen and with adequate snow cover. Understory vegetation is
highly variable and composed of many graminoid, forb, and shrub species.
Common graminoids include elk sedge (C'arexgeyeriBoott), pinegrass
(caiamagrostis rubescens Buckl.),
and Kentucky bluegrass (Poa pratensis L.).
Common forbs include western yarrow (Achillea millefolium L. var. occidental/s
DC.), strawberries (Fragaria spp.), hedstraws (Galium spp.), lupines(Lupinuc
spp.), and heart leafed arnica (Am/ca cord/folia Hook.). Common shrubs are
Thinned
Control
Graze
CExc
TExc
Graze
CExc
TExc
Figure 1. Layout of timber harvest (control and thinned) and
herbivory (Graze cattle and big game grazing; CExc cattle
exciosure, big game grazing only; TExc total exclosure, exclusion
of cattle and big game grazing) treatments for each ponderosa pine
site.
1:1
snowberry and Oregon grap
;
oi'ia tp':
ponderosa pine is the dominant ovcrtory
indley) G. Don). Although
western larch (Larix occidental/s
Nutt.) can often be found.
The following herbivory treatments Wr aoplied to each timber harvest
treatment: 1) grazing by cattle and native Ungutates to achieve 60 percent
utilization (grazed), 2) native ungulate grazing only (cattle exciosure), and 3)
exclusion of cattle and native ungulates (total exc!osure). Sixty percent utilization
is considered heavy relative to current recommendations (Holechek 1995), but
was used because it was considered a typical utilization level for industrial forests.
Grazing treatment fencing was completed in 1986, and cattle were then grazed in
a deferred rotation grazing system. Cattle grazed these sites from mid-June to
mid-July in even years and mid-July to mid-August in odd years. Grazing by cattle
was removed from 2001 through 2003.
Vegetation Sampling
Vegetation was monitored using the same procedures for all collection periods.
Three permanent transect lines, 30 m in length, were established in all treatment
combinations to monitor vegetation chanqes. These permanent transects were
used to determine overstory cover, understory species frequency, and shrub cover
and density. A spherical densiometer (Strickier 1959) was used to determine
percent overstory canopy at 0, 15 arid 30 m along each 30 m transect. Canopy
cover was determined prior to timber harvest in 1985 and again in 2003.
19
Herbaceous product;oH :i
clipping 20,
0.5-rn2
:ieasud in 1989, 1992, 1995 and 2003 by
rectanguLi t1ot' placed ahdornly within each experimental
unit. Plots were clipped to a 2 dn stjbb
hEiqht and 10 plots were grouped into
8 classes: elk sedge, pinegras, K:ntuckv h:c'ss, snowberry, other perennial
grasses, perennial forbs, annuals/biennias and shrubs. An additional 10 plots
were clipped for total production. Production clips were completely dried in a
forced air oven at 55°C, and weighted to the nearest tenth gram.
Species composition of herbaceous species spiraea (Spiraea betu/ifo/fa Palfas),
Oregon grape, and snowberry was determined by frequency counts using 30 cm x
30 cm and 30 cm x 60 cm plot sizes at three-meter intervals along each transect;
thus providing 10 plots per transect and 30 plots for each experimental unit.
Frequency was calculated as th
divided by the total number of pLt
.:iots a given species was present
aunp.ed
er treatment. Species composition
was collected prior to timber harvest in 1985 and then again in 1988, 1991, 1994,
1997, and in 2003. Changes in plant species composition from 1985 were used to
determine differences among treatments and years. Several plant community
indices were calculated: species richness, general diversity using Shannon diversity
index (Shannon and Weaver 1963), and evenness (evenness = Shannon diversity
/ In (richness); Pielou 1969). Richness ws determined by count of unique plant
taxa in the monitoring data.
Shrub cover was measured using the line-intercept method. Shrub canopy
measurements were visually rounded and the intercept recorded to the nearest 10
cm with a minimum hole size of 10 cm. Percentage of cover was calculated as the
20
length of the total intercept
v
specs a...ded by the transect length. Shrub
density, by species, was calculated using a 30 rn x 2 m plot; 1 m on each side of
the 30 m transect. Since many shrubs sprout from their base, an individual was
counted as one stem protrudinc fr11 the so
uface. Shrub canopy and density
were determined prior to timber harvestn 1985 and subsequently in 1988, 1991,
1994, and 2003. Changes in shrub cover and density from 1985 were used to
determine differences among treatments and years.
Statistical Analysis
All data were analyzed using the MIXED procedures in SAS (SAS Inst. Inc.,
Cary, NC) with the block (site replication) effect considered random. The
experimental design was a replicated splitplot design within a randomized
complete block design with repeated measures, using year as the repeated
variable. Timber harvest treatment was analyzed as the whole plot with grazing
treatment analyzed as the sub-plot. Treatment means were separated using
LSmeans procedures of SAS (SAS Inst. Inc., Cary, NC) and were considered
significant at the (P< 0.05) level. Only plant species that occurred in all
treatment combinations were included in the analysis. However, in order to
capture some possible effects on rare species the species richness, Shannon
diversity index, evenness and species turnover rates were calculated using all
unique plant taxa.
PC-ORD (Version 4, MjM Software Design, Gleneden Beach, OR) was used to
analyze the changes in plant community data. Non-metric multidimensional
21
scaling (NMS) ordination w: us::d t cor;p
differences in plant
communities, among timber harvest and hrbivoiy treatments, between 1988,
1991, 1994, 1997 and 2003 with 1935. Resuft; are presented in a series of
diagrams where distances betwn r'oint iidkate the degree of similarity. Multiresponse permutation procedures (MRPP, for determination of year effects, and
blocked MRPP, for determination of timber harvest and herbivory treatment effects
which were blocked by site, were also used to compare the differences in plant
communities between 1988, 1991, 1994, 1997 and 2003 with 1985. Non-metric
multidimensional scaling and MMRP used Sorensen (Bray-Curtis) metric as a
measure of dissimilarity. Prior to analysis, a constant of 100 was added to the
differences in species composition from 1985 in order to remove the negative
values associated with declining species occurrence since 1985. PC-ORD (Version
4) was not capable of conductinci a blocked MRPP using the Sorensen (Bray-
Curtis) metric; therefore, Euclidean (Pythdgoi.an) metric was used instead. Due
to the complex experimental diyn of th
ttidy, PC-ORD required simplification
of the data prior to analysis. As a result, year effects were analyzed first, and
similar years were grouped together and subsequently analyzed for timber harvest
and herbivory main effects. If there were timber harvest main effects, the effects
due to herbivory were then analyzed within each timber harvest treatment.
Differences, for all analyses, were deemed significant at the level of P
0.05.
22
RESULTS
Canopy Cover
In 2003, canopy cover in the thinned tretn
ts was 18% lower than it was in
controlled treatments, and controlled treatrnehts were not different than overstory
canopy in 1985. There was no effect (P> 0.10) of herbivory on overstory canopy
cover.
Understory Production
There was no timber harvest x herbivory x year interaction (P
0.37) among any
of the production classes. Total production was greater (PS 0.01) in the thinned
treatments than the controls for all years (Table 2). However, total production
was greatest (P 0.01) in 1989 and then declined by 580 kg
ha1 in 1992 and
1995. Total production increased again in 2003 but it was still less (P = 0.03)
than was measured in 1989. Total production was also affected by herbivory (Fig.
2) with grazed treatments having 100 kg
ha' less (P 0.001) production than
either the cattle or total exclosures.
Production of Kentucky bluegrass was greatest (P 0.01) in the thinned
treatments in 1989; however, production then declined by 1992 and has remained
similar (P 0.90) through 2003 (Table 2). In 1989, the thinned treatments were
also greater in production than the controls, but this was the only year that this
occurred even though the thinned treatments had numerically greater production
than control treatments.
23
Tab'e 2. The effects of ti:: rst i;i
iderstory production (kg/ha)
çun.
1mber Harvest Treatments
Thfrned
Control
SE
719b12
I 4()8'
168.4
578b1
I
168
4
Total Production
667b1
168:4
873b2
2C(
i!11
168.4
31b
1989
230
50.6
1002
1992
38
50.6
Kentucky bluegrass
1022
1995
36
50.6
1052
2003
48
50.6
70b
269a1
1989
48.2
1442
1992
79
48.2
Other perennial grasses
942
1995
72
48.2
1162
2003
121
48.2
147b
263a
Perennial forbs*
23.4
* there was no timber harvest x year effect (P> 0.05), therefore only main effects
for timber harvest are presented.
a,b
row values with differing superscripts are different at (P < 0.05).
i23
values with differing superscripts within production groups are
within
_p
rosa_p!pefor!:in
terr
different at (P< 0.05).
Production of other perenniai grasses followed a similar trend as Kentucky
bluegrass, with greatest (P < 0.01) production occurring in thinned treatments in
1989. However, in 1992 the production of these other grasses tended (P
0.06)
to be greater in the thinned treatments when compared to the control pastures.
By 1995 and continuing through 2003, there were no differences in production
between timber harvest treatments. Perennial forbs were only affected by timber
harvest with greater (P< 0001) roduct!on occurring in the thinned treatments.
Production of elk sedge was affected by both year and herbivory
treatments (Fig. 2, 3). Production was greater (P
0.01) in 2003 as compared to
1989 and 1992, with 1995 being intermediate, indicating that production of elk
24
1200
0 Grazed
T
1000
OCattle Exc.
{
L!I9a Exc.
T
800
a
.
600
400
rL
:
Total
Production
Elk sedge
Pinegrass
Figure 2. The effects of herbivory treatments on
the understory production (kg/ha) within a
ponderosa pine forest in eastern Oregon. Herbivory
treatments were: Grazed cattle and big game
grazing; Cattle Exc. cattle exciosure, big game
grazing only; Total Exc. -- total exclosure, exclusion
of cattle and big game grazing.
(columns with different letters are different at P<
0.05.)
sedge was increasing across all treatments. However, grazed treatments had less
production of elk sedge than either the cntte or total exciosures. Pinegrass was
also affected by herbivory with lower production occurring in the grazed
treatments when compared to the tota exciosures (Fig. 2).
Snowberry production was affected, but the effect varied by an herbivory
year interaction (P= 0.05). In 1989, shortiy after thinning, production was similar
(P
0.20) among all herbivory treatments (Fig. 4). However, by 2003 production
25
350
300
250
2 200
150
100
50
0
1989 1992 1995 2003
Figure 3. Changes in the production of elk
sedge over time within a ponderosa pine
forest in northeastern Oregon. (a,b,c bars
with different letters are different (P
0.05))
was greater (P 0.01) in the t)
:
c;i; when compared to both the grazed
and cattle exclosures.
Species Composition
These sites supported a relatively rich p!ant community, with a total of 149 plant
species documented during the period of study (listed in Appendix A). Of the 100
forbs identified, only 25 species occurred n every treatment combination during
the period of study. Prior to the initiation of the study, in 1985 (Table 3),
occurrence of plant species showed considerable variation and, as a result, the
effects of timber harvest and herbivory treatments were varied and documented
26
180
160
140
120
.c
c80
60
40
20
0
1989
19921995
2003
zedD&ExcTotal
Figure 4. The effects of herbivory treatments on
snowberry production (kG/ha) within a ponderosa
pine forest in eastern Oregon. Herbivory treatments
were: Grazed cattle and big game grazing; Cattle
Exc. cattle exciosure, big game grazing only; Total
Exc. total exciosure, exclusion of cattle and big
game grazing. (a,b columns with different letters,
within year, were diffrent t P < 0M5)
limited directional responses (Tabies 4, 5, 6; tables only show significant results
for affected species).
Although species ricriness was unaffected by timber harvest and herbivory
treatments, species richness was greatest (P < 0.01) in 1991 as compared to all
other years (Table 7). Species richness in the understory increased from 1985 to
1991 by 28%, from 42 to 54 speces, vthile it decreased from 1991 to 2003 by
42%, from 54 to 31 species on averag. Most of the species gained by 1991 and
subsequently lost by 2003 were forbs. Plant species diversity showed a similar
trend, with diversity being greatest (P
0.01) in 1991 and subsequently declining
27
Table 3. Frequency means fcr the 33 corn.cny occurring species in 1985, prior
_to expenment initiation, in d
in northeastern Oregon
j
t.
SD
Bromus carinatus Hook. & Ari.
Carex concinnoides Macken..
Mean
0
0
0
0
0
0
0
0
CarexgeyeriBoott
7::.0
22.0
11.3
12.0
11.5
23.6
14.2
9.7
16.7
3.3
96.7
40.0
43.3
33.3
73.3
50.0
40.0
Ca/amagrostis rubescens Buck!.
E/ymusg/aucusBuckl.
Meilca bu/bosa Geyer ex Porter & Coult,
PoapratensisL.
Trisetum canescens Buckl.
Achi/lea mi/Iefo/iuin L.
Anemone pieri Britt. ex Rydb.
Antennaria rosea Greene
Am/ca cordifolia Hook.
C/ayton/aperfo//ataDonnexWiild.
Eu/yb/a conspicua (Lindi.) Nesom
Fragariaspp.
Gallumspp
Hferac/um a/b/forum Hook.
Iris missour/ensis N Utt.
I
7
10.2
7.6
30.9
28.0
21.7
0
0
47.8
0
0
0
0
3.3
6.7
10.0
0
0
0
0
0
6.7
Max
0
0
96.7
0
0
1.3
1.6
0
0
0
0
80.0
56.7
50.0
3.3
86.7
30.0
76.7
70.0
23.3
10.0
3.3
6.7
07
2 15
0
67
10.4
12.9
0
50.0
Tha/ictrumfendleriEngelm.exGray
0.6
Trifollum repens L.
Viola adunca Sm.
V/cia americana Muhi. ex Wilid.
1.5
7.4
6.5
0
0
0
0
Mahoniarepens(Lindl.)G.Don
15.7
Spiraeabetu/ifo/iaPalIas
3.5
43.3
1.3
2.3
9.4
10.4
11.9
11.9
16.6
3.3
6.7
30.0
33.3
46.7
50.0
76.7
Lathyrusspp.
Lupinusspp.
Moehringia macrophylla (Hook' Fi
OsmorhizaL'erteroIDC.
Potent/I/a grad/is Dougl.ex Hook.
Prune//a vulgarisL.
Ranuncu/usspp.
Ste//ar/a /ongioesGoldie
Symphyotr/chum spathulatum (LindL)
Nesom var. spathu/atum
Taraxacumoffic/na/eG.H.Weberex
Wiggers
Symphoricarposa/bus(L.) Blake
30.0
19.4
11.3
0.4
42.6
7.8
16.3
20.0
5.2
0
0
35.1
0
0
16.5
18.3
11.4
1.1
22.1
Mm
8.0
1. 4
1.3
20.48
7.0
2.6
0.6
0.4
0
0
0
10.0
0
0
0
0
0
20.0
28
st and r
Table 4 The effects of timb-r
from
the difference in species frequency
1985 in a ponderosa piknnorracm Oregon.
Bromuscarinatus
Hook. &Arn.t
Timber
Harvest
Thin
Cont
Year
1994
1997
19!
jTh7b1
i$8
O.4
0
CarexgeyerIBoottt
2003
SE
2.1
2.1
o
2.6a
02
2.6
122a
..252b1
0.4
74C
0
0
2.2
107
_
Ranunculusspp.
Ste//aria Iongioes
Goldiet
Ca/amagrostis
rubescensBuckl.
Trisetum canescens
Euiybia conspicua
(Lindl.) Nesom
ragariaspp.
Prune//a vu/garisL.
Symphyotrichum
spathu/atum
(Lindl.) Nesom
Cont
Thin
Cont
1.12
____0.7
781
ga7c
1.5
0.72
0.7
3.0
0
0.9a
o.ga
1.la
oa
163b
28.oa
24
176b
193b
189b
oa
8
5b
-'a
.1
oa
06b
93b
Oa
O.9a
15ab
2.02
o
52
6ab
10
gb
14.iI1D
1.2
1.8
1.8
3.2
6.0
14
0.9c
1.3c
6.0
0.8
13 9C
17 8c
28
vajpthulatum
V/cia americana
Muhl. ex Wilid.
2.0
r bc
-u.0
ii.
-,c
Ma/ion/a repens
5b
Oa
22
U6
8
30
(LindL) G. Don
____:_____________ _____________________
* Timber harvest treatment: Thin commercial thinning; Cont control.
Species with a timber harvest x year interaction (P 0.05).
a,b,c
row values with differing superscrints are different (P 0.05).
1.2
column values with differing superscripts within species are different at (P < 0.05).
to its lowest (P< 0.001) levels by 2003. Values for the Shannon diversity index
increased from 3.13 to 3.36 for 1985 to 1991 and then declined to 2.85 by 2003.
Values for species evenness represented good equity in the distribution of plant
species across the sites. There was a decrease (P
0.05) in equity over time; this
decline may be due to the changes in species turnover.
29
Table 5. The effects of timber h vE;t ad herbvory on the change in species
fjr Un northeastern Oregon.
frequency from 1985 in a_pçnderbsa
TimtjerHares
Herbivoryt
Thin
12.4
Con.t
SE
Graze
9.6
64
12.8a
-22.1
-16.1
56
Arnicacordifo/iaHook.
-7.7
Achi//eamillefollumL.
7.8a
-1.6
83b
6.2
2.3
Symphyotrichum
spathu/atum(LindL)
Nesom var.
spathulatum
12.4a
63h
1.4
2.2
Lathyrusspp.
19.5a
22b
Potent/i/a grac//1& Dougl.
34a
04b
0.3
0
45a
24b
E/ymusg/aucusBuckl.
Thsetum canescens
Buckl.
aaytoniaperfof/ata
Donn ex Wilid.
exHook.
Viola aduncaSm.
V/cia americanaMuhl.
ex WilId.
CExc
15.2a
TExc
258b
216b
2.Oa
g2b
57b
5.7
-1.3
-5.1
2.6
2.1
11.2
10.6
6.2
2.3
1.3
33a
16b
06b
4.0
..53a
218b
...54a
4.6
1.9
-0.8
-3.9
2.0
4.2
-1.1
34a
0ab
38b
1.9
3.1
-0.2
3.1
49b
SE
2.5
1.9
* Timber harvest treatment: Thin commercial thinning; Cont control.
Herbivory treatments: Graze cattle and big game grazing; CExc cattle
exciosure, big game grazing onv; T: - tctal exclosure, exclusion of cattle and
big game grazing.
a,b
values with differing superscripts are different at (P < 0.05).
Species turnover rates (Table 8) indicated that until 2003, species lost
were similar to colonization by new spedes. But, in 2003 there was an increase in
species lost from the sites and a reduction in colonization by new species.
Shrub Density and Cover
Shrubs were not a major compoPent of these sites in 1985 (Table 9) and were
highly variable across locations. There was no year x timber harvest x herbivory
30
Table 6. The effects of timber harvest and h:rbvory on the change in understory
species frequency from 1985 in por'derosa 1e forest in northeastern Oregon
limber
Herbivoryt
C'alamagrostis rubescens BuckL
PoapratensisL
Antennaria rosea Greene
Fragariaspp.
Harve5t
Thin
Cont
Thin
Graze
Corfl.
L81
CExc
02ab1
82b2
127ab
TExc
44a1
53a2
87b1
2.7
2.7
2.2a2
10gb
122a2
54
Thin
04
Cont
1.6
Thin
-13.1
Cont____ 55a
56b1
Hieraciuma/bfflorumHoo}cTh
.73a
1.1a
T8abl
Cont
10.72
1.82
24b1
153b2
2.0
0.4
.jga
2.4a
87ab
49b1
Thn0.9a
-3.6
10.22
11b1
07b
Cont
02ab
O.9a2
13b
10.2
73b
5.8'
Cont
5.81
8.9a2
17.6a2
OsmorhizaberteroiDC.
Thn
-16.4
5.8a
-14.9
184b
-10.9
Prune//a vu/gar/sL.
Cont
Thin
Cont
0.2
0.2
Ranuncu/usspp.
1rnr
-0.4
42b1
1.3
167b1
0.22
5.6a
Iris missouriensisNutt.
Thn
Lupinusspp.
Cont
Thin
_çont
Taraxacum officinale G.H. Weber
ex Wiggers
Sycnphoricarposa/bus(L.) Blake
2.41
3.1a2
0.9
Q7a
-2.7
02b
SE
5.8a
2.02
1.1
5.2
6.2
35
0.5
0.5
4.0
4.0
13.2
13.2
0.9
0.9
0.8
0.8
5.5
5.5
6.0
6.0
-6.7
g1b
Thin
104b
4.Oa
2.9
Cont
*Tjmber harvest treatment: Thin commercial thinning; Cont control.
Herbivoiy treatments: Graze cattle and big game grazing; CExc cattJe
exclosure, big game grazing only; TExc total exciosure, exclusion of cattle and
big game grazing.
a,b
row values with differing superscripts are different at (P < 0.05).
1.2
column values with differing superscripts within species are different at (P <
0.05).
7jab
interactions with any species density or cover. The changes in wax current (Ribes
cereum Dougi.) density (Table 9) was affected by a timber harvest x herbivory
interaction (P= 0.02). Number of wax cUrrent shrubs only increased (P 0.05) in
the control treatments where all large ungulate herbivory was excluded.
31
Table 7 Plant species richness, diversity, and evenness responses to timber
harvest and herbivory in a ponderosa pine forest in northeastern Oregon.
Thinned Overstory
Control Overstory
Grazed
Cattle
Total
Grazed
Cattle
Total
Exc.
Exc.
Exc.
Exc.
38
43
54
46
41
34
42
48
56
40
42
33
40
42
52
39
44
41
43
58
42
43
30
51
32
42
42
49
44
46
30
43
55
42
43
29
2.98
3.16
3.11
3.22
3.05
3.18
3.34
3.20
3.18
2.90
3.25
3.17
3.33
3.22
3.21
2.86
3.02
3.14
3.41
3.07
3.12
2.76
3.36
3.18
3.37
3.08
3.04
2.87
334
338
3.29
3.05
2.86
3.09
3.10
2.86
0.81
0.83
0.84
0.82
0.83
0.81
0.86
0.85
0.84
0.82
0.81
0.85
No. of species
1985a
1988a
19g1b
1994
1997a
2003c
Shannon diversity
1985ac
1g88ab
1994a
1997c
2003d
Evenness
1985a
1988a
a,b,c,d
0.82
0.83
0.87
0.83
0.84
0.85
0.85
0.83
iggla
0.84
0.85
0.86
0.84
1994a
0.86
0.84
0.87
0.85
1997b
0.82
0.84
0.83
0.84
2003b
0.83
0.81
0.83
0.84
Years with differing superscripts are different (P < 0.05).
Herbivory treatments also affected (P
0.01) the changes in wild rose
(Rosa gymnocarpa Null.) density (Table 10, Fig. 5), with total exclosures changing
more than either the grazed or cattle exclosures. Rose density increased by 17
plants
ha1 in the total exclosures from 1985 to 2003, but the grazed and cattle
exclosures density of wild rose was not different (P
0.10) than it was in 1985.
Wild rose density also changed (P < 0.01) over time, with an increase (P = 0.001)
of 20 plants
ha1 by 2003.
32
Table 8. Understory species turnver rates from 1985 to 1988, 1991, 1994, 1997
and 2003 in a ponderosa pine forest in northeastern Oregon.
# of species lost since 1985
# of new species since 1985
971
1988
10.8'
14.82
9.6'
1991
SE
1,2,3
11.01
10.81
10.71
10.51
1994
1997
2003
14.1
6.8
1.3
0.9
values within columns with different superscripts are different (P < 0.05)
Table 9. Shrub species density (#/ha) and cover in 1985, prior to experiment
initiation, in a Donderosa Dine forest in northeastern Oreqon.
Ame/anchieralnifolla(Nutt.) Nutt. ex M.
Roemer
.
Crataegus doug/as/i Lindl.
Ho/odiscusdisco/or(Pursh) Maxim.
Physocarpus ma/vaceus (Greene) Kuntze
Prunus virgin/anaL.
RibescereumDougl.
Rosa gymnocarpaNutt.
Density
Cover
Density
Cover
Density
Cover
Density
Cover
Mean
1.7
SD
Mm
1.7
0.7
1.9
0.8
0.1
0.2
0.1
0.1
0.2
2.0
2.4
1.7
0.3
0.5
0.4
0.4
0.8
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Dnsity
Cover
Density
Cover
Density
Cover
0
1.9
1.1
6.5
0.6
0
4.0
2.5
7.0
1.0
Max
6.3
8.3
9.3
5.6
1.3
1.9
1.7
1.4
3.7
0.
13.0
8.8
27.3
3.3
Changes in serviceberry (Ame/anchieralnifo/la (Nutt.) Nutt. ex M. Roemer)
cover (Table 10) of the total exclosures was different (P
grazed or cattle exclosures.
0.05) than either the
Cover within total exclosures increased (P = 0.001)
while cover in both grazed and cattle exclosures has not changed (P
0.25) since
1985. Changes in wild rose cover were also detected (P= 0.01) among the
33
herbivory treatments; however, they were not different (P
0.35) than the cover
measured in 1985.
Table 10. The effects of timber harvest and herbivory on the changes from 1985
of understory shrub density (#/ha) and cover in a ponderosa pine forest in
northeastern Oregon.
Density
Timber
Harv.*
Herbivory Treatmentst
Gr
CExc
TExc
SE
Cover
Herbivory Treatments
Gr
CExc
TExc
SE
Ame/anchier
a/nifolla(Nutt.)
20b 05
06
-0.4
2.0
0.5 0.6a
Nutt. ex M.
Roemer
Rosa gymnocarpa
178b
ab
03b 0.3
6.8a
5.2a
Nutt.
01
Ribescereum
Thin
0.8
2.3
0.1
0.2
-0.1
-0.2 0.9
78b2
0.8a
Doucili'
Cont
1.0
2.3 -1.3
-1.0
-1.6 0.9
*Timber harvest treatment: Thin commercial thinning; Cont control.
Herbivory treatments: Graze cattle and big game grazing; CExc cattle
exclosure, big game grazing only; TExc total exclosure, exclusion of cattle and
big game grazing.
Species with a timber harvest x herbivory interaction (P 0.05).
a,b
row values with differing superscrpts ar dfferent at (P < 0.05).
'2column values with differing superscripts within species are different at (P<
0.05).
Plant Community Responses
Initial NMS ordination and MRPP analysis of pretreatment data, 1985, indicated
that there were differences (P < 0.001, A = 0.24) among replicate sites, therefore
all subsequent analyses were blocked by site. There was an affect (P = 0.003, A
= 0.07) of year on plant community structure, with 1991, 1994 and 1997 being
similar to each other and different from both 1988 and 2003. Analysis of 1988
plant community data revealed that there were no timber harvest effects or
34
30
25
20
15
.10
5
0
-5
Figure 5. The changes in rose (Rosa
gymnocarpa Nutt.) density over time
between 1988, 1991, 1994 and 2003 with
1985. (a,b,c bars with different letters are
different (P 0.05))
herbivory effects. However, NMS analysis of plant community data for 1991
through 1997 revealed that timber harvest may be changing plant communities.
Subsequent blocked MMRP analysis determined that there was a tendency (P =
0.08, A = 0.15) for timber harvest to affect the plant community. However, by
2003 (Fig. 6) the NMS ordination revealed that there was continued separation
among sites since 1985 and there was a continued divergence of plant
communities due to timber harvest treatments, therefore only 2003 data are
presented. Blocked MMRP analysis subsequently determined that there was a
tendency for a timber harvest (P = 0.07, A = 0.20) effect on plant community
structure.
er
'est
Ehin
ont
3
('.1
c'-J
U)
x
Axis I
Herbivory
Grazed
CExc
TExc
("J
U)
x
Figure 6. NMS Ordination of understory species composition in 2003 of (A)
site differences, (B) timber harvest effects, (C) herbivory effects in a
ponderosa pine forest in northeastern Oregon. Lines in charts demonstrate
separation of sites. (Timber harvest treatment: Thin commercial thinning;
Cont control.Herbivory treatments: Grazed cattle and big game grazing;
Cattle Exc cattle exciosure, big game grazing only; Total Exc total
exciosure, exclusion of cattle and big game grazing.)
36
DISCUSSION
The understory vegetation increased with the reduction of overstory canopy cover.
Total understory production was greatest foliowing experiment initiation and was
influenced by timber harvest and herbivory. Many researchers (McConnell and
Smith 1965, 1970; Young et al. 1967; Thompson and Gartner 1971; Long and
Turner 1975; Vora 1993) have also documented that understory production
increased with reduction in canopy cover. McConnell and Smith (1965, 1970)
indicated that understory response to thinning was greater at 8 years compared to
3 years. However, in our study, understory production was greatest 3 years
following thinning. Riegel et al. (1992) determined that increased understory
vegetation production to timber harvest was a result of reduced competition for
resources, mainly nitrogen and water. Therefore, a reason for the early peak in
understory production was likely due to expansion of roots systems from
remaining ponderosa pine trees, again causing increased competition for the
available resources. As a result, differences in understory production in thinned
and control treatments have remained steady at approximately 230 kg
ha1 from
1992 through 2003.
Understory production responses to timber harvest were not proportional
among graminoids, forbs and shrubs. Graminoid species were primarily
responsible for the increased production measured in thinned treatments.
Production of forbs moderately increased within thinned treatments, whereas
shrub production was not different in any period following timber harvest. Pase
37
(1958) and McConnell and Smith (1965, 97O a!o documented that the
responses from grasses were greater than either forbs or shrubs to reduction in
canopy cover.
Cattle and native ungulates shouid benefit from reduction in overstory canopy,
with increased production of graminoids and forbs. Cattle tend to avoid areas of
lower production (Harris 1954; Roath and Krueger 1982; Gillen et al. 1984) and
cattle diets are dominated by graminoids with minimal amounts of forbs and
shrubs (Holechek et al. 1982; Mitchell and Rodgers 1985; Walburger 2005).
Whereas, the composition of mule deer and elk diets, during the summer months,
have greater amounts of forbs and shrubs with lower amounts of graminoids
(Findholt et al. 2004; Beck and Peek 2005), but graminoid consumption by elk
varies depending on time of year
et
.
2004). Gibbs et al. (2004) reported
that elk diets were high in graminoids in May and June, 80% of diet, and then
consumption dropped dramatically in July and August, to < 50% of diet.
Herbivory can also influence the production of understory species. The effects
of cattle grazing on total production were primarily due to the reduction in the
production of elk sedge. Elk sedge has been noted as the most prominent forage
species in the diets of cattle during the summer in northeastern Oregon (Skovlin et
al. 1976). Holechek et al. (1982) determined that elk sedge in cattle diets was as
high as 23% in late summer and 24% fl fall. As documented by this study, elk
sedge may not be able to maintain production following 60% utilization on
ponderosa pine sites.
38
Herbivory by large unguli;
so affct.
production of snowberry within
thinned treatments. Krueger and Wnward (1974) reported that cattle and big
game also reduced snowberry in a Dugsñr\ponderosa pine\Kentucky bluegrass
community in northeastern Oregbn. The eduson of cattle, mule deer, and elk in
the Blue Mountains of Oregon has resulted in
2.7 times greater production of
shrub species inside exciosures than outside (Riggs et at. 2000).
Few plant species in this study demonstrated a significant directional change
to timber harvest or herbivory treatments. This could be due to several factors:
first, plant communities at the beginning of the study, in 1985, were different in all
of the replicates even though they were dassified as the same habitat type. The
continuation of the differences in plant communities by 2003 among the replicates
was still evident (Fig. 6). A closer examkation of the NMS ordination analysis
reveals that the plant communities, within each replicate, are responding
differently to timber harvest.
Therefore1
any directional plant species responses
within this study may be confounded by another replicate.
Secondly, by utilizing a timber harvest technique that minimizes soil
disturbance, such as in this study, secondary succession may not have been
initiated. As a result, residual species were able to dominate these sites. It has
been documented that predisturbance plant species (Halpern 1989; Selmants and
Knight 2003) and level of disturbance (Halpern 1989; Griffis et at. 2001) contribute
to the observed successional responses and resultant plant species following
disturbance. Furthermore, Halpern (1989) documented that understory plant
communities with minimal disturbances were dominated by predisturbance plant
species. Stohigren et al. (13G)) iO deterd that few plant species showed
consistent directional responses to grazinq or the removal of grazing.
Finally, three major soil series occur across these sites (Riegel et al. 1992).
Bennett et al. (1987) reported that soil types a well as canopy cover influenced
understory production. Further, they cornmnted that differences in soils may
have the ability to influence understory vegetation. In addition, Stohigren et al.
(1999) concluded that soil characteristics, climate, and disturbances may have a
greater effect on plant species than grazing.
We found no influences of timber harvest or herbivory on community richness,
Shannon diversity, evenness or species turnover rates. However, species richness
and diversity were greatest 6 years following experiment initiation, whereas,
evenness began to decline 11 years following experiment initiation, possibly
indicating increasing dominance of the site by a few species. Schoonnaker and
Mckee (1988) reported that diversity increased weakly and peaked at 15 years
post-harvest while species richness was greatest 20 years past-harvest in
coniferous forests of the western Cascade Mountains of Oregon. Griffis et al.
(2001) concluded that the increase n species richness foUowing a disturbance was
primarily due to exotic species in a northern Arizona ponderosa pine forest. They
also concluded that exotic species invasion increased with intensity of disturbance.
However, in this study increased species richness was not due to exotic species.
The possible reason for the lack of responses to timber harvest Was that these
ponderosa pine forests were only thinned and intensity of disturbance was
minimized. Stohlgren etal. () and Rig::td. (2000) also found no
differences in richness, diversfty, or evenness attributable to herbivoiy.
IMPLICATIONS
Timber harvest and herbivorj have the potential to change plant community
structure and composition as well as understory production. Timber harvest had
the greatest effect on understory production and plant community structure.
Consequently, by minimizing disturbances on a site the effects on the resultant
species and plant communities are minimal and will probably be determined by the
plant community structure prior to the disturbance and the physical and
microclimatic characteristics of that location.
Utilization rates approaching 60% are not recOmmended by current grazing
plans; therefore, the effects of cattle upon forested rangelands should not be as
dramatic. Deer and elk herbivory influenced the production and occurrence of
several species, but mainly shrubs.
As a result, the impacts of large herds of deer
and elk may be as influential on potential successional pathways of understory
vegetation as cattle grazing.
41
LITERATURE CITED
Augustine, D.J. and S.L. McNaughto.. 1948. Ungulate effects on the functional
species composition of p!ant communitii3s herbivore selectivity and plant
tolerance. Journal of
Management, 64:1 165-1183.
Beck, J.L. and J.M. Peek. 2005. Diet cornçoition, forage selection, and potential
for forage competition among elk, deer, and livestock on aspen-sagebrush
summer range. Range/and Ecology and Management. 58:135-147.
Beisky, LA. and D.M. Blumenthal. 1997. Effects of Livestock Grazing on Stand
Dynamics and Soils in Upland Forests of the Interior West. Conservation
Biology. 11: 315-327.
Bennett, D.L., G.D. Lemme, and P.D. Evenson. 1987. Understory herbage
production of major soils within the Black Hills of South Dakota. Journal of
Range Management. 40:166-170.
Brockway, D.G. and C.L. Lewis. 2003. Influence of Deer, Cattle Grazing and
Timber Harvest on Plant Species Diversity in a Longleaf Pine Bluestem
Ecosystem. Forest Ecology and Management. 175: 49-69.
Findholt, Si., B.K. Johnson, D. Damiran, T. DelCurto, and 1G. Kie. 2004. Diet
composition, dry matter intake and diet overlap of mule deer, elk and
cattle. In: J. Rahm (ED.), iransactions of the Sixty-ninth North American
Wildlife and Natural Resources Conference. 16-20 March 2004; Spokane,
WA: Wildlife Management Institute. p. 670-686.
Gibbs, M.C., J.A. Jenks, C.S. Deperno, B.F. Sowell, and KJ. Jenkins. 2004. Cervid
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Gillen, R. L., W.C. Krueger, and R.F. Miller. 1984. Cattle distribution on mountain
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management treatments in northern Arizona ponderosa pine forests.
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Harris, R.W. 1954. Fluctuations in forage utilization on Ponderosa pine ranges in
42
eastern Oregon. Jomai f Range l7gement. 7:250-255.
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Mixed Coniferous Forest Ranges n Ndrtheastern Oregon. Journal of Range
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and Principles (2 Edition). Prentice Hall. Englewood Cliffs, New Jersey. p.
195-196.
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Understory Vegetation. Journal of Range Management. 20: 247-249.
'!t
Kie, J.G. and J.F. Lehmkuhl. 2001. Herbivory by Wild and Domestic Ungulates in
75: 55-61
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McConnell, B.R. and 1G. Smith. 1965. Understory response three years after
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43
Pase, C.P. 1958. Herbage prcccior and ccmr.:sition under immature ponderosa
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Riggs, R.A., A.R. Tieden,ann, J.G. Cook, T.M. Ballard, PJ. Edgerton, M. Vavra,
W.C. Krueger, F.C. Hall, L.D. Bryant, L.L. Irwin, and T. DelCurto. 2000.
Modification of mixed-conifer forests by ruminant herbivores in the Blue
Mountains ecological province. Research Paper PNW-RP-527. Portland, OR:
U.S. Department of Agriculture, Forest Service, Pacific Northwest Research
Station. 77p.
Roath, L.R. and W.C. Krueger.1982. Cattle grazing and behavior on a forested
range. Journal of Range Management. 35:332-338.
Schoonmaker, P. and A. McKee. 1988. Species composition and diversity during
secondary succession in coniferous forests in the western Cascade Mountains
of Oregon. Forest Science. 34:960-979.
Selmants, P.C. and D.H. Knight. 2003. Understory plant species composition 30-50
years after clearcutting in sthesterr Wyoming coniferous forests. Forest
Ecology and Management. i&:275-2.69.
Shannon, C.E. and W. Weaver. 1963. The mathematical theory of communication.
Urbana, IL: University of Illinois Press. 117p.
Skovlin, J.M., R.W. Harris, G.S. Strickler, and G.A. Garrison. 1976. Effects of cattle
grazing methods on ponderosa pine-bunchgrass range in the Pacific
Northwest. Technical Bulletin 1531., Washington DC: U.S. Department of
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Stohlgren, T.J., L.D. Schell, and B.V. Heuvel. 1999. How grazing and soil quality
affect native and exotic plant diversity in rocky mountain grasslands. Ecological
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Canopy on Permanent Sample Plots. Research Note No. 180. Portland, OR:
USDA, Forest Service, Pacific Northwest Forest and Range Experiment Station.
Thompson, W.W. an F.R. Garther. 1971. Native forage response to clearing low
quality ponderosa pine. Journal of Range Management. 24:272-277.
Vora, R.S. 1993. Effects of timber hrves U-e.drnents on understory plants and
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Walburger, KJ. 2005. The effects of timber harvest and herbivory on understory
vegetation and composition of beef caWe diets on forested rangelands
[dissertation]. Corvallis, OR; Oregon State University. 196p.
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Journal of Forestiy. 65:807-813.
45
CHAPTER 3
The Effects of Herbivory and Timber Harvest on Understory Vegetation
in a Grand Fir (Abiesgrandis) Habitat in Northeastern Oregon.
Kent-ic].
Wa/burger,1
Timothy De/Curto,2 and Martin
Vavra3
Professor,
Authors are: 1 Graduate Research Assistant and 2
Department of Animal Sdences, Eastern Oregon Agriculture Research Center,
Oregon State University, Union, OR 97883; Supervisoty Range/and Scientist,
Pacific Northwest (PNW) Research Station, USDA Forest Service, Forestiy and
Range Sciences Laborato, La Grande, OR 97850
46
ABSTRACT
The objectives of this study were to document the effects of logging and
herbivory, cattle and native ungukt, on understory vegetation within a grand fir
(Abies grandis) habitat. Three replicated sites were established in 1986. Grand fir
sites were arranged as a split-plot design; timber harvest treatments [1) no
harvest (CON), 2) thinning (TH), 3) clearcut (CL)] were whole plots and herbivory
treatments [1) cattle and native ungulate grazing (GR), 2) native ungulate grazing
(BG), and 3) exclusion of cattle and native ungulate grazing (EX)] were the subplots.
Understory production was affected by timber harvest and herbivory.
Total production in CL was 350 kg
275 kg
ha' greater (P < 0.05) than TN, and TH was
ha1 greater (P< 0.05) than CON.
Total production in GR was 102 kg
ha1 greater than EX. A total of 141 spece were identified on these sites,
however, only 29 of these species occurred in all treatment combinations and
were used for analysis. Timber harvest treatment had a greater magnitude of
effect than did herbivory on understory plant species. Differences were observed
among timber harvest and herbivory treatments on individual shrub species
density and cover. The NMS ordination and MMRP analysis revealed that plant
communities were different in 1985 prior to study initiation and these differences
were maintained through 2003. Plant communities were affected by timber
harvest and herbivory treatments. There appeared to be differences among all
timber harvest treatments, whereas, among herbivory treatments, EX appeared to
be different than GR and 8G. This research indicates that the effects of timber
47
harvest may have a greater effec. on unde; so!y vegetation composition,
structure, and diversity than herbivory.
KEYWORDS: Beef cattle, Native unculates, Mule deer, Elk, Mixed conifer
INTRODUCTION
Forests of the interior Northwest on average, receive greater precipitation, have
deeper soil profiles, and have qreiter sol! moisture holding capacity than adjacent
grasslands (Riggs et al. 2004). These forests provide valuable habitat for many
wildlife species and summer grazing allotments for beef cattle. However, over the
last 100 years, many of these forests have developed into relative homogeneous
stands in composition and structure of fire-sensitive and disease-susceptible
species (Beisky and Blumenthal 1997; Hessburg et al. 2005), thereby reducing the
overall understory productivity of the area. Many areas with high potential for
timber harvest and forage production have the lowest output due to dense canopy
cover (Hedrick et al. 1969).
In order to enhance forage production and create a diversity of habitats it
may be necessary to open the canopy. iiriiber harvest on forested rangelands
sets back succession and, in most cases, increases understory forage production.
Forage production response to overstory canopy cover is well documented
(McConnell and Smith 1965 and 1970; Jameson 1967; Young et al. 1967;
Thompson and Gartner 1971) and documents that as overstory canopy cover
decreases understory production (ka/ha) increases.
Large herbivores are attracted to areas of higher forage production and
palatable food resources, and as a result of timber harvest, it can be expected that
they will focus their foraging within these areas (Vavra et at. 2004). Hobbs
(1996), Augustine and McNaughtcin (1998), Riggs et al. (2000), and Kie and
Lehmkuhl (2001) all documeited that hebir cn influence plant community
structure, composition and productictn. Ungulate herbivor' directly affects
vegetation through selective feeding and the zbity of a plant to recover from
herbivory (Augustine and McNauhon 1998). HQwever, plant species diversity is
not consistently affected by grazing (Stohigren et al. 1999, Riggs et al. 2000, and
Brockway and Lewis 2003), but individual species, at the local level, are affected
by herbivory and timber harvest. Herbivory has the greatest effect in altering
plant community structure within clearcuts (Riggs et al. 2000). Stohlgren et al.
(1999) also documented grazing effects on local species, however few plant
species showed consistent directional responses to grazing.
Little is known about how plant community structure responds when
combining the effects of herbivory and reduction of overstory canopy within
a
grand fir forest. Therefore the objective of this study was to document the effects
of timber harvest and herbivory n uriderstory vegetation composition, structure,
diversity and production.
MATERIALS AND METHODS
The study area is located at the Eastern Oregon Agricultural Research
Center's Hall Ranch that is approximately 16 km east of the city of Union in the
Wallowa Mountains of northeastern Oregon. Elevation ranges from 1050 to 1250
m. Summers tend to be dry and warm with temperatures rarely exceeding 38°C,
though freezing or near-freezing temperatures are possible all year. Winters are
50
cold and wet, with the majoty cf the preci ....
Ii.:
(66 cm average) coming in the
form of snow between November and May (Table 1). Elk (cervuse/ap/iusL.) and
mule deer (Odocof/eus hemkwi us Rat.) are idicnous to the area and can be
found throughout the year; howe'it:r, heVie5t. u:e occurs in spring and fall.
Table 1. Precipitation data (cm), for vegetation sampling years, from weather
Center's Hall Ranch.
station located at Eastern
Precipitation (cm)
Year
Jan Feb Mar Apr
May Jun Jul Aug Sep Oct Nov Dec Total
1985
1991
1994
1995
1997
2003
Ava.
1.3
4.6
6.1
9.7
5.8
6.1
5.9
4.3
3.0
7.6
3.0
5.3
6.4
4.9
3.8
6.6
2.3
9.1
1.5
9.9
5.5
4.1
6.1
7.4
9.7
10.4
7.4
6.1
5.8
13.0
7.4
6.9
4.3
7.1
6.6
Three replicated sites, cf an
3.3
7.9
3.3
7.6
3.6
1.0
4.7
1.0
0.5
1.0
2.0
5.3
0.0
1.6
2.5
0.3
0.0
4.1
1.3
3.6
1.8
.hiec qrardLs
5.3
0.0
0.8
1.3
3.3
2.5
2.3
5.6
3.8
3.6
4.1
3.8
2.0
3.6
8.9
11.4
10.4
12.4
4.1
3.6
6.6
2.5
2.0
3.6
9.1
7.6
8.9
5.9
48.5
59.2
53.3
79.0
56.4
58.4
55.6
(Dougl. ex D. Don) Lindl. /
Pach/stima myrsinites (Pursh) Raf. (grand flr) habitat type, were selected to
analyze the effects of herbivory and logging on understory plant communities.
Areas of relatively homogeneous stand structure were initially selected and
experimental sites were subsequently chosen from these areas. The grand fir
sites had three timber harvest treatments applied to them (Figure 1): 1) clear cut,
2) crown thinning (thinned) and 3) uncut (Control). Crown thinning consisted of
removing co-dominant and some dominant trees. Timber harvest began in 1985
and was completed by 1986. To protect herbaceous vegetation and minimize soil
disturbance from the impact of skidding, spacing between skid trails was at least
36.5 m if soils were not frozen and with adequate snow cover. The grand fir
51
Clearcut
Control
/
rr3'
I
.:--.
-4
1-'
_c
-
v'-
-
- -
iii ..*iI
___J-IUEA
a.ui
-r
J\Uj
Graze
CExc
t{;t
TExc
r
Graze
CEc
ag,i
TExc
Figure 1 Layout of the timber harvest (control, thnned and clearcut) and
herbivory (Graze cathe and big game grazing; CExc cattle exciosure, big
game grazing only; TExc total exclosure, exclusion of cattle and big game
grazing) treatments for each grand fir site.
clearcuts were replanted in the spring of 1988 with Ponderosa pine (Pinus
ponderosa P. & C. Lawson), Douglas4Ir (Pseudotsuga menziesll (Mirbel) Franco)
and western larch (Larix occidental/s NulL). Understory vegetation was highly
variable and composed of many graminoid, forb, and shrub species. Common
graminoids included elk sedge (carexgeyeiBoott, pinegrass (ca/amagrostis
rubescens Buckl.), Kentucky bluegrass (Poa pratensis L.) and blue wildrye (Elymus
glaucus Buckl.). Common forbs indude western yarrow (Achi/lea mi/lefol/um L.
var. occidental/s DC.), strawberries (Fragaria spp.), bedstraws (Gal/urn spp.),
lupines (Lupfnusspp.), and heart leafed arnica (Am/ca cordifolla Hook.). Common
shrubs were snowberry (Symphoricarpos a/bus (L.) Blake), Oregon grape
(Ma/ion/a repens (Lindley) G. Don), and ninebark (Physocarpus ma! va ceus
(Greene) Kuntze). Although grand fir was th dominant overstory species, in
thinned and control treatments, ic.-as tr, western larch and Ponderosa pine
were still found.
52
The following grazin t
a:nnts w:
plied to each timber harvest
treatment: 1) grazing by cattle nd native ungUates (to achieve 60 percent
utilization), 2) native ungulate grazirq ory atd
3)
exclusion of cattle and native
ungulates. Grand fir grazing exclosure fehdrg ws completed in 1986, but
perimeter fencing was not completed until 1994; therefore pastures were grazed
in common by allotment cattle, from September to November. Following
completion of perimeter fencing in 1994, pastures were grazed in a deferred
rotation system with pastures grazed from mid-July to mid-August in odd years
and mid-August to the beginning of October in even years.
Grazing by cattle was
removed during 2001, but in 2002 and 2003 pastures were grazed from mid-
August to mid-October by allotment cattle.
Vegetation Sampling
Vegetation was monitored using the same procedures for all collection
periods. Three permanent trans&ct ilnes, 30 m in length, were established in alt
treatment combinations to monitor vegctation changes. These permanent
transects were used to determine overstury cover, understory species frequency,
shrub cover and shrub density. A spherical densiometer (Strickler 1959) was used
to determine percent overstory canopy at 0, 15 and 30 m along each 30 m
transect. Canopy cover was determined prior to timer harvest and again in 2003.
Species composition of each herbaceous species nd low 9rowing shrubs
(spiraea
(Spiraea betuilfo/la Pallas),
Oregon grape, and snowberry) was
determined by frequency counts using 30 cm x 30 cm and 30 cm x 60 cm plot
53
sizes at three-meter interv:
acI t:h thus providing 10 plots per
transect and 30 plots for each eperrnenai Unft. Frequency was calculated as the
number of plots containing a givh spedes dMded by the total number of plots
sampled per treatment. Spedes compoftion was collected prior to timber harvest
in 1985 and then again in 19g4, 1991, arid
in
2003. Changes in plant species
composition from 1985 were used to determine differences among treatments and
years. Several plant community indices were calculated: species richness, general
diversity using Shannon diversity index (Shannon and Weaver 1963), and
evenness (evenness = Shannon diversity
I
In (richness); Pielou 1969). Richness
was determined by count of unique plant taxa in the monitoring data.
Shrub cover was measured using the line-intercept method. Shrub canopy
measurements were visually rounded and the intercept recorded to the nearest 10
cm with a minimum hole size of 10 cm. Percent cover was cacu!ated as the
length of the total intercept by each species divided by the transect length. Shrub
density, by species, was calculated using a 30 m x 2 m plot; 1 m on each side of
the 30 m transect. Since many shrubs sprout from their base, an individual was
counted as one stem protruding from the soil surface. Shrub canopy and density
were determined prior to timber harvest in 1985, 1991, 1994 and 2003. Changes
in shrub canopy cover and density from 1985 (Table 2) were used to determine
differences among treatments and years.
Herbaceous production was measured in 2003 by dipping 0.5 m x LO m
rectangular plots placed randomly within each experimental unit.
Plots
were
clipped to a 2 cm stubble height and grouped into 7 classes: &k sedge, pinegrass,
54
Kentucky bluegrass, other
pdrl L;rase'. ;rnnial forbs, annuals/biennials
and shrubs. Production clips were cmplctely dred, in a forced air oven at 55
degrees Celsius, and weighted to the nearest tenth gram. Production was also
measured in 1995, but only data for tota! prbduction was available.
Statistical Analysis
All data were analyzed using the MIXED procedures in SAS (SAS Inst. Inc.,
Cary, NC) with the block (site replication) effect considered random. The
experimental design was a split-plot within a randomized complete block design
with repeated measures, using year as the repeated variable, and three
replications. Logging treatment was treated as the whole plot with grazing
treatment analyzed as the sub-plot. Treatment means were separated using
LSmeans procedures of SAS (SAS Inst. Inc, Cary, NC) arid were considered
significant at the (P< 0.05) level. Only plant species that occurred in all
treatment combinations were included in the analysis. However, in order to
capture some possible effects on rare spedes the species richness, Shannon
diversity index, evenness and species
turnover
rates were calculated using all
unique plant taxa.
PC-ORD (Version 4, MjM Software Design, Gleneden Beach, OR) was used to
analyze all of the plant community data. Non-metric multidimensional scaling
(NMS) ordination was used to compare the differences in plant communities,
among logging and grazing treatments, between 1988, 1991, 1994, 1997 and
2003 with 1985. Results are presented in a series of diagrams where distances
55
between points indicate the degr
cf sirniki
procedures (MRPP), for determination of year
Multi-response permutation
'
fUects, and blocked MRPP, for
determination of timber harvest and herhivor treatment effects which were
blocked by site, were also used t; compare th differences in plant communities
between 1988, 1991, 1994, 1Y97 and 2003 With 1985. Non-metric
multidimensional scaling and MMRP used Sorensen (Bray-Curtis) metric as a
measure of dissimilarity. Prior to analysis, a constant of 100 was added to the
differences in species composition from 1985 in order to remove the negative
values associated with declining species occurrence since 1985. PC-ORD was not
capable of conducting a blocked MRPP using the Sorensen (Bray-Curtis) metric;
therefore, Euclidean (Pythagorean) metric was used instead. Due to the complex
experimental design of this study, PC-ORD required simplification of the data prior
to analysis. As a result, year effects were analyzed first, and similar years were
grouped together and subsequently analyzed for timber harvest main effects. If
there were timber harvest main effects, the effects due to herbivory were
analyzed within each timber harvest treatment. The data was simplified in this
manner because of the original split-plot design. Differences, for all analyses,
were deemed significant at the level of P 0.05.
56
RESULTS
Canopy Cover
Timber harvest in 1986 affected (P <
0.001) the
overstory canopy in 2003 (Fig. 2).
Overstory canopy cover was greatest in th controls, intermediate in the thinned
and least in the clearcuts, 81, 50, and 30% respectively. The overstory canopy in
the clearcuts was comprised predominantly of ponderosa pine, but grand fir
seedlings were present throughout the treatment.
100
90
80
70
60
50
40
30
20
10
0
Clearcut
Thinned
01985
Control
2003
Figure 2. Overstory canopy cover (%) of
grand fir sites in 1985, prior to timber
harvest, and subsequently in 2003. Columns
are means + SE bars. (a,b,c bars, within
year, with different letters were different at
P< 0.05)
57
Species Composition
Species composition in 1985 was extremely variable (Tab'e 2 and 3), but, there
were differences (P
0.05) in th species c:omposition among sites, timber
harvest and herbivory treatments. Therefore. thanges in species composition
from 1985 were analyzed for 194, 1997 ard 2003.
Table 2. Species frequency means in 1985, prior to experiment initiation, in a
arnd fir forest in northeastern Oreaon.
BromuscarfnatusHook.&Arn.
BromusinermisLeyss.
Carexconcinnoides Mackenzie
CarexgeyeriBoott
Ca/arnagrostis rubescens Buckl.
E/,vmusglaucusBuckl.
Festuca occidental/sHook.
Luzula campestris(L.) DC.
Me//ca bu/bosa Geyer ex Porter & Coult.
PoapratensisL.
TrisetumcanescensBuckl.
Achillea mi/lefolium L.
Anemone pioeri Britt. ex Rydb.
Am/ca cord/folla Hook.
Euiybia conspfcua (Lindi.) Nesom
Fragariaspp.
Gallumspp
Hierac/um albifforum Hook.
Lathyrusspp.
Lupinusspp.
Moehringiamacrophylla(Hook.) Fenzl
Senecio integerrimus N utt.
Stellar/a /ongies Goldie
TaraxacumofficfnaleG.H.WeberexWiggers
Thal/ctrumfend/er/Engelm.exGray
Viola aduncaSm.
Mahoniarepens(LindL)G. Don
Spiraea betuilfolia Pallas
Symphoricarposalbus(L.) Blake
Mean
0.3
0
6.5
53.9
15.2
3.9
23.5
3.9
24.4
7.6
28.0
8.3
22.0
41.9
5.2
48.8
7.9
10.1
44.4
4.5
44.5
SD
Mm
1.3
0
0
0
0
6.9
21.4
9.0
5.6
12.0
3.8
13.1
12.7
13.0
7.6
16.4
16.2
11.6
14.8
15.5
5.9
11.3
7.0
6.7
2.2
0
6.7
0
0
0
6.7
0
0
16.7
0
26.7
0
0
20.0
0
0
17.1
0
10.0
0
0.1
4.1
26.6
14.6
3.4
29.2
13.9
0.6
4.4
19.0
7.0
3.5
9.9
10.5
0
0
0
3.3
0
10.0
0
Max
6.7
0
26.7
86.7
41.1
16.7
53.3
13.3
53.3
44.4
50.0
26.7
66.7
72.2
50.0
80.0
63.3
23.3
70.0
26.7
80.0
0
3.3
15.6
76.7
30
13.3
53.3
36.7
58
Table 3. The average shrub species density (#/ha) and cover (%) in 1985, prior
to exDeriment initiation, in a arand fir forest in northeastern Orecon.
Mean
Acer g/a/.irum Torr. var. glabrum
Ame/anch/era/n/fo//a(Nutt.)Nutt.exM.
Roemer
Crataegusdoug/as//Lindl.
Ho/odiscus disco/or(Pursh) Maxim.
PhlladelphuslewtsllPursh
P/iysocarpus n-ia/vaceus (Greene) Kuntze
Prunus virginiana L.
Ribes cereum Dougi.
Rosa gymnocarpaNutt.
Density
Cover
Density
Cover
Density
Cover
Density
Cover
Density
Cover
Density
Cover
Density
Cover
Density
Cover
Density
Cover
0.02
SD
Mm
0
0.09
0.02
0
0
1.1
1.1
0
0.2
0.2
0.04
0.3
0.5
0.2
2.8
3.0
0.06
0.08
27.4
8.6
0.14
0.03
0.08
0.01
14.2
0.9
0
0
0
0
0
0
0
2.3
0.6
0
0
0
0
0.3
0
3.5
3.4
0.01
0.02
35.5
10.5
0.04
0.01
0.02
0
13.1
1.1
Max
0.44
0.08
3.7
1.5
2.3
0.8
10.7
10.9
0.33
0.43
121.3
30.8
0.67
0.14
0.33
0.08
66.0
3.7
These sites supported a relatively rich plant community, with a total of 141
species (for complete species listing, see Appendix A); however, only 29 species
occurred in alt treatment combinations across all years. Of the 29 species, the
occurrence of 24 species was affected by time, timber harvest and/or herbivory
treatments (Tables 4, 5, 6, 7; tables only show significant results for affected
species). Timber harvest had the greater effect on the magnitude of change in
species composition than did herbivory. In general, the disturbance caused by
timber harvest increased the occurrence of graminoid species. However, forb
species were variable in their response to timber harvest. Western yarrow
increased with timber harvest and lupines increased in timber harvested and
59
Table 4. The effects of timbr harst and h;bivor treatments on the changes in
understory species composit!on ia g-and fir forest in northeastern Oregon.
Timber Hardest Teatmrj
Herbivory Treatments
ClearThuiContrS
Bromuscarinatus
Hook. & Am.
Bromusinerm/sLeyss.
Carexgeyer/Boott
E/ymusglaucusBuckl.
Luzula campestris (L.)
DC.
Ach///eami/efol/umL.
Arnicacord/fo//aHook.
Hierac/um a/b/forum
Hook.
Moehr/ngia
niacrophylla
(Hook.) Fenzl
Tha/ictrum fend/er!
Engelm. ex Gray
Sc'iraeabetu//fo//a
13.Oa
79a
-13.5
32.la
-3.3
6,gb
0b
3,5
2.5
1.4a
429b
3.0
15.7
Graze
CExc
TExc
SE
2.6
8.9
7.5
6.5
2.6
2.9
6.0
3.6
45b
5.1
14.1a
01b
24.9
22.7
29.2
2.9
5,8
3.6
2.4
2.1a
21b
41a
1.7
7.3
15.5
18.8
-11.0
12.7
-12.7
131b
3.7
6.6
5.1
2.0
2.3
-3.1
36.8a
21.4a
374a
68ab
46b
121b
-8.2
-3.7
-4.3
2.5
3,4a
15.4
-9.6
101b
-33.8
-20.6
-16.8
10.2
24.6a
448b
31.8a
9.9
26.5a
73b
12b
6.2
-11.2
-10.2
-13.5
4.9
19b
3,0b
4.1
-9.0
-9.9
-5.6
3.5
13.8
2.3
Pallas
Symphor/carpos a/bus
153b
57C
23.2a
4.1
10.9
12.8
(L.) Blake
a,b
row values with differing superscripts are different at (P< 0.05).
2.8
herbivory treatments, however, peas (Lathyrusspp.) and heartleaf arnica were
negatively affected by timber harvest. Shrubs were also affected by timber
harvest treatments, with snowberry increasing in clearcuts and thinned
treatments, whereas spiraea (Spiraea betuilfolla) decreased in clearcuts and was
not affected in thinned or controls.
Species richness and Shannon diversity were affected (P 0.05) by timber
harvest and herbivory (Table 8). However, since there were no interactions with
years, it was concluded that changes seen in species richness and Shannon
diversity index were evident prior to study initiation and may not be attributed to
Table 5. Changes in species ccmposition, sn::e 1985, in a grand fir forest in
northeastern Oregon.
1997
1994
gb
carexgeyeriBoott
2003
SE
12.9c
5.8
175b
280b
ElymusglaucusBuckl.
45.2'
147b
12.0
Anemone pioeri Britt. ex Rydb.
-I6.9
87b
46b
Am/ca cordifolia Hook.
-i8.9
57b
17.9a
Fragariaspp.
109b
19.3a
23.48
Lathyrusspp.
242b
1o.ga
9.la
Symphoricarposa/bus(L.) Blake
row values with differing superscripts are different at (P< 0.05).
10.9
6.9
8.0
.g
Table 6. Changes in species composition, from 1985, as affected by time and
timber harvest treatments in a grand fir forest in northeastern Oregon.
Year
Timber
1997
Harvest
1994
2003
SE
4a
gbl
40
1a1
Cl arcut
43
-17
11 5
Moehringiamacrophy//a
56b2
41.9a
25.6a1
11:5
Thinned
rtooK.) r-enzi
Stellar/a /ongioesGoldie
Control
Clearcut
Thinned
Control
Crut
47.8a
41b2
15b2
13.3
15.5
g3ab
11.4
104b
1.la
0.7
55a
4.la
73a
11.5
7.1
7.1
7.1
8.2
8.2
8.2
226b
336b1
1g8b1
0.6a
2.3a
Thinned
8.82
-4.7
-11.0
Control
aM
row values with differing superscripts are different at (P < 0.05).
1.2.3
column values with differing superscripts within species are different at (P <
0.05).
Viola adunca Sm.
application of timber harve;t or herbivory treatments. Values for species evenness
represented good equity in the distribution of plant species across treatments.
Species turnover rates, comparing 2003 to 1985 (Table 9), indicated that number
of new species entering these site was only affected by timber harvest (P
0.02)
whereas the number of species disappearing from these sites was affected by a
timber harvest by herbivory interaction (P = 0.006). Thinned harvest treatments
61
Table 7. Changes in specic.3 or' .oition,
timber harvest and herbivoi trtments n
L85 to 2003, as affected by
crand fir forest in northeastern
Oreaon.
Herhivory
______________________________Tratmen
C'alamagrostis rubescens
Bucki.
Timber Harvest Treatments
Cearcut Thinned
Control
SE
:
12.22
25.62
9.2
8.91
7.1
133a12
7.1
TExc
Grdze
-89
14.41
CExcI
l4.8a
81b2
___________________________TExc
119ab
3.0a2
252b2
31.51
16.11
9.9
9.5
-23.0
9.5
-15.7
9.5
Festuca occIdental/s
Hook.
ThsetumcanescensBuckl.
Graze
CExc
TExc
Graze
Lathyrusspp.
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Lupinusspp.
Taraxacumofficfna/eG.H.
Weber ex \iggers
r
Mahonlarepens(Lindl.)
ci
Don
7.2
i0.42
1.02
19.812
..444a1
7412
27.4a1
37b
23.7a2
07b2
g7ab
35.2a1
1g3abl
15b
7.1
6.8
6.8
6.8
6.0
26ab2
41b1
70b2
1.22
1.92
0.4
6.0
5.6a
2.6a
33b1
-2.6
-1.5
-0.7
-3.6
-5.2
107ab1
20.4a1
13.7a1
442
6.0
2.0
2.0
2.0
G
-7.8
Txc
1.2a
85b1
2.6
3.3
*
Herbivory treatments: Grazed cathe and big game grazing; CExc cattle.
exdosure, big game grazing only; TExc total exciosure, exclusion of cattle and
big game grazing.
a,b
row values with differing superscripts are different at (P< 0.05).
12.3
column values with differing superscripts within species are different at (P<
0.05).
had a greater
(P =0.008)
number of new spedes entering the sites than controls,
with clearcuts being intermediate. However, the number of species disappearing
from these sites was greatest (P < 0.05) in the total exctosures of clearcuts.
Other than total exclosures from clearcuts, it appears that the number of new
species were similar to the number of species lost from these sites.
Table 8. Plant species richn.:.
r:rty, :hd
62
ess responses to timber harvest
and herbivory in a grandfirD-cstIr
Species Richness
1985
1994
1997
2003
Average
Average
I
CE 'c
TExc
G-
49
46
44
44
46
41
61
4
31
45
40
39
41
43
41
39
46
'43
3.20
3.28
3.13
3.09
3.18
3.20
3.24
3.14
3.12
3.18
a Overstory
CExc
TExc
Control Overstory
Gr
CExc
TExc
41
43
40
41
49
40
45
43
40
38
42
46
40
32
36
39
36
29
29
33
32
31
31
29
30
30
3.09
3.24
3.25
3.22
3.20
2.98
3.21
3.23
3.30
3.18
3.19
3.29
3.25
3.13
3.22
3.04
3.15
2.96
3.04
3.05
2.99
2.81
2.80
2.89
2.87
3.09
2.91
2.96
2.92
2.97
ab
Shannon Diversity
1985
1994
1997
2003
ClearctflOe 3iL_
Gr*
3
ab
3.16
3.24
3.14
3.21
3.19
Evenness
0.87
0.86
0.80
0.84
0.79
0.83
0.90
0.78
0.86 0.86
0.87
0.87
0.86
0.85
0.83
0.85
0.85
084 0.89 0.87 0.88 0.85 0.83 0.88
0.84 0.85
0.85
0.86
0.86
0.86
0.85
0.83
Averaae
0.85
0.85
0.87
0.86
0.84
0.87
0.83
0.83
* Herbivory treatments: Gr cattle and big game grazing; CExc cattle exciosure, big
game grazing only; TExc total exclosure, exclusion of cattle and big game grazing.
1985
1994
1997
2003
b
0.81
0.85
0.83
0.85
0.84
Herbivory effect (P< 0.05)
Timber Harvest effect (P< 0.05)
Shrub Density and Cover
Shrub cover and density was minimal for most species in 1985 (Table 3), except
for ninebark which had a density ci- 35.5 plants
ha4 and 10.5% cover in 1985.
Changes in serviceberry cover and density were affected by year, timber harvest,
or herbivory treatments. Cover increased l.4% by 2003 in thinned treatments
whereas clearcuts and controls had no cover increases when compared to 1985.
Changes in density were observed, however, these changes were on the order of
1 plant
ha'.
Table 9. The effects of tim
h'vst rJ
ory on understory species
turnover rates from 1985 to 2003 ngridPrFcrestin northeastern Oregon.
Thr1rd
Clearcut
# of new species since 1985*
Control
SE
76b
g4ab
63
1.7
Clearcut
Thnnd
Control
SE
# of species lost since 1985t
14.71
Grazed
12.7'
2.1
9312
672
Cattle Exclosure
11.7'
2.1
87b2
23.3a2
12.31
Total Exciosure
2.1
* Nonsignificant timber harvest x herbivory effect (P= 0.82). Significant timber
harvest treatment effect (P = 0.02).
Significant timber harvest x herbivory effect (P = 0.006).
a,b,c
values within rows with different superscripts are different (P < 0.05).
1,2
values within columns with different superscripts are different (P< 0.05).
Wild rose density was affected by year and herbivory treatments. Wild
rose density was 16 plants
ha
greater (P = 0.03) in 1994 when compared
to 1985 and has remained the same through 2003. The density of wi'd rose has
increased (P = 0.03) in the total éxciosures by 16 plants
ha
but only tended to
increase (P= 0.06) in the cattle exclosures by 12 plants
ha
since 1985.
However, the density of rose in the grazed pastures has not changed (P= 0.26)
since 1985.
Understory Production
There were no interactions (P> 008) between timber harvest and herbivory for
production measures, except for production of Kentucky biuegrass, therefore only
main effects were reported. Total understory production increased (P= 0.02)
from 760 kg
ha
in 1995 to 1108 kg
ha' in 2003. Total understory production
64
br
was also affected (P< O.O: v
atments (Table 10).
Understory production in clearcUts was 3:0 g
ha1 greater (P = 0.02) than
thinned treatments and thinred fteatments were 275 kg . ha' greater (P= 0.04)
than controls. Total undertorv podiction was also affected (P = 0.04) by
herbivory treatments. Grazed traments aeaged 102 kg
ha1 greater (P=
0.01) understory production than the total exclosures. Cattle exclosures were
intermediate but not different
(P 0.13) than the grazed or total exclosures.
Timber harvest had a greater impact on the understory production than did
herbivory. Only Kentucky bluegrass was affected by herbivory (Table 10), and it
exhibited a timber harvest x herbivory nteraction (P= 0.01). Kentucky bluegrass
production was greatest (P < 0.01) in the cattle and total exclosures of clearcuts
Table 10. The effects of timber harvest and herhivory on understory production in a
grand fir forest in northeastern O'i-c.r
T1rnbehvestTreatrnents
Her bivory Treatments*
Clear
Thin
Cont
312
172
211
213ab
307a
127b
142a
ig7a
53b
221
61
242
170
13
205
176
3
js9ac5634c
Total Production
CarexgeyeriBoott
cafamagrostis
rubescens Buckl.
Other perennial
grasses
Perennial forbs
Annuals / biennials
Shrubs
Poa pratensis L.
Grazed
1551
55
224
Gr
CExc
752
çj1 940ab
38.3
219
216
78.7
293
29,7
57.9
22.7
39.3
SE
TExc
SE
8ö'
52.7
261
33.8
167
181
78.7
142
166
85
29.7
195
25
213
218
23
225
154
30
234
50.0
17.6
34.3
63.6
63.6
_____TExc
52'
63.6
* Herbivory treatments: Gr cattle ar,d big came grazing; CExc - cattle exclosure, big
game grazing only; TExc total exclosure, exclusion of cattle and big game grazing.
CEXC
280a2
2g4a2
56b
6
27b
Timber harvest x herbivory interaction (P = 0.01).
row values with differing superscripts are different at (P < 0.05).
1.2.3
column values with differing superscripts within species are different at (P< 0.05).
a,b
65
compared to all other treatrhr
.inatc'h:i;. F)k sedge production tended (P=
0.09) to be greater in the dearcuts comprcd tc the thinned and control
treatments. Production of pinr's was 60% greater in the thinned treatments
compared to the controls, with ie-cuts ntemdiate and not different than either
thinned or control treatment!. Production o
reçrass within grazed treatments
tended (P = 0.08) to be greater than either cattle or total exciosures. Other
perennial grasses also responded to reduction in canopy cover with greater (P <
0.05) production in the clearcuts and thinned treatments when compared to
controls.
Plant Community Responses
Initial NMS and MRPP analysis of pretreatment data, 1985, indicated that there
were differences (P< 0.001, A = 016) among replicate sites, therefore all
subsequent analyses were blocked by site. Upon further investigation, all years
exhibited similar trends; therefore only 2003 data were reported (Fig. 3). The
NMS ordination analysis revealed an apparent difference among sites, with one of
the sites being different than the other two (Fig. 3A). Timber haest (Fig. 3B)
appeared to affect the changes in plant community composition and subsequent
MMRP analysis determined that pl&nt communities changes were different (P<
0.01, A = 0.09).
Changes in plant communities were also affected by herbivory treatments
(P= 0.01, A = 0.19). However, there were no herbivory effects (P 0.33, A
0.01) when it was analyzed within timoer harvest treatment. Closer examination
66
Timber
Harvest
tE
A
1
2
3
Clear
Thin
Cont
U)
Axis 1
Axis 1
A
A
Herbivory
A
Graze
Cattle Exc
Iota! Exc
c..J
C')
x
4:
Axis 1
Figure 3. NMS Ordination of the changes in understory species composition
from 1985 to 2003 of (A) site dfferertces, (B) timber harvest effects, and (C)
herbivory effects in a grand fir forest in northeastern Oregon. Line in charts
demonstrates separation of the differences among sites. (Timber harvest
treatment: Clear clearcut , Thin --crown thinning; Cont - control; Herbivory
treatments: Grazed cattle and big game grazing; Cattle Exc cattle exclosure,
big game grazing only; Total Exc total exdosure, exclusion of cattle and big
game grazing.)
67
of the changes in plant comrnunft
tribL:t:
u hrbivory (Fig. 3C) revealed that
only within Site 1 was there an observabe difference. Changes observed in cattle
exclosures and grazed treatments wre more sftn Var than total exclosures.
Whereas in Sites 2 and 3, it appeared that there were no differences in changes of
plant community composition attnbutablè to herbivory.
DISCUSSION
Understory vegetation composition was affected by timber harvest and herbivory.
Species richness, diversity and evenness were affected by either timber harvest or
herbivory. But, these differences occurred prior to initiation of the study;
therefore, differences observed in subsequent years were probabiy carry over
effects from this time. Species turnover was also affected by timber harvest and
herbivory, but the number of new species was similar to the number of species
lost, except for total exclosures in clearcuts which had greater number of species
lost that new species. The greater number of lost species from total exclosures in
clearcuts was probably the result of this treatment having considerably greater
number of species present than the other clearcut herbivory treatments in 1985
(Table 8). However, by 2003, the differences in species richness in clearcut
herbivory treatments were no longer present.
Many other researchers (Schoonmaker and McKee 1988; Halpern 1989;
Thomas et al. 1999; Selmants et al. 2003) have documented that species richness
and diversity increase following disturbance. Schoonmaker and Mckee (1988)
LI
reported that species richness
-ed t bk at 15 to 20 years after logging
and burning in a Douglas fir fces ir weseri Oregon. They also documented that
diversity is greatest up to 20 years frllov:n dkWrbance.
The lack of changes in spcte:s richness ar.1 diversity observed in our study
could be related to disturbance intensity and nftial plant community composition.
Griffis et al. (2001) documented that low intensity disturbances did not increase
diversity or species richness.
Timber harvest is a significant disturbance for
overstory succession, but if disturbance to soil and vegetation was minimized,
such as in this study, successional trajectories may ultimately be determined by
initial species and site history (Hapern 1989; Selmants et al. 2003). Tilman and
Downing (1994) suggested that communities with high species richness were
more resilient because they probably contain disturbance resistant species.
Halpern (1989) commented further that early succession was characterized by a
shift in abundance of generally persstent species, rather than replacement
species. Riggs et al. (2000) obsere that richness and diversity appeared to be
more dynamic in clearcuts than in selective-cut of uncut areas and varied over
locations.
We observed no differences in plant species richness, diversity, or
evenness that was due to herbivory. Stohlgren et al. (1999) and Riggs et al.
(2000) also found no differences in richness, diversity or eveness as a result of
herbivory. However, we observed that 11
of
the 29 specs, that occurred in aH
treatment combinations over the duration of the study, were affected by
herbivory. Plant community responses (Figure 3) also indicated that herbivory had
minimal effects on plant cornrnunfty structure
dS
compared to timber harvest.
These observed herbivory effects over time were not as great in magnitude when
compared to the timber harvest tretitments but still indicate that ungulates have
the ability to modify plant spedes composition. However, Riggs et al. (2000)
commented that herbivory effects were often equal or even greater in magnitude
than episodic agents, especially in ciearcuts. Irwin et al. (1994) also commented
that herbivory from large ungulates caused moderate to severe reductions in
shrubs and that elk/deer can have important effects in the absence of livestock.
They also noted observed differences in plant communities as a result of native
ungulate herbivory. Many other researchers (Krueger and Winward 1974; Laycock
and Conrad 1981; Milchunas and Lauenroth 1993; Hobbs 1996; Augustine and
McNaughton 1998; Weisberg and Bugmann 2003) have also reported that
ungulate herbivory has the ability to change plant species composition.
Understory production was 98 and 30% greater in the clearcuts and
thinned treatments, respectively, than in the control treatment. Many other
researchers (Pase 1958; McConnell and Smith 1965, 1970; Young et al. 1967;
Thompson and Gartner 1971; Long and Turner 1975; Pyke and Zamora 1982)
have documented similar results. These observed changes in understory
production are primarily due to differences in shading (Naumburg et al. 2001) and
changes in the competition between overstory and understory for available
resources (Riegel et al. 1995).
In this study, increased production of understoiy vegetation was the result
of changes in the production of graminoids; consequently forbs, shrubs and
70
annual/biennials were unaffected by cimber
rvst treatments. Pase (1958), and
McConnell and Smith (1965, 1970) reportad that graminoid production increased
at a greater rate than either forbs or shrubs. However, Thompson and Gartner
(1971) reported that all vegetation types nraed in forage production with
neither groups taking a particular advantage When trees were removed. On
multiple sites across northeastern Oregon, Riggs et al. (2000) reported that
production of forbs was greater than graminoids irrespective of canopy cover.
Timber harvest appears to favor increased livestock and native ungulate
production because of increased understory production. Cattle are attracted to
areas of greater production (Harris 1954; Roath and Krueger 1982; Gilien et al.
1984) and tend to select a diet that is high in grasses (Holechek et al. 1982;
Mitchell and Rodgers 1985, Walburger 2005). Whereas, deer and elk diets are
composed of greater amounts of forbs and shrub species than cattle diets
(Findholt et al. 2004; Beck and Peek 2005), but graminoid consumption by elk
varies depending on time of year (Gibbs et al. 2004). Gibbs et al. (2004) reported
that elk diets were high in graminoids in May and June, 80% of diet, and then
consumption dropped dramatically in July and August, to < 50% of diet.
By only documenting the observed effects on graminoids, forbs, and
shrubs we omitted a significant amount o above ground production. Long and
Turner (1975) reported that 22 years post-harvest mosses comprised <1% of
understory production, but by 30 years post-harvest mosses comprised 11% of
understory production. They also determined that total community production
was greatest 70 years post-harvest. Therefore, maintaining a mosaic of overstory
71
age classes may allow for increased numbe- c habitats from which herbivores and
other wildlife species may occupy and allow for multiple uses across the
landscape.
Herbivory affected tOt8i uñderstor, poduction and Kentucky bluegrass
production. Production was greatest in the grazed treatments compared to the
total exclosures. It appeared that grazing, with1n this study, stimulated greater
forage production. Other researchers in other environments have documented
similar results (McNaughton 1979; Turner et al. 1993; Frank and McNaughton
1993; Fahnestock and Delting 1999; Frank et al. 2002; Loeser et al. 2004). The
reason for the increased production of understory vegetation in the grazed
pastures was probably due to timing of grazing. Pastures were not grazed until
plants were senesced and life cycles were potentially completed in the late
summer. Following grazing, production of the young plant material in the fall and
following spring would have higher photosynthetic rates than the older senesced
plant material (Painter and Detling 1981; Wallace 1990). Grazing would increase
light penetration through the foliage canopy (Monsi et al. 1973) thereby increasing
the amount of light reaching the younger plant material, and subsequently,
resulting in increased plant production. However, Holechek et al. (1995)
concluded that 30% utilization rates by cattle on forested rangelands were
recommended for sustainable grazing. Consequently, extreme care must be taken
when exceeding this recommended utilization rate.
72
IMPLICATIONS
Timber harvest and herbivory arc capabic of influencing plant community structure
and understory production. Afthough,
timber
harvest had the greatest effect on
understory production and spedes composition, herbivory had modifying effects.
If soil disturbances are minimized, understory species occupying the site prior to
harvest will largely determine resultant plant community structure. Grasses and
grass-likes appear to have a greater advantage under these circumstances.
However, herbivory had modifying effects within timber harvest treatments
on species composition. Production of understory vegetation can be maintained,
and possibly increased, with appropriate timing of herbivory upon forested
rangelands. Utilization rates approaching 6O% is not recommended because of
distribution problems with cattle on diverse forested rangelands. Plant community
responses to herbivory and timber were variable and demonstrate the importance
of recognizing that site effects, e.g. dspect, soil depth, soil type, initial plant
community, may have the ability to modify successional trajectories.
73
LITERATURE CITED
Augustine, D.J. and S.L. McNaughtori. 1998 Ungulate effects on the functional
species composition of piht cornut1ities: herbivore selectivity and plant
tolerance. Journal of
Maraqerhtnt 64:1165-1183.
Beck, J.L. and J.M. Peek. 2005. Diet L-ompostioi1, forage selection, and potential
for forage competition among elk, deer, and livestock on aspen-sagebrush
summer range. Range/and Eco/oqy and Management 58:135-147.
Belsky, J.A. and D.M. Blumenthal. 1997. Effects of Livestock Grazing on Stand
Dynamics and Soils in Upland Forests of the Interior West. Conseivation
Biology. 11: 315-327.
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Journal of Forestty. 65:807-8 13.
CHAPTER 4
Effects of Timber Harvest and Previous Herbivory on Subsequent Diets
of Cattle Grazing Forested Rangelands.
Kenric J. Wa/burger,1 77mothy De/Curto,2 and Martin Vavr
Authors are: 1 Graduate Research Assistant, 2 Associate Professor, Department of
Animal Sciences, Eastern Oregon Agriculture Research Center, Oregon State
University, Union, OR 97883; 3 Supervisoty Range/and Scientist, Pacific Northwest
(PNW) Research Station, USDA Forest Service, Forestiy and Range Sciences
Laboratory, La Grande, OR 97850
79
ABSTRACT
The objectives of this study were to document the effects of timber harvest and
herbivory on nutritional quafty ard botarc! cmposition of steer diets in grand fir
(Abies grandis
and ponderos pine (Pin!/sp rderosa forests. Three grand fir
and ponderosa pine sites were established in 1986. Grand fir sites were arranged
as a split-plot design and timber harvest treatments: [1) no harvest (CON), 2)
thinning (TH), 3) clearcut (CL)] were whole plots and herbivory treatments [1)
cattle and big game grazing (GR), 2) big game grazing (BG), and 3) exclusion of
cattle and big game grazing (EX)] were the sub-plots. Ponderosa pine sites were
arranged as a split-plot design arid timber harvest [1) CON, 2) TH] were whole
plots and herbivory treatments [1) GR, 2) BG, and 3) EX] were sub-plots. Diet
samples were collected using four ruminaUy cannulated steers in June and August
of 2001 and 2002. Within each pasture, steels were allowed to graze for 20 mm.
Results from the grand fir habitat revealed that nutritional quality was better in
June than August. The CP and TVOMD were greater (P< 0.05) in June than in
August and ADF and NDF had lower values (P < 0.05) in June compared to
August. The ADF content in the diet was greater (P < 0.05) in the CONT as
compared to CC and TH. Previous herbivory did not (P> 0.10) influence diet
quality. Microhistological analysis of ruminal masticate was used to determine the
botanical composition of diets. Graminoids were the major constituent in the diet
ranging from 65 to 91%, forbs were intermediate ranging from 8 to 31%, and
shrubs were least ranging from 0.2 to 3.5%. Season of use did not affect (P>
80
0.10) the composition of diets. Results fr
t
ponderosa pine sites revealed that
CP, IVOMD, ADF, and NDF of the diets were ary affected by season of use. June
diets were of higher (P < 0.05)
quality
than in August.
Graminoids were the
major constituent in the diet ranqin from 83 t 8%, forbs were intermediate
ranging from 10 to 14%, and shrubs were ft;t ranging from 2 to 3%. Again,
season of use did not affect (P> 0.10) the composition of diets. This study
suggests that timing of grazing had a greater influence on diet quality than did
previous herbivory and(or) timber harvest, however, previous herbivory and(or)
timber harvest had a greater influence on composition of diets than did timing of
grazing.
Key Words: Ab/es grand/s. botanical composition, microhistological analysis, Pinus
poaderosa
81
INTRODUCTION
Grazing cattle and timber harvest re common practices associated with forested
rangelands in North America These areas corr:prlse a significant portion of the
public lands in the west and are productive n producing habitat and forage for
livestock and wildlife, as well as wood products for human use. However, over the
past 100 years many areas with the potential for high forage production have low
outputs due to dense canopy cover (Hedrick et at 1969). Therefore, it may be
necessary to open the canopy to return the understory productivity of these lands.
Timber harvest on forested rangelands sets back succession and, in most
cases, increases understory forage production (McConnell and Smith 1965 and
1970; Jameson 1967; Young et al. 1967; Thompson and Gartner 1971). This
results in an increased opportunity for cattle/wildlife to forage and obtain a higher
quality diet and subsequently iriccease procuc:ivity. Typically, cattle select a diet
that is predominantly grass with limited forbs and shrubs (Holechek et at. 1982;
Mitchell and Rodgers 1985). However, cattle diets vary throughout the grazing
season, with woody vegetation becoming a greater part of the diet as the grazing
season progress (Holechek et al. 1982; Mitchell and Rodgers 1985; Darambazar
2003).
Few studies have evaluated diet quality on forested rangelands over the
grazing season, whereas more is known about changing body condition and
weight change over this same period. Holechek et at. (1981, 1987), Vavra (1984),
and Walburger (2000) have aU documented
1t cittle gain less in the late
summer and fall when compared to ate sprq and early summer.
The combined effects of timber harvest ahd previous herbivory (wild and/or
domestic ungulates) on diet quafltv have rct ber documented. Therefore, the
objectives of this study were to determine hOw timber harvest, previous herbivory
and season of use affect the quality of diets obtained from forested rangelands.
MATERIALS AND METHODS
The study area is located at the Eastern Oregon Agriculture Research Center's Hall
Ranch, which is approximately 16 km east of the city of Union in the Wallowa
Mountains of northeastern Oregon. Elevation ranges from 1050 to 1250 rn and
annual precipitation averages 56 cm with about 65% coming in the winter;
whereas summers are usually dry (Figure 1). Cattle have been grazing the area
since mid-1880. Elk (Cèrvus elaphus
L.)
and mule deer (Odoco//eus hem/onus
Raf.) are indigenous to the area and can be found throughout the year; however,
heaviest use occurs in spring and fail.
The study was conducted as a replicated split-plot design. Three
Abies grand/s
(Dougl. ex D. Don) Lindl. / Pachi:ctima mvrsintes (Pursh) Raf. (grand fir), 22.5 ha
each in size, and three P/nus ponderosa P.& C. Lawson I Symphor/carpos a/bus
(L.) Blake (Ponderosa pine), 15 ha each in size, sites were selected to analyze the
effects of herbivory and overstory canopy cover on botanical composition of diets
and diet quality. Sites were selected within areas of relatively homogeneous stand
I,
E
v7
I!
Jan
Feb
Mar
Apr
May
Jun
Ju.l
Aug
Sep
Oct
Nov
Dec
2001 [32002 4Average (1985-2003)
Figure 1. Precipitation data (cm) from :weather station located at Eastern
Oregon Agriculture Research Center's Hall Ranch.
structure. The grand fir sites had three timber harvest treatments applied: 1)
clear cut, 2) crown thinning and 3) uncut (Control; Figure 2). Crown thinning
consisted of removing co-dominant and some dominant trees. Timber harvest
began in 1985 and was completed in 1986. The grand fir clearcuts were replanted
in the spring of 1988 with Ponderos pine, Douglas-fir (Pseudotsuga menziesi/
(Mirbel) Franco var. glauca (Beissn.) Franco). and western larch (Larix occidental/s
Nutt.). Whereas, the ponderosa pine sites had two timber harvest treatments
applied: 1) commercial thinning and 2) uncut (Control; Figure 3). Thinning within
the Ponderosa pine sites was done to achieve a tree basal area of 24 m2/ha (tree
spacing of approx. 8 m). Timber harvest began in 1985 and was completed in
Control
Thirned
Clearcut
ii;;
__.ni.iii
UTh?'
IxsCirpjx,
.v
Cirp'SIx.
Ix
gr1ftI ,u irrpi
Igwa
SI.
I
I
Figure 2. Layout of the timber harvest (control, thinned and clearcut) and
herbivory (Graze cattle and big game grazing; CExc cattle exciosure, big
game grazing only; TExc total exciosure, exclusion of cattle and big game
grazing) treatments for each grand fir site.
Control
Graze
CExc
Thinned
TExc
iiiiiiiii
Figure 3. Layout of timber harvest (control and thinned) and
herbivory (Graze cattle and big game grazing; CExc cattle
exclosure, big game grazing only; TExc total exciosure, exclusion
of cattle and big game grazing) treatments for each ponderosa pine
site.
..
UU
The following herbivory treatrnei
were ppd within all timber harvest
treatments for both grand flr and ponderosà pine sites: 1) grazing by cattle and
big game (to achieve 60 percent utilization), 2) big game grazing only (Cattle
exclosure), and 3) exclusion of cattle and big game grazing (Total Exclosure).
Sixty percent utilization is considered heavy relative to current recommendations
(Holechek 1995), but was used because it was considered a typical utilization level
for industrial forests. Cattle and total exclosures were approximately 0.5 ha in
size, Grazing by cattle was done in conjunction with allotment grazing from mid-
August through October for the grand fir sites. Whereas, ponderosa pine sites
were grazed in a deferred rotation grazing system. Even years were grazed from
mid-June to mid-July and odd years were grazed from beginning of July to midAugust. Grazing by cattle in ponderosa pine sites was removed from 2001 and
2002 to allow for diet collections in mid-August.
Vegetation on these sites was valed but the dominant grasses were elk sedge
(Carexge,ver/Boott), pinegrass (Ca/amagrostis rubescens Bucki.), and Kentucky
bluegrass (Pea pratensis L.). Numerous forbs were also found which inciude
heartleaf arnica (Am/ca cord/folio Hook.), western yarrow (Achi/lea milefollum L.
var. ocddenta/is DC.), cinquefoil speJes (Potentila spp.), and lupine species
(Lupinusspp.). Several shrub species were typically found which include mallow
ninebark (Physocarpus malvaceus (Greene) Kuntze), common snowberry, Oregon
grape
(Berberis
repens Lindl.), and spirea (Spiraca betulifolia Pallas). Overstory of
the grand fir sites within the controls and thinned timber harvest treatments were
dominated by grand fir, whereas, dominant overstory species within clearcuts was
86
ponderosa pine, Douglas-fir,
nd western
ar
however, grand fir saplings were
numerous. Overstory of the ponderosa pir!e sites was dominated by ponderosa
pine and interspersed with western larch.
Four ruminally cannulated steers were used to determine diets in June and August
of 2001 and 2002. Prior to th grazing but1 steers were transported to site and
ruminally evacuated as described by Lesperance et al. (1960), except rumen walls
were rinsed with a sponge to remove as much material as possible. Steers were
allowed to graze for 20 mm. and grazed masticate samples were removed
immediately following the grazing bout. Multiple collections were made by each
steer within a day, both morning and evening collections. Launchbaugh et al.
(1990) reported no differences in cattle diets between morning and evening
collections, therefore only considerations for possible effects from an empty rumen
were considered. To minimize possible effects of an empty rumen on forage
selectivity by steers, we randomized the order that sites were grazed within each
biock.
Following collection of masticate samples, original rumen contents were
replaced. Masticate samples were completely dried at 55° C in a forced air oven
and were ground through a Wiley Mill (Thomas Scientific, Sweedesboro, NJ) using
a 1 mm screen. Composite samples were created for each experimental unit by
combining 50 g sub-sample of each steers masticate sample. Samples were then
analyzed for CP (AOAC, 1990), ADF, NDF (Ankom20° fiber analyzer, ANKOM
Technology Corporation, Fairport, NY), and IVOMD (Daisy", ANKOM Technology
Corporation, Fairport, NY). Livestock were handled according to the protocol
approved by the Institutionai Anna! Care
r'd
U
Committee at Oregon State
University.
Botanical composition of steer diets was determined using microhistological
analysis. Composite samples were soaked in sodium hydroxide and mounted
using techniques described by Holechek (1982). Three slides for each sample
collected from grand fir sites and four slides for each sample collected from
ponderosa pine sites were prepared and then dried at 55°C in a forced air oven,
for a minimum of 48 hours, prior to analysis. Twenty fields per slide were
systematically observed at lOOx magnification.
Plant fragments were identified
by comparing epidermal characteristics with plant species reference slides and
recorded as frequency counts. Dry weight composition of each sample was
determined by dividing the frequency of each species by the total number of
frequencies for all species (Holechek and Gross 1982).
Herbaceous production was collected in 2CO3 by clipping 0.5 m x 1.0 m
rectangular plots placed randomly within each experimental unit.
Plots
were
clipped by species to a 2 cm stubble height. Production clips were completely
dried in a forced air oven at 55 degrees Celsius and weighted to the nearest tenth
gram. Using the production data, a relative preference index (RPI) was calculated
to determine the relationship between botanical composition and forage
availability. This index was used to account for the differences (data not
provided) in understory forage production ameng the herbivory and timber harvest
treatments. Relative preference index was calculated as: % diet composition / %
forage composition (Krueger 1972).
'I.
All data were analyzed as a split-plot de5i1 within a randomized complete
block design with three replications usinç MIXED procedures in SAS (SAS Inst.
Inc., Cary, NC) with the block (site replication) effect considered random. The
whole-plot experimental unit Was tniber harvest treatment and the sub-plot
experimental unit was herbivory within timber harvest treatments. Treatment
means were separated using LSmeans procedures of SAS (SAS Inst. Inc., Cary,
NC) and were considered significant at P < 0.05.
RESULTS
Grand Fir Sites
There were no interactions (P> 0.40) among season of use, timber harvest and
herbivory treatments for any measures of diet quality determined; therefore, only
treatment means are reported. Crude protein, IVOMD, NDF and ADE were all
affected by season of use (Table 1). Crude protein and IVOMD of steer diets were
4.5 and 6.1 percentage points, respectively, higher (P< 0.001) in June than in
August. Only minor differences (P
0.02) in NDF and ADF content occurred
between June and August (2.4% and 1.4%; respectively).
Even though August
diets were of lower quality, the nutritional requirements for a lactating cow were
met (NRC 1996).
Timber harvest and herbivory treatments had little effect on the quality of
steer diets; except for ADF. The clearcuts and thinned treatments had lower (P
0.03) ADF values than control treatments. This increase of ADF in the control
89
Table 1. The effect of seash of use and tihr harvest on the subsequent
aualitv of steer diets in a arand fir forested habitat:.
CP, % OM
Season of Use
June
SE
Auaust
)3b 0.3
14.ga
7i4b
78.5a
0.5
580b
55.6a
0.7
3g7a
411b
s
Timber Harvest Treatment
Clearcut Thinned Control
12.6
12.5
75.4
55.8
76.6
IVOMD (%)
NDF, % OM
57.3
38.4a
40.Oa
ADF, % OM
a,b
values with differing superscripts are different at (P< 0.05)
12.6
74.5
57.4
42gb
SE
0.3
0.9
0.6
0.8
treatments could be due to greater amount of forbs consumed by steers and/or
the effects of increased canopy cover on structural characteristics of consumed
forages.
Botanical composition of steer diets did not exhibit season of use x timber
harvest x herbivory treatment interactions (P 0.70). However, graminoids and
forbs exhibited a timber harvest x herbivory treatment interaction (P= 0.04).
Graminoids were also affected by season of use, with June diets having greater
amount of graminoids than August diets (83.53'o and 80.4%, respectively).
Amount of shrubs consumed was not affected (P 0.11) by either season of use,
timber harvest or herbivory treatments,
Consumption of graminoids was least (P
0.02) in controls, across all
herbivory treatments, compared to clearcuts and thinned treatments (Table 2). In
addition, total exciosures within the controls, had the lowest amount of graminoids
in steer diets compared to grazed and cattle exciosures. Conversely, consumption
of forbs was greatest (P
0.04) in controls, across all herbivory treatments,
compared to clearcuts and thinned treatments.
90
Table 2. The effects of timber harvest and herbivory treatments on the botanical
composition (%) of steer diets in a grand fir forested habitat.
Herbivory
Timber Harvest Treatments
SEM
Treatmentst
Thinned
Control
Clearcut
827ab12
76.1a2
Grazed
88.3'
88.0a1
758a2
Graminoids
3.17
91.5'
Cattle Exc
794b2
654b3
9031
Total Exc
Forbs
Shrubs
Grazed
Cattle Exc
io.'
156ab1
8.3'
10.7a1
Total Exc
93'
Grazed
Cattle Exc
Total Exc
0.9
0.2
0.4
22.5a2
21.1a2
311b3
1.7
1.3
1.3
3.1
3.5
0.9
3.10
0.96
Herbivory treatments: Grazed cattle and big game grazing; Cattle Exc cattle
exciosure, big game grazing only; Total Exc total cxci osure, exclusion of cattle
and big game grazing
a,b
column values with differing superscripts within forage type are different at (P <
0.05)
1,2,3
row values with differing superscripts are different at (P < 0.05)
Also, RPI did not exhibit a season of use x timber harvest x herbivory
treatment interaction (P 0.75). Flc.wcver, t
within steer diets was affected (P
RPI of graminoids and forbs
0.04) by a timber harvest x herbivory
treatment interaction. Steers showed a strong preference for graminoids in all
timber harvest and herbivory treatments (Table 3). However, preference did vary
within timber harvest treatments. Within clearcuts, preference for graminoids was
greatest (P 0.04) in cattle exciosures. Within controls, graminoids were
preferred and not different among all grazing treatments. In clearcuts, steers
selected forbs in proportion to availability, but in the thinned treatment, steers
preference for forbs was greatest (P
0.01) in the total exclosure compared to
either the grazed or cattle exclosures. Steers also showed a preference for forbs
in the cattle and total exclosures of control (no timber harvest) treatments (1.53
91
rhivory treatments on the relative
Table 3. The effects of timhr harvest an
preference index (RPI)* of grarninoids, forbs, and shrubs for steer diets in a grand
fir_forested_habitat.
Herbivory
Treatments
Graminoids
Forbs
Grazed
Cattle Exc
Timber Harvest Treatments
-__CleatcUt
Thinned
1.39a
Total Exc
1.57c
1.32a
Grazed
Cattle Exc
0.90
0.55
Total Exc
0.701
1.14a
0.84a
194b2
Control
1.50
1.42
1.51
SEN
0.12
1.08a
153b
0.34
2.00c2
0.09
0.05
0.04
0.08
0.11
0.05
0.10
* RPI = % diet composition / % forage composition (Krueger 1972)
Herbivory treatments: Grazed cattle and big game grazing; Cattle Exc cattle
exciosure, big game grazing only; Total Exc total exclosure, exclusion of cattle
and big game grazing
a,b
column values with differing superscripts within forage type are different at (P <
0.05)
Shrubs
1,2,3
Grazed
Cattle Exc
Total Exc
0.03
0.02
0.03
row values with differing superscripts are different at (P< 0.05)
and 2.00, respectively). Shrubs, on th othr hand, were not preferred (P
0.23)
in the timber harvest or herbivory treatments.
Total understory production and graminoid production in 2003, 18 years post-
harvest, was only affected (P 0.05) by timber harvest treatments (Table 4).
Total understory production was greater in the clearcuts compared to both thinned
and controls, 353 and 591 kg
was 540 kg
ha1 greater respectively. Graminoid production
ha' greater (P= 0.02) in clearcuts compared to controls. Thinned
treatments tended (P = 0.09) to be greater than controls but were not different (P
= 0.22) from clearcuts. Production of forbs and shrubs were not affected (P
0.61) by either timber harvest or herbivory.
92
Table 4. The effects of timb:r hrvcst on urderstory production (kg . ha4), 18
years post-harvest, n a gra'iiit Iorestira northeastern Oregon
SE
Thnned
Control
Cleircut
Total Production
Graminoids
Forbs
Shrubs
a,b
17b
947a
736b
832b
407b
246
242
17S
178
205
224
1423
118
64
39
values with differing superscripts are different at. (P < 0.05)
Ponderosa Pine Sites
There were no interactions (P> 0.11) among season of use, timber harvest and
herbivory treatments for any measures of diet quality determined; therefore, only
treatment means are reported. Neither timber harvest nor herbivory treatments
influenced (P> 0.10) diet quality. Crude protein, IVOMD, NDF and ADF were only
affected by season of use (Table 5), with the ftgher quality diets occurring in June
compared to August. Crude protein and IVOMD were 3.9 and 5.7 percentage
points, respectively, higher (P <0.001) n June than in August. In contrast, ADF
and NDF content were 3.6 and 5.7 percentage points, respectively, lower (P
001) in June than in August.
Table 5. The effects of season of use on the diet quality of steers grazing a
onderosa oine forested habitat.
Season of Use
June
Crude protein
IVOMD
13.7a
80.2a
NDF
ADF
35.2
a,b
535a
August
98b
745b
592b
388b
values with differing superscripts are different at (P<0.05)
SE
0.28
045
0.65
0.50
93
There were no interacthns (P
0.21) aohg season of use, timber harvest
and herbivory treatments on the hotanic
ccmpoition of diets; therefore, only
treatment means are reported. Neither sesbn of use nor timber harvest
treatments affected (P
0.28) the botaric
composition of steer diets, but the
amount of graminoids and forbs ir the steer diets was affected (P
0.04) by
herbivory treatments. Greater than 80% of the steer diets (Table 6) was
graminoids, however, the diets from cattle exclosures contained 5.2% and 4.3%
more (P
0.04) graminoids than the grazed and total exclosures, respectively.
Composition of forbs in the diets was greater (P = 0.01) in the grazed pasture
than in the cattle exclosure, and the total exclosure tended to be greater (P
0.08) than the cattle exclosure.
There were no interactions (P 0.11) among season of use, timber harvest
and herbivory treatments on the RPI for graminoids, forbs and shrubs of steer
Table 6. The effects of herbivory on the botanical composition (%) and relative
preference index (RPI)* of steeL diets within a ponderosa pine forest typ
Herbivor/ Treatmentst
Total Exc
SE
Cattle Exc
Grazed
Botanical Composition
884b
84.la
83.2a
2.75
Graminoids
ggb
14.3a
2.64
Forbs
2.9
0.40
1.7
Shrubs
2.5
RPI
Graminoids
Forbs
Shrubs
1.28a
0.54a
0.38
1.28a
0.51a
027ab
150b
381b
015b
0.15
0.15
0.10
* RPI = % diet composition / % forage composition (Krueger 1972)
Grazing treatments: Grazed cattle and big game grazing; Cattle Exc cattle
exclosure, big game grazing only; Total Exc total exclosure, exclusion of cattle
and big game grazing
a,b
values with differing superscripts are different at (P<0.05)
94
diets; therefore, only treatn!r: 15 ae
rti.d. Neither season of use nor
timber harvest treatments afte:td (P 0.13) the RPI for grarninoids, forbs or
shrubs in steer diets, but the RPI for grarninod and forbs in steer diets were
affected (P
0.02) by herbivory
.r
tments. Steers preferred a diet that was
dominated by graminoids (Table 6); however, preference was greater (P< 0.001)
in the total exclosure than in either the grazed or cattle exciosures. Overall, steer
diets were proportionally lower in forbs or shrubs than available in the pastures.
However, steer diets in the total exclosures had greater (P
0.03) RPI for forbs
than either the grazed or cattle exclosures. In contrast, cattle grazing the total
exclosures had lower (P
0.02) RPI for shrubs than the grazed treatments and
tended to have lower RPI (P
0.07) than the cattle exclosures.
Total understory production tended (P = 0.08) to be greater in thinned
treatments compared to controls (Table 7). Aiso, production of forbs was 144 kg.
ha1 greater (P = 0.001) in thinned treatment3 compared to controls. Herbivory
treatments only influenced the productton of shrubs. Shrub production of total
exclosures was greater than grazed and cattle exclosures, 128 and 105 kg
respectively. Graminoid production ws not affected (P
ha
0.18) by either timber
harvest or herbivory treatments.
DISCUSSION
Quality of diets collected from grand fir and ponderosa pine sites declined in late-
June to mid-August irrespective of timber harvest and herbivory treatments. Even
95
on U dtry production (kg . ha1), 18
Table 7. The effects of tim
years post-harvest, in a pode'-un pne tirst in 'ortheastern Oregon.
Herbivory Treatments*
Timber Harvst
Treattnent
Total Exc
SE
Contr
craze Cattle Exc
Thin
SE
Total
Production
Graminoids
Forbs
Shrubs
1111
873
237
974
1016
988
237
634
5
131h
115
0
652
187
535
120
206
216
52
85
92a
115a
220b
84
275a
160
125
* Grazing treatments: Grazed cattle and big game grazing; Cattle Exc cattle
exciosure, big game grazing only; Total. Exc total exclosure, exclusion of cattle
and big game grazing
a,b
values with differing superscripts are different at (P<0.05)
though nutritional quality of these diets was declining, the quality of diets obtained
in August was of sufficient quality to meet the requirements for lactating cattle
during this period (NRC 1996). Declining quality of cattle diets is a result of
declining forage quality and(or) declining quantity of desirable forages. Cook arid
Harris (1968), $kovlin (1967), and Ciark (2U3) have reported that plant quality
declines with increasing plant phenology and as the grazing season progresses,
with grass quality declining the greatest. However, shrubs and forbs typically
remain higher in quality and were able to maintain that quality throughout the
summer.
Walburger et al. (2000) reported that aspect, north vs. south, with its
accompanying differences in
overstory1
soils and moisture availability can influence
the quality of plants throughout gra2lng season, with north aspects having higher
quality later in the grazing season than south aspects. Svejcar and Vavra (1985)
speculated that decreasing canopy cover increases the amount of sunlight
96
reaching the soil surface, therb1
creasnc. t'.' zoil temperature and reducing the
available soil moisture later in the summer; as a result, accelerating plant
phenology and reducing forage quality. Howver, McEwen and Dietz (1965),
Dealy (1966), and Severson nd Uresk (19S) were unable to detect differences in
CP due to changes in overstory canopy cpvr in ponderosa pine forests. As well,
Regelin et al. (1974) found no differences in CP content in various understory
species within a mixed-conifer forest. Our results indicate that the change in
overstory canopy cover did not affect the quality of diets because there were no
interactions with season of use and timber harvest treatment, nor were there any
interactions with season of use and timber harvest treatment in the botanical
composition of diets. Therefore, cattle in this study were selecting compositionally
similar diets in June and August and quality of diets was not changing due to
timber harvest treatment in August. We speculate that decreased canopy cover
may slow plant phenology of consumed fora: species thereby allowing them to
maintain higher forage qualities later in the year.
Within this study, steer diets, for grand fir and ponderosa pine sites, were
dominated by grasses, 65-90% of the diet, forb composition was intermediate, 8-
31% of the diets, and shrub consumption was minimal, 3.5% of diets. The
changes in composition within these diets, especially for grand fir sites, were likely
due to changes in understory production. The ability of steers to obtain a diet
high in grasses, a preferred constituent, in contro treatments of grand fir sites
would be more difficult than in clearcuts because graminoid production was 591
kg
ha1 less compared to thinned and clearcut treatments. Therefore,
97
composition of diets, to a gra
idfL:x:i he steer's ability to obtain
desired forage components
Unexpectedly, season
or
use Was not an friflUencing factor on botanical
composition. Beck and Peek (2005)j in northeastern Nevada, also reported that
cattle diets are dominated by grässe and forbs with minimal inputs from shrubs.
However, Mitchell and Rodgers (1985), in a Douglas-fir/ninebark habitat type in
northern Idaho, reported cattle had similar composition of diets from mid-June
through August, but the diets were dominated by grasses and browse. Other
researchers (Holechek 1982; Uresk and Painter 1985) have reported changes in
diet composition during this time period of mid-June through August. Hoechek et
aT. (1982), in eastern Oregon, reported that cattle diets were composed primarily
of grass but it varied among seasons with consumption of browse increasing with
progression of the grazing season, They also reported that consumption of forbs
declined through the grazing period and were minor constituents in the diet.
Uresk and Painter (1985), in the Black Hilts, reported that consumption of grasses
and shrubs were similar throughout the grazing period and constituted
approximately 54% and 28% of the dt, respectively, whereas consumption of
forbs declined throughout the grazing period and only averaged 17% over the
grazing season. The probable reason for the differences in diets was that the
other studies (Holechek 1982; Mitchell nd Rodgers 19S5; Uresk and Painter
1985) allowed grazing to continue throughout the duration of the seasonal
collection periods which, in turn, may reflect changes in forage availability. In fact,
Launchbaugh et al. (1990) reported that cattle diets differed depending on when
98
during the grazing period they
Ere ampet1. Whereas this study only allowed
grazing after collections were completed, therefore, these diets represent what
cattle could select if forage availability was not limited.
Even though the composithn of diets was changing, cattle still showed a
preference towards grasses. Forbs were of simar proportions in the diet as was
available in the pasture and shrubs were' not preferred. Beck and Peek (2005)
found similar preferences for grasses, forbs and shrubs. However, site did
influence their results with cattle using grasses and forbs in proportion to their
availability in aspen habitats, but cattle preferred grass in a sagebrush habitat.
Holechek at al. (1982) provided percent cover data which allowed for comparison
to percent within diets, however, this may not be a surrogate for production.
They reported that during mid-June cattle preferred a grass dominated diet with
forbs in similar proportion of diet composition to percent cover, and shrubs not
preferred. But by mid-August, forbs were not preferred and shrubs composition
within diets was of similar proportion to percent cover. However, these
differences may be the result of continued grazing of the plant communities and
representconfounded by changes n forage availability.
MANAGEMENT IMPLICATIONS
Cattle grazing on forested rangelands have the ability to select a diet that has
sufficient quality to meet their nutritional requirements. In this study, timber
harvest and previous herbivory had no effects on the quality of diets that cattle
were able to select. Seasoti of u
was the orfy influence on diet quality; even
though diet quality was declining ft was suffident to prevent weight loss in a
lactating cow. Cattle grazing forested rangelands in northeastern Oregon
preferred a diet that was dorninatd by grarnirmids. However, as graminoid
production decreases, such as in heavily timbered areas, cattle will increase
consumption of forbs. Shrubs occurred considerably less in the diets and forbs
occurred of similar proportion as available on the rangeland. These results were
determined when forage availability was not limited. As a result, managers may
want to monitor cattle use and determine rotational schedules based on when
cattle start consuming shrubs in order to meet management goals.
100
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15th ed. Assoc. Offic. Anal. Chem.,
Beck, J.L. and J.M. Peek. 2005. diet compositon1 forage selection, and potential
for forage competition among 1k, deer, and livestock on aspen-sagebrush
summer range. Range/and Ecology and Management. 58: 135-147.
Cook, C.W. and L.E. Harris. 1968. Nutritional value of seasonal ranges. Utah
Agricultural Experiment Station Bulletin 472.
Darambazar, E. 2003. Factors influencing diet composition of beef cattle grazing
mixed conifer mountain riparian areas. M.S. Thesis. Oregon State University.
Corvallis.
Dealy, J.E. 1966. Bitterbrush nutritional levels under natural and thinned
ponderosa pine. USDA Forest Service Research Note PNW-33. Pacific
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Hedrick, D.W., B.R. Eller, J.A.B. McArthur, and R.D. Pettit. 1969. Steer grazing on
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Holechek, J.L., M. Vavra, and 3. Skovlin. 19i. Diet quality and performance of
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Holechek, J.L., M. Vavra, 3. Skovlin, and W.C. Krueger. 1982. Cattle diets in the
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101
Jameson, D.A. 1967. The Relationship of Tre Overstory and Herbaceous
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McConnell, B.R. and J.G. Smith. 1965. Understory response three years after
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103
CHAPTER S
CONCLUSION
104
The overall objectives o tHs eseirJ V:re to evaluate the relationship
between overstory canopy cover ard ungulate herbivory on understory production,
understory species composition, and diversity in a ponderosa pine and grand fir
forest in northeastern Oregon. Ti primary fectors evaluated were species
frequency, shrub cover and derislt, and understory production.
Changes in species frequency was affected by both herbivory and timber
harvest. The magnitude of species changes was greater with timber harvest than
herbivory, thereby indicating that timber harvest had a greater affect on
influencing successional trajectories of understory vegetation. However, due to
large variations in species occurrence across ponderosa pine treatments, we were
unable to detect consistent directional responses attributed to either timber
harvest or herbivory treatments. These non-directional responses by the
understory vegetation were probably due to initial composition of plant
communities and subsequent differences in pnt community changes among the
ponderosa pine sites. Timber harvest and herbivory had little effect on shrub
cover and density.
Grand fir sites were not as variable in plant community composition
compared to the ponderosa pine sites. As a result, we were able to document
consistent responses by the understorv vegetation. Graminoid species increased
with increased levels of overstory removal, whereas, forbs responses were
determined by life history traits. Again, timber harvest and herbivory had little
effect on shrub cover and density.
105
Total production of u derstuy vs,
increased with decreasing
overstory canopy cover. Treatmits grazed by cWe, elk and mule deer reduced
understory production in ponderosa Dine sftes but increased understory production
in the grand fir sites. These dificrcmes were most likely attributed to soil
characteristics.
Understory vegetation dynamcs f011owing timber harvest is of major
concern, because herbivores are attracted to these areas and they have the
potential for altering successional pathways. However, this research
demonstrated that herbivory following timber harvest had minor effects on the
resulting plant communities. Specifically, the plant communities among the
ponderosa pine sites all responded differently. Across a landscape this could be
very important in maintaining biological diversity and, in some cases, in mitigating
effects of animal congregation areas. Ponderosa pine sites were also grazed by
cattle during the time when many of native peennial grasses are most susceptible
to damage from grazing (Stout and Quinton 1986) which could have influenced
community responses.
Irwin et al. (1994) and Riggs et al. (2000) documented that herbivores
reduced production and cover of shrubs compared to exclosures. However, in our
study, there was not a response of either shrub density or cover to removal of
herbivores. This unexpected result could be the result of lack of disturbance (in
the case of the ponderosa pine sites), herbivory by cattle, elk and deer over the
past 40 years has suppressed shrubs to the point that they are now removed from
106
iU these systems to simulate shrub
this system and(or) fire needs to be
reg rowth.
The lack of strong directcn, changes lti understory vegetation may also
relate to our design structure
Ev
aving the.
xperimerit structured this way,
limits our analytical abilities. Rare pnts and plants that occupy a single
treatment were removed from all analyses. As a result, we could be
underestimating the effects of herbivory and timber harvest treatments. However,
the plants used for analyses typically represented the dominant species in
frequency and production. By reducing the rare species; the ability to detect
differences in plant communities is increased because of the reduction in noise
(McCune and Grace 2002).
Further studies need to be determined on the levels of disturbance,
specifically, understory vegetation responses to fire, timber harvest and herbivory.
Also, studying the effects of different stocking rates and season of use would
provide valuable insight into estabishng grazing systems on forested rangelands.
The second objective of this research was to determine the effects of
timber harvest, ungulate herbivory, and season of use on the nutritional quality
and botanical composition of cattle diets. The primary factors evaluated for
nutritional quality of diets were CP, IVOMD, ADF, and NDF. The primary factors
evaluated for botanical composition of diets was the microhistological
determination of the proportions of grarnnoids, forbs arid shrubs from the diets.
Season of use was the primary factor in determining nutritional quality of
steer diets in ponderosa pine and grand fir sites. Higher quality diets were
107
harvest and herbivory had very
measured in June compared ta Agust.
limited effect on the nutritional auaty of sttr diets. Graminoid species
dominated the diets of steer across all sftes
But., consumption of forbs increased
with increased overstory crpy cvr in grand
fir
sites. Shrubs were not an
important component of steer diets.
Cattle grazing on forested rangelands are constantly faced with changes in
species availability, changing plant communities across the rangeland, variation in
production from feeding site to feeding site, and seasonal changes in moisture
content and forage quality. Even wfth all these changes and variability in potential
forage species, cattle typically selected a diet that is high in grarninoids when
given the opportunity.
Rotational grazing systems are regularly used to better distribute cattle
across rangelands and to provide vegetation recovery periods. Therefore, further
study is needed in determining tne effects u
rnited forage availability on cattle
diet selection for various time periods throughout the grazing season. Also,
documenting the effects of timber harvest, herbivory and progression through the
grazing season on the available forage quality.
108
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115
APPENDICES
116
A
LiSt of Species Identf1d
Grn Fir nd Ponderosa Pine Sites
117
Table 1. List of all species idnUfld y eqncy counts for ponderosa pine sites.
CODE
(grasslike)
PLANT SPECtS
COMMON NAME
CACO
Carex cor,dnnokfes
CAGE
ARO
LUCA
Carexgerevfi
NW sedge
Elk sedge
Ross sedge
Woodrush
Carex ross/i
Luzuld 7zg
(shrubs)
MARE
SPBE
SYAL
Ma/ion/a repens
Spirea betul/folia
Symp/iccarpos e//iu
Oregon grape
Spirea
Snowberry
(grasses)
AGCA
AGID
AREL
BRBR
BRCA
BRHO
BRIN
BRRI
BRTE
BRVU
CARU
DAIN
DAGL
Agropyron can/urn
Agrost/s idahoensis
Arr/ienat/ierum elat/us
Bromus briziformis
Bromus carinatuis
Brornus hordeaceus
Brornus inermus
Bromus ri'/dus
Bromus tectorum
Brornu vu/garLc
c/amagrostr/s rubescens
Danthonia hiterrnedi
Dactyls gioci-iert
DECA
ELGL
FEID
FEOC
FEOV
FERU
FESU
KOCR
MEBU
PHPR
POCO
POPR
POSA
PSSP
STOC
TRCA
Deschamps/a coespitosa
Elyrnus glaucus
Festuca idahoensis
Festuca occidenta/is
VUMI
Festucaovina
Festuca rubra
Festuca sub/ara
Koeleria cr5tata
Meilcabu/bosa
Ph/eurnpratense
Poa compressa
Pea pretense
Poa secunda
Pseudoroegnerfa spicata
Bearded wheatgrass
Bentgrass
Tall oatgrass
Rattle brome
frTtn. brome
Soft brome
Smooth brome
ripgut brome
Cheatgrass
Co!umbia
Pirtegrass
timber oatgrass
Orchard grass
tufted hairgrass
Blue wildrye
Idaho fescue
Western fescue
fescue
red fescue
Bearded fescue
Prairie Junegrass
Onion grass
Timothy
Canadian bluegrass
Kentucky bluegrass
Sandberg's bluegrass
Bluebunch wheatgrass
5tia occidental/s
'N. needlegrass
Tr/setum canescens
Tall trisetum
Vu/pi microstachys
Small fescue
Ach/ilea milefo//um
Adenocaulon b/color
Yarrow
Pathfinder
(forbs)
ACMI
ADBI
118
Table_1._continued.
CODE
PLANT SECIE3
AGGL
ANPA
ANMA
Agoseris g/äuca
ANPI
ANRO
APAN
Anemone piJ.iri
Antenndf Ia is-ea
Apocyrlum ahdrcsaem/fnIiom
Aquilegia Foimosa
AQFO
ARCO
ARSE
ASCA
BLSC
CAAP
CAQU
CEGL
CHAN
CHUM
CIAR
C1VU
CLDO
CLHI
CLPE
CLRH
COGR
COLI
COPA
CROC
D]TR
EPBR
ERSU
EUCO
GAHU
GETR
GEVI
HEUN
HIAL
HICY
IRMI
LASE
LASE
LIBO
UNU
LIRU
LOTR
Anemorw pan'if1'ra
Anapha/is ;ñagritaci
Am/ca cord/To//a
Arenaria serpy//ofolia
Astragalous canadensis
Blepharpappus scaber
Casti/eja appfegate/
Cainassia quamash
Cerastium glomeratum
Chamer/on angustifo//um
ch/inaph//a urnbe//ate
c/ms/urn aivense
C/ms/urn vulgare
Cl/nopodium douglas/i
c/ematis /i/rsutissima
clayton/a perfoliata
C/ark/a rhomboidea
co/tom/a grandiflora
Co/tom/a linear/s
Co//ins/a par/:f
crepis occidental/s
D/sporurn trachycarpum
Ep//obium brachycarpurn.
Er,ieron subtrinervis
COMMON NAME
pale agoseris
smallfiowered anemone
pearly everlasting
Piper's anemone
Rosy pussytoes
spreading dogbane
Western columbine
Heartleaf arnica
Thymeleaf sandwort
Canadian milkvetch
Rough eyelashweed
Indian paintbrush
small camas
Sticky chickweed
Fireplant
Pipsissewa
Thistle
Bull thistle
yerba buena
clernatis
Minr's lettuce
diamond clarkia
grand colomia
tiny trumpet
Bluelips
Largeflower hawksbeard
fairybells
tall annual willowherb
Ga/turn spp.
Fleabane
Showy aster
Strawberries
Bedstraw
Gayophytuni hum/fe
dwarf grcundsmoke
Geum tr/flortm
Prairiesmoke avens
St:icky geranium
oneflower helianthella
Hawkweed
hou ndstongue hawkweed
Rocky Mountain iris
Pricly lettuce
Peavine
prickly lettuce
Eurybia consp/cua
Fragaria spp.
Geranium viscosissium
He//anthella uniflora
Hieradum spp.
Hieradum cyrioglossoides
Iris missouriensis
Lactuca serriola
Lathyrus spp.
Lactuca serrio/a
Linnaea borealis
Linanthusnuttal/li
Twinflower
linanthus
Lithospermum ruberale
Lomat/urn triternatum
Stoneseed
nine leaf biscuitroot
119
Table 1. continued.
CODE
MAMI
MARA
MAST
MELU
MINU
MODI
MOMA
MYVE
OSBE
PACA
PAPS
PAST
PLANT SPECXFS
COMMON NAME
Lupinus spp
Mad/a sop.
Ma1&nt/7emcrfl icemosw
Ma/a ntfl emLr ste//a turn
Medicago Iuuufi,a
Lupine
Microser, n;tans
Month dichotoma
Moehringia macrophylla
Myosotis verna
Osmorhiza berteroi
Packera cana
Packera pseudaurea
Packera streptanthifo/la
Pedicu/aris spp.
PH LO
Perkierid/a gairdoeri
Ph/ox grad/is
Ph/ox hood/i
Ph/ox longifo/la
PIEL
PODO
Piper/a e/egans
Polygonum doug/as/i
POGL
PRVU
Potent/la glandulosa
Potent/la grazilis
Prunella vu/agaris
R.AAC
Ranuncu/us acriform,:c
RAUN
RUAC
SELA
Ranuncu/us iinc,atus
PEGA
PHGR
PHHO
POGR
SEIN
SESE
SIDO
SIM E
SIOR
SOMI
STAM
STLO
Rumex acetasella
Sedum Ianceo/atum
Senido inergerrirnu
Senecio serra
Silene doug/as/i
Si/ene menz/esfi
Sida/cea oregana
So/idago mi.ssour/ensii
Streptopus amp/exfo/ius
TAOF
THFE
TRDU
TRRE
Ste//aria /ongies
Symphyotrichum spathulatum
Taraxacum offic/na/e
Thai/drum fenderi
Tragopodon dub/us
Trifo//um repens
VESE
Veronica serpv///fo/ia
VAED
VESE
Va/eriana edu/Ls
SYSP
VIAD
VIAM
ZIPA
Veronica serpylilfolia
Viola abunca
V/cia americana
Zi'adenus paniciilatus
tarweed
Western False Solomon's Seal
Starry False Solomon's Seal
black medick
Nodding microceris
dwarf minerslettuce
Large leaf sandwort
spring forget-me-not
sweetcicely
woolly groundsel
falsegold groundsel
Rocky Mountain groundsel
lousewort
Gardner's yampah
slender phlox
spiny phlox
longleaf phlox
elegant piperia
Knotweed
Gland cinqufoil
NW cinquefoil
Common self-heal
sharpleaf buttercup
Woodland buttercup
Sheep's sorrel
spearleaf stonecrop
Western groundsel
Tall ragwort
catchfly
Menzies' campion
Oregon checkerb!oom
Missouri goldenrod
claspleaf twistedstalk
longstalk starwort
Western Mountain aster
Dandelion
Meadow rue
Yellow salsify
White clover
Thyme-If sandwort
tobacco root
thyrneleaf speedwell
Violet
American vetch
foothill deathcamas
120
Table 2. List of all species dert
counts in grand fir sites.
PLANT SPECIES
COMMON NAME
cACO
C'arex condnno/ces
CAGE
carex gerayif
CARO
LUCA
Luzula carnpet7is
NW sedge
Elk sedge
Ross sedge
Wood rush
CODE
(grasslike)
Carex tsi
(shrubs)
MARE
SPBE
SYAL
Ma/ion/a repens
Spfrea betu/i7o/h
Oregon grape
Spirea
Symphocarpos a/bus
Snow berry
Agropyron can/urn
Agrostis gIqantea
Agrostis idahoensis
Arrhenatherum elatius
Bronius brizifornils
Bromus cariqatu/s
Bromus hordeaceus
Brornus inermus
Bearded wheatgrass
redtop
Bentgrass
Tall oatgrass
Rattle brome
Mtn. brome
Soft brome
Smooth brome
Bromusri'idus
ripgutbrome
Brornus tectc'rurn
Brornus vulgar/s
Ca/amagrostris rubescens
Cheatgrass
Columbia
Pinegrass
Orchard grass
Onespike danthonia
Slender hairgrass
Blue wildrye
Quackgrass
Idaho fescue
Western fescue
fescue
Bearded fescue
Prairie Junegrass
Onion grass
(grasses)
AGCA
AGGI
AGID
AREL
BRBR
BRCA
BRHO
BRIN
BRRI
BRTE
BRVU
CARU
DAGL
DAUN
DEEL
ELGL
ELRE
FEID
FEOC
FEOV
FESU
KOCR
MEBU
PHPR
POCO
POPR
POSA
PSSP
SIHY
Dactyl/s gkmrila
Danthonfa unispicata
Descharnpsi elongate
E/ymusglaucus
Elymusrepens
Festuca idahoensis
Festuca occidenta/,&
Festuca ovine
Festuca sub/eta
Koeferfa cristata
Me//ca bu/bosa
Phleurnpratense
Pea coinpressa
Poe pretense
Poe secunda
Pseudoroegnerfa spicata
5/tan/on hystrA'
THIN
5tia occidenta/L
Thinopyrurn intermed/um
TRCA
VUMI
Trisetum canescens
Vu/p/a microstachys
STOC
Timothy
Canadian bluegrass
Kentucky bluegrass
Sandberg's bluegrass
BI uebunch wheatgrass
Bottlebrush squirreltail
W. needlegrass
Intermediate wheatgrass
Tall trisetum
Small fescue
121
Table 2. continued.
CODE
PLANT 5PECE5
COMMON NAME
Ac/i//lea msie?O/iam
Yarrow
Pathfinder
pale agoseris
field pussytoes
smallflowered anemone
Piper's anemone
Rosy pussytoes
Narrowleaf pussytoes
Western columbine
Heartleaf arnica
Thymeleaf sandwort
Canadian milkvetch
Rough eyelashweed
Indian paintbrush
Sticky chickweed
(forbs)
ACMI
ADBI
AGCL
ANNE
ANPA
ANPI
ANRO
ANST
AQFO
ARCO
ARSE
ASCA
BLSC
CMP
CEGL
CHAN
CHUM
CIAR
CIVU
Adenocau/C77 b/color
Agose.r's giaut
Antennaria Peg&cta
Anemone parviflora
Anemone p#oer/
Antennaria rosea
Antennaria stenophylla
Aquilegia Formosa
Am/ca cord/fe//a
Arenar/a serpy//ofo/la
Astragalous canadensfs
Blepharpappus scaber
Casti//eja applegatel
Cerastium glomeratum
C'hamerion angustif cl/urn
C'himaph//a umbe//ate
Cr5/urn arvense
Clrsium vulgare
Clayton/a perfoliata
Firepla nt
Pipsissewa
COGR
Cb/lomia grand/flora
COLI
Co//omia linear/s
Thistle
Bull thistle
Miner's lettuce
diamond ciarkia
grand collomia
tiny trumpet
COPA
CROC
CYOF
EPBR
ERHE
EPLA
Cb//ins/a paTxa
Bluelips
crepis occidental/s
Largeflower hawksbeard
Houndstongue
tall annual willowherb
Creamy buckwheat
willowherb
CLPE
CLRH
ERSU
EUCO
c/ark/a rhornboia
cynoglossum offlciia/e
Epiobium brach,vcaipurn
Er/ogonum /ierac/eoides
Epiob/um tact/forum
Eriieron subtrinervis
Euiybia conspicua
Fragaria spp.
Ga//urn spp.
GETR
GEVI
GOOB
HIAL
IRMI
LASE
UBO
URU
AMI
MARA
Geuin tr/florum
Geranium v/.sces/ssium
Goodyera oblongifolia
Hieracium spp.
Iris niissouriensis
Lactuca serriola
Lat/iyrus spp.
L/nnaea borea/i
Lithospermurn ruhera/e
Lupinus spp.
Mad/a spp.
Ma/anthemum racemosum
Flea bane
Showy aster
Strawberries
Bedstraw
Prairiesmoke avens
Sticky geranium
Rattlesnake plantain
Hawkweed
Rocky Mountain iris
Pricly lettuce
Peavine
Twinflower
Stoneseed
Lupine
tarweed
Western False Solomon's Seal
122
Table 2. continued.
CODE
PLANT SPECIES
COMMON NAME
MAST
MINU
MOMA
ORSE
OSBE
PAPS
fr1ahnthemum ste//atisn
Microseris nutans
fr1oehrimiia
cJvphy/
Orthi/la secimda
Osmorhiza berterol
Packerapseudaurea
PEGA
Peridendi gairdneri
PHGR
Ph/ox graci/i
Po/ygonuin doug/as/i
Potentii/a glandu/osa
Starry False Solomon's Seal
Nodding microceris
Large leaf sandwort
Sidebells wintergreen
sweetcicely
falsegold groundsel
Gardner's yampah
slender phlox
Knotweed
Gland cinqufoil
NW cinquefoil
Common self-heal
Bracken fern
Woodland buttercup
Sheep's sorrel
Western groundsel
P000
POGL
POGR
PRVU
PTAQ
RAUN
RUAC
SEIN
SESE
SIDO
SIME
SOMI
SPRO
STAM
STLO
STME
SYSP
TAOF
THFE
TRDU
TRGR
TRRE
VESE
VETH
VIAD
VIAM
VIGL
Potentilla graz//Ls'
Prune//a vu/agaris
Pter/dfucn aqu/ilnurn
Ranuncu/us unc/natus
Rumexacetose//a
Sen/do /nergerrfrnus
Senecio serra
Si/ene doug/as/f
Silene menzfesf/
Solidago missourfensil
Spiranthes romanzoffiana
Streptopus ampfexfo/ius
Stellarfa /ongioes
Ste//aria med/a
Symphyotrfdium
Taraxacum officinafe
Thalictrumfenderi
Tragopodon dub/us
Trite/eia grandiflora
Trifollum repens
Veronica serpyllifolfa
Verbascum thapsus
Viola abunca
Vicia americana
Viola glabella
Tall ragwort
Catchfly
Menzies' campion
Missouri goldenrod
Whirled Orchid
claspleaf twistedstalk
longstalk starwort
Common chickweed
Western Mountain aster
Dandelion
Meadow rue
Yellow salsify
Largeflower triteteia
White clover
Thyme-If sandwort
Mullein
Violet
American vetch
pioneer violet
123
Appendix 3
Ponderosa Pine Understory Vegetation Changes
124
Table 1. The changes in species cornpoitior'
i
a por.derosa pine forest from 1985 to
1988 using a 30 cm x 30 cm ptftnie.
ControOvrstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
0.0
-20.0
0.0
20.0
0J
0.0
0.0
26,7
3.3
3.3
0.0
3.3
13.3
6.7
26.7
13.3
-3.3
56.7
0.0
0.0
-26.7
6.7
23.3
0.0
0.0
-20.0
-26.7
33.3
0.0
0.0
-13.3
-10.0
-13.3
33.3
-23.3
20.0
0.0
0.0
-10.0
-33.3
-23.3
-26.7
0.0
0.0
10.0
10.0
-10.0
3.3
0.0
0.0
0.0
-10.0
-36.7
13.3
0.0
0.0
0.0
3.3
-36.7
0.0
0.0
0.0
6.7
43.3
-13.3
6.7
0.0
0.0
0.0
0.0
6.7
16.7
0.0
0.0
3.3
0.0
0.0
23.3
0.0
0.0
-36.7
0.0
0.0
-20.0
-40.0
-10.0
0.0
30.0
0.0
-3.3
0.0
-30.0
-6.7
-6.7
3.3
0.0
6.7
400
-3.3
0.0
30.0
-20.0
3.3
6.7
-10.0
10.0
-10.0
0.0
-3.3
-40.0
0.0
-23.3
-6.7
-16.7
0.0
-6.7
-40.0
-36.7
-6.7
-10.0
-40.0
-23.3
-60.0
-13.3
3.3
6.7
-3.3
0.0
-3.3
0.0
3.3
0.0
0.0
0.0
0.0
0.0
3.3
0.0
6.7
-3.3
-3.3
13.3
6.7
10.0
3.3
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-20.0
3.3
-26.7
-6.7
3.3
3.3
0.0
0.0
0.0
0.0
10.0
-23.3
-33.3
20.0
0.0
3.3
3.3
-10.0
6.7
0.0
0.0
26.7
33.3
-10.0
3.3
-30.0
10.0
-3.3
3.3
3.3
10.0
6.7
0.0
3.3
0.0
6.7
0.0
0.0
23.3
Site I
Bromus carinatus Hook. & Ar
C'arex concinnoides Mackenzie
carexgeyeriBoott
Ca/amagrostirubescensBuck.
Elymusg/aucusBuckl.
Me//ca bu/bosa Geyer ex Porter &
Coult.
Poapratens/-L.
Trisetum canescens Buckl.
Ach//learni//efollumL.
Anemone piperiBritt. ex Rydb.
Antennar/aroseaGreene
Am/ca cordifolia Hook.
Clayton/a perfollata Donn ex
Eu,ybfaconsp/cua(Lindl.) Nesom
Fragar/aspp.
Gal/umspp
Hieracium a/b/forum Hook.
Irimisour/ensiNuft.
Lathyrusspp.
Lupinusspp.
Moehring/a macrophy//a (Hook.)
Fenzl
Osmorhiza bettero/DC.
Potent/Na grad/is Dougl. ex
Hook.
Prune//a vu/gar/sL.
Ranuncu/usspp.
Ste/far/a /ongies Goldie
Symphyotrichum spathulatum
(Lindi.) Nesom var.
spathu/atum
Taraxacurn offic'/na/eG.H. Weber
ex Wiggers
Tha//ctrum fend/eriEngelm. ex
Gray
Tr/follumrepensL.
Viola adunca Sm.
V/cIamericanaMuhl.exWiJId,
Mahonia repens (Lindi.) G. Don
Spiraeabetul/fo/iaPallas
Symphoricarposa/bus(L.) Blake
20.0
-20.0
26.7
-10.0
3.3
-10.0
OO
0
3.3
0.0
3.3
0.0
13.3
-20.0
13.3
125
Table 1. continued.
Contrct)versQty
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
0.0
0.0
2(0
0.0
23.3
0.0
-3.3
3.3
0.0
20.0
3.3
0.0
0.0
0.0
20.0
3.3
13.3
0.0
Sit*?
BromuscarfnatllsHook. &Arn.
carex concinno/des Mackenzie
carexge,vei-/Boott
ca/amagrosti rubescens Buck!.
E/ymusg/aucusBuckl.
Me//ca bulbosa Geyer ex Porter &
10.0
-6.7
26,7
0.0
0.0
0.0
26.7
13.3
3.3
0.0
0.0
0.0
10.0
0.0
0.0
6.7
-10.0
-6.7
0.0
0.0
6.7
20.0
-50.0
0.0
0.0
3.3
-6.7
6.7
0.0
0.0
-3.3
10.0
-26.7
0.0
0.0
16.7
0.0
13.3
Coult.
Poapratens/sL.
26.7
Thetum canesceas Buckl.
-20.0
Achi/Ieami/Iefo//umL.
-16.7
Anemone ptoer/Britt. ex Rydb.0.0
AntennariaroseaGreene
3.3
13.3
Arn/cacordffo/iHook.
aaytoniapen'o//ataDonn ex
3.3
-30.0
-10.0
6.7
0.0
0.0
-10.0
0.0
0.0
0.0
0.0
0.0
0.0
conspicua (Lindi.) Nesom
Fragar/aspp.
Gallumspp
Hierac/urn a/b/fforum Hook.
Iris m/ssour/ens/sNutt.
Lat/7yrusspp.
Lupinusspp.
0.0
6.7
0.0
-6.7
0.0
13.3
0,0
10.0
CD
26.7
0.0
13.3
3.3
0.0
0.0
-26.7
-6.7
0.0
0.0
0.0
13.3
0.0
-6.7
13.3
0.0
6.7
0.0
16.7
0.0
6.7
-6.7
13.3
-43.3
0.0
-30
0.0
3.3
0.0
-16.7
0.0
3.3
-6.7
-6.7
-6.7
-6.7
-3.3
-3.3
-3.3
0.0
0.0
3.3
3.3
0.0
3.3
0.0
0.0
0.0
-3.3
0.0
0.0
Prune//a vu/garisL.
0.0
0.0
0.0
0.0
0.0
16.7
Ranuncu/usspp.
0.0
0.0
0.0
0.0
0.0
Ste//aria /ongioes Goldie
0.0
0.0
3.3
3.3
0.0
0.0
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-3.3
-3.3
0.0
-3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
10.0
0.0
3.3
-3.3
0.0
0.0
16.7
-6.7
16.7
0.0
0.0
16.7
0.0
0.0
0.0
3.3
-3.3
20.0
-3.3
0.0
3.3
3.3
-3.3
3.3
0.0
0.0
0.0
0.0
13.3
Euiyb/a
Moe/irfngimacrophy//a(Hook.)
Fenzl
Osmor/ii.abeitero/DC.
Potent/I/a gradilic Dougl. ex
Hook.
0
Symphyotr/chum spat/iu/atum
(Lindi.) Nesom var.
spatliulatuni
Taraxacum offidna/eG.H. Weber
cx Wiggers
ThaI/ctrum fend/er/Engelm. ex
Gray
Tr/fol/umrepensL.
Viola adunca Sm.
V/cia americana Muhl. ex WilId.
Ma/ion/a
repens(Lindl.) G. Don
Spiraea betuflfoli'a Pallas
S,vrnphoricarposalbus(L.) Blake
0.0
-6.7
-3.3
-13.3
-10.0
0.0
26.7
126
Table_1._continued.
CorroOyert
Thinned Overstory
CEXC
TExC
GRAZED
CEXC
TEXC
GRAZED
0.i
16.7
0,0
20.0
-IOJJ
10.0
23.3
0.0
26.7
-10.0
-6.7
20.0
0.0
20.0
-10.0
13.3
-3.3
0.0
20.0
-10.0
0.0
3.3
0.0
16.7
-16.7
10.0
20.0
6.7
-3,3
-3.3
6.7
0.0
-3.3
13.3
6.7
-43.3
6.7
0.0
0.0
10.0
20.0
-43.3
10.0
0.0
-36.7
-6.7
0.0
3.3
-3.3
-10.0
0.0
0.0
0.0
20.0
50.0
3.3
0.0
3.3
6.7
0.0
0.0
0.0
3.3
0.0
0.0
0.0
10.0
3.3
0.0
0.0
-20.0
23.3
-6.7
Iiismissouriens,'sNutt.
0.0
-6.7
0.0
0.0
-3.3
Lathyrusspp.
-6.7
Lup/nusspp.
frloehr/ng/a macrophylla (Hook.)
3.3.
0.0
-10.0
-3.3
6.7
0.0
-6.7
3.3
Site 3
Bromus carinatus Hook. & Am.
carex concinnoides Mackenzie
Carexgeyer/Boott
C'a/amagrost&rubescensBuckl.
ElymusglaucusBuckl.
Me//cabu/bosaGeyerexPorter&
Coult.
Poapratens/sL.
TrisetumcanescensBuckl.
,4chi//eamf//efol/umL.
Anemone pioer/Brift. ex Rydb.
AntennariaroseaGreene
Am/ca cordifo/la Hook.
C/ayton/aperfolfataDonnex
Eulybia conspicua (LindI.) Nesom
Fragariaspp.
Galiumspp
Hierac/um a/b/forum Hook,
Fenzl
Osmorhizabertero/DC.
20.0
-10.0
-10.0
-30.0
-6.7
0.0
0.0
3.3
-10.0
3.3
6.7
13.3
-33.3
-6.7
0.0
13.3
13.3
0.0
-20.0
-30.0
-3.3
0.0
-6,7
16.7
10.0
0.0
3.3
3.3
-20.0
-76.7
-40.0
-10.0
3.3
-3.3
10.0
0.0
3.3
-6.7
0.0
0.0
36.7
16.7
20.0
3.3
6.7
0.0
-3.3
0.0
6.7
0.0
6.7
0.0
3.3
3.3
0.0
-6.7
0.0
3.3
6.7
3.3
10.0
0.0
10.0
13.3
13.3
00
0.0
0.0
0.0
0.0
0.0
6.7
6.7
0.0
-10.0
3.3
16,7
3.3
0.0
-3.3
0.0
0.0
-3.3
TrifoliumrepensL.
16.7
V/a/a aduncaSm.
V/cia americana MuhI. ex Willd.
Mahon/a repens (LindL) G. Don
0.0.
10.0
16.7
-3.3
3.3
-3.3
0.0
30.0
-10.0
20.0
Potentiiagraci/iDougI. ex
Hook.
Prune//a vu/garisL.
Ranuncufusspp.
Ste//arh/ongioesGoIdie
Symphyotrichum spathulatum
(Lindl.) Nesom var.
spat/lu/atum
TaraxacuniofflcinaleG,H. Weber
ex Wiggers
Tha/Ictrum fend/er/En gelm. ex
Gray
0.0
13.3
23.3
0.0
-10.0
0.0
0.0
-20.0
13.3
-3.3
-16.7
13.3
3.3
-6.7
-10.0
Spiraeabetullfol/aPallas
16.7
16.7
3.3
0.0
0.0
0.0
Symphor/carposa/bus(L.) Blake
-6.7
-6.7
-26.7
-20.0
23.3
-3.3
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big
game grazing only; TEXC total exclosurè, exclusion of cattle and big game grazing.
127
Table 2. The changes in spedts
r
oCsitio.' ir
1988 using a 30 cm x 60cjplotfrme
:.cnderosa pine forest from 1985 to
çc.ntistory
Thined Overstory
CEXC
TE)C
GRAZED
CEXC
TEXC
GRAZED
0.0
0.0
0.0
-3.3
3.3
0.0
0.0
3.3
6.7
23.3
0.0
3.3
13.3
6.7
13.3
0.0
0.0
13.3
-3.3
53.3
0.0
0.0
-23.3
3.3
36.7
0.0
0.0
-16.7
-23.3
20.0
3.3
-20.0
-6.7
-10.0
6.7
23.3
6.7
0.0
0.0
-20.0
-43.3
16.7
0.0
0.0
-3.3
33.3
-13.3
20.0
0.0
0.0
-3.3
3.3
0.0
20.0
16.7
3.3
3.3
-20.0
-3.3
site I
Bromus car/paWs Hook. & Am,
carexconcinno/des Mackenzie
carexgeyeriBoott
caIamagrostisrubescensBuck.
E/ymusg/aucus8uckl.
ft/let/ca bulbosa Geyer ex Porter &
Coult.
PoapratensisL.
TrLcetum canescensBuckl.
Ach/lleamitlefofiumL.
Anemone p4oeriBritt. ex Rydb.
AntennariaroseaGreene
Am/ca cord/lot/a Hook.
c/ayton/apeifo//ataDonnexwilld.
Ewybia conspicua (Lindi.) Nesom
Fragar/aspp.
Ga//umspp
Hieracium a/b/forum Hook.
.Trim,ssour/ensLcNutt.
Lathyrusspp.
Lupinusspp.
ftloehr/ngfa macrop/iylla (Hook.)
Fenzl
Osmorhiza berteroiDC.
Potent//ta graci7iDougI. ex Hook.
Prune//a vulgarisL.
Ranunculusspp.
Ste//ar/a /ongioesGoldie
Symphyotrichum spathulatum
(LindL) Nesom var. spathu/atum
Taraxacum off/c/nate G.H. Weber ex
Wiggers
Tha/ictrurn fendlerfEngelm. ex Gray
TrifoliumrepensL.
Viola adunca Sm.
V/c/a americana Muhi. ex WiUd.
ft/la/ion/a repens(LindL) G. Don
Spiraeabetul/follaPallas
Symp/ioricarposa/bus(L.) Slake
Site 2
BromuscarfnatusHook.&Arn.
C'arexcoaci'nnofdes Mackenzie
C'arexgeyeriBoott
0.0
-36.7
-3.3
0.0
3.3
3.3
6.7
0.0
-3:3
-133
3.3
-30.0
-6.7
0.0
0.0
-3.3
0.0
0.0
3.3
0.0
3.i
-10.0
-3.3
0.0
6.7
-50.0
3.3
0.0
3.3
20.0
0.0
0.0
-16.7
-6.7
-3.3
3.3
10.0
6.7
-3.3
16.7
0
0.0
20.0
-10.0
3.3
3.3
0.0
43.3
3.3
3.3
6.7
ftO
-6.7
0.0
-10.0
-13.3
-23.3
43.3
-16.7
0.0
-6.7
-43.3
-6QO
3.3
-3.3
0.0
0.0
-10.0
-3.3
0.0
0.0
3.3
-6.7
-6.7
0.0
-20.0
0.0
-3.3
26.7
-60.0
0.0
0.0
3.3
-43.3
-3.3
-3.3
16.7
0.0
6.7
10.0
10.0
10.0
00
00
00
00
00
00
-13.3
3.3
-20.0
6.7
6.7
0.0
6.7
-36.7
-6.7
-3.3
3.3
-3.3
0.0
6.7
-13.3
10.0
0.0
0.0
26.7
0.0
33.3
-13.3
13.3
-6.7
3.3
-3.3
0.0
13.3
-36.7
20.0
6.7
0.0
-16.7
0.0
10.0
3.3
16.7
6.7
0.0
3.3
0.0
26.7
26.7
0.0
36.7
6.7
0.0
40.0
0.0
0.0
26.7
0.0
0.0
26.7
-3.3
0.0
0.0
3.3
0.0
10.0
-20,0
0.6
6.7
0.0
-20,0
00
23.3
10.0
128
Table 2. continued.
O:,ntrOeito
C'alamagrostis rubescens BuckL
E/ymusqlaucusBuckl.
Me//ca bu/bosa Geyer ex Porter &
TEXC
GRAZED
CEXC
TEXC
GRAZED
40.0
13.3
0.0
0,0
0.0
0.0
0.0
3.3
0.0
0.0
3.3
0.0
10.0
0.0
0.0
30.0
-50.0
6.7
-26.7
-6.7
0.0
.o
Coult.
PoapratensisL.
Thined Overstory
EXC
40.0
-26.7
-13.3
0.0
-363
10.0
-16.7
6.7
0.0
Lathyrusspp.
0.0
-6.7
0.0
-10.0
0.0
-40.0
-23.3
-3.3
0.0
0.0
-3.3
0.0
0.0
20.0
0.0
3.3
0.0
36,7
Lup/nusspp.
20.0
-10.0
6.7
3.3
0.0
13.3
0.0
0.0
0.0
10.0
3.3
13.3
0.0
-23.3
0.0
6
-10.0
-6.?
3.3
0.0
0.0
0.0
0.0
0.0
CLO
0.
3.3
0.0
0.0
0.0
0.0
0.0
0.0
00
00
0.0
Trisetum canescensBuckl.
3.3
-16.7
20.0
0.0
3.3
6.7
3.3
0.0
0.0
6.7
0.0
0.0
6.7
0.0
13.3
0.0
16.7
6.7
-3.3
-6.7
-3.3
0.0
-16.7
0.0
0.0
3.3
0.0
0.0
0.0
0.0
0.0
6.7
0.0
23.3
0.0
3.3
0
00
00
00
-3.3
-3.3
16.7
-3.3
3.3
0.0
10.0
0.0
0.0
-3.3
-6.7
-13.3
0.0
16.7
-6.7
0.0
0.0
20.0
6.7
0.0
26.7
0.0
0.0
3.3
0.0
23.3
0.0
33.3
0.0
0.0
0.0
6.7
16.7
0.0
26.7
-3.3
0.0
6.7
6.7
3.3
0.0
6.7
-6.7
0.0
0.0
0.0
0.0
CalarnagrostisrubescensBuckl.
0.0
23.3
-6.7
-10.0
E/ymusg/aucusBucki.
200
0.0
33,3
-3.3
23.3
23.3
0.0
26.7
-10.0
0.0
23.3
0.0
26.7
0.0
6.7
-3.3
0.0
23.3
-16.7
-3.3
13.3
0.0
23.3
-13.3
6.7
23.3
3.3
-3.3
3.3
-10.0
-3.3
10.0
10.0
-53,3
-3.3
-56.7
20.0
-46.7
16.7
-36.7
-6.7
-43.3
AchJl/eamilefo//umL.
Anemone pioeriBritt. ex Rydb.
AntennanaroseaGreene
Am/ca cord/folia Hook.
C%ytoniapetfo/iataDonn ex Wifld.
Euiyb/aconspicua(Lindl.) Nesom
Fragar/aspp.
Ga//urn spp
H/eracfum a/b/forum Hook.
Iris m/ssour/ensisNutt.
Moehr/ngiarnacrophylla(Hook.)
Fenzl
Osrnorhizabertero/DC.
Potenti//agraci/isDougl. ex Hook.
Prune/Ia
vulgar/sL.
Ranunculusspp.
Stellar/a /ongices Goldie
Symphyotrichum spathu/aturn
(Lindi.) Nesom var. spathu/aturn
Táraxacum officinale G.H. Weber ex
Wiggers
Thafictrum fendleriEngelm. ex Gray
TrifollumrepensL.
Viola adunca Sm,
V/cia americana Muhi. ex WilId.
Ma/ion/a
repens(Lindl.) G. Don
Spiraeabetu//fo//aPallas
Symphoricarposalbus(L.)
Blake
3,3
-3.3
00
3.3
13.3
0.0
0.0
-3.3
0.0
-23.3
0.0
16.7
16.7
-3.3
0.0
0.0
13.3
0.0
Site 3
Bromus car/natus Hook. &Arn.
crexcoiicinnoidesMackenzie
carexgeyer/Boott
Me//ca bu/bosa Geyer cx Porter C
Coult.
Poapratens/sL.
Trisetum canescensBuckl.
-60.0
129
Table 2. continued.
CtrC;e
Ac/il/lea milefo//um L.
Anemone pioerlBritt. ex Rydh
AntennariaroseaGreene
Am/ca cord(o/h Hook.
Clayton/a perfoliata Donn ex WkJ.
Euiybfa consplcua (LindL) Neom
Fragariaspp.
G'allumspp
Hierac/um a/b/forum Hook.
IrmmissouriensLsNutt.
Lat/iyrusspp,
Lup/nusspp.
flloe/ir/nglamacrop/iy//a(Hook.)
CEXC
tEX
'10.0
0.0
0.0
6.7
3,3;
0.0
13.3
0.0
0.0..
00
0.0.
GRft.ZED
CEXC
TEXC
GRAZED
-10.0
0.0
13.3
-3.3
-6±
6.7
-6.7
6.7
-10.0
0.0
3.3
3.3
.3.3
6.7
-3.3
3.3
0.0
-3.3
-10.0
-30.0
-3.3
20.0
0.0
6.7
3.3
0.0
0.0
0.0
13.3
10.0
0.0
10.0
-3.3
3,3
0.0
3.3
6.7
0.0
0.0
10.0
3.3
-6.7
0.0
6.7
-3.3
0.0
0.0
-3.3
6.7
-33.3
-20.0
-80.0
-43.3
-23.3
-3.3
0.0
-6.7
46.7
-3.3
6.7
00
10.0.
3.3
10.0
33.3
3.3
0.0
10.0
13.3
13.3
-3.3
30.0
0.0
13.3
00
00
00
00
-10.0
10.0
13.3
Fenz
Osmorhiza berteroiDC.
Potent/I/a grad/is Dougl. ex Hook.
3.3
Prune//a vulgarLcL.
3.3
(!ti
0.0
Ranunculusspp.
6.7
Ste//aria /ongioes Goldie
0.0
0.0
6.7
6.7
3.3
0.0
Symp/iyotr/c/ium spat/iu/atum
(UndL) Nesom var. spathu/aturn
0
00
Taraxacumofflc/naleG.h. Webcex
Wiggers
Thallctrum fend/er/Engelm. ex Gray
Thined Overstory
.3
3.3
0.0
-23.3
33.3
-10.0
-3.3
10.0
0.0
6.7
-3.3
..
33
6.7
13
0,0.
0.0
-3.3
0.0
0.0
-3.3
40.0
33.3
0.0
Viola adunca Sm.
0.0
23.3
-3.3
-16.7 -10.0
0.0
VIa americana MuhI. ex Wifid.
-6.7
-25.7
3.3
16.7
3.3
16.7
Mahon/a repens(Lindl.) G. Don
OM
-20.0
6.7
3.3
0.0
3.3
5p/raeabetu//fo/iPallas
23.3.
16.7
6.7
0.0
0.0
0.0
Symphor/carposalbus(L.) Blake
-26.7
-6.7
-13.3
-13.3
23.3
-3.3
Herbivory treatrnent:s: Graze cattle and bq game grazing; CEXC cattle exclosure, big
game grazing only; TEXC total exciosure, exclusion of cattle and big game grazing.
Tr/follumrepensL.
6.7
130
Table 3. The changes in spees comc;itior
1991 using a 30 cm x 30
i pderosa pine forest from 1985 to
Cônt:):r
Thined Overstory
CEXC TEC GRAZED
CEXC
TEXC
GRAZED
3.3
0.0
3.3
-3.3
53.3
26.7
0.0
3.3
0.0
0.0
13.3
0.0
--6.7
0.0
0.0
36.7
3.3
36.7
-26.7
-3.3
33.3
0.0
0.0
3.3
-23.3
-30.0
-26.7
-13.3
0.0
-3.3
0.0
-6.7
6.7
16.7
0.0
0.0
10.0
13.3
0.0
-10.0
3.3
0.0
0.0
-10.0
3.3
Lupinusspp.
10.0
13.3
20.0
-20.0
3.3
13.3
3.3
0.0
0.0
0.0
-13.3
-10.0
3.3
3.3
-23.3
0.0
10.0
Lathyrusspp.
6.7
-3.3
-3.3
0.0
0.0
3.3
6.7
0.0
-13.3
3.3
0.0
0.0
-13.3
-10.0
3.3
-3,3
-3.3
0,0
0.0
3.3
-6.7
Sfte I
Bromuscar/natusHook.&Arp.
Carexconc/nnoides Mackenzie
C'arexgeyer/Boott
C'a/amagrostLcrubescensBuckl.
Elyrnusg/aucusBuckl.
Me//ca bu/bosa Geyer ex Porter &
Poapratens/sL.
TrietumcanescensBuckL
Achil/eam///efo//umL.
Anemone pioeriBritt. ex Rydb.
Antennaria rosea Greene
Ar,i/ca cord/lot/a Hook.
c'/ayton/petfo/itaDonnexWilld.
Euiybfaconspicua(Lindl.) Nesom
Fragariaspp.
Gal/urn spp
Hieraciuma/bfflorumHook.
IrLmisouriensisNutt.
0.0
0.0
-3.1
6.7
6.7
Osmorh/zaherteroiDC.
Prunellavu/garisL.
Ranuncu/usspp.
Stellari'a/ongioesGoldie
Symphyotrichum spathulatum
(LindL) Nesom var. spat/iu/aturn
Taraxacumoff/dna/eG.H. Weberex
Wtggers
Tha/ictrum fend/er/Engelm. ex Gray
TrifollumrepensL.
Viola aduncaSm.
V/c/a americana MuhI. ex WilId.
Mahon/arepens(Lindl.)G. Don
Sp/raeabetu/ilo/i'aPaflas
Symphor/carposatbus(L.) Blake
Site 2
Bromuscar/natusHook.&Arn.
CarexcondnnoidesMackenzie
CarexgeyeriBoott
0,0
-13.3
3.3
3.3
-3.3
3.3
0.0
-23.3
-13.3
-10.0
-16.7
3.3
40.0
0.0
10.0
0.0
0.0
-63.3
23.3
-6.7
23.3
0.0
0.0
3.3
3.3
0.0
-23.3
10.0
-3.3
0.0
-3.3
-40.0
0.0
0.0
-6.7
-16.7
-6.7
0.0
0.0
0.0
0.0
-53.3
-13.3
0.0
-3.3
3.3
-33.3
-16.7
-3.3
0.0
13.3
-56.7
-3.3
0.0
-3.3
0.0
-23,3
-6.7
-43.3
-13.3
-3.3
-3.3
0.0
-13.3
0.0
3.3
0.0
3.3
3.3
0.0
-3.3
-10.0
3.3
0.0
10.0
0.0
0.0
6.7
20.0
0.0
0.0
33.3
0.0
0.0
3.3
13.3
3.3
0.0
-3.3
0.0
0.0
6.7
23.3
0.0
0.0
13.3
0.0
0.0
6.7
6.7
3.3
0.0
3.3
0.0
20.0
-6.7
0.0
33.3
16.7
0.0
40.0
6.7
0.0
23.3
-16.7
0.0
3.3
-3.3
0.0
0.0
0.0
Moehthgimacrop/iy//a(Hook.)
Potent//ta gradllLs Dougi. ex Hook.
33
3.3
-6.7
-13.3
6
-40.0
6.7
-3.3
6.7
--6.7
0.0
40.0
131
Table 3. continued.
ChtftOerto
Thined Overstory
EXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
Ca/amagrostis rubescens Buckl.
Elymus glaucus Buckl.
t'le/lca bulbosa Geyer ex Porter &
Coult.
Poa pratensic L.
Tnetum canescens Bucki.
Ac/i//lea mi/lefollurn L.
Anemone piøer/Britt. ex Rydb.
Antennarla rosea Greene
Am/ca cord/fat/a Hook.
Clayton/a perfollata Donn ex Wilid.
Ewyb/a conspicua (Lindi.) Nesorn
:400
0.0
10.0
0.0
3.3
0.0
0.0
6.7
16.7
0.0
13.3
0.0
0.0
0.0
0.0
3.3
0.0
3.3
0.0
-20.0
-20.0
-367
-6.7
-6.7
40.0
13.3
10.0
0.0
0.0
0.0
16.7
3.3
0.0
fragarla spp.
-40.0
0.0
-10.0
6.7
10.0
0.0
3.3
0.0
Ga//urn spp
Hierac/um aib/flarum Hook.
Zr/s m/ssour/ens/s N utt.
Lathyrus spp.
Lup/nus spp.
Moehring/a macrophy/la (Hook.)
Fenzl
Osrnarh,a berterol DC.
Potent//la grad/is Dougl. ex Hook
Prune//a vulgar/s L.
Ranunculus
Stellar/a /ongioes Goldie
Symphyotrichurn spatliulaturn
(Lindl.) Nesom var. spathulatum
Taraxacum officinale G.H. Weber ex
Wiggers
Tha/ictrum fend/er! Engelm. ex Gray
Tr/fol/urn repens L.
Viola adunca Sm.
V/cia arner/cana MuhI. ex WiUd.
t'lahon/arepens(Lindl.) G. Don
5a/raea betu/Ifolia Pa Has
Syrnphoricarpos a/bus (L.) Blake
q.o
OO
0.0
6.7
-13.3
0.0
-10.0
0.0
-46.7
6.7
0.0
10.0
0.0
0.0
6.7
0.0
-6.7
0.0
33.3
-26.7
0.0
3.3
10.0
3.3
0.0
0.0
0.0
-46.7
0.0
-16.7
-10.0
20.0
-30.0
10.0
0.0
6.7
-3.3
0.0
0.0
6.7
13.3
-6.7
0.0
-26.7
0.0
.57
-6.7
-6.7
6.7
-3.3
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.3
0.0
0.0
0.0
0.0
0.0
-6.7
0.0
0.0
3.3
3.3
3.3
0.0
3.3
3.3
3.3
0.0
0.0
0.0
0.0
23:3
0.0
3.3
0.0
0.0
0.0
0.0
-3.3
-3.3
-3.3
-3.3
3,3
0.0
3.3
3,3
0.0
-3.3
-3.3
0.0
-6.7
0.0
0.0
-6.7
6.7
10.0
0.0
0.0
10.0
0.0
0.0
0.0
0.0
10.0
3.3
13.3
0.0
6.7
3.3
0.0
-3.3
0.0
13.3
0.0
-3.3
0.0
0.0
-3.3
0.0
0.0
0.0
16.7
0.0
13.3
6.7
13.3
10.0
3.3
23.3
-16.7
-60.0
10.0
-3.3
16.7
6.7
-40.0
-3.3
:3.3
6.7
6.7
13.3
30.0
20.0
-30.0
0.0
0.0
6.7
-6.7
13.3
10.0
0.0
-6.7
-6.7
0,0
-13.3
-10.0
-16.7
10.0
-20.0
6.7
-26.7
30.0
-36.7
3.3
c.o
.0
-10.0
3.3
-3.3
0.0
3.3
-p3.3
-3.3
-3.3
0.0
0.0
-23.3
0.0
0.0
10.0
0.0
3.3
0.0
13.3
Site 3
Brornus car/natus Hook. & Am.
Carex con dnnoides Mackenzie
Carex geyer/ Boott
Ca/arnagrost/s rubescens Buckl.
Elyrnus glaucus Buckl.
Meilca bulbosa Geyer ex Porter &
Co u It.
Poa pratens/s L.
Triseturn canescens Buckl.
0.0
i0.()
-iO.0
-3,3
-:m.o
132
Table 3. continued.
Thined Overstory
Ach/ilea millefoliurn L.
Anemone pier/ Britt. ex Rydb.
Antennarh rosea Greene
A rn/ca cordifolia Hook.
clayton/a petfo/iata Donn ex WiId.
Eutybia conspicua (Lindl.) Nesorn
Fragaria spp.
Gal/urn spp
Hieradum a/b/forum Hook.
Iris missour/ensis Nutt.
Lat/iyrus spp.
Lupinusspp.
Moehringia macrophylla (Hook.)
Fenzl
Osmor/itzaberteroiDC.
Potent//Ia grad/is Dougl. ex Hook.
Prune/la vulgar/sL.
Ranuncuiusspp.
SteiarilongiesGoldie
Symp/iyotric/ium spathu/atum
(LindL) Nesom var. spathu/atum
Taraxacum officinaleG.H. Weber ex
Wiggers
Thai/drum fend/eriEngelrn. ex Gray
Trifo/iumrepensL.
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-6.7
-13
0.0
0.0
0.0
0.0
0.0
-10.0.
0.0
0.0
0.0
3.3
-3.3
0.0
3.3
-6.7
0.0
0.0
0.0
-3,3
0.0
-16.7
10.0
-10.0
3.3
26.7
0.0
3.3
0.0
6.7
0.0
-6.7
20.0
23.3
0.0
-13.3
6.7
10.0
13.3
0.0
16.7
6.7
0.0
-6.7
0.0
10.0
0.0
0.0
-20.0
6.7
20.0
6.7
-16.7
16.7
0.0
0.0
23.3
-3.3
0.0
-26.7
23.3
-16.7
-3.3
6.7
0.0
-6.7
16.7
0.0
-13.3
16.7
-13.3
36.7
6.7
-20.0
0,0
0.0
0.0
0.0
-13.3
3.3
20.0
-3.3
-3.3
-3.3
16.7
-6.7
0.0
0.0
0.0
-3.3
0.0
-10.0
0.0
0.0
0.0
6.7
16.7
20.0
-6.7
-3.3
0.0
13.3
00
00
00
00
00
00
6.7
3.3
3.3
-13.3
0.0
0.0
20
10.0
3.3
0.0
0.0
0.0
-3.3
3.3
3.3
6.7
0.0
10.0
0.0
-67
0.0
-3.3
0.0
0.0
Viola adunca Sm.
-3.3
0.0
6.7
3.3
V/cia americana MuhI. ex WilId.
20.0
20.0
0.0
16.7
frlahon/arepens(LindL)G. Don
1DM
-6.7
-3.3
0.0
3.3
SpfraeabetullfoflaPallas
0.0
16.7
-10.0
3.3
3.3
Symphoricarposalbus(L.) Blake
-26.7
-3.3
-10.0
-3.3
13.3
-6.7
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big
game grazing only; TEXC total exciosure, exclusion of cattle and big game grazing.
0.0
-3,3
3.3
3.3
13.3
133
Table 4. The changes in species composition ir a
nderosa pine forest from 1985 to
1991 using a 30 cm x 60 cm plot frame
Contrc!Cverstory
Thined Overstor
.
Site 1
BromuscarinatusHook. &Arn.
Carex condnno/des Mackenzie
Carexgeyer/Boott
Ca/amagrost/crubesceasBucki.
E/ymusg/aucusBuckl.
frle//cabu/bosaGeyerexPorter&
Pea pratensLcL.
Tr/setumcanescensBuckl.
Ac/iil/eamf/lefolfumL.
Anemone pieriBritt. ex Rydb.
AntennarfaroseaGreene
Arnicacord/fo/laHook.
Clayton/a peifoliata Donn ex WilId.
Euiybiaconspicua(LindL) Nesom
Fragariaspp.
CEXC
IEXC
GRAZED
CEXC
TEXC
GRAZED
3.3
3.3
0.0
6.7
3.3
3.3
(LO
0.0
0.0
33.3
3.3
23.3
3.3
0.0
-3.3
-3.3
53.3
26.7
0.0
6.7
0.0
3.3
13.3
0.0
-33.3
-3.3
30.0
3.3
-33.3
-30.0
-30.0
13.3
16.7
-23.3
10.0
20.0
3.3
-20.0
-13.3
56.7
0.0
-3.3
0.0
0.0
3.3
0.0
6.7
3.3
-30.0
0.0
6.7
0.0
0.0
-60.0
16.7
-3.3
0.0
-3.3
-3.3
10.0
0.0
33.3
0.0
0.0
0.0
10.0
0.0
-23.3
10.0
0.0
-3.3
-13.3
3.3
-6.7
0.0
0.0
-3,3
-10.0
0.0
0.0
-13.3
-6.7
-6.7
0.0
3.3
0.0
3.3
0.0
-13.3
-10.0
-6.7
20.0
0.0
0.0
Lathyrusspp.
6.7
0.0
-3.3
6.7
0.0
0.0
-16.7
Lupinusspp.
6.7
-16.7
10.0
13.3
0.0
-13.3
13.3
-6.7
0.0
-10.0
13.3
0.0
-133
-3.3
0.0
-6.7
-20.0
Potent/I/a grad/is Dougl. ex Hook.
Prune//a vulgarisL.
-50.0
3.3
-3.3
-3.3
-3.3
0.0
0.0
-10.0
0.0
-33.3
-13.3
-3.3
Ranuncu/usspp.
10.0
0.0
0.0
-53.3
-23.3
-3.3
-3.3
6.7
3.3
3.3
10.0
0.0
13.3
-63.3
0.0
0.0
-3.3
0.0
00
00
00
00
00
00
-26.7
-6.7
-40.0
-16.7
-6.7
3.3
-26.7
-3.3
-6.7
0.0
0.0
0.0
16.7
3.3
0.0
-6.7
0.0
0.0
16.7
30.0
6.7
0.0
0.0
0.0
6.7
6.7
3.3
0.0
-3.3
0.0
0.0
3.3
23.3
3.3
0.0
26.7
0.0
33.3
20.0
0.0
50.0
-3.3
0.0
26.7
-16.7
Ga//urn spp
Hieracium a/b/forum Hook.
Iris rni'ssouriensisNutt.
Moe/irfrgiamacrophy//a(Hook.)
OsmorhizaberteroiDC.
Ste//aria IongioesGoldie
Symp/iyotrichum spat/7uiatum
(Lindl.) Nesom var. spatliulaturn
Taraxacum officia/e G.H. Weber ex
Wggers
Thai/drum fend/er/Engelm. ex Gray
Trifo/iurnrepensL.
Viola adunca Sm.
V/cia americana Muhi. ex Wilid.
Mahoniarepens(Lindl.)G. Don
Spiraeabetuifo/faPallas
Syrnphor/carposa/bus(L.) Blake
10.0
3.3
-3.3
-6.7
O0
0.0
0.0
0.0
-16.7
-10.0
3.3
-6.7
-36.7
0.0
10.0
13.3
0.0
3.3
0.0
10.0
-10.0
0.0
10.0
-6.7
Site 2
Bromus car/natus Hook. &Arn.
Carex concinnoides Mackenzie
CarexgeyeriBoott
0.0
26.7
-10.0
134
Table 4. continued.
ControWier
Ca/amagrostLrubescensBuckl.
E/ymusglaucusBuckl.
Me/icabuIbosaGeyerexPorte'&
Couft.
PoapratensisL.
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-13.3
0.0
61
0.0
0.0
0.0
6.7
-13.3
0.0
3.3
0.0
3.3
3.3
0.0
6.7
16.7
-16.7
-3.3
-3.3
0.0
3.3
-33.3
3.3
0.0
13.3
3.3
0.0
0.0
-3.3
-46.7
TrLsetumcanescensBuckl.
-2.6.7
-6J
Ac/if/lea mi//efo/fum L.
Anemone pioeri Britt. ex Rydb.
-20.0
0.0
3.3
0.0
3.3
0.0
-13.3
0.0
-10.0
0.0
0.0
0.0
AntennarhroseaGreene
Am/ca cordifolia Hook.
C/atonipeifo/itaDonn exWilId.
conspfcua (Lindi.) Nesom
Fragariaspp.
Ga/fumspp
Hieracium a/b/forum Hook.
Eutyb/a
Thined Overstory
10.0
0.0
0.0
3.3
0.0
-3.3
-6.7
-6.7
-13.3
0.0
6.7
13.3
6.7
0.0
-3.3
0.0
-53.3
-30.0
10.0
0.0
10.0
-3.3
0.0
0.0
-3.3
13.3
-3.3
-3.3
0.0
0.0
13.3
3.3
16.7
0.0
10.0
0.0
-10.0
6.7
6.7
0.0
0.0
0.0
0.0
0.0
0.0
Lathyrusspp.
-46.7
10.0
-10.0
-16.7
-23.3
Lup/nusspp.
40.0
-36.7
3.3
-30.0
-6.7
3.3
10.0
-6.7
-10.0
-6.7
6.7
-6.7
3.3
0.0
0.0
0.0
3.3
0.0
0.0
3.3
Prune/la vulgariL.
0.0
0.0
0.0
-13,3
0.0
Ranunculusspp.
0.0
0.0
0.0
0.0
Stellar/a longioes Goldie
S,vmphyotrichum spathu/atum
0.0
0.0
-3.3
3.3
3.3
3.3
0.0
3.3
6.7
6.7
0.0
0.0
0.0
6.7
-3.3
3.3
6.7
-3.3
3.3
0.0
-6.7
0.0
-3.3
0.0
20.0
0.0
-3.3
0.0
0.0
-3.3
0.0
6.7
0.0
-3.3
6.7
10.0
3.3
0.0
23.3
0.0
0.0
0.0
0.0
10.0
3.3
16.7
0.0
6.7
3.3
0.0
6.7
0.0
-13.3
0.0
13.3
-3.3
6.7
6.7
13.3
20.0
-50.0
6.7
6.7
13.3
10.0
-6.7
-3.3
10.0
0.0
33.3
-20.0
20.0
-3.3
16.7
30.0
20.0
-30.0
3.3
0.0
3.3
6.7
13.3
10.0
-10.0
-10.0
-10.0
-16.7
0.0
-13.3
-20.0
-20.0
6.7
-20.0
6.7
-13.3
30.0
-40.0
IrLrn/ssouriens/sNutt.
Moehring/amacrophy//a(Hook.)
Fenzl
OsmorhizaberteroiDC.
Potent//la gracf/isDougl. ex Hook.
(LindL) Nesom var. spat/wiatum
Taraxacum officinale G.H. Weber ex
Wiggers
Thafictrum fend/er/Engelrn. ex Gray
TrifollumrepensL.
Viola aduncaSm.
Vi'da americana MuhI. ex Willd.
Mahoniarepens(Lindl.)G. Don
Spfraeabetu//fo/hPallas
Symphoricarposalbus(L.) Blake
Site 3
BromuscarinatusHook. &Arn.
C'arex con cinnoides Mackenzie
carexgeyeriBoott
Calainagrost/srubescensBuckl.
E/ymusglaucusBuckl.
Meilca bulbosa Geyer ex Porter &
Coult.
Poapratens/sL.
Tr/s'etum canescens Buckl.
0.0
3.3
6.7
0.0
-13.3
0.0
-6.7
0.0
13.3
-3.:3
-40.0
-3.3
135
Table 4. continued.
Control Overstory
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-13.3
0.0
0.0
0.0
0.0
0.0
6.7
-3.3
0.0
0.0
0.0
-6.7
0.0
0.0
6.7
13.3
0.0
0.0
3.3
0.0
-20.0
13.3
-13.3
3.3
0.0
0.0
3.3
3.3
0.0
0.0
-10.0
3.3
3.3
0.0
-13.3
-30.0
3.3
23.3
10.0
-6.7
20.0
0.0
-6.7
20.0
-6.7
0.0
-20.0
26.7
16.7
0.0
3.3
-3.3
10.0
0.0
-3.3
20.0
20.0
0.0
3.3
16.7
3.3
13.3
10.0
0.0
-56.7
6.7
10.0
0.0
0.0
20.0
0.0
-16.7
20.0
-10.0
40.0
3.3
-20.0
Prune/la vulgar/sL.
3.3
0.0
0.0
3.3
13.3
23.3
0.0
-10.0
33.3
-3.3
3.3
Ranunculusspp.
0.0
0.0
0.0
-3.3
Stellar/a /ongies Goldie
Symphyotr/chum spathu/atum
(Lindi.) Nesom var. spathulatum
Taraxacum off/cinale G.H. Weber ex
Wiggers
Thalictrum fendler/Engelm. ex Gray
TrifollumrepensL.
0.0
0.0
3.3
26.7
3.3
0.0
0.0
6.7
20.0
00
00
00
00
00
0
3.3
0.0
-16.7
-3.3
-6.7
Ach//lea mi//efollum L.
Anemone pioeriBritt. ex Rydb.
Antennaria rosea Greene
A rn/ca cordifo/ia Hook.
clayton/a peifoliata Donn ex WilId.
Euiybia consp/cua (Lindl.) Nesom
Fragaria spp.
Ga//urn spp
Hieracium a/b/forum Hook.
Iris misouriens/s N utt.
Lathyrus spp.
Lupinus spp.
Moehringiamacrophylla(Hook.)
Fenzl
Osmorh,'abe,terofDC.
Potent//la gradliDougl. ex Hook.
-16.7
-3.3
13.3
-16.7
3.3
0.0
13.3
20.0
-3.3
3.3
0.0
-3.3
-6.7
6.7
0.0
0.0
0.0
Viola adunca Sm.
10.0
3.3
3.3
10.0
10.0
V/cia americana Muhi. ex WilId.
20.0
20.0
3.3
0.0
16.7
Mahoniarepens(Lindl.) G. Don
-6.7
13.3
-3.3
-3.3
0.0
10.0
SpiraeabetullfollaPallas
-16.7
-10.0
0.0
16.7
6.7
3.3
Symphorfcarposa/bus(L.) Blake
-6.7
-13.3
-26.7
-3.3
0.0
-3.3
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exciosure, big
game grazing only; TEXC total exclosure, exclusion of cattle and big game grazing.
6.7
-3.3
3.3
-10.0
136
Table 5. The changes in species composition in a ponderosa pine forest from 1985 to
1994 using a 30 cm x 30 cm plot frame.
Control Overstory
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
0.0
-13.3
10.0
30.0
0.0
0.0
23.3
-3.3
40.0
0.0
3.3
33.3
-3.3
33.3
3.3
0.0
0.0
0.0
50.0
3.3
0.0
-3.3
10.0
43.3
3.3
0.0
-10.0
-26.7
56.7
0.0
0.0
0.0
-23.3
-33.3
-23.3
6.7
3.3
6.7
3.3
0.0
0.0
3.3
10.0
-6.7
3.3
6.7
0.0
10.0
-3.3
0.0
-36.7
0.0
0.0
3.3
10.0
0.0
10.0
6.7
-13.3
0.0
3.3
-26.7
0.0
-10.0
-26.7
3.3
-20.0
-16.7
-10.0
0.0
0.0
-6.7
0.0
36.7
-36.7
-6.7
-10.0
0.0
-10.0
-3.3
13.3
30.0
3.3
36.7
6.7
0.0
3.3
6.7
0.0
-16.7
33.3
6.7
-3.3
-26.7
3.3
-23.3
-6.7
-16.7
0.0
-6.7
-40.0
-36.7
-3.3
-3.3
0.0
0.0
-13.3
-10.0
0.0
0.0
6.7
-6.7
-3.3
0.0
0.0
0.0
-53.3
-16.7
3.3
-3.3
-6.7
-30.0
-23.3
6.7
0.0
0.0
-56.7
-6.7
3.3
-3.3
0.0
67
00
40 0
00
00
33 3
0.0
-23.3
-6.7
-43.3
-10.0
-13.3
-3.3
0.0
-20.0
0.0
6.7
0.0
-3.3
6.7
0.0
-13.3
-6.7
0.0
0.0
10.0
0.0
3.3
6.7
0.0
0.0
0.0
20.0
0.0
0.0
-6.7
0.0
16.7
0.0
10.0
0.0
-3.3
-6.7
0.0
0.0
16.7
3.3
13.3
0.0
-6.7
0.0
16.7
-3.3
0.0
33.3
30.0
0.0
0.0
6.7
0.0
26.7
-3.3
0.0
13.3
Site 1
Bromus carinatus Hook. &Arn.
CarexconcmnnoidesMackenzie
Carexgeyer/Boott
Calamagrostis rubescens BuckL
E/ymusg/aucusBuckl.
Meilca bu/bosa Geyer ex Porter &
Coult.
Poapratens/sL.
Tr/setumcanescensBuckL
Ac/ii/feam//IefoffumL.
Anemone pioeriBrftt. ex Rydb.
Antennar/aroseaGt-eene
Am/ca cordIfolia Hook.
aaytoni peifo//ata Donn ex WilId.
Euiyb/aconspicua(LindL) Nesom
Fragariaspp.
Ga//urn spp
Hieradum a/b/forum Hook.
Irfcm/ssour/ens/sNutt.
Lathyrusspp.
Lupinusspp.
Moehring/a macrophylla (Hook.)
Fenzl
OsmorhizaberteroiDC.
Potent//Ia grad//c Dougi. ex Hook.
Prune//a vulgar/cL.
Ranunculusspp.
Stellar/a /ongi;oes Goldie
Symphyotr/chum spathu/atufl?
(Lindl.) Nesom var. spathulatum
Taraxacum off/c/nate G.H. Weber ex
Wiggers
Thal/ctrum fendler/Engelm. ex Gray
TrifollumrepensL.
Viola adunca Sm.
V/c/a amer/cana Muhl. ex Wifld.
Mahoniarepens(Lindl.)G. Don
Sp/raea betuilfolia PaJias
Symphoricarposa/bus(L.) Blake
-20.0
10.0
0.0
3.3
10.0
0.0
0.0
-16.7
-13.3
0.0
0.0
-20.0
Q.0
13.3
0.0
0.0
3.3
0.0
30.0
-40.0
0.0
-6.7
3.3
-20.0
13.3
16.7
0.0
0.0
Site 2
Bromuscam/natusHook. &Arn.
Carex condnno/des Mackenzie
Carexgeyer/Boott
-13.3
0.0
0.0
3.3
137
Table 5. continued.
Contro' Overstory
Ca/amagrostisrubescensBuckl.
ElymusglaucusBuckl.
Meilca bu/bosa Geyer ex Porter &
Couft.
PoapratensiL.
TrtgetumcanescensBuckl.
Ach///eam/llefol/umL.
Anemonepier/Britt.exRydb.
Antennar/aroseaGreene
Am/ca cord/fo/la Hook.
aayton/a peifo/hta Donn ex WilId.
Euryb/a consp/cua (Lindi.) Nesom
FragarIaspp.
Gal/urn spp
H/erac/um a/b/forum Hook.
Lathyrusspp.
Lupinusspp.
Moehr/ng/a inacrop/iylla (Hook.)
Fenzl
OsrnorhizaberteroiDC.
Potent/I/a grad/is Dougl. ex Hook.
Prune//a vu/garIsL.
Ranunculusspp.
Ste//ar/a Iongioes GoIdie
Syrnphyotrichumspathulatum
(Lindi.) Nesom var.
spathu/aturn
Taraxacurnoffic/na/eG.H.
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
6.7
233
10.0
-6.7
0.0
6.7
30.0
16.7
23.3
10.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
10.0
-3.3
6.7
-3.3
3.3
3.3
30.0
-46.7
13.3
133
-16.7
-3.3
13.3
0.0
0.0
3.3
0.0
-10.0
0.0
0.0
0.0
-10.0
0.0
0.0
10.0
10.0
-6.7
3.3
3.3
6.7
0.0
0.0
13.3
0.0
16.7
13.3
16.7
0.0
0.0
13.3
6.7
26.7
33.3
-36.7
-10.0
0.0
0.0
0.0
-6.7
0.0
0.0
3.3
0.0
3.3
0.0
36.7
-26.7
0.0
-43.3
3.3
0.0
-63.3
6.7
-6.7
-6.7
-6.7
-3.3
-3.3
-3.3
0.0
0.0
0.0
0.0
3.0
0.0
0.0
0.0
0.0
0.0
-6.7
3.3
t).O
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
6.7
0.0
0.0
0.0
3.3
20.0
0.0
0.0
33.3
6.7
0.0
0.0
-3.3
0.0
13.3
-3.3
6.7
0.0
0.0
3.3
0.0
0.0
-3.3
-3.3
0.0
0.0
0.0
0.0
-3.3
0.0
-6.7
3.3
0.0
-3.3
0.0
-6.7
3.3
-6.7
3.3
0.0
-3.3
0.0
0.0
0.0
10.0
10.0
0.0
16.7
0.0
0.0
3.3
0.0
0.0
0.0
3.3
10.0
6.7
-46.7
13.3
26.7
20.0
20.0
-20.0
-10.0
0.0
10.0
13.3
0.0
0.0
-20.0
10.0
-3.3
Iris rnissouriensi Nutt.
Weberex
Wiggers
Tha/ictrurn fend/er/En geim. ex Gray
Trllb//urnrepensL.
Viola adunca Sm.
VidaarnericanaMuhl.exWilld.
Mahonia repens(Lindl.) G. Don
Sp/raeabetu//fo//aPallas
Syrnphoricarposalbus(L.) Blake
Thined Overstory
CEXC
0.0
-50.0
-6.7
-3.3
-16.7
10.0
16.7
0.0
10.0
-6.7
0.0
-6.7
0.0
16.7
-6]
Site 3
Bromus car/natus Hook. & Am.
Carexconcinno/desMackenzie
CarexgeyeriBoott
CalamagrostisrubescensBuckl.
E/yrnusg/aucusBuckl.
Me//ca bulbosa Geyer ex Porter &
Co u It.
PoapratensfsL.
Trisetum canescensBuckl.
0.0
0.0
3.3
-16.7
6.7
20.0
26,7
-43.3
6.7
3.3
0.0
0.0
-13.3
6.7
-3.3
-23.3
-6.7
-3.3
10.0
3.3
-40.0
-10.0
-13.3
-30.0
3.3
16.7
-20.0
-16.7
6.7
-16.7
-10.0
6.7
-6.7
0.0
-10.0
0.0
-30.0
-3.3
-36.7
10.0
-3.3
138
Table 5. continued.
:olOei:sto
Achi/lea millefo//um L.
Anemone pioeriBritt. ex Rydb.
Antennar/aroseaGreene
Am/ca cordifolia Hook.
C/aytoniapeifoliataDonnexWilld.
Euiyb/a conspicua(Lindl.) Nesom
Fragariaspp.
Galiumspp
Hierac/um albiflorum Hook.
IrimLs-souriensiNuft.
Lathyrusspp.
Lupinusspp.
Moehring/amacrophylla(Hook.)
Fenzl
OsmorhLzaberteroiDC.
Potent//Ia grad/is Dougl. exHook.
Prune/Ia vulgarLcL.
Ranuncuiusspp.
Stellar/a Iongi;oes Goldie
Symphyotrichum spathu/atum
(LindL) Nesom var. spathulatum
Taraxacum officina/e G.H. Weber ex
Wggers
Tha/ictrum fend/eriEngelm. ex Gray
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-6.7
0.0
0.0
-10.0
0.0
0.0
-16.7
-10.0
-3.3
-3.3
-10.0
-3.3
-16.7
0.0
0.0
13.3
-3.3
16.7
10.0
-56.7
3.3
3.3
-3.3
36.7
-6.7
0.0
-43.3
-16.7
13.3
20.0
6.7
3.3
-10.0
0.0
13.3
-10.0
13.3
-6.7
0.0
-26.7
23.3
16.7
0.0
3.3
0.0
3.3
33.3
-10.0
-3.3
-20.0
3.3
-26.7
0.0
-33.3
0.0
0.0
-33.3
0.0
0.0
-6.7
-6.7
0.0
0.0
0.0
-23.3
-20.0
-76.7
-40.0
-10.0
-6.7
0.0
6.7
0.0
0.0
-6.7
0.0
3.3
0.0
0.0
0.0
0.0
-10.0
0.0
0.0
-3.3
3.3
-3.3
0.0
13.3
-6.7
10.0
0.0
0.0
40.0
-6.7
3.3
0.0
0.0
13 3
67
00
20 0
30 0
00
0.0
3.3
-16.7
0.0
-3.3
3.3
0.0
0.0
00
-13.3
10.0
-3.3
0.0
0.0
10.0
-3.3
3.3
13.3
3.3
3.3
0.0
0.0
3.3
-6.7
0.0
0.0
0.0
Viola adunca Sm.
6.7
0.0
6.7
0.0
-6.7
3.3
V/c/a americana Muhi. ex WilId.
-6.7
3.3
-26.7
10.0
-3.3
0.0
I'lahoniarepens(Lindl.) G. Don
-13.3
10.0
3.3
-3.3
-16.7
-6.7
Sp/raea betu/ifolia Pallas
-6.7
0.0
-50.0
0.0
0.0
0.0
Symphor/carposa/bus(L.) Blake
-10.0
-3.3
6.7
-6.7
-10.0
0.0
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big
game grazing only; TEXC total exciosure, exclusion of cattle and big game grazing.
TrffollurnrepensL.
139
Table 6. The changes in species composition in a ponderosa pine forest from 1985 to
1994 using a 30 cm x 60 cm plot frame.
Control Ovestory
Thined Overstory
_____________________________CEXC
Site 1
Bromus carinatus Hook. &Arn.
arexconcinno/desMackenzie
carexge,veriBoott
Ca/amagrostisrubescensBuckl.
E/ymusg/aucusBuckl.
frlellcabu/bosa Geyer ex Porter &
Poapratens/sL.
Thetumcaaesceas5uckl.
Ac/i/I/eami/IefoliumL.
Anemone pieri Britt. ex Rydb.
Antennaria rosea Greene
Am/ca cordifolla Hook.
aataniapeifo/i'ataDonn ex WilId.
Eur/biaconspicua(Lindl.) Nesom
Fragariaspp.
Gal/urn spp
Hieradum a/b/forum Hook.
IrLcmigsauriensiNutt.
Lathyrusspp.
Lupinusspp.
fr/oehring/amacrophylla(Hook.)
Fenzl
Osmorh,a befteraiDC.
Potent//Ia grac///sDougl. ex Hook.
Prunella vulgarisL.
Ranunculusspp.
Stellar/a Iongioes Goldie
Syinphyotric/iuin spat/lu/atum
(Lindi.) Nesom var. spathulatum
Taraxacum officinale G.H. Weber ex
Wiggers
Thallctrum fendler/Engelm. ex Gray
Tr/follumrepensL.
V/olaaduncaSm.
V/c/a americana Muhi. ex WilId.
Mahoniarepens(Lindl.) G. Don
Spiraeabetu//follaPallas
S,vmphoricarposa/bus(L.) Blake
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
3.3
33.3
-3.3
26.7
3.3
0.0
-3.3
0.0
60.0
3.3
0.0
13.3
0.0
33.3
6.7
0.0
23.3
0.0
50.0
-3.3
13.3
43.3
-6.7
-26.7
46.7
0.0
0.0
3.3
-33.3
-33.3
-23.3
-6.7
-33.3
3.3
0.0
6.7
13.3
0.0
0.0
-16.7
-10.0
0.0
0.0
-20.0
3.3
-6.7
-20.0
-16.7
0.0
0.0
-3.3
0.0
50.0
-33.3
-6.7
-43.3
6.7
0.0
3.3
6.7
0.0
13.3
-6.7
0.0
-10.0
-10.0
3.3
0.0
3.3
3.3
0.0
3.3
20.0
0.0
-13.3
10.0
0.0
0.0
13.3
-3.3
26.7
3.3
40.0
10.0
0.0
0.0
6.7
0.0
-23.3
36.7
6.7
6.7
-33.3
0.0
0.0
-36.7
-3.3
0.0
0.0
0.0
0.0
33.3
-33.3
0.0
0.0
3.3
-16.7
26.7
-23.3
-13.3
-16.7
0.0
-6.7
-43.3
-53.3
-6.7
-3.3
0.0
0.0
-20.0
-10.0
-60.0
-30.0
0.0
-3.3
-6.7
-26.7
-23.3
10.0
0.0
-3.3
-66.7
-6.7
3.3
0.0
10.0
-6.7
-16.7
0.0
0.0
0.0
20 0
00
46 7
00
00
43 3
0.0
-33.3
-10.0
-36.7
-13.3
-20.0
-3.3
-3.3
-23.3
0.0
3.3
0.0
-3.3
6.7
-26.7
-23.3
-10.0
3.3
13.3
0.0
3.3
3.3
-3.3
-6.7
10.0
16.7
0.0
6.7
0.0
0.0
20.0
3.3
-13.3
0.0
3.3
13.3
0.0
0.0
0.0
20.0
0.0
30.0
3.3
0.0
36.7
43.3
0.0
0.0
6.7
0.0
30.0
-3.3
0.0
16.7
-16.7
0.0
0.0
-3.3
0.0
0.0
46.7
6.7
6.7
3.3
0.0
13.3
-3.3
0.0
-3.3
-10.0
0.0
-3.3
-23.3
10.0
10.0
-3.3
0.0
16.7
0.0
0.0
Site 2
Bromus carinatus Hook. & Am.
Carex conc/nnoides Mackenzie
CarexgeyeriBoott
140
Table 6. continued.
ContrciOverstory
Thned Overstory
CEXC
TEXC
GRAZED
10.0
cExc
3.3
6.7
TEXC
GRAZED
20.0
13.3
-6.7
0.0
46.7
0.0
33.3
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Poapratens/sL.
13.3
Trisetum canescens Buckl.
-26.7
-10.0
0.0
-3.3
0.0
-6.7
6.7
3.3
20.0
10.0
0.0
-3.3
0.0
-13.3
0.0
6.7
0.0
33.3
-46.7
-6.7
0.0
-40.0
6.7
10.0
-6.7
-6.7
3.3
3.3
-3.3
0.0
0.0
13.3
0.0
23.3
0.0
-56.7
0.0
13.3
10.0
10.0
0.0
0.0
13.3
10.0
-10.0
10.0
-13.3
0.0
-50.0
40.0
-46.7
-20.0
-6.7
0.0
0.0
3.3
0.0
0.0
6.7
0.0
-3.3
0.0
50.0
-36.7
30.0
-6.7
0.0
-6.7
0.0
20.0
-6.7
-6.7
-10.0
-6.7
-3.3
-6.7
-3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-3.3
-20.0
3.3
10.0
0.0
-3.3
0.0
0.0
0.0
-3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.3
20 0
33
00
43 3
67
00
0.0
-3.3
3.3
16.7
-3.3
6.7
0.0
0.0
3.3
0.0
-10.0
-6.7
-3.3
0.0
-3.3
-6.7
0.0
0.0
0.0
20.0
0.0
-3.3
6.7
0.0
0.0
0.0
-6.7
0.0
-6.7
6.7
-6.7
3.3
6.7
23.3
0.0
0.0
0.0
13.3
0.0
0.0
-3.3
0.0
20.0
0.0
26.7
3.3
0.0
-13.3
0.0
3.3
-13.3
0.0
23.3
0.0
26.7
-33.3
20.0
10.0
0.0
0.0
-13.3
3.3
-3.3
10.0
6.7
-43.3
10.0
30.0
26.7
6.7
-13.3
-20.0
10.0
26.7
23.3
-3.3
-13.3
6.7
-26.7
-6.7
-3.3
-13.3
-10.0
-16.7
6.7
3.3
-50.0
-16.7
-16.7
0.0
-53.3
3.3
-40.0
13.3
-10.0
Calamagrostis rubescens Buck!.
Elymus g/aucus Buck!.
Mefica bu/bosa Geyer ex Porter &
Coult.
AchiI/eamilefollumL.
Anemone pieriBrift. ex Rydb.
AntennariaroseaGreene
Am/ca cordi'fo/i Hook.
aaytoniapetfo/iataDonnexwilld.
Eurybia consp/cua (Lindi.) Nesom
Fragariaspp.
Ga//urn spp
Hieracium a/b/forum Hook.
Iris missour/ensisNutt.
Lathyrusspp.
Lupiriusspp.
Moehringia macrophy/la (Hook.)
Fenzl
Osmorhiza
berterof DC.
PotentillagradlisDougi. ex Hook.
Prune//a vu/garisL.
Ranunculusspp.
Ste//aria /ongioes GoIdie
Symphyotrfchum spat/lu/atum
(Lindi.) Nesom var. spathu/atum
Taraxacumofflcina/eG.H. Weberex
Wiggers
Tha/ictrum fend/er/En geim. ex Gray
TrifollumrepensL.
Viola adunca Sm.
V/cia americana Muhi. ex WilId.
Mahon/arepens(LindL) G. Don
Spiraeabetu//follaPallas
Symphoricarposafbus(L.) Blake
Site 3
BromuscarinatusHook. &Arn.
Carex conc/nno/des Mackenzie
CarexgeyeriBoott
Ca/amagrosticrubescensBuckl.
ElymusglaucusBuckl.
Meilca bulbosa Geyer ex Porter &
Coult.
PoapratensisL.
Triseturn canescens
Buckl.
10.0
3.3
0.0
0.0
-36.7
C0
13.3
0.0
10.0
-6.7
0.0
0.0
16.7
10.0
141
Table 6. continued.
Caro'Overt
Thired Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-13.3
0.0
0.0
-3.3
0.0
0.0
-33.3
0.0
0.0
-33.3
0.0
0.0
-3.3
-6.7
3.3
0.0
6.7
-26.7
6.7
16.7
10.0
13.3
10.0
3.3
-46.7
-6.7
0.0
20.0
13.3
0.0
3.3
0.0
3.3
43.3
-16.7
-16.7
-3.3
0.0
-10.0
-3.3
-26.7
0.0
0.0
0.0
0.0
0.0
-13.3
-6.7
-23.3
6.7
-23.3
-3.3
0.0
-33.3
Potent/I/a gradllLsDougl. ex Hook.
Prune//a vulgarLsL.
-6.7
0.0
6.7
Ranuncuiusspp.
Stellarla longioes Goldie
Symphyotrichum spathulatum
(Lindi.) Nesom var. spathu/atum
Taraxacum offic/nale G.H. Weber ex
Wiggers
Thallctrum fend/er/Engelm. ex Gray
Achileami/efollumL.
Anemone pieriBritt. ex Rydb.
Antennar/aroseaGreene
Am/ca cord/fo//a Hook.
Clayton/a petfol/ata Donn ex WilId.
Euiyb/a consp/cua (LindL) Nesom
Fragar/aspp.
Gallumspp
Hieradum albiflorum Hook.
Iris misouriensiNutt.
Lathyrusspp.
Lupfnusspp.
Moehringia macrophy/la (Hook.)
Fenzl
OsmorhLza berteroiDC.
3.3
6.7
-3.3
40.0
-13.3
0.0
3.3
13.3
-20.0
-40.0
23.3
-6.7
0.0
-16.7
26.7
-6.7
0.0
6.7
-20.0
-80.0
-43.3
-23.3
-13.3
0.0
3.3
10.0
0.0
-10.0
0.0
0.0
0.0
0.0
0.0
0.0
6.7
-3.3
13.3
-3.3
0.0
16.7
-13.3
16.7
0.0
0.0
16.7
-20.0
3.3
0.0
0.0
16 7
10 0
00
26 7
30 0
00
0.0
0.0
3.3
-167
0.0
-13.3
3.3
13.3
3.3
3.3
-3.3
3.3
30.0
0.0
0.0
0.0
0.0
-6.7
0.0
3.3
Viola aduncaSm.
-6.7
6.7
3.3
-6.7
13.3
6.7
V/cia americana Muhi. ex WilId.
3.3
-40.0
-10.0
0.0
-3.3
0.0
Mahoniarepens(Lindl.)G. Don
3.3
0.0
0.0
3.3
0.0
-3.3
Sp/raeabetuifollaPalias
-56.7
0.0
-13.3
0.0
0.0
0.0
Symphoricarposa/bus(L.) Blake
3.3
-23.3
3.3
3.3
16.7
OM
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big
game grazing only; TEXC total exclosure, exclusion of cattle and big game grazing.
Tr/follumrepensL.
142
Table 7. The changes in species composition in a ponderosa pine forest from 1985 to
1997 usincj a 30 cm x 30 cm olot frame.
Thined Overstory
ContrtOverstory
CEXCTEXC
Site 1
SromuscarinatusHook. &Arn.
GRAZED
CEXC
TEXC
GRAZED
0.0
0.0
-43.3
-3.3
23.3
0.0
0.0
-33.3
-3.3
-10.0
0.0
0.0
-30.0
-13.3
33.3
3.3
3.0
-13.3
-3.3
26.7
10.0
-13.3
6.7
0.0
0.0
6.7
-3.3
40.0
0.0
0.0
0.0
-23.3
-33.3
-30.0
PoapratensiL.
13.3
Tr/setumcanescensBuckl.
Achillea millefo/lum L.
Anemone pioeriBritt. ex Rydb.
-16.7
-6.7
6.7
-20.0
-6.7
6.7
0.0
-6.7
6.7
6.7
-40.0
-16.7
-13,3
-10.0
0.0
0.0
-3.3
16.7
0.0
0.0
20.0
3.3
-16.7
-10.0
6.7
0.0
0.0
-3.3
23.3
-33.3
6.7
10.0
0.0
-3.3
0.0
0.0
3.3
-50.0
0.0
0.0
-16.7
10.0
6.7
-33.3
3.3
3.3
0.0
0.0
0.0
0.0
-40.0
-10.0
0.0
0.0
-10.0
-3.3
33.3
-13.3
30.0
0.0
0.0
3.3
0.0
3.3
-30.0
0.0
6.7
-3.3
-46.7
0.0
-20.0
-6.7
0.0
0.0
-6.7
-40.0
Ste//ari'a/ongi,esGoIdie
10.0
-3.3
-13.3
0.0
0.0
0.0
-16.7
0.0
0.0
0.0
6.7
-56.7
-10.0
Ranuncu/usspp.
-43.3
0.0
-3.3
0.0
0.0
-3.3
-3.3
-36.7
-13.3
-3.3
0.0
6.7
-66.7
-6.7
0.0
-3.3
3.3
10.0
20.0
13.3
10.0
20.0
36.7
-6.7
-23.3
-10.0
-36.7
-16.7
-13.3
-3.3
0.0
-16.7
0.0
6.7
0.0
-6.7
0.0
0.0
-20.0
-13.3
-3.3
0.0
36.7
0.0
0.0
-10.0
0.0
0,0
0.0
23.3
16.7
10.0
0.0
13.3
0.0
-3.3
-23.3
13.3
23.3
0.0
13.3
0.0
0.0
0.0
0.0
0.0
0.0
13.3
0.0
6.7
6.7
0.0
13.3
40.0
0.0
40.0
0.0
13.3
13.3
-23.3
0.0
3.3
-6.7
13.3
16.7
Carexconcinnoides Mackenzie
CarexgeyeriBoott
Ca/amagrostisrubescensBuckl.
ElymusglaucusBuckl.
Me//ca bu/bosa
Geyer ex Porter &
Coult.
Antennaria rosea Greene
Am/ca cordifo//a Hook.
C/ayton/apeifo//ataDonnexWilld.
Euiybiaconspicua(LindL) Nesom
Fragar/aspp.
Gallumspp
Hieracium a/b/forum Hook.
IrLs'misouriensisNutt.
Lathyrusspp.
Lupinusspp.
Moehri'ngi'a macrophy/la (Hook.)
Fenzl
Osmorhi'aberteroiDC.
Potentilagraci/isDougi. ex Hook.
Prune/la vu/gar/sL.
Symp/iyotrichumspathu/atum
(Lindi.) Nesom var. spathu/atuni
Taraxacum officinale G.H. Weber ex
Wiggers
Tha/ictrum fend/er/En geim. ex Gray
TrifoliumrepensL.
Vio/aaduncaSm.
V/c/a americana Muhi. ex WilId.
Mahoni'arepens(Lindl.) G. Don
Spiraeabetu/ifollaPallas
Symphoricarposa/bus(L.) Blake
13.3
0.0
0.0
3.3
0.0
-20.0
-3.3
-3.3
0.0
0.0
0.0
0.0
00
.3
-i0
10.0
3.3
-10.0
Site 2
Bromus carinatus Hook. &Arn.
CarexconcinnoidesMackenzie
CarexgeyeriBoott
0.0
143
Table_7._continued.
Thined Overstory
CcmtroiOverstory
calamagrostisrubescensBuckL
Elymusg/aucusBuckl.
Me//ca bu/bosa Geyer ex Porter &
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
0.0
26.7
0.0
3.3
0.0
3.3
-3.3
30.0
0.0
23.3
0.0
o.o
0.0
0.0
0.0
50.0
-16.7
10.0
0.0
6.7
-20.0
0.0
0.0
13.3
16.7
-6.7
0.0
-36.7
6.7
13.3
13.3
3.3
0.0
10.0
0.0
13.3
Coult.
PoapratensLsL.
TrisetumcanescensBuckl.
Achll/eami/efollumL.
Anemone pioeriBritt. ex Rydb.
AntennariaroseaGreene
Am/ca cordifolia Hook.
aayton/aperfo/iataDonnexwilld.
consp/cua (LindL) Nesom
Fragar/aspp.
Gallumspp
Hieracium a/b/forum Hook.
Eu/yb/a
Ir/smicsouriensiNutt.
Lathyrusspp.
.Lup/riusspp.
frloehringiamacrophy/la(Hook.)
Fenzl
Osmor/iLzaberteroiDC.
Potent///agrac//i.DougI. ex Hook.
Prune//a vulgarisL.
Ranuncu/usspp.
Stellar/a /ongioesGoldie
Svmp/iyotrichum spathulatum
(Lindl.) Nesom var. spat/lu/atum
Taraxacumoff,dna/eG.H.
-3.3
-16.7
-3.3
0.0
6.7
16.7
0.0
-6.7
0.0
0.0
0.0
0.0
0.0
-3.3
0.0
-3.3
0.0
63.3
-30.0
0.0
3.3
-10.0
0.0
3.3
20.0
3.3
0.0
0.0
0.0
-30.0
0.0
-13.3
0.0
00
-3.3
3.3
0.0
0.0
0.0
0,0
3.3
0.0
0.0
0.0
0.3
0.3
0.0
0.0
0.0
0.0
3.3
16 7
33
10.0
3.3
TrifollurnrepensL.
Viola adunca Sm.
V/cia americana Muhi. ex WilId.
Mahon/arepens(Lindl.) G. Don
Sp/raea betuilfo/la Pallas
Symphoricarposa/bus(L.) Blake
Site 3
Bromuscar/natusHook. &Arn.
Carwc' concinno/des Mackenzie
CarexgeyeriBoott
Ca/amagrost/srubescensBuckl.
E/ymusglaucusBuckl.
Me/fcabu/bosaGeyerexPorter&
Coult.
-3.3
0.0
3.3
0.0
-16.7
0.0
0.0
0.0
-33.3
6.7
0.0
0.0
-13.3
0.0
-3.3
0.0
-56.7
-6.7
-3.3
-3.3
0.0
0.0
0.0
0.0
3.3
0.0
0.0
0.0
0.0
3.3
0.0
0.0
0.0
0.0
10.0
33
33 3
13 3
13 3
-3.3
-3.3
-3.3
0.0
0.0
0.0
0.0
0.0
0.0
-13.3
-3.3
-16.7
0.0
-3.3
-3.3
0.0
-16.7
0.0
0.0
0.0
-3.3
0.0
0.0
-13.3
0.0
6.7
0.0
-6.7
3.3
3.3
-3.3
0.0
6.7
0.0
0.0
0.0
0.0
3.3
3.3
3.3
0.0
0.0
0.0
0.0
-6.7
0.0
10.0
0.0
-26.7
-16.7
20.0
10.0
6.7
6.7
0.0
-3.3
0.0
0.0
16.7
-3.3
-3.3
0.0
3.3
-26.7
3.3
26.7
0.0
6.7
-60.0
6.7
-3.3
0.0
3.3
-20.0
-10.0
-13.3
3.3
3.3
W.
-6.7
10.0
-6.7
-13.3
-30.0
-6.7
-36.7
3.3
-16.7
6.7
-33.3
43.3
-33.3
50.0
-50.0
0.0
-13.3
0.0
0.0
0.0
-33.3
Weberex
Wiggers
Thalictrum fendleriEngelm. ex Gray
Poapratens/sL.
Trisetum canescensBuckl.
10.0
-6.7
13.3
0.0
-6.7
16.7
L
144
Table 7. continued.
COn oOv
Ach//lea millefollum L.
Anemone pi;oer/ Britt. ex Rydb.
AntennariaroseaGreene
Am/ca cord/folia Hook.
C'/aytoni'apeifo/iataDonnexWiild.
Eurybia consp/cua (Lindi.) Nesom
Fragar/aspp.
Ga//umspp
H/ct-ac/urn albiflorum Hook.
Iris mLcsouriensisNutt.
Lath,yrusspp.
Lupinusspp.
Moehring/a macrophylla (Hook.)
Fenzl
OsrnorhLza berteroiDC.
Potent//Ia grac//isDougl. ex Hook.
Prunella vu/garisL.
Ranuricu/usspp.
Stellar/a /ongioesGoldie
Symphyotrichum spathu/aturn
(LindL) Nesom var. spathulatum
Taraxacumoff/cinalec3.H. Weber cx
Wiggers
Tha/ictrum fend/eriEngelm. ex Gray
Tr/fo/iumrepensL.
Viola aduncaSm.
V/c/a americana Muhi. ex Willd.
Mahonia repens (Lindi.) G. Don
Sp/raeabetu//fo/iaPalIas
Thined Oversto
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
40.0
-67
-10.0
10.0
0.0
-6.7
0.0
13.3
-16.7
23.3
00
1.3
0.0
0.0
-20.0
-16.7
-6.7
-3.3
-3.3
-3.3
0.0
0.0
-23.3
-13.3
-16.7
0.0
-13.3
0.0
6.7
6.7
0.0
0.0
-10.0
-6.7
-6.7
0.0
13.3
-13.3
3.3
23.3
3.3
3.3
-13.3
0.0
0.0
-26.7
23.3
-3.3
0.0
-23.3
6.7
20.0
6.7
0.0
-6.7
0.0
0.0
-20.0
-6.7
-3.3
0.0
3.3
10.0
0.0
-23.3
3.3
-66.7
-30.0
10.0
-16.7
6.7
6.7
0.0
0.0
-10.0
0.0
0.0
0.0
0.0
-13.3
0.0
0.0
3.3
0.0
16.7
-3.3
-3.3
13.3
-16.7
-3.3
0.0
0.0
13.3
-10.0
3.3
-3.3
0.0
3.3
00
00
3
26 7
16 7
23 3
0.0
3.3
0.0
-10.0
-3.3
-10.0
00
40.0
3.3
-36.7
0.0
0.0
-6.7
30.0
-13.3
0.0
-30.0
-3.3
0.0
0.0
-3.3
0.0
-6.7
0.0
0.0
-6,7
-3.3
-6.7
-6.7
0.0
6.7
-6.7
-26.7
10.0
0.0
20.0
-23.3
10.0
6.7
-3.3
-20.0
-10.0
6.7
23.3
0.0
3.3
0.0
0.0
Symphor/carposa/bus(L.) Blake
-20.0
10.0
-20.0 -10.0
-16.7
3.3
Herbivory treatments: Graze cathe and big game grazing; CEXC cattle exciosure, big
game grazing only; TEXC total exclosure, exclusion of cattle and big game grazing.
3,3
-3.3
0.0
3.3
145
Table 8. The changes in species composition in a pohcerosa pine forest from 1985 to
1997 usina a 30 cm x 60 cm clot frarne
ContreOverj
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
3,3
0.0
-10.0
-3.3
0.0
0.0
10.0
-3.3
23.3
0.0
0.0
23.3
0.0
0.0
3.3
-13.3
13.3
-43.3
-3.3
40.0
0.0
0.0
-10.0
-6.7
-13.3
0.0
0.0
-33.3
-10.0
23.3
0.0
0.0
0.0
-33.3
-26.7
-33.3
3.3
-30.0
-10.0
3.3
-13.3
6.7
23.3
0.0
-3.3
30.0
3.3
-16.7
-6.7
-3.3
0.0
6.7
-3.3
20.0
-36.7
0.0
-3.3
0.0
3.3
-23.3
-16.7
6.7
0.0
-6.7
10.0
6.7
-36.7
-13.3
-13.3
0.0
-10.0
-?0.0
10.0
10.0
-46.7
-6.7
10.0
0.0
-3.3
0.0
0.0
-33.3
-13.3
3.3
3.3
-3.3
3.3
23.3
-10.0
33.3
0.0
0.0
0.0
0.0
6.7
-26.7
10.0
13.3
-3.3
-50.0
6.7
-13.3
-13.3
0.0
0.0
-6.7
-40.0
-63.3
0.0
-3.3
0.0
10.0
-3.3
-10.0
0.0
0.0
0.0
-20.0
-6.7
0.0
0.0
-40.0
-6.7
-3,3
0.0
6.7
'-80.0
13.3
-63.3
-23.3
-3.3
-3.3
-3.3
13.3
26.7
23.3
30.0
26.7
50.0
-33.3
-10.0
-40.0
-16.7
-16.7
0.0
0.0
0.0
-10.0
0.0
-3.3
0.0
0.0
-26.7
30.0
16.7
0.0
6.7
0.0
3.3
0.0
0.0
10.0
0.0
6.7
0.0
16.7
-13.3
0.0
6.7
-10.0
0.0
13.3
6.7
Site I
Bromuscar/natusHook.&Arn.
Carex conc/ano/des Mackenzie
CarexgeyeriBoott
C'a/amagrostLrubescensBuckL
E/ymusg/aucusBuckl.
Me//ca bu/bosa Geyer ex Porter &
Co u It.
Pea pratens/sL.
Thsetuci canescens Buckl.
Ac/i//lea m//lefoliumL.
Anemone pieriBritt. ex Rydb.
Antennarla rosea Greene
Am/ca cord/fe/ía Hook.
Cla,Vton/apemfo/i'taDannexWilld.
Euiyb/a consp/cua (LindL) Nesom
Fragariaspp.
Galiumspp
Hierac/um a/b/forum Hook.
Iris m/ssourfensisNutt.
Lathyrusspp.
Lupinusspp.
Moehring/amacrophy//a(Hook.)
Fenzl
Osmor/ilza bertero/DC.
Potenti/lagrac///sDougl. ex Hook.
Prune//a vu/garísL.
Ranunculusspp.
Stel/ari'a/ongioesGoldie
Symphyotr/chumspathulatum
(Lindi.) Nesom var. spathu/atum
Taraxacum offic/nale G.H. Weber ex
Wiggers
Tha/ictruin fendleriEngelm. ex Gray
Tr/follumrepensL.
V/o/aaduncaSm.
V/c/a americana MuhI. ex Wilid.
Mahoniarepens(Lindl.) G. Don
Sp/raeabetu//foliaPallas
Symphor/carposa/bus(L.) Blake
Site 2
Bromuscar/natusHook. &Arn.
C'arexconc/nnoidesMackenzie
CarexgeveriBoott
13.3
0.0
0.0
3.3
0.0
-16.7
-10.0
-33
-3.3
-3.3
-16.7
0.0
0.0
-3.3
00
6.7
0.0
0.0
-13.3
16.7
3.3
0.0
26.7
0.0
13.3
6.7
0.0
26.7
43.3
0.0
46.7
6.7
00
-26.7
-30.0
-10.0
0.0
13.3
13.3
0.0
-6.7
0.0
0.0
6.7
-46.7
0.0
0.0
-23.3
16.7
3.3
-6.7
0.0
-3.3
10.0
146
Table 8. continued.
cbQLc.)jto
Ca/amagrostLcrubescensBuckL
E/,vmusg/aucusBuckl.
Me//ca bu/bosa Geyer ex Porter &
CEXC
GRAZED
CEXC
TEXC
GRAZED
3.3
267
0.0
0.0
3.3
-3.3
13.3
0.0
13.3
0.0
3.3
0.0
0.0
0.0
:33
-3.3
i33
6.7
3.3
-3.3
6.7
0.0
0.0
0.0
13.3
0.0
3.3
0.0
-23.3
-6.7
26.7
0.0
3.3
10.0
0.0
10.0
0.0
0.0
-6.7
0.0
0.0
0.0
-40.0
6.7
.0
0)
CouJt.
Poapratens/sL.
Tr/setumcanescensBuckl.
Ac/i///eam//lefa//umL.
Anemone piveriBritt. ex Rydb.
Antennar/aroseaGreene
Am/ca cord/lot/a Hook.
clayton/a peifoliata Donn ex WilId.
Eu/yb/a conspicua (Lindi.) Nesom
Fragar/aspp.
Ga//urn spp
Hieracium a/b/forum Hook.
Jr/sm/s-sour/ens/s Nutt.
Lath,vrusspp.
Lup/nusspp.
Moehringia macrophyfla (Hook.)
Osmorhi-abefterofDC.
(Lindi.) Nesom var. spathu/atum
Taraxacum offic/naleG.H. Weber ex
Wiggers
Tha/ictrum fend/eriEngelm. ex Gray
TrifollumrepensL.
Viola aduncaSm.
V/cia americana Muhi. ex Wild.
Ma/ion/a repens(Lir.dl.) G. Don
Sp/raea betuli, b//a PaIIas
Symphor/carposa/bus(L.) Blake
-40.0
-10.0
33
67
0.0
6.7
0.0
3,3
0.0
-20.0
0.0
-3.3
0.0
-30.0
16.7
0.0
0.0
3.3
0.0
0.0
0.0
0.0
-6.7
0.0
73.3
-40.0
0.0
10.0
26.7
10.0
0.0
-10.0
0.0
-30.0
0.0
-10.0
0.0
56.7
-6.7
20.0
0.0
6.7
-10.0
0.0
0.0
10.0
16.7
0.0
0.0
-40.0
16.7
-6.7
3.3
0.0
-6.7
10.0
0,0
0.0
0.0
0.0
10.0
0.0
0.0
0.0
0.0
-16.7
0.0
0.0
0,U
-3.3
0.0
0.0
0.0
6.7
6.7
0.0
3.3
0.0
0.0
3.3
0.0
0.0
0.0
0.0
20.0
20:0
3.3
3.3
40.0
20.0
26.7
-3.3
-3.3
0.0
3.3
3.3
0.0
-3,3
-6.7
0.0
-6.7
3.3
6.7
0.0
3.3
16.7
0.0
-3.3
0.0
0.0
-3.3
0.0
6.7
16.7
0.0
0.0
3.3
0.0
16.7
3.3
16.7
0.0
0.0
-6.7
0.0
-6.7
0.0
-3.3
0.0
3.3
13.3
-3.3
0.0
0.0
6.7
-13.3
-3.3
33.3
6.7
6.7
-53.3
3.3
-3.3
0.0
10.0
-16.7
-16.7
-13.3
13.3
-16.7
Fenzl
Potent//la graci//s- Dougl. ex Hook.
Prune//a vu/gar/sL.
Ranuncu/usspp.
Stellar/a Iongioes Goldie
Symp/lyotr/c/7um spat/lu/atum
Thined Overstory
TEXC
0.0
0.0
3,3
0.0
-6.7
-6.7
-16.7
O.
00
-6.7
0.0
-16.7
Sfte3
Bromus carinatus Hook. &Arn.
carexconc/nnoidesMackenzie
W0
carexgeyeriBoott
-10.0
-13.3
23.3
13.3
6.
-6.7
13.3
-3.3
0.0
6.7
3.3
-13.3
0.0
-13.3
-3.3
-36.7
-6.7
-43.3
-3.3
-20.0
3.3
-56.7
46.7
-33.3
46.7
-56.7
ca/amagrostisrubescensBuckl.
E/ymusg/aucusBuckl.
Me//ca bulbosa Geyer ex Porter &
Coult.
Poe pratens/sL.
Trisetum canescens Buckl.
0.0
147
Table 8. continued.
mined Oversto'
Achillea mi//efolium L.
Anemone pioerf Britt. ex Rydb.
Antennaria rosea Greene
A rn/ca cord/fo//a Hook.
clayton/a perfol/ata Donn ex WiUd.
Euryb/a consp/cua (Lindi.) Nesom
Fragaria spp.
Ga//urn spp
Hierac/um a/b/forum Hook.
Zri m/ssouriens/s N utt.
Lathyrus spp.
Lupinus spp.
Moeliringia macropfiylla(Hook.)
Fenzl
Osmor/ii'a berteroi DC.
Potent/I/a grac//is Dougl. ex Hook.
Prune//a vulgarlsL.
.Ranuncu/usspp.
Ste/far/a longoes Goldie
Symphyotrichum spathufatum
(Lindl.) Nesom var. spathu/atuni
Taraxacum offic/nale G.H. Weber ex
Wiggers
Thafictrum fend/er/Engelm. ex Gray
Tr/foliumrepensL.
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-16.7
-10.0
20.0
-20.0
26.7
10.0
6.7
0.0
0.0
-13.3
-3.3
3.3
6.7
-13.3
0.0
O0
6.7
0.0
0.0
-30.0
-10.0
-20.0
6.7
-13.3
3.3
16.7
6.7
6.7
3.3
0.0
0.0
0.0
-20.0
36.7
-10.0
3.3
-26.7
16.7
20.0
10.0
3.3
0.0
0.0
0.0
-6.7
0.0
3.3
0.0
10.0
13.3
0.0
-33.3
13.3
-60.0
-33.3
3.3
-20.0
6.7
-16.7
16.7
-3.3
-3.3
20.0
-26.7
-10.0
10.0
0.0
0.0
-20.0
0.0
-10.0
0.0
3.3
-6.7
0.0
0.0
0.0
0.0
0.0
0.0
16.7
-30.0
-6.7
-3.3
0.0
6.7
00
00
33
36 7
26 7
33 3
3.3
3.3
-10.0
0.0
-20.0
16.7
0.0
0.0
0.0
-10.0
-16.7
3.3
3.3
10.0
-133
0.0
3.3
-20.0
-30.0
0.0
0.0
-13.3
30.0
-10.0
0.0
-23.3
0.0
-3.3
0.0
0.0
3.3
-3.3
3.3
-3.3
-6.7
0.0
3.3
Viola adunca Sm.
0.0
-6.7
-3.3
-13.3
-3.3
0.0
V/c/a americana Muhi. ex Wifid.
40.0
3.3
-10.0
6.7
3.3
26.7
Mahoni'a repens (Lindl.) G. Don
-23.3
3.3
10.0
3.3
-20.0
0.0
Sp/raea betul/fol/a Pallas
0.0
0.0
6.7
30.0
0.0
0.0
Symphor/carposalbus(L.) Blake
-?3.3 -16.7
-23.3
16.7
-23.3
16.7
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exciosure, big
game grazing only; TEXC total exciosure, exclusion of cattle and big game grazing.
10.0
Table 9. The changes in species composition in a ponderosa pine forest from 1985 to
2005 using a 30 cm x30 cm ppfrme._
citd. CvesjQry
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
BromuscarinatusHook. &Arn.
thO
00
Carex concinnoides Mackenzie
0.0
16.7
-23.3
36.7
43.3
0.0
36.7
-3.3
-26.7
36.7
0.0
0.0
-20.0
-3.3
3.3
0.0
16.7
-3.3
23.3
0.0
-33.3
26.7
0.0
26.7
-26.7
33.3
00
3.3
43.3
-23.3
-33.3
-30.0
60.0
-23.3
0.0
13.3
0.0
-13.3
-10.0
0.0
3.3
0.0
-3.3
0.0
23.3
-46.7
26.7
-23.3
-16.7
16.7
0.0
-6.7
-16.7
0.0
-6.7
0.0
3.3
-13.3
-6.7
33.3
-36.7
16.7
3.3
0.0
-3.3
-36.7
0.0
0.0
-3.3
-36.7
-3.3
0.0
0.0
0.0
-3.3
0.0
-3.3
00
-13.3
6.7
0.0
-56.7
3.3
10.0
-167
-10.0
-10.0
-13.3
0.0
0.0
0.0
-6.7
0.0
3.3
0.0
6.7
-3.3
-16.7
-23.3
-10.0
-6.7
-36.7
33.3
-13.3
36.7
0.0
0.0
3.3
40.0
0.0
0.0
0.0
-6.7
3.3
-40.0
-70.0
o.o
o.d
0.0
0.0
0.0
0.0
3.3
-3.3
3.3
0.0
16.7
-10.0
0.0
0.0
0.0
20.0
0.0
0.0
6.7
0.0
30.0
3.3
0.0
-3.3
-6.7
33.3
-10.0
-3.3
13.3
0.0
20.0
-6.7
0.0
0.0
0.0
33.3
-333
-167
-233
67
-367
-233
-13.3
-26.7
-10.0
-46.7
-20.0
-10.0
-3.3
0.0
6.7
0.0
6.7
0.0
3.3
0.0
0.0
0.0
-23.3
10.0
0.0
23.3
0.0
0.0
-3.3
0.0
3.3
0.0
26.7
0.0
0.0
3.3
0.0
40.0
0.0
16.7
0.0
-3.3
-30.0
-3.3
36.7
0.0
16.7
0.0
0.0
6.7
0.0
16.7
0.0
0.0
3.3
16.7
0.0
0.0
53.3
0.0
6.7
26.7
0.0
0.0
3.3
0.0
0.0
10.0
6.7
0.0
Site I
Carexgeyer/Boott
C'alamagrost/srubescensBuckl.
E/ymusglaucusBuckl.
Mellcabu/bosaGeyerexPorter&
Coult.
PoapratensiL.
TrLcetumcanescensBuckl.
Achulleami/Iefo/iumL.
Anemone pioeriBrift. ex Rydb.
Antennaria rosea Greene
Am/ca cord/fo/la Hook.
aaytoniiperfa//ataDonri ex WiHd.
Eu/yb/a consp/cua(LindL) Nesom
Fragariaspp.
Gallumspp
Hieracium albifloruni Hook.
Iris' mis'sour/ensisNutt.
Lathyrusspp.
Lupinusspp.
Moehring/a macrophylla (Hook.)
Fenzl
Osmorh/za berteroiDC.
Potent//Ia grad/is' Dougl. ex Hook.
Prune//a vu/gar/sL.
Ranuncu/usspp.
Stellar/a /ongioes Goldie
Symphyotrichumspathu/atum
(Lindl.) Nesorn var. spathulatum
Thraxacum offidnale G.H. Weber ex
Wiggers
Thafictrum fend/er/Engelm. ex Gray
Tr/fo/iumrepensL.
Viola adunca Sm.
V/c/a americana Muhl. ex WilId.
Mahoni'a repens(Lindl.) G. Don
Sp/raeabetu//fol/aPaHas
Symp/ior/carposa/bus(L.) Blake
.0
13.3
10.0
Site 2
BromuscarinatusHook. &Arn,
carexconc/nnoidesMackenzie
C'arexgeyer/Boott
13.3
149
Table_9._continued.
nOLQvsto
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
Calamagrost/srubescensBuckl.
E/ymusg/aucusBuckl.
MeIica/ulbosa Geyer ex Porter &
-13.3
0.0
-13.3
0.0
3.3
0.0
-10.0
-3.3
-23.3
0.0
13.3
0.0
0.0
0.0
0.0
0.0
0.0
PoapratensiL.
-3.3
-20.0
-10.0
0.0
3.3
-3.3
0.0
0.0
-13.3
0.0
-36.7
23.3
-6.7
0.0
-367
-10.0
3.3
0.0
0.0
-10.0
0.0
0.0
-3.3
0.0
10.0
-23.3
-6.7
0.0
-6.7
-10.0
-3.3
0.0
0.0
26.7
0.0
0.0
-23.3
0.0
-6.7
0.0
-6.7
0.0
-10.0
-40.0
-13.3
-26.7
-3.3
0.0
-23.3
20.0
0.0
-3.3
0.0
-66.7
0.0
-3.3
-3.3
-23.3
0.0
0.0
-3.3
-3.3
-6.7
3.3
0.0
0.0
10.0
0.0
0.0
-6.7
0.0
-53.3
-6.7
-3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Ranunculusspp.
0.0
0.0
0.0
0.0
Stellar/a /ongioesGoldie
Symphyotrichum spathulatum
(Lindl.) Nesom var. spathu/atum
Taraxacum officinale G.H. Weber ex
Wiggers
Thafictrum fend/er/Engelm. ex Gray
15.7
32
0.0
36.7
0.0
0.0
0.0
0.0
16.7
0.0
0.0
0.0
0.0
20.0
-13 3
33
00
67
67
-6 7
-3.3
-6.7
-6.7
-3.3
3.3
0.0
0.0
0.0
0.0
-16.7
-3.3
-16.7
0.0
-3.3
-3.3
0.0
0.0
-3.3
0.0
0.0
-10.0
0.0
23.3
0.0
0.0
-6.7
0.0
0.0
23.3
0.0
0.0
-3.3
0.0
0.0
0.0
26.7
0.0
0.0
0.0
0.0
0.0
0.0
-3.3
10.0
13.3
0.0
-30.0
-33.3
3.3
0.0
-43.3
-23.3
-6.7
0.0
0.0
-23.3
-10.0
-3.3
0.0
0.0
-13.3
-6.7
36.7
10.0
0.0
-3.3
-10.0
0.0
30,0
0.0
-20.0
-20.0
3.3
-10.0
10.0
-3.3
-6.7
0.0
-10.0
56.7
-30.0
40.0
-43.3
36.7
-36.7
30.0
-33.3
33.3
-43.3
43.3
-50.0
Trisetum canescensBuckl.
Ac/iilleamll/efo/iumL.
Anemone pieriBritt. ex Rydb.
Antennar/aroseaGreene
Am/ca cord/folia Hook.
aaytoniapeifo/iataDonn exWilld.
Euiybiaconspicua(Lindl.) Nesom
Fragar/aspp.
Ga//urn spp
Hieracium a/b/forum Hook.
ir/mfssour/ensisNutj.
Lathyrusspp.
Lupinusspp.
frloehringiamacrophy/la(Hook.)
Fenzl
OsmorhtabefteroiDC.
Potent//la grac//iDougl. ex Hook.
Prunella vu/garisL.
Th'fo//urnrepensL.
Viola aduncaSrn.
Wc/a americana MuhI. ex Willd.
Mahonia repens (Lindi.) G. Don
Spiraeabetullfo//aPallas
Symphor/carposa/bus(L.) Blake
Site 3
BromuscarinatusHook.&Arn.
Carex concinnoides Mackenzie
CarexgeyeriBoott
Ca/amagrostisrubescensBuckl.
Efymnusg/aucussuckl.
Me//ca bu/bosaGeyer ex Porter &
Couft.
Poapratens/sL.
Th-etum canescensBuckl.
3.3
0.0
16.7
13.3
0.0
6.7
-6.7
0.0
0.0
-86.7
13.3
-3.3
0.0
-3.3
150
Table 9. continued.
cQnQLYo
Ach/Ilea mi/lefo//um L.
Anemone pieri Britt. ex Rydb.
Antennar/a rosea Greene
A rn/ca cord/folia Hook.
clayton/a perfol/ata Donn ex WUId.
Eu/yb/a conspicua (LindI.) Nesom
Fragarla spp.
Thined Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-13.3
6.7
0.0
-60.0
-13.3
3.3
0.0
-63.3
0.0
23.3
0.0
-33.3
16.7
3.3
0.0
6.7
16.7
0.0
-56.7
16.7
0.0
13.3
0.0
-10.0
16.7
-40.0
-16.7
16.7
26.7
0.0
-53.3
-3.3
0.0
0.0
0.0
0.0
3.3
-3.3
3.3
6.7
0.0
-23.3
0.0
-13.3
0.0
-20.0
6.7
-23.3
-13.3
0.0
10.0
0.0
10.0
-13.3
-20.0
-13.3
-96.7
36.7
0.0
-10.0
0.0
-13.3
26.7
-76.7
-20.0
0.0
0.0
0.0
0.0
0.0
0.0
Potent//Ia gracilisDougl. ex Hook.
Prunella vulgarisL.
6.7
0.0
3.3
0.0
0.0
0.0
0.0
-10.0
6.7
0.0
-3.3
6.7
0.0
0.0
16.7
-6.7
-3.3
6.7
RanuncuIusspp.
0.0
0.0
0.0
0.0
0.0
0.0
3.3
3.3
6.7
13.3
36.7
23.3
-6 7
-10 0
-10 0
-16 7
-6 7
-20 0
0.0
0.0
0.0
-3.3
-3.3
10.0
Ga//urn spp
Hierac/um a/b/f lorum Hook.
Iris misouriens/s N utt.
Lathyrus spp.
Lupinus sop.
frloehr/ng/amacrophy//a(Hook.)
Fenzl
Osmorhi'abertero/DC.
Ste//ar/a /ongiesGoldie
Symphyotrichurn spat/7u/atum
(Lindi.) Nesom var. spathu/atum
Taraxacum offic/nale G.H. Weber ex
Wiggers
Thalictrum fend/er/Engeim. ex Gray
Trifo//umrepensL.
i.).J
.)J
-16.7
3.3
-10.0
-16.7
-3.3
0.0
0.0
-3,3
0.0
0.0
0.0
0.0
0.0
-6.7
-3.3
16.7
50.0
20.0
23.3
Viola adunca Sm.
46.7
60.0
6.7
-10.0
3.3
-3.3
V/cia americana Muhi. ex Wild.
-20.0 -10.0
-13.3
-6.7
6.7
13.3
-10.0
Ma/ion/a repens(Lindl.) G. Don
16.7
0.0
-3.3
0.0
0.0
Sp/raeabetullfol/aPallas
-3.3
0.0
10.0
6.7
-20.0
3.3
Symphor!carposa/bus(L.) Blake
-3.3
-6.7
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big
game grazing only; TEXC total exclosure, exclusion of cattle and big game grazing.
t)
151
Tab'e 10. The changes in species composition in a ponderosa pine forest from 1985 to
2005 usinci a 30 cm x 60 cm IDiot frame.
Control Overstory
Thinned Overstory
CEXC TEXC GRAZED CEXC
TEXC GRAZED
Site 1
BromuscarinatusHook.&Arn.
Carex concinno/des Mackenzie
CarexgeyeriBoott
Ca/amagrostis rubescens Buckl.
E/ymusglaucusBuckl.
Me//ca bulbosa Geyer ex Porter & Coult.
Pea pratenstsL.
TrLsetumcanescensBuckl.
Ac/ii7/eamf//efo//umL.
Anemone pipert Britt. ex Rydb.
Antennaria rosea Greene
Am/ca cordifolia Hook.
aaytonia peifoliata Donn ex WilId.
Ewybiaconspicua(Lindl.) Nesom
Fragarfaspp.
Ga/turn spp
Hieradurn a/b/forum Hook.
IrimissouriensisNutt.
Lathyrusspp.
Lup/nusspp.
tv/eel )r/ng/a macrop/iy//a (Hook.) Fenzl
OsmorhizaberteroiDC.
Potenti//agraa7isDougl. ex Hook.
Prune/ta vulgariL.
Ranuncu/usspp.
Ste//aria Iongioes GoIdie
Symphyotrichum spathulatum (Lindl.)
Nesom var. spathulatum
Taraxacumofficina/eG.H. Weberex
Wiggers
Thallctrum fend/eriEngelm. ex Gray
Trilb/iumrepensL.
Viola adunca Sm.
V/cia americana Muhi. ex WilId.
Mahonia repens (LindL) G. Don
Spfraeabetu/ifoh'aPallas
Symphor/carposa/bus(L.) Blake
0.0
0.0
6.7
-26.7
36.7
0.0
46.7
-36.7
3.3
0.0
-6.7
-20.0
-3.3
3.3
0.0
-30.0
3.3
0.0
16.7
-3.3
10.0
50.0
-13.3
-23.3
-6.7
-26.7
-16.7
13.3
0.0
-13.3
0.0
0.0
-36.7
-13.3
-3.3
0.0
0.0
0.0
-23.3
-70.0
0.0
-3.3
6.7
0.0
23.3
0.0
-6.7
0.0
0.0
-13.3
-10.0
-6.7
0.0
3.3
-26.7
-13.3
-23.3
-6,7
0.0
0.0
0.0
0.0
0.0
-3.3
0.0
0.0
-33.3
-3.3
-6.7
0.0
10.0
-3.3
-16.7
-33.3
-13.3
0.0
10.0
0.0
-3.3
-26.7
-33.3
23.3
-43.3
6.7
3.3
0.0
-6.7
0.0
0.0
6.7
-30.0
0.0
3.3
-6.7
3.3
0.0
-63.3
-3.3
-3.3
-3.3
-6.7
16 7
26 7
40 0
-10.0
-33.3
-3.3
-3.3
6.7
0.0
3.3
0.0
0,0
3.3
3.3
10.0
-16.7
0.0
53.3
0.0
33.3
50.0
0.0
23.3
-3.3
-40.0
-33.3
3.3
-50.0
33.3
0.0
23.3
-26.7
30.0
-33.3
30.0
-13.3
-20.0
0.0
0.0
-3.3
0.0
0.0
-33.3
-6.7
0.0
6.7
43.3
16.7
-10.0
-6.7
-40.0
-6.7
-3.3
3.3
0.0
0.0
0.0
0.0
-26.7
36.7
6.7
0.0
13.3
-3.3
-6.7
6.7
-43.3
-83.3
-6.7
0.0
0.0
0.0
36 7
23 3
43 3
-10.0
-50.0
-26.7
-13.3
0.0
-26.7
-3.3
-23.3
6.7
0.0
3.3
0.0
0.0
-6.7
0.0
3.3
0.0
23.3
0.0
-3.3
0.0
0.0
40.0
0.0
3.3
0.0
-3.3
-36.7
-6.7
33.3
0.0
10.0
0.0
0.0
3.3
0.0
23.3
0.0
-6.7
0.0
0.0
46.7
-16.7
0.0
10.0
13.3
16.7
0.0
0.0
0.0
0.0
0.0
-3.3
-3.3
3.3
0.0
3.3
-36.7
0.0
6.7
0.0
6.7
3.3
0.0
Site 2
Sromuscari'natusHook. &Arn.
Carex concinnoides Mackenzie
CarexgeyeriBoott
Ca/amagrostLcrubescensBuckl.
E/yrnusg/aucusBuckl.
152
Table 10. continued.
cgntcToLvesto
Mel/ca bu/bosa Geyer ex Porter & Coult.
PoapratensiL.
Tr/setumcanescensBuckl.
Achi/eamilefollumL.
Anemone pioer/Britt. ex Rydb.
AntennariaroseaGreene
Am/ca cord/fo/la Hook.
Clayton/a perfol/ata Donn ex WilId.
Euiyb/a consp/cua (Lindi.) Nesom
Fragar/aspp.
Gallumspp
Hieracium a/b/forum Hook.
IrLcm/ssouriensisNutt.
Lathyrusspp.
Lup/nusspp.
Moetiringla macrophylla (Hook.) Fenzl
Osmorhizabertero/DC.
Potent/llagraci/isDougL ex Hook.
Prunellavu/gar/sL.
Ranunculusspp.
Stellar/a /ongi;oesGoldie
Symphyotriclium spathu/atum (LindL)
Nesom var. spathu/atum
Taraxacum officinaleG.H. Weber ex
Wiggers
Thai/drum fend/er/Engelm. ex Gray
TrifoI/umrepensL.
Viola adunca Sm.
V/c/a amer/cana Muhi. ex WilId.
Mahoniarepens(LindI.) G. Don
Spiraeabetulffo/iaPallas
Symphoricarposa/bus(L.) Blake
CEXC
TEXC
0.0
-6.7
-26.7
-10.0
0.0
6.7
-3.3
0.0
0.0
-20.0
0.0
-20.0
0.0
-33.3
30.0
-6.7
0.0
0.0
0.0
0.0
0.0
0.0
-46.7
-23.3
-6.7
0.0
0.0
0.0
0.0
0.0
6.7
0.0
-13.3
0.0
10.0
-33.3
-10.0
0.0
0.0
0.0
0.0
0.0
33 3
Thinned Overstory
GRAZED CEXC
0.0
-10.0
-33.3
20.0
0.0
10.0
-13.3
0.0
0.0
23.3
20.0
-3.3
0.0
-73.3
6.7
TEXC
GRAZED
0.0
-3.3
-16.7
-3.3
0.0
0.0
16.7
0.0
0.0
-6.7
0.0
-6.7
0.0
-53.3
-3.3
-3.3
-3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-3.3
-6.7
-13.3
0.0
0.0
0.0
0.0
-20.0
-26.7
-6.7
0.0
0.0
-23.3
0.0
0.0
6.7
0.0
0.0
0.0
-86.7
6.7
-6.7
0.0
0.0
0.0
0.0
0.0
33
00
56 7
16 7
26 7
0.0
-3.3
-6.7
3.3
0.0
3.3
0.0
0.0
0.0
0.0
-20.0
-6.7
-13.3
0.0
-3.3
-6.7
0.0
16.7
0.0
30.0
0.0
-3.3
0.0
0.0
0.0
0.0
13.3
0.0
-6.7
0.0
0.0
-3.3
0.0
10.0
0.0
30.0
0.0
0.0
-6.7
0.0
-6.7
0.0
-20.0
13.3
0.0
-23.3
-40.0
3.3
-13.3
13.3
0.0
3.3
0.0
10.0
0.0
-36.7
-23.3
-6.7
-23.3
-16.7
-3.3
-3.3
26.7
-43.3
33.3
0.0
-23.3
-26.7
6.7
-16.7
43.3
-60.0
10.0
30.0
0.0
-56.7
0.0
0.0
-6.7
-16.7
-3.3
0.0
0.0
23.3
0.0
0.0
-6.7
0.0
-30.0
0.0
-6.7
-3.3
-6.7
-3.3
-13
3.3
-3.3
16.7
0.0
36.7
Site 3
Bromus car/natus Hook. & Am.
Carexconcinno/desMackenzie
Carexgeyer/Boott
CaIamagrost/ rubescens Buckl.
E/ymusglaucusBuckl.
Meilca bulbosa Geyer ex Porter & Coult.
PoapratensicL.
Trisetum canescens Buckl.
Ach//leamll/efoliumL.
Anemone pieriBritt. ex Rydb.
Antennar/aroseaGreene
Am/ca cord/fol/a Hook.
aaytonia peifol/ata Donn ex Willd.
60.0
-36.7
-23.3
10.0
0.0
-56.7
0.0
10.0
40.0
-53.3
-23.3
6.7
0.0
-70.0
0.0
3.3
36.7
0.0
-33.3
0.0
0.0
0.0
-13.3
-20.0
36.7
-13.3
26.7
-56.7
20.0
10.0
0.0
-96.7
0.0
-10.0
-13.3
0.0
-10.0
30.0
-46.7
10.0
16.7
0.0
-60.0
0.0
153
Table 10. continued.
Clverstory
CEXC
Euiybiaconspicua(Lindl.) Nesom
Fragariaspp.
Ga/iumspp
H/eraciuma/biflorumHook.
Iris missourienstsNutt.
Lathyrusspp.
Lupinusspp.
Moehrinqi'amacrophyl/a (Hook.)
Fenzl
Osmorhi'aberterofDC.
Potent/I/a grac/l.& Dougl. ex Hook.
Prune/la vu/gar/sL.
Ranuncu/usspp.
Stellaria/ongioesGoldie
Symphyotrichuinspathu/atum(Lindl.)
Nesom var. spathu/atum
TaraxacumofficinaleG.H. Weberex
Wiggers
Tha/ictrum fend/en Engeim. ex Gray
TEXC
TEXC
GRAZED
0.0
-3.3
-20.0
-20.0
-20.0
0.0
-10.0
6.7
0.0
0.0
-6.7
20.0
6.7
0.0
0.0
16.7
-43.3
-26.7
-6.7
0.0
26.7
0.0
3.3
10.0
16.7
40.0
26.7
0.0
0.0
-6.7
-3.3
0.0
3.3
-3.3
6.7
3.3
0.0
-26.7
6.7
0.0
3.3
0.0
3.3
-3.3
43.3
0.0
-13.3
0.0
0.0
-23.3
36.7
-6.7
0.0
-10.0
43.3
-80.0
-26.7
0.0
-3.3
10.0
0.0
0.0
-13.3
-6.7
-15.7
0.0
-16.7
3.3
-33.3
-20.0
0.0
0.0
0.0
0.0
0.0
-6.7
Thinned Overstory
GRAZED CEXC
16.7
0.0
-23.3
-3.3
3.3
0.0
-3.3
10.0
-23.3
-43.3
-20.0
-3.3
6.7
0.0
0.0
0.0
-3.3
0.0
0.0
-3.3
-3.3
-6.7
-3.3
0.0
0.0
0.0
Viola aduncaSm.
60.0
30.0
23.3 26.7
63.3
50.0
V/cia americana Muhi. ex WilId.
-16.7 10.0
-33.3 -16.7
-3.3
6.7
Mahoniarepens(Lindl.) G. Don
-6.7
13,3
23.3
-3.3
6.7
-6.7
Spi'raeabetu/ifoli'aPallas
0.3
6.7
-13.3
0.0
0.0
0.0
Syinphoricarposalbus(L.) Blake
-13.3 -33.3
0.0
10.0
-3.3
3.3
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exciosure, big
game grazing only; TEXC total exciosure, exclusion of cattle and big game grazing.
ThifollumrepensL.
154
Appendix C
Ponderosa Pine Shrub Density and Cover Changes
155
Table 1. Changes in shrub density from 19
1988, 1991, 1994, and 2003.
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
-1.0
1.3
-6.3
-3.3
1.0
-1,3
0.3
0.3
0.0
-2.0
-2.7
-1.0
-1.3
0.0
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
-1.7
0.0
-5.3
-6.7
1.3
3.0
-2.3
-1.7
-3.0
0.0
-0.3
Rosa gymnocarpa Nutt.
0.0
22.7
3.0
0.0
R/bescereumDougl.
1.3
0.0
3.3
-1.0
-1.0
0.3
0.3
-0.3
0.3
0.0
-1.0
-1.3
-1.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-23.7
0.0
0.0
8.3
0.0
0.0
-7.0
0.0
0.0
0.0
0.0
0.0
2.0
0.0
0.0
-0.3
-0.3
-1.3
-0.3
-2.3
-0.7
-0.3
-2.7
-5.3
-0.3
-1.3
7.7
-1.7
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.3
-0.3
0.0
0.0
-5.0
0.0
0.0
-6.0
0.0
0.0
-4.0
0.0
-0.3
-0.3
0.0
0.0
-5.7
0.0
0.0
2.7
1.3
1.3
2.0
3.0
5.0
2.0
0.0
10
0.0
00
-0.7
-0.7
-0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-1.7
0.0
-2.3
4.0
0.0
10.7
0.0
-0.7
2.3
-3.7
0.0
19.7
0.0
-0.7
8.3
0.0
1.7
9.7
Site 1
1988
Ame/anchier a/n/foli (Nutt.)
Nutt. ex M. Roemer
crataegus doug/as/i LindL
Holod/scus disco/or (Pursh)
Maxim.
Physocarpus malvaceus
(Greene) Kuntze
Prunus virginiana L.
Site 2
a/n/fat/a (Nutt.)
Nutt. ex M. Roemer
Crataegusdoug/as//Lindl.
Holodiscus disco/or (Pursh)
Maxim.
Physocdrpus ma/vaceus
(Greene) Kuntze
Prunus virgin/anaL.
R/bescereumDougl.
Rosa gymnocarpaNutt.
Arne/anchier
Site 3
Ametanchiera/n/fo/ia(Nutt.)
Nutt. ex M. Roemer
Crataegusdoug/asiiLindl.
Hotodiscus disco/or (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virgin/anaL.
R/bescereurnDougl.
Rosa gymnocarpaNutt.
1991
Site 1
Ame/anch/er a/n/fo/ia (Nutt.)
Nutt. ex M. Roemer
Crataegusdouglasi/Lindl.
Ho/od/scus disco/or (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virgin/anaL.
R/bescereumDougl.
Rosa gymnocarpaNutt.
3.0
156
ble 1. continued.
Thinned Overstory
______________________cExcitxc
GRAZE
CEXC
TEXC
GRAZE
Site 2
Amelanchier alnifolia (Nutt.)
Nutt. ex M. Roemer
C'rataegus doug/as/fund!.
Holod/scus discolor (Pursh)
0.0
1.7
0.3
0.7
0.3
0.0
1.0
0.0
1.0
2.3
1.7
0.7
0.0
0,0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-15.3
0.0
0.0
35.3
0.0
0.3
11.7
0.0
0.0
0.3
0.0
0.0
19.3
0.0
0.0
0.3
3.7
45.0
1.0
-0.7
1.3
1.3
0.3
-5,3
-0.3
1.0
1.3
3.7
0.0
0.0
0.0
0.3
1.3
0.0
0.0
0.0
0.0
0.0
-0.3
-0.3
0.0
0.0
2.3
0.0
0.0
-2.3
0.0
0.0
8.3
0.0
-0.3
4.3
0.0
0.0
3.3
0.0
0.0
2.7
4.0
.3
34.7
24.7
22.3
7.7
C'rataegus doug/as/f Lind!.
Ho/odiscus disco/or (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
0.3
0.7
0.0
9.3
1.3
-0.3
9.3
0.0
0.0
00
0.0
0.0
0.3
0.0
0.0
0.0
0.0
-1.7
PrunusvfrginianaL.
Ribes cereun Doug!.
Rosa gym.'ccarpaNutt.
0.0
0.3
13.0
0.0
35.7
5.0
0.0
-3.7
13.3
5.0
25.3
0.0
0.3
53.7
0.0
0.7
14.3
2.7
9.0
1.3
1.7
8.3
0.0
0,0
-1.3
1.0
-0.3
2.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
.o
0.0
0.0
0.0
0.0
0.0
0.0
0.0
40.0
0.0
0.0
4.3
Maxim.
Physocarpus malvaceus
(Greene) Kuntze
Prunus vfrginfana L.
Rfbes cereum Doug!.
Rosa gymnocarpaNutt.
Site 3
Ame/anchier alnifolfa
(Null.)
Null. ex N. Roemer
Crataegusdoug/asf/Lindl.
Ho/od/scus discolor (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus vfrginina L.
Rfbes cereurn Doug!.
Rosa gymnocarpaNutt.
Ste1
1994
Ame/anchier ain/folla
(Null.)
Null. ex M. Roemer
1.3
Site 2
Amelanchier alnifolla
(Null.)
Null. ex M. Roemer
Crataegus douglas/f Lindl.
Ho/odiscus disco/or (Pursh)
Maxim.
Physocarpusma/vaceus
(Greene) Kuntze
Prunus virgin/anaL.
Ribes cereum Doug!.
Rosa gymnocarpaNutt.
0.0
0.0
-11.7
0.0
0.0
45.7
0.0
0.0
30.0
1.0
157
Table 1. continued.
Control Overstory
IEXC GRAZE
CEXC
Thinned Overstory
CEXC
TEXC
GRAZE
Site 3
Amelanchiera/nifo/ia(Nutt)
Nutt. ex M. Roemer
Crataegus douglas/i Lindi.
Ho/odiscusdisco/or(Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus
virginiana L.
RibescereurnDougl.
2003
Rosa gymnocarpNuft.
Site 1
Amelanchier alnifo/la (Nutt.)
Nutt. ex M. Roemer
crataegus doug/as/i Lindl.
Ho/odiscus disco/or (Pursh)
Maxim.
Ptiysocarpus ina/vaceus
(Greene) Kune
Prunus virgii7iana L.
RibescereumDougl.
Rosa gyrnnocarpaNutt.
1s6.:
360.7
-1.3
-0.3
3.0
-0.7
7.3
tO
-0.7
3.0
0.0
1.0
0.0
0.0
0.0
-0.3
6.7
0.0
0.0
0.0
57.3
0.0
7.3
1.0
0.0
0.0
16.3
0.0
0.0
5.3
0.0
0.0
5.7
0.0
0.3
0.0
0.0
Q.0
0.0
0.0
13.0
3.7
2.3
5.7
5.7
11.0
16.7
2.7
0.3
0.3
1.3
0.7
0.0
7,0
0.0
0.0
0.0
0.0
0.7
0.0
0.0
0.0
0.0
0.0
-1.7
0.0
-2.0
17.3
0,0
25.0
16.3
0.0
5.3
3.7
25.3
0.7
23.0
0.0
0.0
78.7
6.0
3.7
17.7
1.0
21.3
1.0
7.0
1.0
1.7
4.0
0.3
0.3
-0.3
2.3
0.3
0.0
0.0
0.0
0.3
0.0
0.0
b.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-5.3
0,0
0.0
47.3
0.0
0.0
31.0
0.0
0.0
2.7
0.0
0.0
19.7
0.0
0.0
0.7
4.3
4.0
3.3
1.0
2.0
7.7
2.3
-4.3
0.7
-0.7
0.7
7.7
0.0
0.0
0.0
-0.3
-0.3
0.0
0.0
0.0
0.0
0.7
1.0
-0.3
1.3
Site 2
Aine/anchier alaifolia (N utt.)
Nutt. ex M. Roemer
C'rataegus doug/as/i Lindl.
Holodiscus discolor (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virginiana L.
RibescereumDougl.
Rosa gyrnnocarpaNutt.
Site 3
Ame/anchier airilfolia
(Null.)
Null. ex M. Roemer
C'rataegus douglas/i Lindl.
Ho/odiscus discolor (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virginiana L.
0.0
0.0
0.0
0.3
0.0
0.0
17.3
22.3
57.0
_pgyrnnocarpaNutt.
Herbivory treatments: Graze cattle arid big game grazing; CEXC
game grazing only; TEXC total exclosure, exclusion of cattle and
RibescereumDougl.
0.0
0.0
0.7
0.3
1.3
9.0
5.0
cattle exctosure, big
big game grazing.
0.0
2.3
158
Table 2. Changes in shrub coy
from i99to191991, 1994, and 2003.
çontro:ccrstory
CEXC
Site
1988
.EXC
GRAZE
Thinned Overstory
CEXC
TEXC
GRAZE
I
Ame/anchfer alnifo/fa (Nutt.)
Nutt. ex M. Roemer
CrataegusdougIasf/Lindi.
Ho!od/scusdLsco/or(Pursh)
Maxim.
P/iysocarpus ma/vaceus
(Greene) Kuntze
Prunus vfrgfnfana L.
RibescereumDougl.
Rosa gymnocarpaNutt.
Site 2
Ame/anchier alnifo/fa (Nutt.)
Nutt. ex M. Roemer
CrataegUsdoug/asf/Lindt.
Ho/odiscusdiscolOr(Pursh)
04
4
4
-7 1
08
0.0
00
0.0
-4.1
-0.1
-0.1
0.0
0.0
0.0
0.5
2.
-o
3
00
0
00
00
00
-1 4
0.0
-3.3
0.0
-2.8
0.9
0.1
0.0
-0.7
-0.1
0.0
-2.2
0.0
1.1
-0.1
-1.1
0.0
0.6
0.0
0
00
00
00
04
00
0.0
0.0
0.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
00
00
00
00
0
00
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virginfana L.
R/bescereurnDougJ.
0.0
0.0
0.0
0.0
0.0
0.0
0.()
0.0
0.0
0.0
0.0
0.0
Rosa gyrnnocarpaNutt.
-0.8
0.0
-2.1
0.0
0.0
0.1
o.
O)
0.0
0.3
0.0
0.0
2.5
-5.3
0.0
0.2
0.6
-0.2
O.
0.0
0.0
0.6
0.0
0.0
00
00
00
00
-0 8
00
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.2
0.0
0.0
0.1
-0.6
-0.1
-0.1
-0,1
0.0
3.b
-
3.0
-3.6
2.7
-0.8
0.0
0.0
0.0
-4.2
0.0
-0.1
2.1
00
0.0
0.0
0.0
0.5
0.0
0.0
0.0
0.0
-1.4
0.0
Site 3
Amelanchieralnifo/ia(Nutt.)
Nutt. ex M. Roemer
Crataequsdouq/asiiLindi.
/-Io/odiscusdisco/or(Pursh)
Maxim.
Physocarpus ma! vace us
(Greene) Kuntze
Prunus virgin/anaL.
RibescereumDougl.
Rosa gymnocarpaNutt.
Site I
1991
Ame/anc/iler a/nib/ia (Nutt.)
Nutt. ex M. Roemer
CrataegusdouglasllLindl.
Ho/odiscusdisco/or(Pursh)
Maxim.
Physocarpusma/vacoL's
(Greene) Kuntze
Prunus virgin/anaL.
RibescereumDougl.
Rosa gymnocarpaNutt.
0..
.
0.0
0.0
0.0
0.0
0.0
-2.7
-5.0
-3.9
0.0
0.0
0.5
-0.2
08
0.4
1.7
0.7
0.5
159
Table 2. continued.
CoitrO'rstory
CEXC 1XC
Site
Ame/anchiera/nifo/ia(Nutt'j
Nutt. ex M. Roemer
Crataegusdougfasf/undL
Ho/od/scusdiscolor(Pursh'
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virgin/anaL.
RibescereumDougj.
Rosa gymnocarpaNutt.
Site 3
Aine/anchiera/nifol/a(Nutt.)
Nutt. ex M. Roemer
Ciataegusdoug/asi/Lindi.
Ho/od/s'cusdtsco/or(Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
PrunusvirginiaraL.
Ri'bescereumDougl.
Rosa gymnocarpaNutt.
1994
Thinned Overstory
GRAZE
CEXC
TEXC
GRAZE
0.0
1.3
0.0
0.0
0.4
0.0
0.0
0.0
-0.2
0.1
0.0
0.0
0.0
U.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-1.0
0.0
0.0
0.5
0.0
0.0
-2.2
0.0
0.0
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.2
4.8
0.2
0.0
0.2
0.0
2.8
-4.9
0.0
-0.1
0.0
0.5
0.0
0.0
0
-0.9
1.2
0.0
0.0
0.0
0.0
0.0
-0.8
0.0
0,0
0.0
0.2
0.0
0.0
-0.7
0.0
0.0
0,0
-0.2
0.0
0.0
0.1
0.1
0.0
0.0
0.2
2.0
6.3
4.1
-4.1
2.8
-1.1
0.2
1.0
0.0
-4.5
0.1
1.4
0.0
0.0
0.0
0.0
0.1
0.0
0.0
0.0
0.0
0.0
-1.4
0.0
-3.2
0.9
0.0
-5.9
0.4
0.0
-4.1
0.0
0.2
0.4
1.5
0.0
-0.7
2.2
0.0
0.0
0.5
0.2
0.0
0.0
0.0
0.2
0.0
0.0
0.0
-0.1
0.0
0.6
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-2.0
0.0
0.0
0.0
0.0
0.0
0.7
0.0
0.0
0.0
0.0.
0.1
Site I
Anie/anchieralnifo//a(Nutt.)
Nutt. ex M. Roemer
Crataegus douglas/i Lindl.
Holodicusdiscolor(Pursh)
Maxim.
Physocarpus malvaceus
(Greene) Kuntze
Prunus virgin/anaL.
RibescereumDougl.
Rosa gymnocarpa Nutt.
Site 2
Ame/anchieralnifo//a(Nutt.)
Nutt. ex M. Roemer
Crataegusdoug/asiiLindl.
Ho/odi5cusd/scolor(Pursh)
Maxim.
Physocarpusmalvaceus
(Greene) Kuntze
Prunus virgin/anaL.
R/bescereumDougl.
Rosa gymnocarpaNutt.
0.0
0.0
-0.8
0.0
0.0
0.2
160
Table 2. continued.
OtroOverjy
Site 3
Ame/anchier a/n/Ia/ia (Nutt.)
Nutt. ex M. Roemer
Crataegus douglas/i L.ndL
l-lo/odiscus disco/ar (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
PrunusvirginianaL.
RibescereurnDougL
Rosa gymnocarpaNutt.
2003
Thinned Overstory
CEXC
IEXC
GRAZE
CEXC
TEXC
GRAZE
3.7
IOM
0.0
-0.2
0.0
0.0
5.5
-S.3
0.0
-0.2
0.0
-1.3
0.0
0.0
0.0
-0.9
0.1
0.0
0.0..
0.0
1.6
0.0
0.7
0.0
0.0
0.0
0.4
0.0
0.0
-0.6
0.0
0.0
0.0
-0.2
-0.1
-0.1
0.0
0.0
0.8
0.0
0.0
0.0
1.1
3.7
3.8
-2.6
6.8
7.0
0.2
0.1
0.0
-4.4
4.5
-0.1
1.5
0.0
0.0
0.0
0.0
-0.4
0.0
0.0
0.0
0.0
0.0
-1.4
0.0
-3.0
.0.6
0.0
0.0
0.7
0.0
-0.7
0.7
0.0
0.0
2.4
0.0
-4.9
0.0
1.9
-E.5
-0.2
0.0
0.0
0.0
0.1
0.6
0.0
0.0
0.0
-0.2
0.0
4.7
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-1.1
-0.1
0.0
0.0
-2.8
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.2
1,2
0.0
-0.2
1.4
0.1
2.1
-1.0
0.0
-0.6
0.3
0.7
0.0
0.0
0.0
-0.9
0.0
0.0
0.0
0.0
0.0
0.0
-0.8
0.0
0.0
00
0.0
0.0
0.0
0.0
Site I
Amelanchiera/nifolia(Nutt.)
Nutt. ex M. Roemer
Crataegusdoug/asllLindl.
Ho/odiscus discolor (Pursh)
Maxim.
Physocarpus malvaceus
(Greene) Kutze
Prunus virgin/anaL.
RibescereuniDougl.
Rosa gyrnnocarpaNutt.
Site 2
Amelanchiera/nifol/a (Nutt.)
Nutt. ex M. Roemer
Crataegusdoug/as//Lindl.
Holodiscus disco/or (Pursh)
Maxim.
Physocarpus malvaceus
(Greene) Kuntze
Prunus virginiana L.
RibescereumDougl.
Rosa gymnocarpaNutt.
Site 3
Ame/anchier a/nib//a (Nutt.)
Nutt. cx M. Roemer
crataegusdouglasllLindl.
Ho/odLccus disco/or (Pursh)
Maxim.
Physocarpus ma/vaceus
(Greene) Kuntze
Prunus virgin/anaL.
0.0
0.0
0.0
-0.2
0.0
RibescereumDougl.
0.0
1.7
Rosa gymnocarpaNutt.
-0.4
0.0
-0.1
-0.2
0.0
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exciosure, big
game grazing only; TEXC total exclosure, exclusion of cattle and big game grazing.
161
Appendix D
Grand Fir and Ponderosa Pine Understory Production
Table 1. The effects of year, timber harvest, and
Forest
Type
Year
Site
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
ABGR
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
Timber
Harvest
Clearcut
Clearcut
Clearcut
Control
Control
Control
Thinned
Thinned
Thinned
Clearcut
Clearcut
Clearcut
Control
Control
Control
Thinned
Thinned
Thinned
Clearcut
Clearcut
Clearcut
Control
Control
Control
Thinned
Thinned
Herbivoy
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Grazed
CExc
CGExc
Total
Prod.
1231.8
1261.9
1399.5
876.0
925.4
738.2
1060.8
967.1
948.2
1180.5
1329.6
1322.7
754.9
665.6
770.2
1354.2
1175.0
899.0
1769.8
1449.7
1934.2
1022.5
902.0
830.4
1167.8
934.2
production.
CAGE
CARU
POPR
SYAL
Pgrass
Pforb
Annual
Shrub
124.0
289.2
288.4
336.0
178.0
198.8
113.0
235.8
193.6
171.8
473.8
208,8
283.0
296.4
191.8
69.6
312.2
90.2
296.4
422.0
535.8
264.6
36.0
116.8
281.2
104.4
54.4
33.0
172.6
8.8
16.6
6.8
135.4
218.0
269.6
113.8
210.8
320.4
89.0
64.2
249.6
464.0
154.6
78.2
64.4
206.0
740.8
212.2
200.0
292.4
364.2
528.0
273.2
371.2
158.4
52.8
23.8
7.6
132.2
123.0
90.8
82.0
71.2
11.4
148.4
132.2
138.2
63.8
109.4
25.2
21.4
21.8
25.8
53.4
165.4
96.2
8.6
31.0
24.0
49.0
63.0
60.4
50.0
83.4
85.2
21.0
6.0
21.4
26.2
3.2
227.4
63.4
165.8
74.4
113.8
52.4
496.4
94.0
159.8
188.0
91.0
128.2
35.0
45.6
43.0
70.8
196.0
345.6
217.4
68.0
132.6
63.2
24.6
29.2
121.0
65.0
230.2
296.0
466.8
244.6
277.2
316.2
180.8
50.0
216.4
143.8
6.0
4.4
2.6
29.0
24.8
13.4
12.2
2.6
8.4
0.6
0.6
5.4
0.6
4.8
2.2
62.8
12.0
40.0
320.6
212.6
171.8
220.4
267.2
192.2
95.4
251.2
117.2
398.2
1.2
5,8
10.0
0.8
27.6
28.6
483.6
439.2
294.0
27.2
0.0
00
33.6
4.2
105.2.
231.6
97.8
103.2
109.2
47.8
29.6
70.6
490.0
82.4
114.6
410.4
117.0
155.8
197.0
246.4
151.2
68.4
123.6
5.6
0.2
3.6
38.8
0.4
21.&
432.2
322.8
1.18
120.8
154.8
268.8
327.6
81.8
245.4
107.8
209.8
358
208.6
216.4
169.8
Table 1. continued.
Forest
1
Timber
Harvest
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
2
COntrol
2
Control
Control
Thinned
Thinned
Thinned
Control
Contro'
Control
Thinned
Thinned
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
Control
Control
Control
jype
Year
Site
ABGR
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
2003
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1989
1992
1992
1992
1992
1992
1992
1992
1992
1992
3
P1PO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
1
1
1
1
1
2
2
2
2
3
3
3
3
3
3
1
1
1
1
1
1
2
2
2
Herbivoy
Grazed
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
Total
CAGE
CARU
POPR
SYAL
Pgrass
Pforb
Annual
Shrub
865.6
868.7
1047.7
1744.7
1788.9
1883.4
494.0
467.9
145.8
197.0
152.8
182.0
164.8
182.2
182.8
131.2
134.2
2470
46.6
65.2
27.2
118.2
367.8
528.6
473.4
1.6
0.0
0.0
903.2
1209.6
1239.2
690.8
699.5
746.0
1225.0
1319.5
1355.5
757.5
852.2
723.9
935.7
1215.7
1102.2
338.4
311.9
558.0
145.4
298.4
251.2
231.6
24?.2
116.2
260.8
216.2
270.0
314.0
307.2
117.4
259.6
131.4
88.4
142.2
251.6
153.8
20.6
215.8
123.2
108.8
141.8
261.6
152.8
26.6
23.6
8.8
84.0
60.2
51.4
64.4
19.8
38.8
17.0
29.8
31.4
80.6
146.8
64.6
63.6
143.0
161.8
47.2
38.8
24.0
223.6
130.0
91.4
147.2
342.2
472.8
524.6
42.6
25.2
9.4
121.8
109.8
104.6
95.0
56.6
36.0
157.6
414.6
174.4
127.4
107.6
147.0
137.8
261.2
161.0
57.4
38.0
33.2
136.2
264.6
309.0
277.2
341.0
290.2
463.0
198.4
253.4
193.6
370.8
582.0
441.4
236.4
268.8
329.6
448.6
320.4
403.0
125.6
50.6
137.6
157.0
176.6
176.6
116.6
17.0
27.6
4.8
32.4
12.8
74.4
76.6
11.2
68.4
4.6
5935
509.6
45.6
60.0
80.6
13.8
0.0
0.0
0.0
2.4
49.2
63.8
25.4
164.2
92.8
38.0
75.2
44.8
68.6
71.8
0.8
0.0
13.0
11.0
0.0
16.6
13.0
45.6
115.2
242.2
11.6
30.4
35.6
37.6
84.2
11.2
22.6
25.6
19.0
20.4
15.4
40.4
51.2
57.8
107.6
71.0
41.4
3.2
0.0
43.4
0.0
68.8
116.0
145.0
27.6
0.0
14.0
Prod.
1119.3
12.6
46.0
16.2
57.0
8.0
3.8
188.0
172.0
266.8
96.8
109.2
73.0
172.0
267.2
141.8
3.2
21.0
0.0
1.0
12
4.8
5.6
.6
LO
0.0
20
5.2
3.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
12.0
6.0
9.0
i
L.J
Table 1. continued.
Forest
Type
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
pIPO
PIPO
PTPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
Year
Site
1992
1992
1992
1992
1992
1992
1992
1992
1992
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
1995
2003
2
2
2
3
3
3
3
3
3
1
1
1
1
1
1
2
2
2
2
2
2
3
3
3
3
3
3
1
Timber
Harvest
Thinned
Thinned
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
Control
Herbivoy
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
GRAZED
CEXC
CGEXC
CGEXC
Total
Prod.
543.9
731.1
751.7
498.2
608.2
558.1
623.1
775.0
753.2
906.6
1025.7
980.2
1048.1
1067.1
1112.9
378.7
436.2
689.2
494.9
771.7
867.5
380.5
600.7
602.8
588.0
874.7
1049.2
1287.8
CAGE
CARU
POPR
SYAL
Pgrass
Pforb
Annual
Shrub
261.4
243.8
222.2
210.0
253.6
482.0
101.0
123.4
173.4
251.4
304.4
173.6
174.2
330.2
4.0
83.6
170.8
13.6
43.6
23.2
51.8
69.4
5.0
0.0
267.2
385.0
9.2
0.0
80.8
2.2
18.4
16.2
6.2
28.0
274.0
12.0
11.0
3.2
48.6
138.8
10.4
134.4
20.0
70.4
61.0
2.0
34.8
0.0
31.6
87.1
50.6
153.6
94.2
39.2
145.4
103.8
205.2
0.0
0.4
8.6
46.0
69.6
27.4
0.0
22.8
8.8
7.2
12.6
3.8
16.4
14.0
71.4
113.4
99.2
161.2
84.8
105.2
95.4
7.4
10.2
12.4
25.6
15.8
2.2
3.6
18.2
23.4
139.8
170.8
39.4
111.4
67.6
25.8
131.6
111.8
147.0
136.4
145.6
100.8
219.6
124.6
176.8
21.2
2.0
22.4
109.4
225.4
87.0
84.0
77.0
42.4
117.8
127.6
217.4
211.8
0.0
7.2
0.0
1.2
0.0
3.0
35.4
28.8
67.4
35.4
8.4
82.4
0.0
0.0
41.6
0.6
35.8
36.8
38.6
3.0
0.0
17,8
122.2
271.8
518.6
185.4
321.8
389.8
78.4
478.4
224.8
92.4
211.4
482.8
295.2
1.0
9.6
18.4
21.8
200.4
95.4
106.4
17.2
1.2
54.8
301.4
49.0
63.4
56.2
127.8
39.8
52.8
29.2
50.2
76.4
72.2
136.2
207.8
260.4
300.4
461.2.
96.6
94.2
93.2
77.4
146.0
152.2
131.8
62.4
82.8
240.6
154.8
300.4
239.6
1.0
8.6
28.8
19.0
4.0
42.6
11.4
59.6
27.6
58.0
0.0
7.8
6.2
23.2
19.6
23.2
23.4
3.2
6.2
1.0
3.4
16.0
9.6
13.2
1.2
6.0
0.0
0.0
2.8
44.8
147.0
0.6
25.2
4.4
55.0
Table 1. continued.
Forest
Type
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
PIPO
ABGR
Year
Site
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
2003
1
1
Timber
Harvest
Control
Control
Thinned
Thinned
Thinned
Control
Control
Control
Thinned
Thinned
Thinned
Thinned
Thinned
Thinned
Control
Control
Control
Herbivoy
CEXC
GRAZED
CEXC
GRAZED
CGEXC
GRAZED
CGEXC
CEXC
GRAZED
CGEXC
CEXC
GRAZED
CGEXC
CEXC
CGEXC
CEXC
GRAZED
Total
Prod.
1438.1
1260.5
CAGE
CARU
POPR
SYAL
Pgrass
Pforb
Annual
Shrub
507.6
137.2
479.6
330.8
281.2
137.0
297.4
236.0
238.6
324.4
307.4
258.0
269.2
0.8
0.0
53.6
116.8
36.0
40.8
65.4
61.8
151.0
168.0
10.2
95.6
106.0
168.0
181.0
115.6
0.0
0.0
0.6
25.8
10.2
46.8
120.8
129.8
88.6
150.4
110.8
198.2
22.2
29.8
11.4
25.6
20.6
21.0
21.4
111.2
38.0
25.8
8.4
20.6
127.0
546.6
170.0
323.8
56.6
41.2
52.4
26.4
76.4
21.4
96.0
127.8
27.0
144.0
35.8
56.2
131.0
171.6
200.6
300.4
317.6
418.6
131.8
79.0
105.4
268.6
189.4
244.6
181.4
184.6
230.8
50.2
60.4
127.4
0.0
0.2
22.0
90.4
118.0
0.4
0.0
0.0
3.2
92.2
76.2
135.2
145.2
347.4
2.0
24.2
8.4
14.0
63,2
19.2
4.6
8.0
2.2
133.0
1532.9
1654.7
1
1
1582.4
570.8
2
2
694.9
2
462.5
2
906.5
2
868.6
1.0
2
994.2
5.8
3
713.0
15.0
3
903.2
507.6
86.4
125.0
11.6
3
845.9
184.2
77.8
151.8
19.4
3
576.2
2T,?.8
54.4
3.8
5.0
3
824.5
378.2
76.6
35.6
7.2
73.6
3
741.1
293.4
56.6
82.0
0.2
20.4
Abies grand/s (Dougi. ex D. Don) Lindl.; PIPO Pinus ponderosa P.& C. Lawson
Herbivory treatments: Graze cattle and big game grazing; CExc cattle exclosure, big game grazing only; TExc total exclosure,
exclusion of cattle and big game grazing.
Total Prod. -Total Production; CAGE CarexgeyerfBoott; CARU Calamagrost/srubescensBuckl.; POPR PoapratensLcL.; SYAL
Symphor/carpos a/bus (L.) Blake; PGrass other perennial grasses; PForb perennial forbs; Annuals annuals and biennials; Shrubs
other shrubs.
1
I.-'
166
Appendix E
Grand Fir Understory Vegetation Changes
Table 1. Thç
a grand fir forest from 1985 to 1994 upgp cm x 3Q_cplot frame.
Clearcut Ovety
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
Bromus carinatus Hook. & Am.
Bromus inermis Leyss.
0.0
3.3
-1.1
-27.8
33.3
-11.1
-1.1
3.3
0.0
23.3
6.7
-3.3
40.0
30.0
20.0
-6.7
0.0
3.3
13.3
-13.3
80,0
10.0
13.3
0.0
0.0
-23.3
23.3
20.0
-26.7
-3.3
0.0
0.0
3.3
-16.7
66.7
10.0
-43.3
-10.0
0.0
0.0
-36.7
3.3
46.7
6.7
0.0
-1.1
3.3
-2.2
-28.9
-12.2
40.0
-16.7
-4.4
0.0
0.0
C'arex concinno,des Mackenzie
Carex geyer/ Boott
Ca/amagrostis rubescens Buckl.
Elymus glaucus Buckl.
Festuca occidental/s Hook.
Luzu/a campestris (L.) DC.
0,0
3.3
10.0
-33.3
6.7
60.0
6.7
-13.3
-38.9
4.4
62.2
-21.1
-5.6
-16.7
-46.7
25.6
-30.0
-43.3
-14.4
-36.7
-16.7
-32.2
73.3
-20.0
46.7
-56.7
-26.7
3.3
-40.0
-33.3
0.0
6.7
10.0
36.7
-40.0
50.0
-63.3
-20.0
6.7
-56.7
-36.7
-10.0
-36.7
0.0
42.9
13.3
3.3
-46.7
-13.3
-20.0
-26.7
-13.3
-26.7
-20.0
-20.0
-30.0
0.0
-5.6
-1.1
0.0
3.3
-36.7
-40.0
-16.7
-16.7
3.3
16.7
-10.0
0.0
-10,0
6.7
-6.7
-10.0
-50.0
0.0
-16.7
16.7
0.0
-23.3
-13.3
43,3
-38.9
-23.3
-4.4
-63.3
0.0
-3.3
-38.9
2.2
-33.3
-6.7
-13,3
-13.3
-13.3
-36.7
0.0
-10.0
-36.7
0.0
26.7
-22.2
133
-36.7
-18.9
-58.9
0.0
-21.1
-13.3
0.0
-47.8
4.4
-60.0
-70.0
-63.3
-50.0
-80.0
-53.3
-50.0
-56.7
-50.0
0.0
33.3
0.0
53.3
0.0
23.3
0.0
10.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.3
0.0
3.3
Site 1
Mel/ca bu/bosa
Geyer ex Porter &
Cout.
Poapratensis L.
Tilsetum canescens Buckl.
Ac/i/flea mil/efollurn L.
Anemone pieri Britt. ex Rydb.
Am/ca cord/lot/a Hook.
Eurybia consp/cua (LindL) Nesom
Fragaria spp.
Gal/urn spp
Hierac/um a/b/forum Hook.
Lathyrus spp.
Lup/nus spp.
Moehririgfa macrophy/la (Hook.)
Fenzl
Seneclo integerr/mus N utt.
Ste//ar/a Iongi;oes Goldie
-44.4
3.3
3.3
'2,2
-8.9
18.9
-21.1
-20.0
-12.2
3-3
00
17.8
Table 1. continued.
Cearcut
Gray
Viola adunca Sm.
Mahoniarepens(LindL) G. Don
SpiraeabetullfollaPaUas
Symphoricarposa/bus(L.) Biake
Thinned Overstory
CEXC
TEXC
GRAZED
GRAZED
-3.3
-15.6
-3.3
-10.0
0.0
0.0
-13.3
-6.7
-3.3
-30.0
-11.1
6.7
3.3
-10.0
0.0
3.3
-5.6
3.3
3.3
-10.0
0.0
-23.3
-10.0
0.0
-22.2
36.7
-13.3
6.7
-23.3
-6.7
0.0
30.0
13.3
-26.7
-4.4
-4.4
15.6
3.3
-6.7
10.0
16.7
-6.7
23.3
10.0
33.3
-3.3
0.0
-26.7
-4.4
L0.0
0.0
0.0
10.0
10.0
0.0
-3.3
0.0
0.0
3.3
3.3
0.0
16.7
13.3
0.0
16.7
5.7
0.0
-8.9
Taraxacurnoffidna/eG.H. Weber
cx Wiggers
ThallctrumfendlerfEnge!m. ex
Contro' Overstoj
CEXC
TEXC
GRAZED
TEXC
CEXC
-36.7
20.0
13.3
1.1
Site 2
BromuscarinatusHook. &Arn.
6.7
5rornuinerrnILeyss.
00
'carex concinnaide9 Mackenzie
26.7
carexgeyeriBooi
0.0
ca/amagrostLcrubescensBucki.
EIymusgfaucusutki.
Festuca occidental/sHook.
Luzula campstrLc(L) DC,
Mel/ca bulbosaGeyer ex Porter &
Couft.
PoapratensisL.
TrLcetum canescerzsBuckl.
Ac/if/lea mil/efoliurn L.
Anemone pioeri Britt. ex Rydb.
Am/ca cordifolia Hook.
Eutybia conspicua (LindL) Nesom
Fragarfaspp.
Galiumspp
Hieracium a/b/forum Hook.
00
10.0
0.0
18.1
48.5
-0.1
58.8
-10.8
-3.3
-3.3
-26.7
-16.7
-23.3
-26.7
16.7
3.3
22.5
-13.9
0.0
-23.3
-3.3
3.3
0.0
0.0
15.6
26.7
-10.0
16.7
30.0
10.0
26.7
-43.3
-6.7
0.0
11.1
-6.7
-3.3
21.1
16.7
26.7
13.3
3.3
20.0
46,7
0.0
14.4
26.7
-6.7
-1.1
10.0
6.7
13.3
0.0
26.7
30.0
6.7
-3.3
26.7
2.1
16.7
-20.0
20.0
-13.3
-16.7
-16.7
0.0
-33.3
0.0
-33.3
3.3
-10.0
-38.1
-0.7
-18.5
3.3
0.0
1.3
-53.3
3.3
-6.7
3.3
-8.1
13.3
-6.7
0.0
0.0
10.0
0.0
13.3
-6.7
21.1
50.0
-16.7
-2.2
-21.1
-36.7
-53.3
-32.2
3.3
'8.9
1.1
-2.2
'0.0
-23.3
3.3
-30.0
-3.3
0.0
13.3
6.7
-3.3
16.7
-26.7
-6.7
-30.0
43.3
-26.7
-53.3
0.0
-3.3
-6,7
-3.3
-25.6
27.8
-8.9
-11.1
-3.3
-5.6
-16.7
0.0
-6.7
5&
-2.2
-15.6
0.0
8.9
40.0
13.3
23.3
Table 1. continued.
Lathyrus spp.
Lup/nus spp.
Moeliringia macrc'phylla (Hook.)
Fenzl
Senecic integerrimus Nutt.
Stellar/a longies Goldie
Taraxacum officinale G .H. Weber
ex Wiggers
Tha/ictrum fend/en Engeim. ex
Gray
Viola adunca Sm.
Mahonia repens (Lindi.) G. Don
Spiraea betulifolia Paltas
Symohonicorpos a/bus (L) B'ake
Clearcut Overstory
CEXC
TEXC
GRAZED
Contro' Overstory
CEXC
TEXC
GRAZED
-40.0
-42.8
-46.7
-6.7
26.7
-7.8
-3.3
-36.7
-41.1
0.0
-6.9
-3.3
-20.0
3.3
-6.7
-3.3
3.3
11.1
-33.3
-42.4
-33.3
-26.7
-60.0
-52.2
-20.0
-73.3
-40.0
Thinned Overstory
CEXC
TEXC
GRAZED
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.4
0.0
-3.3
0.0
0.0
0.0
3.3
0.0
0.0
-1.0
0.0
0.0
0.0
6.7
0.0
0.0
-1.1
-40.0
-21.9
-3.3
0.0
-6.7
2.2
-16.7
-36.7
8.9
-6.7
27.1
56.7
-13.3
-16.7
-3.3
6.7
-6.7
10.0
-3.3
-1.5
-3.3
0.0
-6.7
-2.2
-13.3
10.0
-3.3
-16.7
-23.9
-46.7
-20.0
-10.0
-1.1
6.7
-6.7
-6.7
20.0
12.6
3.3
13.3
-3.3
-2.2
-6.7
-3.3
20.0
0.0
6.7
13.3
3.3
0.0
6.7
0.0
0.0
0.0
6.7
0.0
10.0
0.0
0.0
3.3
0.0
3.3
0.0
-3.3
2.8
-13.3
0.0
0.0
2.2
20.0
3.3
20.0
-40.0
12.2
-30.0
6.7
13.3
-11.1
23.3
6.7
2.2
-13.3
8.5
20.0
23.3
16.7
15.6
30.0
33.3
22.2
30.0
26.3
23.3
16.7
33.3
23.3
46.7
30.0
40.0
-36.7
-25.8
-6.7
20.0
-13.3
-13.3
20.0
23.3
0.0
-6.7
3.2
-6.7
16.7
0.0
-3.3
0.0
3.3
-1.1
-10.0
-17.2
-33.3
-36.7
-23.3
-14.4
0.0
-10.0
-10.0
100.0
72.6
80.0
3.3
0.0
27.8
16.7
3.3
6.7
30.0
-21.0
-46.7
-16.7
-3.3
-2.2
-10.0
-10.0
-8.9
-10.0
35.7
-13.3
6.7
0.0
-1.1
26.7
10.0
30.0
Site 3
Bromus carinatus Hook. & Am.
Bromus inermis Leyss.
Carex concinno/des Mackenzie
Carexgeyeni Boott
Calamagro5t/s rubescens Buckh
Elymus glaucus Buckl.
Festuca occidentali Hook.
Luzula campestris(L.) DC.
Me//ca bulbosa Geyer ex Porter &
Couft.
Poa pratensis L.
Tnisetum canescens Bucki.
A chit/ca millefollum L.
'.0
Table 1. continued.
Anemone pieriBritt. ex Rydb.
Am/ca card/fo/la Hook.
Euryb/aconsp/cua(Lindl.) Nesom
Fragariaspp.
Gallumspp
/-fieracium albifforum Hook.
Lathyrusspp.
Lupinuss.
Moehr/ng/amacrophy/Ia(Hook.)
Fenzl
Sencio/ntegerrimusNutt.
Ste//ar/a Iongies Goldie
Tdraxa-um off,c r,d e G i-I Weber
ex Wiggers
Thalictrumfepd/erfEngelm. cx
Clearcut Overstory
CEXC
TEXC
GRAZED
-10.0
-11.8
-30.0
-40.0
-39.6
-36.7
0.0
-1.3
0.0
-3.3
-12.2
-53.3
0.0
-0.3
0.0
-13.3
-13.5
-13.3
-36.7
-29.4
-70.0
33.3
2.8
10.0
CEXC
-13.3
-20.0
0.0
-3.3
0.0
-23.3
6.7
0.0
3.3
10.0
0.0
17.8
-31.1
33.7
]Thinned Overstory
CEXC
TEXC
GRAZED
-16.7
3.3
-3.3
-10.0
23.3
11.1
3.3
3.3
5.6
0.0
-13.3
-4.4
0.0
0.0
3.3
0.0
-3.3
-2.2
-16.7
-3.3
-41.1
0.0
6.7
42.2
-10.0
-31.8
-46.7
-30.0
-40.0
-37.8
-26.7
-23.3
-36.7
0.0
6.7
0.0
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-6 7
-4 3
-6 7
00
-6 7
20 0
67
00
11
-6.7
-39.4
-70.0
-16.7
-30.0
-5.6
-23.3
6,7
-1(3.0
Gray
Viola adunca Sm,
-3.3
Mahoniarepens(Lindl.) G. Don
0.0
-4.6
-3.3
0.3
3.3
5p/raea betu/ifolia Pallas
-26.7
-16.7
-20.0
S,vmphoricarpos albus(L.) Blake
13.3
9.0
30.0
Herbivory treatments: Graze cattle and big game grazing; CEXC
exclusion of cattle and big game grazing.
control Overstory
TEXC
GRAZED
-16.7
8.9
-20.0
5.6
0.0
-2.2
6.7
-3.3
3.3
-2.2
-10.0
-3.3
22.2
3.3
3.3
11.1
0.0
-3.3
0.0
-6.7
-6.7
5.6
0.0
-6.7
-8.9
3.3
-20.0
15.6
6.7
10.0
-1.1
0.0
10.0
0.0
cattle exclosure, big game grazing only; TEXC total exclosure,
I-
Table 2. The changes in spedes composition in a grand fir forest from 1985 to 1994 using a 30 cm x 60 cm plot frame.
Clearcut Overstory
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
Site I
Bromus carinatus Hook. & Am.
Bromus inermic Leyss.
Carex concinnoides Mackenzie
Carex geyeri Boott
Ca/amagrostLs rubescens Buckl.
Elymus glaucus Bucki.
Festuca occidentaIi Hook.
Luzufa campestris (L.) DC.
Me//ca bulbasa Geyer ex Porter &
Coult.
Poa pratensis L.
Trisetum canescens Buckl.
Ac/il/lea mi//efo//urn L.
Anemone pioeriBritt. ex Rydb.
Am/ca cord/b/la Hook.
Euiybla conspicua (Lindi.) Nesom
Fragarla spp.
Gal/urn spp
Hieracium a/b/forum Hook.
Lathyrus spp.
Lupinusspp.
Moehringia macrophy/la (Hook.)
Fenzl
Senec/o integerrimus Nutt.
Stellar/a Jongioes Goldie
0.0
3.3
3.3
-36.7
13.3
53.3
16.7
0.0
0.0
6.7
20.0
-30.0
10.0
60.0
10.0
-16.7
3.3
-4.4
-23.3
-17.8
43.3
-23.3
-23.3
-60.0
70.0
-23.3
50.0
-60.0
-36.7
-3.3
..333
-36.7
0.0
-10.0
10.0
40.0
-53.3
50.0
-83.3
-23.3
3.3
-56.7
-40.0
-16.7
-40.0
0.0
-70.0
0.0
43.3
3.3
0.0
3.3
-20.0
20.0
23.3
-26.7
-3.3
-3.3
0.0
6.7
-20.0
60.0
13.3
-56.7
-16.7
0.0
3.3
0.0
-27.8
38.9
-20.0
-30.0
-33.3
47.8
0.0
-43.3
-1.1
-3.3
0.0
3.3
6.7
-6.7
80.0
6.7
10.0
0.0
0.0
0.0
-55.6
-20.6
71.7
-31.1
1.1
3.3
0.0
16.7
3.3
3.3
50.0
16.7
20,0
-56.7
-21.1
-50.0
-20.0
-40.0
23.3
-30.0
-3.3
-3.3
26.7
-16.7
13.3
-40.0
-13.3
-16.7
-16.7
-6.7
-54.4
-24.4
-5.6
-58.9
0.0
-5.6
-48.9
4.4
-46.7
3.3
-16.7
-30.0
0.0
-10.0
-26.7
-6.7
0.0
-53.3
-23.3
-36.7
-26.7
-6.7
-26.7
-36.7
-33.3
-26.7
3.3
-73.3
-67.8
-66.7
0.0
63.3
0.0
36.7
0.0
10.0
-51.1
20.0
-1.1
-3.3
-11.1
3.3
0.0
-4.4
-3.3
3.3
3.3
10.0
0.0
-30.0
-16.7
35.0
-42.8
32.8
-31.7
-77.8
-2.2
-22.8
-20.0
-1.7
-57.2
6.7
-90.0
-63.3
-50.0
-63.3
-62.2
0.0
3.3
0.0
0.0
0.0
6.7
0.0
3.3
0.0
0.0
21.1
-12.2
-22.2
-1.1
G
-13.3
-23.3
-43.3
-3.3
-13.3
Table 2. continued.
çntrol Overstory
Clearcut Overstory
Taraxacum officinale G .H. Weber ex
Wiggers
Tha/ictrum fendleriEngelm. ex Gray
Viola adunca Sm.
Mahonia repens (Lindi.) G. Don
Spiraea betulifolia Pallas
Syrnphoricarpos a/bus (L.) Blake
Site 2
Bromus carinatus Hook. & Am.
Bromus inermis Leyss.
arex con cinnoides Mackenzie
Carex geyeri Boott
Ca/amagrostis rubescens Bucki.
Elyrnus g/aucus Bucki.
Festuca occidentaiLc Hook.
Luzula campestris(L.) DC.
Me//ca bulbosa Geyer ex Porter &
Coult.
Poa pratensLc L.
Thetum canescens Bucki.
Achi//ea mi//efo//um L.
Anemone pIoeriBritt. ex Rydb.
Am/ca cordifolia Hook.
Eutybia conspfcua (Lind!.) Nesom
Fragaria spp.
Gal/urn spp
/-/ierac/um a/b/florum Hook.
Lathyrus spp.
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
-20.0
-22.2
-13.3
-13.3
-1.1
0.0
-23.3
-10.0
-6.7
13.3
10.0
-36.7
13.3
-33.3
3.3
0.0
-16.7
20.0
-8.9
0.0
-22.2
32.2
6.7
-6.7
16.7
-30.0
-6.7
6.7
-10.0
30.0
6.7
-23.3
-13.3
-8.9
-5.6
-14.4
0.0
-3.3
-10.0
-20.0
20.0
3.3
-16.7
23.3
16.7
46.7
-10.0
13.9
-3.3
-30.6
-2.2
6.7
0.0
33.3
23.3
-3.3
40.0
33.3
3.3
16.7
0.0
13.6
-30.0
-5.7
53.8
-10.8
-5.8
13.3
10.0
33.3
-33.3
26.7
56.7
20.0
0.0
0.0
0.0
6.7
-6.7
23.3
20.0
-3.3
-3.3
10.0
0.0
3.3
-10.0
33.3
33.3
-6.7
13.3
0.0
0.0
6.7
12.2
24.4
30.0
-8.9
2.2
3.3
3.3
16.7
-20.0
10.0
53.3
10.0
0.0
13.3
0.0
26.7
-3.3
40.0
56.7
13.3
3.3
6.7
0.0
..144
-6.7
-36.0
-26.7
-36.7
-40.0
-28.9
-43.3
-60.0
-42.2
33.3
6.7
30.0
-26.7
-40.0
0.0
-30.0
6.7
-13.3
-46.7
22.2
-15.7
16.9
-14.3
26.7
-20,0
16.7
-16.7
-60.0
6.7
-26.7
0.0
-3.3
10.0
-33.3
-3.3
-6.7
0.0
0.0
-16.7
0.0
-10.0
23.3
8.9
-5.6
3.3
8.9
25.6
-4.4
-22.2
0.0
10.0
0.0
3.3
-23.3
3.3
-33.3
-3.3
0.0
13.3
6.7
-6.7
-10.0
26.7
-40.0
46.7
-30.0
-46.7
0.0
-10.0
-6.7
0.0
-36.7
2.2
-30.0
32.2
-13.3
-6.7
-3.3
2.2
24.4
32.2
-33.3
-41.4.
-1.0
-23.8
0.6
-9.7
-47.1
-1.1
3.3
-10.0
13.3
16.7
-53.3
16.7
0.0
-3.3
0.0
-3.3
-3.3
11.1
21.1
20.0
63.3
-17.8
-2.2
NJ
Table 2. continued.
Clearcut Overstory
Lupinusspp.
Moe/iringia ínacrophylla (Hook.)
Fenzl
Senec/ointegerrirnusNutt.
Stellar/a long,;oes Goldie
TaraxacumofficinaleG.H. Weberex
Wiggers
ThalictrumfeadleriEngelm. ex Gray
Viola adunca Sm.
Mahonia repens (Lindi.) G. Don
SpiraeabetullfoliaPallas
Syniphoricarposalbus(L.) Blake
Site 3
BromuscarinatusHook. &Arn.
5romusinern7iLeyss.
Carex concinnoides Mackenzie
CarexgeyeriBoott
Ca/aniagrosticrubescensBuckl.
fflyrnusglaucusBuckl.
FestucaocddentalLHook.
Luzulacampestr/s(L.) DC.
frle/icabu/bosaGeyerex Porter&
PoapratensisL.
Trisetum canescensBuckl.
Achileamlllefo/iurnL.
Anemone pieri Britt. ex Rydb.
Am/ca cord/fe/ia Hook.
ccntrol Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
3.3
-9.7
-6.7
-30.0
-43.3
-52.2
-50.0
0.0
0.0
0.0
-0.4
0.0
Thinned Overstory
CEXC
TEXC
GRAZED
6.7
GRAZED
-11.1
-3.3
0.0
13.3
-36.7
-66.7
-68.9
-23.3
-76.7
-52.2
0.0
0.0
0.0
-3.3
0.0
0.0
0.0
6.7
0.0
0.0
0.0
6.7
0.0
0.0
-1.4
0.0
0.0
0.0
6.7
0.0
-3.3
2.2
-43.3
-13.3
-6.7
0.0
23.3
-29.6
6.9
-0.8
3.3
63.3
-6.7
-70.0
-3.3
0.0
-20.0
0.0
-23.3
26.7
-10.0
-16.7
-6.7
-6.7
-3.3
1.1
-16.7
-5.6
2.2
-2.2
-16.7
0.0
-20.0
-10.0
-6.7
-40.0
-36.7
6.7
-20.0
-6.7
5.6
12.2
-4.4
-16.7
20.0
6.7
6.7
0.0
-36.7
-20.0
40.0
-40.0
-10.0
6.3
0.0
0.0
0.0
1.9
14.2
14.0
49.6
-33.2
5.0
16.7
20.0
-16.7
-50.0
23.3
33.3
-13.3
-6.7
-13.3
3.3
36.7
33.3
23.3
20.0
0.0
0.0
-3.3
13.3
26.7
50.0
-13.3
-3.3
0.0
3.3
6.7
10.0
40.0
0.0
0.0
23.3
7.8
23.3
-25.6
-5.6
0.0
0.0
20.0
36.7
43.3
63.3
20.0
6.7
40.0
30.0
3.3
46.7
-6.7
5.6
-10.0
-22.1
-40.0
-40.0
-33.3
-17.8
-6.7
-13.3
-15.6
93.3
33.3
-20.0
-13.3
-50.0
78.5
-22.6
48.9
-19.7
-44.4
83.3
-60.0
-16.7
-46.7
-40.0
6.7
-26.7
10.0
-16.7
-16.7
0.0
-16.7
0.0
-20.0
-16.7
27.8
-4.4
3.3
8.9
0.0
23.3
-10.0
30.0
-20.0
-6.7
3.3
-16.7
0.0
-10.0
33.3
16.7
-10.0
-35.1
26.4
6.7
6.7
-1.1
-11.1
11.1
21.1
34.4
-6.7
7.8
cA)
Table 2. continued.
Clearcut Overstory
Eurybiaconspicua(Lindl.) Nesom
Fragar/aspp.
G'alluinspp
Hieracium
a/bifloru,n
Lathyrusspp.
Lupinusspp.
Moehringiainacrophy//a(Hook.)
Fenzl
SeneciointegerrimusNutt.
Ste//aria longipes Goldie
Taraxacu,nofficina/eG.H.
Weberex
Wiggers
Thalictrum fend/eriEngelm. ex Gray
Viola aduncaSm.
Thinned Overstory
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
0.0
0.0
-40.0
0.0
-23.3
-80.0
16.7
0.0
-16.7
-6.7
-26.7
3.3
3.3
0.0
10.0
6.7
0.0
6.7
0.0
-2.2
-13.3
-26.7
-33.3
40.0
-1.4
-20.7
2.2
-19.2
-35.6
4.7
20.0
-22.2
3.3
6.7
-16.7
0.0
3.3
-13.3
3.3
-10.0
-39.4
-63.3
-33.3
-50.0
-47.8
-36.7
-30.0
-41.1
0.0
10.0
0.0
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-13.3
-2.4
-10.0
0.0
-6.7
20.0
-6.7
0.0
-13.3
0.0
Hook.
Control Overstory
CEXC
1.1
TEXC
3.3
-13.3
0.0
-3.3
0.0
13.3
GRAZED
4.4
-12.2
3.3
-3.3
-28.9
41.1
-2.2
-20.0
-47.9
-80.0
-20.0
-26.7
-13.3
-23.3
6.7
0.0
-6.7
-4.9
-3.3
-20.0
-10.0
26.7
10.0
3.3
24.4
Mahoniarepens(Lindl.)G. Don
-3.3
-1.7
6.7
0.0
-3.3
-1.1
-10.0
-6.7
4.4
Spiraeabetu/ifo/iaPallas
-43.3
-25.3
-26.7
23.3
0.0
-7.8
3.3
-10.0
12.2
Syinphoricarposa/bus(L.) Blake
13.3
3.8
36.7
6.7
13.3
0.0
3.3
10.0
0.0
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big game grazing only; TEXC
total exclosure,
exclusion of cattle and big game grazing.
Table 3 The chanoes in soecies comDosition in a grand fir forest from 1985 to 1997 using a 30 cm x 30 cm plot frame.
Clearcut Overstory
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
3.3
0.0
0.0
-56.7
0.0
30.0
0.0
6.7
23.3
0.0
0.0
-33.3
-3.3
10.0
-6.7
-13.3
28.9
0.0
-2.2
-38.9
-8.9
3.3
-16.7
0.0
0.0
0.0
0.0
23.3
-3.3
-46.7
-10.0
3.3
0.0
0.0
-14.4
-11.1
16.7
-4.4
-1.1
0.0
0.0
0.0
-20.0
10.0
3.3
-20.0
-6.7
-1.1
3.3
0.0
3.3
-20.0
-6.7
13.3
16.7
16.7
0.0
0.0
0.0
-20.0
-23.3
23.3
-6.7
3.3
10.0
0.0
0.0
-55.6
4.4
22.2
-24.4
-5.6
167
-46.7
-25.6
-30.0
-43.3
-14.4
-36.7
-16.7
-32.2
76.7
Trisetum canescens Buckl.
-26.7
Achf/IeamillefolfumL.
33.3
Anemone pi;oeriBritt. ex Rydb.
-50.0
Am/ca cord/Ia//a Hook.
-23.3
Euiybiaconspicua(Lindl.) Nesom
-3.3
Fragar/aspp.
-50.0
6a1/umspp
-33.3
Hierac/um a/b/forum Hook.
0.0
52.2
-40.0
37.8
-44.4
-26.7
-4.4
-53.3
-10.0
-6.7
-18.9
12.2
30.0
-33.3
-13.3
-10.0
3.3
-10.0
-30.0
26.7
-10.0
-16.7
0.0
-6.7
-43.3
-13.3
6.7
-13.3
-3.3
-26.7
-10.0
-16.7
-20.0
0.0
13.3
-25.6
-12.2
-2.2
18.9
-14.4
-3.3
27.8
-6.7
-8.9
-3.3
0.0
-13.3
16.7
-16.7
0.0
-23.3
0.0
-6.7
-13.3
-13.3
33.3
-32.2
-1.1
30.0
-3.3
0.0
-30.0
0.0
-13.3
-10.0
23.3
0.0
-3.3
0.0
-22.2
-25.6
0.0
-34.4
-6.7
-10.0
-7.8
4.4
Sate 1
BromuscarfnatusHook.&Arn,
BromusinermiLeyss.
CarexconcinnoidesMackenzie
CarexgeyeriBoott
Ca/aniagrostisrubescensBuckl.
E/ymusglaucusBuckl.
Festucaoccidenta/iHook.
Luzula ca,npestris(L.) DC.
Mefica bu/bosa Geyer ex Porter &
Coult.
PoapratensisL.
Lathyrusspp.
-10.0
Lup/nusspp.
10.0
53.3
-46.7
20.0
-60.0
-23.3
0.0
-76.7
-56.7
-10.0
-23.3
0.0
-60.0
-66.7
-60.0
-10.0
-50.0
-6.7
-20.0
-40.0
-43.3
0.0
20.0
0.0
33.3
0.0
53.3
0.0
20.0
0.0
3.3
0.0
13.3
0.0
23.3
0.0
16.7
0.0
26.7
Moehring/amacrophylla(Hook.)
Seneciointegerr/musNutt.
5tellarh/ongiaesGoldie
11.1
I-'
Ui
Table 3. continued.
Clearcut Overstory
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
0.0
-6.7
-15.6
-3.3
-3.3
0.0
3.3
-13.3
-6.7
67
-30.0
-11.1
0.0
6.7
-10.0
-3.3
0.0
-5.6
-20.0
-3.3
-36.7
10.0
-20.0
0.0
-30.0
10.0
-16.7
0.0
-32.2
30.0
-16.7
6.7
-13.3
13.3
0.0
16.7
10.0
-3.3
-11.1
-3,3
-6.7
-3.3
-3.3
0.0
0.0
10.0
-10.0
3.3
30.0
20.0
-11.1
16.7
0.0
23.3
3.3
3.3
13.3
-3.3
0.0
10.0
0.0
28.1
2.5
-3.3
33.3
0.0
30.0
-23.3
16.7
26.7
-6.7
0.0
0.0
0.0
-3.3
-3.3
43.3
0.0
0.0
-3.3
0.0
0.0
-16.7
13.3
26.7
6.7
13.3
-6.7
0.0
0.0
0.0
-8.9
24.4
0.0
30.0
-4.4
0.0
0.0
-3.3
-26.7
33.3
26.7
6.7
0.0
6.7
0.0
0.0
-3.3
30.0
36.7
3.3
-10.0
13.3
0.0
-8.9
-5.6
27.8
26.7
-6.7
-2.2
-3.3
-26.7
-16.7
-23.3
-26.7
-21.1
-36.7
-53.3
-32.2
29.2
22.8
Anemone pioeriBritt. ex Rydb.
Am/ca cordifolla Hook.
Euiyb/a conspicua(Lindl.) Nesom
Fragar/aspp.
Ga/iumspp
Hieracium a/b/forum Hook.
-13.3
-16.7
3.3
-16.7
-3.3
-13.3
-16.7
23.3
-10.0
26.7
-13.3
-60.0
0.0
-3.3
6.7
0.0
0.0
-20.0
-3.3
0.0
33.3
0.0
0.0
-3.3
-6.7
3.3
36.7
0.0
-6.7
0.0
0.0
-1.1
Ac/i//lea mi//ePa//urn L.
10.0
0.0
26.7
0.0
0.0
6.7
-13.3
-10.0
6.7
33.3
-33.3
46.7
-23.3
-60.0
0.0
-10.0
-3.3
-10.0
27.8
-22.2
27.8
-12.2
Taraxacumoffic/na/eG.H. Weber
ex Wiggers
ThallctrurnfendleriEngelm. ex
Gray
Viola aduncaSm.
Mahoniárepens(LindI.)G. Don
Spiraeabetu/ifollaPallas
Sy,nphoricarposalbus(L.) Blake
Site 2
Bromus cariiatus Hook, &Arn.
BromusinermLsLeyss.
Carex concinno/des Mackenzie
CarexgeyeriBoott
Ca/amagrostfsrubescensBuckl.
Elymusg/aucusBuckl.
Festuca occidental/s Hook.
Luzulacampestris(L.) DC.
Meilca bulbosaGeyer ex Porter &
Couft.
PoapratensisL.
TrisetumcanescensBuckl.
-21.8
19.9
22.1
22.1
-38.1
-0.7
4.9
-2.1
-8.1
10.0
0.0
0.0
-4.4
-2.2
-2.2
-5.6
2.2
-4.4
26.7
1.1
-12.2
0.0
2.2
-16.7
0.0
-3.3
-21.1
-3.3
-5.6
0.0
-10.0
a'
Table 3. continued.
Clearcut Overstory
Lathyrus spp.
Lupinus spp.
Moebringia macrophy/fa (Hook.)
Fenzl
Senecio integerrfrnus Nutt.
Ste//aria longipes Goldie
Taraxacum officinale G.H. Weber
ex Wiggers
Tha//ctrum fendleriEngelm. ex
Gray
Viola adunca Sm.
Ma/ion/a repens (Lindi.) G. Don
Sp/raea betuI/fo/h PaUas
Symphoricarpos a/bus (L.) Blake
Site 3
Brotnus carinatus Hook. & Am.
Bromus inermis Leyss.
Carex concinno/des Mackenzie
Carex geyer/ Boott
Ca/amagrost/s rubescens Bucki.
E/ymus g/aucus Bucki.
Festuca occ/denta/i Hook.
Luzula campestric(L.) DC.
Me//ca bulbosa Geyer ex Porter &
Co u It.
Poa pratensL L.
Tri'setuni canescens Bucki.
Ac/i/flea ini//efo/fum L.
ntrol Overstory
CEXC
TEXC
GRAZED
CEXC
-43.3
0.0
-42.8
-6.9
-40.0
-3.3
-16.7
-26.7
-20.0
-42.4
-26.7
0.0
0.0
0.0
-0.4
-3.3
TEXC
3.3
Thinned Overstory
GRAZED
CEXC
TEXC
GRAZED
0.0
-4.4
0.0
3.3
0.0
-33.3
6.7
-34.4
17.8
26.7
6.7
7.8
-10.0
-70.0
-26.7
0.0
0.0
0.0
-3.3
0.0
3.3
0.0
0.0
0.0
0.0
0.0
6.7
0.0
6.7
-1.0
0.0
0.0
0.0
0.0
0.0
0.0
-1.1
-46.7
-25.3
-13.3
3.3
-3.3
8.9
-13.3
-30.0
-17.8
20.0
-3.3
0.0
30.0
40.4
-4.9
-17.2
12,6
53.3
0.0
-40.0
30.0
-13.3
0.0
-23.3
10.0
-20.0
-10.0
-6.7
0.0
-13.3
-2.2
2.2
-5.6
6.7
-10.0
-3.3
-10.0
6.7
3.3
-16.7
0.0
-13.3
-3.3
-16.7
26.7
10.0
0.0
-16.7
-23.3
-3.3
13.3
-53.3
-3.3
13.3
0.0
-7.2
2.2
3.3
0.0
-3.3
3.3
23.3
0.0
-16.7
-3.3
0.0
0.0
6.7
0.0
33.3
0.0
-26.7
-6.7
0.0
0.0
-11.1
-17.8
8.9
0.0
-10.0
0.0
3.3
0.0
0.0
10.0
50.0
36.7
-10.0
0.0
0.0
0.0
-3.3
-23.3
16.7
16.7
20.0
10.0
0.0
0.0
23.3
32.9
-25.8
-3.5
0.0
0.0
-13.3
-43.3
10.0
23.3
-3.3
-6.7
-10.0
-17.2
-33.3
-36.7
-23.3
-14.4
0.0
-io.o
-10.0
100.0
-16.7
36.7
99,3
-21.0
25.7
93.3
-40.0
30.0
3.3
-10.0
10.0
0.0
0.0
3.3
-2.2
-2.2
-4.4
0.0
23.3
20.0
0.0
3.3
3.3
26.7
5.1
-11.1
38.9
6.7
-13.3
-1.1
21.1
20.0
I-'
e 3. continued.
Clearcut Overstory
CEXC
Anemone piperiBritt. ex Rydb.
Arnicacordifo/iaHook.
Eurybiaconspicua(Lindl.) Nesom
Fragariaspp.
Gallumspp
Hieracium albiflorum Hook.
Lat/iyrusspp.
Lupinusspp.
Moehringia macrophylla (Hook.)
Fenzl
SeneciointegerrimusNutt.
Stellar/a longiesGoldie
TaraxacumofflcinaleG.H. Weber
ex Wggers
ThalictrumfendleriEngelm. ex
Gray
Viola adunca Sm.
-3.3
-43.3
-3.3
TEXC
-15.1
Control Overstory
GRAZED
CEXC
0.0
-6.7
0.0
-13.3
0.0
-20.0
-10.0
0.0
TEXC
0.0
Thinned Overstory
GRAZED
CEXC
TEXC
GRAZED
15.6
31.9
10.0
46.7
0.0
-23.3
0.0
-16.7
-20.0
-3.3
-3.3
7.8
2.2
0.0
-10.0
6.7
3.3
-6.7
0.0
-3.3
-6.7
0.0
3.3
0.0
-13.3
-33.3
13.3
-18.9
-0.3
-10.1
-46.1
19.4
-30.0
-53.3
0.0
-50.0
0.0
-20.0
-60.0
16.7
-10.0
-31.8
-43.3
-13.3
-6.7
8.9
-16.7
13.3
-16.7
0.0
6.7
0.0
6.7
0.0
20.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.3
-10.0
-4.3
-6.7
0.0
-6.7
0.0
-6.7
-3.3
-1.1
-13.3
-42.8
-73.3
-16.7
-6.7
11.1
-16.7
-6.7
0.0
-39.6
2.1
26.7
0.0
13.3
0.0
-3.3
13.3
0.0
-2.2
-13.3
-2.2
1.1
-1.1
-11.1
0.0
-8.9
-27.8
42.2
-4.6
3.3
-13.3
-3.3
-7.8
10.0
.3.3
4.4
0.3
0.0
0.0
3.3
-3.3
-6.7
-6.7
12.2
SpiraeabetulifoliaPallas
-20.0
-23.3
-3.3
-20.0
-12.2
0.0
-23.3
15.6
Symphoricarposalbus(L.) Blake
25.7
16.7
16.7
3.3
-1.1
3.3
10.0
6.7
Herbivory treatments: Graze cattle and big game grazing; CEXC cattle exclosure, big game grazing only; TEXC total exclosure,
exclusion of cattle and big game grazing.
Mahoniarepens(Lindl.)G. Don
10.0
0.0
-26.7
10.0
03
Table 4. The chancies in sDecies comDosition in a grand fir forest from 1985 to 1997 using a 30 cm x 60 cm plot frame.
Clearcut Overstory
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
Site 1
Bromuscar/natusHook. &Arn.
BromusfnernifsLeyss.
10.0
0.0
0.0
-53.3
0.0
30.0
0.0
6.7
33.3
0.0
0.0
-40.0
-3.3
10.0
-6.7
-16.7
38.9
0.0
-4.4
-50.0
-14.4
0.0
-23.3
-2.2
3.3
0.0
0.0
-20.0
20.0
6.7
-13.3
-6.7
-3.3
0.0
0.0
-6.7
20.0
0.0
-56.7
-16.7
2.2
0.0
0.0
-16.7
2.2
22.2
-13.3
-23.3
-60.0
-30.0
-33.3
80.0
-30.0
43.3
-56.7
-23.3
50.0
-63.3
23.3
-83.3
-30.0
44.4
-54.4
42.2
-57.8
-27.8
10.0
-26.7
-23.3
-33.3
-10.0
-3.3
Fragariaspp.
&allu,nspp
Hieracium a/b/forum Hook.
Lat/iyrusspp.
Lupinusspp.
Moeliringia macrophylla (Hook.)
-56.7
-40.0
SeneciointegerriniusNutt.
Carexconc/nno/desMackenzie
CarexgeyeriBoott
Ca/amagrostirubescensBuckl.
E/ymusg/aucusBuckl.
Festuca occidental/s Hook.
Luzulacampestris(L.) DC.
Meilca bulbosa Geyer ex Porter
& Coult.
PoapratensisL.
TrLs-etumcanescensBuckl.
Achilleamil/efo/iurnL.
Anemone pieri Britt. ex Rydb.
Am/ca cordifolia Hook.
Euiybhconspicua(Lindl.)
Nesom
Stellar/a IongioesGoldie
0.0
0.0
3.3
-23.3
-23.3
20.0
13.3
0.0
0.0
-62.2
-6.7
-31.1
1.1
3.3
0.0
3.3
-10.0
-6.7
20.0
23.3
20.0
0.0
-6.7
-56.7
-21.1
-50.0
-20.0
-40.0
10.0
-33.3
-10.0
0.0
14.4
30.0
-16.7
-3.3
-3.3
33.3
-31.1
10.0
10.0
-16.7
-53.3
-20.0
0.0
-6.7
-30.0
6.7
-13.3
13.3
-13.3
-26.7
-17.8
-5.6
-13.3
-3.3
-12.2
-13.3
-3.3
-2.2
3.3
-23.3
10.0
-76.7
-56.7
-16.7
-20.0
0.0
-52.2
-6.7
-8.9
-28.9
21.1
-33.3
36.7
-10.0
-10.0
-3.3
-13.3
-13.3
-30.0
-13.3
0.0
-5.6
28.9
-3.3
-14.4
-4.4
-6.7
26.7
-13.3
10.0
10.0
-16.7
3.3
-10.0
-23.3
-16.7
-34.4
-3.3
-16.7
-18.9
3.3
-66.7
-66.7
-64.4
-6.7
-50.0
0.0
-10.0
-46.7
-45.6
0.0
30.0
0.0
46.7
0.0
66.7
0.0
20.0
0.0
6.7
0.0
33.3
0.0
36.7
0.0
23.3
0.0
30.0
1.1
23.3
21.1
Tabie 4. continued.
Clearcut Overstory
Taraxacumofficina/eG.H.
Weber ex Wiggers
ThallctrumfendlerfEngelm. ex
Gray
Viola adunca Sm.
Ma/ioniarepens(Lindl.) G. Don
SpiraeabetulifoliaPalias
Symphoricarposa/bus(L.) Blake
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
6.7
-20.0
-22.2
-13.3
-3.3
5.6
3.3
-23.3
-3.3
-13.3
-33.3
-12.2
0.0
6.7
-10.0
-3.3
-3.3
-10.0
-20.0
-3.3
-33.3
-6.7
-20.0
0.0
-33.3
0.0
-14.4
0.0
-35.6
28.9
-16.7
10.0
-33.3
16.7
-6.7
20.0
23.3
0.0
-15.6
-5.6
-5.6
-3.3
-10.0
-10.0
16.7
-13.3
6.7
33.3
33.3
-4.4
-6.7
-8.9
-8.9
16.7
0.0
23.3
3.3
-10.0
10.0
-6.7
3.3
10.0
0.0
36.9
-23.3
17.6
23.8
-4.2
36.7
0.0
33.3
3.3
24.4
-7.8
3.3
0.0
-3.3
-36.7
26.7
33.3
6.7
0.0
10.0
0.0
-6.7
-20.0
30.0
40.0
-58
0.0
0.0
-20.0
3.3
33.3
6.7
10.0
-6.7
0.0
0.0
0.0
-17.8
16.7
33.3
-6.7
0.0
0.0
0.0
-10.0
-3.3
33.3
0.0
10.0
-13.3
-16.7
13.3
0.0
-7.8
-5.6
33.3
33,3
-14.4
-2.2
-6.7
-36.0
-26.7
-36.7
-40.0
-28.9
-46.7
-60.0
-42.2
10.0
3.3
46.7
-20.0
2.3
-13.3
26.7
-20.0
-70.0
0.0
-26.7
-3.3
-3.3
36.7
0.0
-3.3
-6.7
-3.3
36.7
-1.1
-12.2
0.0
-7.8
32.2
0.0
-3.3
-23.3
28.9
24.3
33.6
-24.3
-44.7
-16.7
-13.3
40.0
-43.3
50.0
-26.7
-56.7
25.6
-30.0
35.6
-3.3
-3.3
33
-1.0
0.0
0.0
0.0
-4.4
10.0
0.0
-3.3
-6.7
-3.3
6.2
-2.8
3.3
6.7
0.0
0.0
-6.7
3.3
-15.6
0.0
-16.7
-3.3
0.0
-7.8
7.8
-1.1
Site 2
BromuscarinatusHook. &Arn.
BromusinermiLeyss.
CarexconcinnoidesMackenzie
CarexgeyeriBoott
ca/amagrostisrubescensBuckl.
E/ymusglaucusBuckl.
FestucaoccidentaIiHook.
Luzulacampestris(L.) DC.
Mefica bu/bosa Geyer ex Porter
& Cou!t.
PoapratensisL.
TrisetumcanescensBuckl.
Achil/eamillefollumL.
Anemone pier/Britt. ex Rydb.
Am/ca cordifolla Hook.
Euiybiaconsp/cua(Lindl.)
Fragariaspp.
Gal/urn spp
-16.7
31.1
10.0
3.3
.57
Tab'e 4. continued.
Clearcut Overstory
CExC
Hieracium a/b/forum Hook.
Lathyrusspp.
Lup/nusspp.
Moehringiamacrophylla(Hook.)
Fenzl
SeneciointegerrimusNutt.
Ste//aria /ongioes Goldie
Taraxacumofflcina/eG.H.
Weber ex Wiggers
Thalictrurn fendleriEngelm. ex
Gray
Viola adunca Sm.
Mahoniarepens(Undl.) G. Don
SpiraeabetulifoliaPallas
Syrnphoricarpc'safbus(L.) Blake
Site 3
Bromus car/natus Hook. &Arn.
BromusinerinisLeyss.
CarexconcfnnofdesMackenzie
CarexgeyeriBoott
Ca/amagrostirubescensBuckl.
E/yinusg/aucusBuckl.
Festuca occidental/sHook.
Luzu/acampestrLc(L.) DC.
Me/k'abu/bosaGeyer
exPorter
PoapratensisL.
TrfsetutncanescensBuckl.
TEXC
-13.1
Control Overstory
Thinned Overstory
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-50.4
-9.7
-3.3
-46.7
-6.7
-3.3
-10.0
-36.7
3.3
10.0
0.0
3.3
3.3
-4.4
-3.3
3.3
6.7
-20.0
-43.3
13.3
-7.8
-33.3
20.0
-26.7
-48.9
-43.3
26.7
13.3
4.4
-6.7
-66.7
-18.9
0.0
0.0
0.0
-0.4
0.0
0.0
0.0
-3.3
0.0
3.3
0.0
0.0
0.0
0.0
0.0
6.7
0.0
6.7
-3.3
-1.4
0.0
0.0
0.0
6.7
0.0
-3.3
2.2
-60.0
-32.9
-13.3
20.0
-3.3
11.1
-13.3
-26.7
-27.8
13.3
-6.7
-6.7
23.3
43.6
-7.5
-21.8
9.7
70.0
-3.3
-53.3
16.7
-20.0
0.0
-23.3
16.7
-16.7
-10.0
-10.0
0.0
-26.7
-5.6
2.2
6.7
-13.3
0.0
-6.7
-16.7
3.3
-23.3
3.3
-11.1
133
-33.3
-23.3
-13.3
20.0
-56.7
-10.0
12.9
0,0
-14.7
-2.5
4.0
32.9
-33.2
-5.0
3.3
0.0
-16.7
-60.0
13.3
26.7
-6.7
-6.7
0.0
0.0
-16.7
-10.0
23.3
3.3
-16.7
-3.3
0.0
0.0
3.3
0.0
33.3
0.0
-20.0
-10.0
0.0
0.0
-17.8
-24.4
-15.6
-2.2
6.7
0.0
0.0
16.7
46.7
43.3
-16.7
0.0
0.0
0.0
-3.3
-16.7
23.3
16.7
20.0
10.0
0.0
0.0
20.0
-12.2
34.4
10.0
-23.3
-ic.o
-22.1
-40.0
-40.0
-33.3
-17.8
-6.7
-13.3
-15.6
93.3
-23.3
98.5
-29.3
96.7
-46.7
3.3
-16.7
0.0
-6.7
-2.2
-7.8
0.0
30.0
0.0
0.0
30.0
20.0
-20.0
-53.3
0.0
0.0
1.1
11.1
3.3
-4.4
-23.3
30.0
-1,1
Table 4. continued.
CIearcutOversty
Ac/il/lea mi//efo//umL.
Anemone pieriBritt. ex Rydb.
Am/ca cord/folio Hook.
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
50.0
-6.7
-50.0
35.6
10.0
0.0
-6.7
10.0
10.0
20.0
-6.7
15.6
30.0
36.7
-13.3
10.0
0.0
-23.1
-41.1
33.3
-46.7
-63.3
3.3
50.0
11.1
19
0.0
0.0
0.0
-2.2
6.7
0.0
1.1
-6.7
0.0
-26.7
-30.0
13.3
-17.4
2.2
-15.8
-52.2
21.4
-43.3
0.0
-30.0
-70.0
23.3
-16.7
-6.7
-23.3
-16.7
0.0
6.7
0.0
-6.7
20.0
0.0
-26.7
-2.2
3.3
-2.2
0.0
-20.0
0.0
-3.3
-6.7
3.3
-26.7
0.0
-20.0
-23.3
6.7
-12.2
0.0
-10.0
-18.9
-10.0
-394
-60.0
-6.7
-16.7
-1.1
-13.3
20.0
-2i.1
0.0
13.3
0.0
13.3
0.0
33.3
0.0
0.0
0.0
0.0
0.0
3.3
0.0
3.3
0.0
0.0
0.0
6.7
-16.7
-5.7
.100
0.0
-6.7
0.0
-6.7
-3.3
-2.2
-26.7
-54.6
-80.0
-6.7
-3.3
6.7
-23.3
-13.3
3.3
16.7
0.0
-8.2
-1.7
-28.6
Nesom
Gal/urn spp
Hieracium albifloruin Hook.
Lathyrusspp.
Lupinusspp.
Macbring/a macrophy/la (Hook.)
Fenz
Senecio/ntegerrimusNutt.
Stellar/a longiesGoldie
Taraxacumoffldiia/eG.H.
Weber cx Wiggers
T/iallctrumfendieriEngelm. ex
Gray
Viola adunca Sm.
Th1nnethQvei:stiy
TEXC
Euiybiaconspicua(LindL)
Fragariaspp.
Control Overstory
CEXC
GRAZED
31.1
-6.7
41.1
6.7
-23.3
-3.3
-6.7
10.0
-13.3
21.1
Mahonfarepens(Lindl.)G. Don
0.0
0.0
0.0
-4.4
-10.0
-6.7
14.4
-43.3
Spiraea/ietulifoiaPallas
-33.3
0.0
13.3
-11.1
10.0
-23.3
15.6
Symphoricarposalbus(L.) Blake
20.0
27.1
26.7
10.0
6.7
-3.3
13.3
'10.0
10.0
Herbivory treatments: Graze cattle and big game grazing; CEXC c3ttle exclosure, big game grazing only; TEXC
total exclosure,
exclusion of cattle and big game grazing.
Table 5. The changes in species composition in a grand fir forest from 1985 to 2003 using a 30
Clearcut Overstory
ntrol Overstorv
cm x 30cpfrre.
Thinned Oversgj
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
33.3
3.3
6.7
3.3
10.0
30.0
-6.7
3.3
36.7
70.0
10.0
0.0
-3.3
40.0
-6.7
-13.3
28.9
36.7
3.3
16.7
0.0
16.7
-46.7
1O0
0.0
6.7
0.0
5.6
-7.8
40.0
-4.4
5.6
33.3
26.7
-2.2
-8.9
20.0
-16.7
-4.4
23.3
10.0
0.0
13.3
6.7
23.3
-26.7
-6.7
10.0
40.0
16.7
23.3
-20.0
73.3
3.3
00
38.6
23.3
0.0
-8.9
24.4
52.2
5.6
7.8
-13.3
-46.7
-25.6
10.0
26.7
45.6
-6.7
-16.7
-32.2
76.7
-26.7
60.0
-33.3
3.3
-3.3
-36.7
-33.3
0.0
53.3
-46.7
36.7
-50.0
-10.0
0.0
-63.3
-33.3
-10.0
-23.3
0.0
48.9
-40.0
-6.7
-50.0
-13.3
33.3
-6.7
-50.0
13.3
-12.2
-8.9
-27.8
-26.7
-4.4
-36.7
10.0
-6.7
-25.6
18.9
13.3
-43.3
-3.3
10.0
6.7
-13.3
-13.3
40.0
-6.7
0.0
-6.7
33
-6.7
-16.7
6.7
-20.0
-20.0
-16.7
0.0
-1.1
-16.7
-26.7
10.0
3.3
0.0
16.7
40.0
-33.3
6.7
46.7
-26.7
0.0
6.7
10.0
0.0
-16.7
-10.0
53.3
-47.5
34.4
-8.9
-52.2
0.0
-7.8
-10.0
-6.7
-4.4
4.4
-53.3
-46.7
-20.0
-56.7
-20.0
-6.7
0.0
-33.3
30.0
26.7
33.3
46.7
0.0
36.7
53.3
40.0
40.0
43.3
6.7
36.7
6.7
10.0
3.3
0.0
Site 1
Bromus carinatus Hook. &Arn.
BromusinermisLeyss.
carexconcinnoidesMackenzie
GarexgeyeriBoott
Ca/amagrostirubescensBuckL
E/yrnusglaucusBuckh
Festucaoccidenta/iHook.
Luzu/acarnpestris(L.) DC.
Me/icabuibosaGeyerex
Porter&
Coult.
Poapratens/cL.
TrisetumcanescensBuckl.
Achi//earn///efo//umL.
Anemone pioer/Britt. ex Rydb.
Arnicaco.rd/fo/iHook.
Eur,vbiáconspicua(UndL) Nesom
Fragarfaspp.
Ga/turn spp
1-/feracium a/b/forum Hook.
Lathyrusspp.
Lupinusspp.
Moe/7ring/a macrnp/iyl/a (Hook.)
SenedointegerriflusNutt.
Ste//ar/a /ongioes GoIdie
0.0
13.3
3.3
6.7
1.1
;1.1
-20.0
3.3
67
21.1
18.9
-34,4
-6.7
-2.2
6.7
1.1
.13.3
0.0
0.0
36.7
23.3
36.7
.200
Table 5. continued.
Clearcut Overstory
Taraxacum officinale G.H. Weber
ex Wiggers
Thallctrum fend/en Engeim. ex
Gray
Viola adunca Sm.
Ma/ion/a repens(LindL) G. Don
Spiraea betu/ifolia Pallas
S.vniphonicarpos a/bus (L.) Blake
Site 2
Bromus can/natus Hook. & Am.
Bromus frierinLs Leyss.
Carex conci'inoide Mackerzie
Carex geyeri Boott
Calamagrostis rubescens Buckl.
Ely/mis glaucus Buckl.
Festuca occidenta/, Hook.
Luzula campestnis (L) DC.
Me//ca bu/bosa Geyer ex Porter &
Coult.
Poa pratensi L.
Pisetum canescens BuckL
Ac/il/lea mi/lefo//uni L.
Anemone pieniBritt. ex Rydb.
A rn/ca cord/Ia//a Hook.
Eu,ybia conspicua (Lindi.) Nesom
Fragar/a spp.
G'aliumspp
1-fieradum a/b/forum Hook.
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
3.3
-6,7
-5.6
-3.3
-6.7
3.3
0.0
-13.3
-3.3
3.3
-26.7
-7.8
0.0
10.0
0.0
10.0
0.0
-5.6
26.7
-3.3
-6.7
20.0
20.0
0.0
0.0
26.7
23.3
0.0
-25.6
40.0
-16.7
6.7
10.0
13.3
0.0
6.7
20.0
16.7
-1.1
3.3
-6.7
-3.3
20.0
-6.7
46.7
66.7
53.3
3.3
6.7
10.0
-7.8
0.0
0.0
20.0
33.3
13.3
16.7
-16.7
0.0
0.0
3.3
31.4
11.5
29.9
55.4
-14.2
-3.3
0.0
60.0
0.0
0.0
-3.3
-20.0
0.0
-23.3
26.7
20.0
10.0
-30.0
-6.7
6.7
-30.0
-4.4
0.0
0.0
-6.7
16.7
46.7
43.3
-10.0
0.0
0.0
-13.3
23.3
0.0
0.0
-3.3
26.7
36.7
0.0
-30.0
0.0
-6.7
50.0
23.3
50.0
-23.3
-10.0
00
0.0
-5.6
37.8
27.8
30.0
-33.3
-2.2
-3.3
-13.3
-3.3
-6.7
13.3
22.2
-36.7
-53.3
-32.2
16.7
25.8
9.4
15.4
26.7
16.7
-3.3
46.7
23.3
-60.0
0.0
20.0
0.0
0.0
10.0
-13.3
3.3
53.3
43.3
0.0
6.7
0.0
0.0
33.3
16.7
-3.3
46.7
43.3
0.0
-3.3
3.3
-3.3
45.6
-6.2
10.0
20.0
0.0
33.3
6.7
0.0
-3.3
0.0
-6.7
43.3
6.7
50.0
11.1
11.1
.33
23.3
-3.3
-16.7
0.0
-43.3
-3.3
-13.3
-28.1
-0.7
-5.1
-2.1
-8.1
2OG
-4.4
21.1
1.1
24.0
0.0
0.0
31.1
17.8
-1.1
45.6
40.0
-2.2
14.4
0.0
-1.1
0.0
-6,7
-30.0
0.0
23.3
0.0
-13.3
27.8
17.8
-1.1
-3.3
1.1
3.3
-6.7
Table 5 continued.
Clearcut Overstory
TEXC
GRAZED
-36.7
-32.8
-40.0
0.0
-6.9
-3.3
CEXC
Lathyrusspp.
Lupinusspp.
Moehring/amacrophy//a(Hook.)
Fenzl
Senec/ointegerrimusNutt.
Stellar/a /ongiesGoIdie
Taraxacurnoff/c/na/eG.H. Weber
ex Wiggers
Tha//ctrumfend/eriEngelm. ex
Gray
Via/a adunca Sm.
Nahaniarepens(Lindl.)G. Don
SpiraeabetulifollaPaHas
'Syrnphoricarposabus(L.) Blake
Site 3
Bromus carinatus Hook. &Arn.
BromusfnermfsLeyss.
Carexconcinnoide5Mackenzie
CarexgeyeriBoott
ca/amagrosti:crubescenssuckl.
E/ymusg/aucusBuckl.
Festuca occidental/s Hook.
Luzu/acampestris(L.) DC.
MeI/cabulbosaGeyerexPorter&
PoapratensisL.
TrfsetumcanescensBuckl.
Achilleam/I/efollumL.
CEXC
-6.7
-23.3
ntroI Overstory
TEXC
GRAZED
16.7
0.0
5.6
10.0
Thinned Overstory
CEXC
TEXC
GRAZED
36.7
-6.7
-7.8
0.0
10.0
14.4
224
10.0
33.3
3.3
14.4
40.0
-20.0
20.0
0.0
0.0
0.0
-0.4
0.0
0.0
6.7
10.0
6.7
36.7
0.0
16,7
3.3
3.3
6.7
23.3
10.0
6.7
-3.3
-1.0
6.7
0.0
0.0
0.0
3.3
0.0
-1.1
-46.7
21,9
.6.7
20.0
0.0
12.2
3.3
-23.3
2.2
26.7
-3.3
-6.7
40.0
47.1
73.3
-3.3
-23.3
43.3
-10.0
0.0
0.0
-16.7
-10.0
3.3
13.3
26.7
-13.3
-10.0
13.3
46.7
10.0
0.0
13.3
23.3
3.3
-13.3
-2.2
8.9
4.4
0.0
0.0
-6.7
30.0
20.0
13.3
-16.7
-3.3
0.0
-16.7
23.3
36.7
0.0
-40.0
-6.7
1.4
0.0
-11.1
-17.8
32.2
0.0
-30.0
-3.3
45.5
0.0
3.3
50.0
53.3
33.3
-13.3
0.0
0.0
0.0
-10.0
23.3
43.3
16.7
-16.7
0.0
2.6
0.0
6.7
5.6
65.6
16.7
-20.0
7
-1.5
-27.2
26.0
-33
20.0
20.0
10.0
3.3
-16.7
-3.3
10.0
10.0
-53.3
-6.7
-25.8
-3,5
20.0
0.0
-13.3
10.0
36.7
40.0
-6.7
-6.7
-10.0
-17.2
-33.3
-33.3
-23.3
-14.4
0.0
-10.0
-10.0
80.0
-33.3
83.3
86.0
-21.0
42.4
73.3
-46.7
63.3
16.7
-13.3
3.3
15.6
0.0
1.1
10.0
-8.8
36.7
3.3
23.3
6.7
10.0
-1.5
46.7
36.7
0.0
-7.2
48.9
1.8
19.6
3.3
1.1
-10.3
-4.4
-1.1
cx
Table 5.
Co
aearcut Overstory
Anemone p/eriBritt. ex Rydb.
Am/ca cord/b/la Hook.
Euiyb/aconsp/cua(Lindl.) Nesom
Fragarlaspp.
Ga//urn spp
H/erac/urn a/b/forum
Hook.
Lathyrusspp.
Lup/nusspp.
Moehr/ng/amacrophyl/a(Hook.)
Fenzl
Senec/o/ntegerrlrnusNutt.
Stellar/a iongi-1Jothe
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
16.7
-1.8
-36.3
-1.3
16.7
-20.0
-3.3
0.0
0.0
-23.3
16.7
0.0
33.3
13.3
0.0
13.3
0.0
6.7
10.0
0.0
18.9
38.9
-2.2
-13.3
-2.2
4.4
18.9
3.3
30.0
16.7
0.0
3.3
0.0
-3.3
6.7
10.0
56.7
43.3
0.0
26.7
-0.3
-13.5
-39.4
9.4
-20.0
-43.3
0.0
-20.0
0.0
-20.0
-60.0
46.7
-3.3
0.0
-16.7
-20.0
-3,3
8.9
10.0
-8.9
-34.4
42.2
0.0
1.5
-23.3
0.0
30.0
28.9
20.0
23.3
6.7
6.7
10.0
0.0
0.0
0.0
13.3
3.3
10.0
0.0
3.3
0.0
6.7
16.7
0.0
3.3
13.3
3.3
0.0
3
4 .i
-6 7
33
-6 7
00
-6 7
33
11
-13.3
-32.8
.73.3
-30.0
16.7
1.1
-10.0
3.3
-10.0
50.0
6.7
-26.7
26.7
22.1
-3.1
-40.0
-3.3
13.3
0.0
-10.0
-20.0
43.3
iggers
T/a/lctrurnfend/erlEngelrn. ex
Gray
V/olaaduncaSm.
Mahonia repens(Lindl.) G. Don
Sp/raeabetuilfo//aPallas
Thinned Overstory
TEXC
ra-axacurp otficinale ' H Weber
ex
Control Overstory
CEXC
11.1
13.3
0.0
-26.7
-30.0
Symnp/iorfcarposalbus(L.) Blake
49.0
13.3
Herbivory treatments: Graze cattle and big game grazing; CEXC
exclusion of cattle and big game grazing.
11.1
-1.1
-10.0
-3.3
-7.8
36.7
10.0
41.1
0.0
-3.3
-3.3
0.0
-6.7
2.2
-3.3
6.7
-22.2
6.7
-16.7
22.2
26.7
3.3
15.6
13.3
36.7
6.7
cattle exclosure, big game grazing only; TEXC total exclosure,
I.
co
Table 6. The changes in species composition in a grand flr forest from 1985 to 2003 using a 30 cm x 60 cm plot frame.
Clearcut Overstory
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
Site 1
Bromus car/natus Hook. & Am.
Bromus inermis Leyss.
Carex con cinnoides Mackenzie
Carex geyer/ Boott
Ca/amagrost/s rubescens Buck!.
Elymus g/aucus Buck!.
Festuca occidental/s Hook.
Luzula campestris (L.) DC.
Me//ca bu/bosa Geyer ex Porter &
Cou!t.
Poa pratencic L
7Trfsetum canescens Buck!.
Ach/Ilea mi//efoliurn L,
Anemone pipe/-f Britt. ex Rydb.
Am/ca cord/fat/a Hook.
Eurybia conspicua (LindL) Nesom
Fragaria spp.
Gallum spp
Hierac/um a/b/forum Hook.
Lathyrus spp.
Lupinus spp.
Moehr/ngia macrophytla (Hook.)
Fenzl
Senec/o integerrinius N utt.
Stellar/a longioes Goldie
50.0
13.3
6.7
10.0
13.3
36.7
-6.7
3.3
40.0
83.3
20.0
6.7
-3.3
40.0
-10.0
-16.7
28.9
40.0
5.6
-10.0
-14.4
20.0
-23.3
-5.6
23.3
13.3
0.0
10.0
10.0
23.3
-26.7
-6.7
0.0
23.3
0.0
13.3
-26.7
3.3
-60.0
-16.7
-20.0
-60,0
-30.0
133
73.3
-30,0
50.0
-36.7
10.0
-10.0
-36.7
-43.3
3.3
-3.3
13.3
50.0
-63.3
46.7
-66.?
-13.3
-3.3
-70.0
-43.3
-16.7
-6.7
0.0
41.1
-54.4
0.0
-53.3
62.2
-13.3
-41.1
-27.8
-5.6
-38.9
20.0
-5.6
-32.2
24.4
-46.7
-53.3
6.7
16.7
30.0
30.0
-1.1
10.0
16.7
53.3
23.3
16.7
-20.0
66.7
-3.3
-3.3
45.1
0.0
0.0
-7.8
45.6
-13.3
7.8
40.0
33.3
23.3
0.0
0.0
53.3
30.0
40.0
26.7,
55.6
-16.7
-20.0
-40.0
-16.7
13.3
-16.7
-13.3
3.3
-3.3
-13.3
-3.3
-10.0
-60.0
-23.3
33.3
0.0
-60.0
16.7
-30.0
-33.3
-10.0
0.0
16.7
-20.0
-6,7
23.3
-25.6
-42.2
-5.6
2.2
3.3
-4.4
53:3
-23.3
3.3
3.3
-20.0
-16.7
-30.0
6.7
-6.7
13.3
16.7
36.7
-43.3
3.3
43.3
-10.0
-3.3
3.3
3.3
0.0
-26.7
-16.7
53.3
-56.2
37.8
-6.7
-47.8
-2.2
-21.1
-13.3
-13.3
-2.2
23.3
-51.1
-30.0
-60.0
-16.7
0.0
3.3
-38.9
36.7
56.7
3.3
46.7
56.7
46.7
50.0
46.7
10.0
50.0
6.7
16.7
3.3
3.3
.
-1.1
36.7
0.0
-2.2
24.4
53.3
-1.1
16.7
I,
Table 6. continued.
Clearcut Overstory
Taraxacurnofflcina/eG.H. Weber
ex Wggers
Thalictrumfend/eriEngelm. ex
Gray
Viola aduncaSm.
Mahoniarepens(Lindl.)G. Don
SpiraeabetullfoliaPallas
Symphoricarposalbus(L.) Blake
Site 2
BrornuscarinatusHook. &Arn.
Brornus:nermsLeyss.
Cdrex condnnoids Mackenzie
GrexgeyeriBoott
ca/ainagrostfsrubescens6uckL
E/yrnusglaucusBuckl.
Festucaocc/dentaii-Hook.
Luzula campestris(L.) DC.
Mefica bulbosa Geyer ex Porter &
Coult.
Poapratens/sL.
TrfsetumcanescensBuckl.
Achi/Ieamil/efollumL.
AriemonepioeriBritt. ex Rydb.
Arnica cordifolla Hook.
Eurybiaconspicua(Lindl.) Nesom
Fragariaspp.
Ga/iumspp
Hieracium a/b/forum Hook.
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
6.7
-23.3
-8.9
-13.3
-10.0
5.6
3.3
-23.3
-3.3
-3.3
-26.7
-2.2
0.0
13.3
0.0
200
-3.3
-10.0
23.3
-6.7
3.3
13.3
20.0
0.0
-6.7
23.3
38.9
0.0
-28.9
35.6
-13.3
16.7
-3.3
20.0
-6.7
10.0
33.3
20.0
-5.6
-5.6
17.8
2.2
-3.3
-10.0
-16.7
26.7
0.0
50.0
70.0
8.9
0.0
14.4
4.4
0.0
0.0
26.7
43.3
3.3
26.7
-26.7
0.0
0.0
10.0
30.3
3.3
24.3
0.0
0.0
-13.3
16.7
30.0
3.3
-36.7
-10.0
0.0
0.0
-20.0
20.0
26.7
10.0
-50.0
-3.3
26.0
0.0
3.3
25.6
17.8
6.7
-42.2
-7.8
0,0
0.0
-10.0
-17.5
5.8
0.0
66.7
26.7
-6.7
20.0
-6.7
-23.3
0.0
0.0
0.0
0.0
-6.7
30.0
33.3
56.7
-40.0
-16,7
-6.7
-19.3
-10.0
-13.3
10.0
24.4
-46.7
-60.0
-42.2
20.0
0.0
33.3
6.7
-23.3
0.0
-33.3
-3.3
-20.0
35.6
31.0
23.6
29.0
-31.4
-1.0
20.0
-3.3
46.7
40.0
-60 0
0.0
26.7
0.0
-3.3
16.7
6.7
3.3
43.3
20.0
-3.3
50.0
40.0
0.0
-3.3
6.7
-3.3
58.9
-11.6
0.0
35.6
42.2
-4.4
20.0
26.7
3.3
36.7
6.7
0.0
20.0
0.0
-3.3
50.0
3.3
56.7
-10.0
-30.0
0.0
20.0
3.3
-26.7
22.2
23.3
28.9
30.0
10.0
-3.3
8.9
57.1
-7.1
-2.8
-13.1
633
43.3
0.0
3.3
0.0
-3.3
11.1
0.0
-3.3
0.0
43.3
50.0
-13.3
600
0.0
0.0
-4.4
34.4
2.6.7
36.7
-41.1
-2.2
1.1
-7.8
Table 6. continued.
Clearcut Overstory
CEXC
TEXC
GRAZED
Lat/iyrus spp.
Lupinus spp.
frloehringfa rnacrophyfla (Hook.)
Fenzl
Seneclo integerrfinus Nutt.
Ste//aria /ongioes Goldie
Taraxacurn off/cinale G. H. Weber
ex Wiggers
Tha/ictrum fend/eufEngelm. ex
Gray
V/c/a adunca Sm.
Ma/ion/a repens (.Lindl.) G. Don
5/reea betu/ifo/fa Pall as
Symphoricarpos a/bus (L.) Blake
Site 3
Bromus carinatus Hook. & Am.
Bromus inermi5 Leyss.
Carex concinnoides Mackenzie
Carex geyerf Boott
Ca/amagrostis rubescens Buckl.
E/ymus glaucus Bucki.
Festuca occidental/s Hook.
Luzula campestrLc (L.) DC.
Me//ca bu/bosa Geyer ex Porter &
Coult.
Poa pratensis L.
Trfcetum canescens Buckl.
Ac/i/flea rniI/efo/ium L.
Control Overstory
CEXC
TEXC
GRAZED
Thinned Overstory
CEXC
TEXC
GRAZED
-43.3
-40.4
-43.3
-6.7
16.7
6.7
36.7
-3.3
3.3
0.0
-9.7
-6.7
-36.7
0.0
-11.1
0.0
-16.7
-3.3
10.0
-32.2
3.3
33.3
0.0
11.1
40.0
-10.0
21.1
0.0
0.0
0.0
6.7
6.7
0.0
6.7
6.7
10.0
6.7
6.3
3.3
16.7
40.0
23.3
6.7
26.7
10.0
-3.3
-1.4
10.0
0.0
0.0
6.7
3.3
-3.3
-1.1
-60.0
-26.3
0.0
26.7
-3.3
17.8
3.3
-23.3
-1.1
30.0
53.6
80.0
-16.7
-10.0
-20.0
30.0
23,3
25.6
-3.3
-4.2
-3.3
0.0
-10.0
-5.6
-16.7
6.7
-1.1
3.3
-38.5
-40.0
-3.3
0.0
8.9
0.0
-3.3
6.7
56.7
29.7
33.3
0.0
10.0
4.4
26.7
16.7
23.3
13.3
39.6
36.7
-3.3
1.4
2.1
55.2
0.0
4.6
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-33.3
-11.4
-16.7
-20.0
-26.7
-17.8
10.0
-10.0
0.0
0.0
40.8
-6.7
20.0
26.7
-17.8
50.0
23.3
1.1
3.3
-2.6
36.7
23.3
43.3
27.8
53.3
46.7
57.8
13.3
19.6
43.3
20.0
0.0
0.0
36.7
20.0
20.0
-56.7
-33.2
-13.3
-23.3
-46.7
-42.2
-20.0
-20.0
-30.0
-13.3
-5.0
-6.7
-3.3
-10.0
-5.6
0.0
0.0
-1.1
-10.0
-22.1
-40.0
-36.7
-33.3
-17.8
-6.7
-13.3
-15.6
76.7
85.1
76.7
16.7
3.3
1.1
10.0
3.3
13.3
-43.3
-29.3
-60.0
-20,0
8.6
-13.2
-8.6
33.3
2.1
80.0
48.9
66.7
6.7
0.0
-3.3
36.7
0.0
47.8
(.Q
Table 6. continued.
Clearcut Overstory
CEXC
TEXC
-3.1
Control Overstory
Thinned Overstory
GRAZED
CEXC
TEXC
GRAZED
CEXC
TEXC
GRAZED
-30.0
-56.7
0.0
-20.0
0.0
-30.0
-66.7
30.0
30.0
-3.3
25.6
30.0
-2.2
-20.0
-2.2
6.7
24.4
6.7
36.7
33.3
0.0
-10.0
3.3
-3.3
3.3
10.0
53.3
50.0
0.0
-3.3
0.0
-20.0
-10.0
3.3
30.0
7.8
-2.2
-6.7
-6.7
-26.7
10.0
0.0
40.0
13.3
0.0
13.3
0.0
0.0
20.0
0.0
-10.0
-32.2
47.8
Anemone pieri Britt. ex Rydb.
Am/ca cord/fe/ia Hook.
Eu,ybia conspicua (Lindi.) Nesom
Fragariaspp.
Ga/iumspp
/-//erac/um a/biflorum Hook.
16.7
-46.7
-3.3
3.3
0.0
-23.3
Lathyrusspp.
-200
Luoinussoo.
Moehr/ng/a macrop/7y/Ia (Hook.)
50.0
-19.2
-42.2
-1.9
3.3
0.6
-36.7
23.3
33.3
28.9
26.7
23.3
12.2
6.7
13.3
3.3
0.0
0.0
13.3
3.3
16.7
0.0
10.0
0.0
6.7
16.7
6.7
3.3
13.3
6.7
0.0
-6 7
-5 7
-10 0
33
-6 7
00
-6 7
-
3
-2 2
-20.0
-44.6
76.7
-20.0
13.3
-3.3
-13.3
6.7
-6.7
Fenzl
Senecio/ntegerrimusNutt.
5teiaiia/ongi,esGoldie
Taraxacum officinale G 9 Weuer
ex W!ggers
Tha/ictrum fènd/eriEngelm. ex
bray
Viola aduncaSm.
Mahon/a repens(Lindl.) G. Don
-27.8
-1.4
9.3
-1.1
-3.3.
.
1.1
10.0
46.7
31.8
13.3
-16.7
-3.3
-10.0
46.7
3.3
47.8
6.7
-5.0
0.0
0.0
-6.7
-4.4
-3.3
-6.7
4.4
Sp/raeabetu/ifo//aPallas
-43.3
-35.3
-43.3
6.7
3.3
-31.1
23.3
-10.0
32.2
Symphor/carposalbus(L.) Blake
63.3
50.4
10.0
33.3
6.7
13.3
16.7
36.7
16.7
Herbivory treatments: Graze cattle and big game grazing; CEXC - cattle exctosure, big game grazing only; TEXC total exclosure,
exclusion of cattle and big game grazing.
'0
191
Appendx F
Grand Fir Shrub Density and Cover Change
Table 1. The changes
shrub
1991
to 1991, 1994, and 2003.
Clearcut Overstory
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
0.4
-0.1
-0.6
-0.1
0.9
-1.0
-1.1
-0.8
-2.7
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
17.0
0.0
0.7
5.3
0.6
0.0
5.0
4.0
0.0
-0.3
0.0
-0.7
0.0
-2.3
0.0
0.0
0.0
0.3
0.0
-0.3
0.0
0.0
0.0
0.0
0.0
1.0
-4.0
-0.3
-2.7
3.7
0.0
-0.3
-0.7
-0.3
-1.1
-0.6
-0,1
0.6
0.3
-0.1
0.4
-0.1
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.2
0.0
0.0
0.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.4
0.0
7.0
23.4
7.7
1.3
1.7
3.1
18.7
-0.7
9.8
0.0
0.0
-1.0
0.0
1.0
65.8
0.0
-0.3
-4.7
0.0
0.0
1.3
0.0
0.0
2.0
0.0
0.0
3.4
0.0
0.0
3.3
0.0
0.0
3.7
0.0
0.0
-2.6
0.0
-1.1
-0.1
-0.4
0.9
-0.4
-0.8
-1.7
-0.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Site 1
Acer glabrum Torr. var. glabruín
Ame/anchiera/nffo//a(Nutt.) NutI. ex
M. Roemer
Crataegusdoug/asi/Lindl.
Ho/odiscusdfsco/or(Pursh) Maxim.
Phllade/phus/ewi//Pursh
Physocarpusmalvaceus(Greene)
Kuntze
Prunus virginiana L.
RibescereumDougl.
Rosagymno:arpaNutt.
2.1
Site 2
Acer glabrurn Torr.
var. glabrum
Nutt. ex
Arnelanchiera/n/fo/ia(Nutt.)
M. Roerner
Crataegus doug/as/f LindL
Holodfscusdfscolor(Pursh) Maxim.
Phllade/phus/ewLc//Pursh
Physocarpus malvaceus (Greene)
Prunus virginfana L.
R/bescereumDougl.
Rosa gymnocarpa Nutt.
Site 3
Acer glabrum Torr. var. glabrum
Ame/anch/eralpifo/fa (Nutt.) Nutt. ex
M Roem
Crataegusdoug/asi/Lindl.
0.0
0.0
Tabte 1. continued.
Clearcut Overstory
Holodiscus discolor(Pursh) Maxim.
Phi/ade/phus lewis/i Pursh
Physocarpus ma/vaceus (Greene)
Kuntze
Prunus virginiana L.
Ribes cereum Dougi.
Rosa gymnocarpa Nutt.
1994
Control Overstory
Thinned Overstory
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
0.0
0.0
0.0
-0.1
0.0
0.0
0.0
0.0
0.0
0.0
3.2
-0.4
0.5
0.0
0.5
0.0
2.1
0.0
-1.3
3.6
-0.7
3.7
3.0
-0.8
5.3
0.0
2.4
0.0
0.0
-2.0
0.0
0.9
-4.0
0.0
0.0
-3.0
0.0
0.0
-0.7
0.0
0.0
16.3
0.0
0.0
2.8
0.0
-0.3
16.0
0.0
0.0
1.7
0.0
0.0
9.3
5.6
-0.3
3.2
-0.4
2.7
-0.5
-1.1
4.5
1.8
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
.0.0
0.0
0.0
0.0
1.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
16.0
a.7
8.3
7.7
19.3
9.9
10.3
3&7
45.7
0.0
-0.3
19.3
0.0
.1.7
5.0
0.0
4.3
2,0
0.0
-0.3
0.0
-0.3
-0.3
0.0
0.0
1.3
0.0
2.7
-0.3
00
12.3
0.0
0.3
0.3
-1.1
0.2
-0.3
1.3
0.3
1.0
0.9
0.1
-0.5
-0.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.0
0.0
0.0
2.5
0.0
8.0
14.1
35.7
4.0
5.3
5.8
23.0
9.3
19,8
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Site 1
Acer glabrum Torr. var. glabrum
A,ne/anc;5iera/nifo/ia(Nutt.) Nutt.
ex
M. Roerner
Crataegusdouglas//LindL
Hoiodiscuso7sco/or{Pursh) Maxrn.
Phi/ade/p/ius/ewisllPursh
Ph,vcocarpus malvaceus (Greene)
Kuntze
Prunus virginiana L.
RibescereumDougl.
Rosa gymnocarpaNutt.
Site 2
Acer glabrum Torr. var. glabruni
Ame/anchieralnifo/la (Nutt.) Nutt. ex
M. Roemer
Crataegusdouglas//Lindl.
Holodiscusdiscolor(Pursh) Maxim.
Phllade/phuslewisiiPursh
Physocarpus ma/vaceus (Greene)
Kuntze
Prunusvi'rg/nianaL.
1.0
(J
Table 1. continued.
Clearcut Overstory
CEXC
TEXC
GRAZE
0.0
0.0
-0.3
23.3
15.4
10.3
RibescereumDougl.
Rosa gyrnnocarpaNutt.
cqntoirt
CEXC
TEXC
GRAZE
0.0
22.3
0.0
18.7
0.0
29.4
Thinned Oversto
CEXC
TEXC
GRAZE
0.3
0.0
0.0
22.7
43.7
33.1
Site3
Acer glabrum Torr. var. glabruin
Amelanchieralnifolla(Nutt.) Mutt. ex
M. Roemer
crataegus douglas/f LindL
Holadiscusdisco/or(Pursh) Maxim.
Phi/adeip/itis lewisfiPursh
Physocarpusma/vaceus(Greene)
kuntZe
Prunus virgin/anaL.
RibcscereurnDougL
Rosa gyrnnocarpa Mutt.
2003
0.2
-0.5
0.4
-0.5
2.0
1.5
-2.0
-1.7
-0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.6
-0.1
0.0
0.0
0.0
0.0
0.6
0.0
0.0
0.0
0.3
0.0
0.0
0.0
1.3
1.2
0.0
0.0
4.3
-0.4
0.0
0.0
4.0
-0.3
10.7
7.3
23.6
13.0
13.0
-0,6
0.0
0.0
13.0
0.0
-0.1
26.0
0.0
0.0
0.0
0.0.
.0.0
0.0
0.0
1.3
2.3
0.0
0.0
0.0
-0.3
0.0
0.0
25.3
27.7
4.4
17,0
68.7
0.2
0.9
0.6
-0.8
-0.2
-LU
-0.9
0.7
-0,7
0.6
0.0
0.1
1.4
1.8
1.8
2.0
2.9
1.3
5.9
0.0
0.0
12.8
0.0
0.0
9,0
0.0
0.0
3.6
0.0
0.0
8.9
0.0
0.0
2.2
0.0
0.0
2.3
1.3
0.0
9.6
0.3
00
0.0
0.0
19.0
24.3
30.7
20.3
18.3
10.2
23.7
34.7
36.0
8.7
7.0
-0.7
1.7
1.3
0.0
5.3
26.3
140.0
-0.7
44.0
80.3
0.3
0.0
0.0
0.3
-0.3
2.3
0.3
1.3
50.7
3.0
-0.3
38.0
0.7
-1.1
1.1
Site I
Acerg/8brumTorr. var. g.Iabrum
Ame/anchieraln/folfa(Nutt.)Nutt.ex
M. Roemer
ratagusdouglasiiLindL
Ho/odiscusdisco/or(Pursh) Maxim.
PhlladelphuslewisiiPursh
Physocarpusma/vaceus(Greene)
Prunus virgin/anaL.
R/bescereumDougl.
Rosa gymnocarpaNutt.
Site 2
3.3
1.6
Tab'e 1. continued.
Clearcut Overstory
Acer glabrum Torr. var. g/abrum
Amelanchieralnifo/ia(Nutt.) Nutt. ex
M. Roemer
Crataegusdouglasf/Lindl.
Ho/odicusd/sco/or(Pursh) Maxim.
Phlladeiohus/ewL/iPursh
Physocarpus ma/vaceus (Greene)
Prunus
Control Overstory
Ihinned Overstory
CEXC
TEXC
GRAZE
GRAZE
CEXC
TEXC
GRAZE
-0.5
TEXC
0.1
0.1
0.6
-0.3
-0.2
0.2
0.3
-0.6
0.0
0.0
0.3
0.3
1.8
1.3
0.0
0.9
0.1
7.3
0.0
0.0
11.8
0.0
0.0
3.7
0.0
0.0
2.6
0.0
0.0
0.8
0.1
0.0
0.7
0.0
0.0
5.1
0.0
0.0
5.3
0.0
0.0
2.2
0.0
0.0
2.0
8.7
29.3
2.3
4.7
5.4
30.3
19.3
15.1
20.7
0.0
10.3
11.0
0.0
15.8
1.7
38.7
0.0
31.3
20.0
0.0
38.0
32.8
CEXC
0.0
38.7
4.3
R/bescereuinDougl.
0.0
0.0
0.0
Rosa gymnocarpaNutt.
9.0
23.4
-9.3
4.0
0.0
14.0
Site 3
Acer glabrurn Th. v-. giabrum
1.1
0O
0.0
-0.5
0.8
0.0
2.0
-1.7
-0.4
0.8
0.2
3.2
1.2
0.4
0.1
12
0.0
4.9
0.0
0.0
12.1
4.8
0.0
0.0
4,1
0.1
0.0
0.3
0.2
0.0
4.1
0.3
1.6
0.3
-0.4
0.0
3.2
0.9
0.0
2.2
0.0
-0.1
7.7
2.6
-0.3
1.3
3.0
2.6
virginiana L.
Ame/aac/iIra/n/fo//a(Nutt.) Nutt. cx
M. Roemer
C'rataegusdouglasfiLindL
Ho/odiscusd/sco/or(Pursh) Maxim.
Phf/ade/phus/ew/siiPursh
Physocarpusrna/vaceus(Greene)
OJ
0.0
12.3
4.7
4.8
Prunus virgin/anaL.
4.7
32.7
0.0
00
30.7
0.0
18.3
0.0
4.0
RfbescereumDougl.
1.3
-0.1
1.0
0.0
0.0
0.0
0.0
0.0
0.0
Rosa avmnocarvaNutt.
10.7
15.0
20.3
4.0
12.0
1.1
31.3
53.3
1.7
Herbivory treatments: Graze catiJe and big game grazing; CExc - cattle exclosure, big game grazing only; TExc total exciosure,
exclusion of cattle and big game grazing.
0
Lu
Table 2. The changes in shrub densi' from 1995 to 1991, 1994, and 2003.
Control Overstoy
Clearcut_Overstot
Thinned Overstory
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
Acer glabrurn Torr. var. glabrum
Ame/anchiera/nifolia (Nutt.) Nutt.
ex M. Roemer
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.2
-0.1
0.0
4.4
1.3
Q-ataegus doug/as/i LindI.
Ho/od/scus d/sco/or(Pursh) Maxni.
Philade/phus lewis/i Pursh
Physocarpus ma/vaceus (Greene)
Kuntje
Prunus v/rg/n/ana L.
0.0
0.0
-5.2
0.0
0.0
-1.1
0.0
-0.8
-1.3
0.0
-0.2
-7.8
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0,4
0.0
0.0
-0.2
0.0
0,0
0.0
0.2
0.0
0.0
0.0
0.0
0.0
-10.9
0.0
0.0
-30.8
0.0
0.0
-6.5
0.0
:. -0.1
-8.9
0.0
0.0
-8.8
0.0
0.0
-4.2
-0.1
0.0
-8.8
0.0
0.0
-0.6
0.0
-1.0
0.0
0.
0.0
co
0.0
0.0
0.0
0.0
0.0
o,t
0.0
0.0
-02
-0.7
0.0
0.0
0.2
1.2
-0.3.
-0.4
0.0
0,0
0.0
0.0
0.0
-2.8
0.0
0.0
-0.3
0.0
0.0
0.0
o.o
0.0
0.0
0.0
o.o
o..o
o.o
-3.0
-5.0
1.5
-3.2
-10
0.0
0.0
-0.6
0.0
0.0
0.0
0,0
0.0
0.0
0.0
0.0
0.0
-8.9
0.0
0.0
-7.0
0.0
0.0
-2.0
0.0
0.0
9.8
0.0
0.0
1.0
0.0
0.0
-0.1
0.0
0.0
3.6
0.0
0.0
6.6
0.0
0.0
-5.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.1
0.0
0.0
0.0
0.0
-0.2
-0.1
0.2
0.0
-1.5
0.0
0.0
0.1
1991
Site 1
P,cs cercum Douqi.
Rosa gymr,ocarpaNutt.
G.0
Site 2
Acer g/abrurn Torr. var. g/abrum
Ame/anchier a/nub/ia (Nutt.) Nutt.
ex M Roemer
crataegus doug/as/i Lindl.
Ho/od/scus d/sco/or(Pursh) Maxim.
Phi/ade/phus /ew/s/i Pursh
Physocarnus ma/vaceus (Greene)
Kuntze
Prunus virginiana L.
Ribes cereum Dougl.
Rosa gymnocarpa Nutt.
Site 3
Acer g/abrurn Torr. var. glabrum
Ame/anchierafnifo//a (Nutt.) Nutt.
ex M. Roemer
Tab'e 2. continued.
Crataegusdoug/as/iLindI.
Holodiscusdisco/or(Pursh) Maxim.
PhlladeIphusIewi/iPursh
Physocarpusma/vaceus(Greene)
Kuntze
Prunus virgin/anaL,
.R/bescereurnDougL
Rosa gyrnnocarpaNutt.
1994
Clearcut Overstory
CEXC
TEXC
GRAZE
0.0
0.0
0.0
0.0
0.6
0.0
-3.4
-3.7
-5.2
0.0
0.0
-1.8
0.0
0.0
-6.7
0.0
0.0
0.0
CEXC
0.0
0.0
0.7
nrQ1 Qverstory
TEXC
GRAZE
0.0
0.0
0.0
0.0
2.2
-1.8
Thinned Overstory
CEXC
TEXC
GRAZE
0.0
0.0
0.0
-0.1
0.0
0.0
-2.4
0.0
-4.1
0.0
0.0
0.0
0.0
0.0
-1.6
0.0
0.0
3.0
0.0
0.0
1.9
0.0
0.0
-6.3
0.0
0.0
-5.2
0.0
0.0
0.0
0.0
3.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.4
0.5
1.8
0.1
0.2.
2t
1.9
0.0
0.0
-3.3
0.0
0.0
.0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.8
-0.2
0.0
0.0
0.0
Q.0
0.0
-02
-0.4
-8.0
0.0
0.0
0.0
--0/p
0.0
0.0
0.0
-0.4
0.0
0.0
0.Q
0.0
0.3
-12.7
-1.0
0.0
-0.1
-9.1
0.0
0.0
5.2
0.0
0.0
-23.6
1.8
-0.1.
0.0
0.0
0.0
5.5
0.0
.0.6
-30.8
0.0
.0.5
0.0
-1.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
00
0.0
0.0
0.0
0.5
-,1.
0.2
0.0
0.5
1.2
-0.3
0.9
0.0
0.0
0.0
0.0
0.0
0.0
-3.0
0.0
0.0
-0.1
0.0
0.0
0.3
0.0
0.0
1.9
0.0
0.0
-6.4
0.0
0.0
1.5
0.0
0.0
0.1
1.2
1.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.6
Site 1
Acer glabrurn Torr. var. glabrum
Ame/anch/era/nifolla(Nutt.) Nutt.
cx M.
Roerner
cr3taegusdoug/asllLnd!.
Ho!od/scusd/sco/or(Pursh)
Maxim.
Ph//adeiphus/ewisi/Pursh
Physocarpusmaivaceus(Greene)
Kuntze
Prunus vtrginiana L.
0.0
Ribescereum0ougl.
Ro.cagymnocarpaNutt.
Site 2
Acer glabrum Torr. var. giabrum
Amelanc/iier:fn/folia(Nutt.) Nutt.
Crataegus doug/as/i Lindt.
Ho/odLccusdicolor(Pursh) Maxim.
Phlladelphuslewisi/Pursh
Physocarpusmalvaceus(Greene)
.
D.2..
Table 2.
nued.
CiarcuLQ:sto
Prunus virginiana L.
RibescereumDougl.
Rosa gyrnnocarpaNutt.
Control Overstoy
Overstory
ThJnn
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
6.3
0.0
0.0
-4.3
0.0
0.0
3.7
0.0
0.0
9.7
0.0
0.0
-1.2
0.0
0.0
2.0
0.0
0.0
-6.5
0.0
0.0
-2.7
0.0
0.0
6.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.1
0.0
0.0
0.0
0.8
-0.2
0.1
0.5
0.3
-1.2
1.0
0.5
-0.1
0.0
0.0
-3.4
0.0
0.0
2.2
0.0
0.0
-5.2
0.0
0.0
0.4
0.0
0.0
0.4
0.0
0.0
-3.3
0.0
-0.1
-1.1
0.0
0.0
10.2
0.0
0.0
-6.6
0.0
0.0
0.0
0.0
0.0
Q.0
0.0
0,0
0.0
-1.7
0.0
0.0
-6.3
0.0
16
-11.3
0.0
0.0
0.6
0.0
0.0
5.3
0.0
0.0
15.1
00
-1.6
0.0
0.0
0.0
0.0
-1.3
0.0
0.0
10
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
00
0.7
3.6
0.8
0.1
3.0
0.1
0.9
11.1
1.3
0.0
0.0
-5.2
0.1
0.0
1.0
1.3
-0.2
-1.0
0.1
0.6
0.0
0.0
0.0
0.0
0.0
0.0
0.3
0.1
0.0
-0.9
1.2
-0.1
-0.4
0.0
0.0
-0.2
0.0
0.0
0.0
-0.4
0.0
0.0
0.0
0.0
0.0
-9.0
0.0
0.2
-24.3
0.0
2.1
-6.1
0.0
0.5
-9.1
0.0
0.0
2.7
0.0
0.0
3.6
-0.1
0.0
-2.7
0.0
0.0
-0.6
0.0
0.0
-0.5
0.0
0.0
Site 3
Acer glabrum Torr. var. glabruin
Amelanchier alnifolia (Nutt.) Nutt.
ex M. Roemer
crataegus douglas/i Lindi.
Ho/odiscus disco/or (Pursh) Maxim.
Ph//ade/p/ius /ewi// Pursh
Physocarpus nia/vaceus (Greene)
Kuntze
Prunus vlrqin/ana L.
Rites cereuni Dougl.
Rosa gymnocarpa Nutt.
2003
Site 1
Acer g/abrurn Torr. var. glabrum
Arnelanchier alnifolia (Nutt.) Nutt.
ex M. Roemer
C'rataegus douglas/f LindI.
Ho/odiscus disco/or(Pursh) Maxim..
Philadeiphus lewis/i Pursh
Physocarpus ma/vaceus (Greene)
Kuntze
Prunus virginiana L.
Ribes cereum Dougl.
Rosa gymnocarpa Nutt.
Site 2
Table 2. continued.
Clearcut Overstory
Control Overstorq
Thinned Overstory
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
CEXC
TEXC
GRAZE
Acer glabrum Torr. var. glabrum
Ame/anchiera/nifolla(Nutt.) Nutt.
ex N. Roemer
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
-0.1
0.2
1.2
0.1
0.3
1.3
2.7
1.0
C'rataegus douglas/i Lindl.
0.0
0.0
0.7
0.0
0.0
0.6
0.0
0.0
-0.3
0.0
0.0
-0.3
0.0
0.0
-10.3
0.0
0.0
-1.2
0.0
0.0
0.2
0.1
0.0
1.0
0.0
0.0
-0.6
0.0
0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
15.0
0.9
0.0
6.3
0.0
0.0
5.1
0.0
0.0
-5.3
0.0
0.0
0.5
0.0
0.0
0.6
0.0
0.0
-1.7
0.0
2.8
0.0
0.0
O0
-4.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.1
0.0
0.0
0.0
0.7
0.3
-0.1
-0.1
0.9
-1.5
0.2
.0
2.9
0.0
0.0
-1.7
0.0
0.0
0.9
0.0
0.0
-5.1
0.0
0.0
0.7
0,2
0.0
2.4
0.0
0.0
-1.0
0.0
-0.1
0.0
0.0
0.0
0.0
0.0
0.0
Holodiscusdisco/or(Pursh) Maxim.
lewis/i Pursh
Physocarpus malvaceus (Greene)
Kuntze
Prurius v/rgh'iiana L.
Phi/ade/phus
RlbescereuinDougl.
Rosa gymnocarpaNutt.
Site 3
Acer glabrum Torr. var. g/abrum
Ame/anchiera/nifolla(Nutt.) Nutt.
ex N. Roemer
crataegus doug/as/i Lindl.
/-Io/odiscusdiscolor(Pursh)
Maxim.
Phi/adelphuslewisllPursh
Physocarpusma/vaceus(Greene)
Kuntze
PrunusvirginianaL.
-2.1
-6.7
RibescereumDougl.
0.0
2.6
Rosa gyinn
paNutt.
0.0
0.0
Herbivory treatments: Graze cattle and big game grazing; CExc
exclusion of cattle and big game grazing.
1.6
0,0
0.0
11.0
-5.9
0.0
0.0
0.0
0.1
0.0
-1.6
1.6
4.6
-7.5
11.8
4.7
1.5
0.0
0.0
0.0
0.0
0.0
1.3
1.1
0.0
0.0
0.0
0.0
0.0
0.0
cattle exclosure, big game grazing only; TExc total exciosure,
_0.0
I'D
I'0
200
Appendix G
Nutritional Quality of Steer Diets
201
Table 1. Nutritional quality of diets coeted in June and August of 2001 and 2002 for
ponderosa pine sites.
YEAR
TIME
Site
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2002
2002
2002
2002
2002
2002
2002
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
1
1
1
1
1
1
2
2
2
2
2
2
3
3
3
3
3
3
AUG
AUG
1
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
1
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
1
1
1
1
2
2
2
2
2
2
3
3
3
3
3
3
1
1
1
1
1
1
2
Timber
Harvest
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
ont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Hrbivcrv
IVOMD
ADF
NDF
CP
Grazc
0,80
0.77
0.77
0.82
0.79
0.82
0.82
0.80
0.82
0.74
0.77
0.79
0.80
0.76
0.79
0.76
0.74
0.77
0.73
0.73
0.72
0.75
0.73
0.76
0.73
0.72
0.76
0.72
0.74
0.72
0.75
0.74
0.74
0.72
0.73
0.73
0.87
0.84
0.79
0.84
0.78
0.83
0.86
0.32
0.33
0.35
0.35
0.34
0.29
0.36
0.36
0.34
0.38
0.39
0.38
0.30
0.36
0.31
0.36
0.38
0.35
0.37
0.35
0.37
0.35
0.36
0.38
0,37
0.39
0.37
0.43
0.38
0.40
0.37
0.36
0.36
0.39
0.36
0.37
0.37
0.36
0.31
0.33
0.37
0.37
0.32
0.56
0.57
0.51
0.58
0.58
0.47
0.48
0.54
0.52
0.54
0.59
0.59
0.48
0.54
0.51
0.57
0.60
0.56
0.61
0.61
0.60
0.60
0.58
0.63
0.59
0.58
0.57
0.59
0.60
0.64
0.60
0.58
0.59
0.62
0.61
0.55
0.54
0.52
0.42
0.45
0.57
0.61
0.44
12.61
12.48
14.49
14.95
13.45
13.16
12.33
13.39
12.68
12.71
12.64
12.68
11.40
16.09
13.61
14.50
13.63
13.21
9.78
8.97
8.62
8.69
10.36
9.78
7.07
11.50
9.14
11.20
9.09
8.52
9.00
10.79
10.05
10.32
7.65
8.86
14.22
13.19
12.44
14.15
14.47
16.77
13.98
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc.
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
tExc
Graze
CExc
TExc
Graze
202
Table 1. continued.
YEAR
TIME
Site
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
JUNE
JUNE
JUNE
2
Tiniber
Harvest
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Herbivorv
IVOMD
0.81
0.80
0.80
0.78
0.85
0.86
0.82
0.79
0.77
0.84
0.82
0.73
0.79
0.75
0.74
0.73
0.74
0.75
0.74
0.76
0.77
0.71
0.77
ADF
NDF
CP
0.33
0.51
14.43
0.32
0.52
1315
2
0.34
0.48
14.33
JUNE
2
CExc
0.35
0.53
14.12
JUNE
2
TExc
0.41
0.59
12.10
JUNE
3
Graze
0.37
0.54
15.87
JUNE
3
CExc
0.38
0.55
14.25
JUNE
3
TExc
0.37
0.60
15.11
JUNE
3
Graze
0.37
0.50
12.99
JUNE
3
CExc
0.39
15.22
0.58
JUNE
3
TExc
0.38
0,52
13.49
AUG
1
Graze
0.43
0.57
12.10
AUG
1
CExc
0.39
0.61
10.78
AUG
1
TExc
0.43
0.60
10.57
AUG
1
Graze
0.39
0.59
9.35
AUG
1
CExc
0.40
0.63
9.35
AUG
1
TExc
0.39
13.62
0.60
AUG
2
Graze
0.40
0.53
10.50
AUG
2
CExc
0.40
0.59
9.36
AUG
2
TExc
0.41
0.58
9.72
AUG
2
Graze
0.39
0.58
9.32
AUG
2
CExc
0.39
0.55
7.69
AUG
2
TExc
0.40
0.63
9.20
AUG
3
Graze
077
0.40
0.58
10.07
AUG
3
CExc
0.72
0.38
0.60
10.88
AUG
3
TExc
0.77
0.39
0.64
9.94
AUG
3
0.77
Graze
0.43
0.59
10.92
AUG
3
CExc
0.78
0.41
0.61
11.21
AUG
3
TExc
0.79
0.39
0.51
8.97
Timber harvest treatments: Thin commercial thinning; Cont control.
Herbivory treatments: Graze cattle and big game grazing; CExc cattle exciosure, big
game grazing only; TExc total exclosure, exclusion of cattle and big game grazing.
2
CExc
TExc
Graze
203
Table 2. Botanical composition and relative preference index for steer diets collected in
June and August of 2001 and 2002 for the ponderospine sites.
Botanical Cçnposition
Relative Pref. Index
Timber Herb.
YEAR TIME Site
Grass
Forbs Shrubs Grass
Forbs
Shrubs
Harvest
Trt,
2001 JUNE
1
Cont
CExc
92.39
5.67
1.95
1.29
0.46
0.12
2001 JUNE
1
Cont
TExc
88.34
3.11
8.55
1.76
0.42
0.11
2001 JUNE
1
Cont
Graze
94.63
4.38
1.00
1.38
0.25
0.07
2001 JUNE
1
Thin
CExc
86.42
994
3.64
1.47
0.47
0.18
2001 JUNE
1
Thin
TExc
73.93
18.11
7.96
2.11
0.57
0.24
2001 JUNE
1
Thin
Graze
96.77
3,23
0.00
1.59
0.15
0.00
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 JUNE
2001 AUG
2001 AUG
2001 AUG
2001 AUG
2001
AUG
2001
AUG
2001
AUG
2001
AUG
2001
AUG
2001
AUG
2001
AUG
2001
AUG
2001 AUG
2001 AUG
2001 AUG
2001
AUG
2001 AUG
2001 AUG
2002 JUNE
2002 JUNE
2002 JUNE
2002 JUNE
2002 JUNE
2002 JUNE
2
2
2
2
2
2
3
3
3
3
3
3
1
1
1
1
1
1
2
2
2
2
2
2
3
3
3
3
3
3
1
1
1
1
1
1
Cont
Cont
Cont
TExc
Graze
Thin
Thin
Thin
TExc
Graze
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
CExc
CExc
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
Thin
Thin
TExc
Graze
Cont
Cont
Cont
Thin
CExc
TExc
Graze
CExc
Thin
Thin
Cont
TExc
Graze
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
TExc
Graze
CExc
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
87.82
85.21
72.26
85.04
82.69
67.57
91.86
89.15
85.51
80.10
74.38
79.59
94.08
88.12
88.12
94.37
87.44
88.20
76.63
86.64
77.98
77.97
85.14
82.00
96.09
68,34
88.85
85.67
92.84
88.94
94.48
92.19
77.21
95.22
90.12
98.45
10.41
13.27
23.50
12.83
16.64
31.41
7.22
6.96
9.29
15.54
18.31
16.69
3.50
7.97
7.52
3.39
7.89
8.57
18.39
11.10
16.97
19.51
13.60
14.70
1.90
24.64
9.39
12.42
5.98
10.64
4.82
6.75
19.75
4.78
9.48
1.55
1.77
1.52
4.24
2.14
0.67
1.02
0.93
3.90
5.20
4.36
7.30
3.72
2.42
3.91
4.36
2.24
4.68
3.22
4.98
2.26
5.05
2.52
1.26
3.30
2.01
7.02
1.76
1.91
1.18
0.42
0.70
1.06
3.03
0.00
0.39
0.00
1.21
1.14
1.25
1.25
1.27
1.20
1.16
1.40
1.11
1.18
1.06
1.11
1.31
1.75
1.29
1.61
2.50
1.45
1.06
1.16
1.35
1.15
1.31
1.45
1.21
1.07
1.15
1.26
1.32
1.24
1.32
1.83
1.13
1.62
2.58
1.61
0.45
0.88
0.66
0.47
0.69
0.83
0.81
0.74
0.53
0.57
1.00
0.68
0.28
0.39
0.43
0.16
0.25
0.39
0.79
0.74
0.48
0.71
0.56
0.39
0.21
2.62
0.54
0.46
0.33
0.43
0.39
0.33
1.14
0.22
0.30
0.07
0.41
0.15
0.65
0.48
0.06
0.18
0.08
0.14
0.93
0.92
0.65
1.06
0.15
0.13
0.31
0.11
0.14
0.19
1.14
0.22
0.77
0.57
0.12
0.60
0.17
0.26
0.31
0.40
0.11
0.12
0.04
0.04
0.21
0.00
0.01
0.00
204
Table_2._continued.
$niaLQmJositJon
YEAR
TIME
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
Site
limb
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cont
Cont
Thin
Thin
Thin
Cont
Cant
Cont
Thin
Thin
Herb.
Grass
Forbs
Shrubs
Relative Pref. Index
Grass
Forbs
15.60
0.76
1.16
0.67
1.17
12.04
0.88
0.80
2
4.03
1.18
Graze
28.02
0.79
2
4.30
0.37
1.40
CExc
0.16
2
TExc
4.35
0.40
1.46
0.18
2
Graze
19.35
5.34
1.33
0.51
3
0.98
0.73
CExc
6.50
1.17
3
1.40
TExc
9.53
1.76
1.01
3
Graze
4.77
0.37
1.23
0.27
3
CExc
13.15
1.06
1.26
85.79
0.48
3
TExc
L54
1.30
91.26
7.19
0.39
3
1.24
1.13
Graze 81.12
17.63
0.72
AUG
1
CExc
6.56
0.71
1.29
92.73
0.53
AUG
1
2.50
1.77
TExc
8.67
0.42
88.84
AUG
1
Graze 93.17
6.45
0.37
1.36
0.37
AUG
1
0,38
1.61
CExc
94.66
4.95
0.23
AUG
1
2.35
TExc
2.29
0.54
80.23 17.43
AUG
1
Graze 80.62
3.95
1.32
15.42
0.71
AUG
2
1.12
CExc
17.08
2.00
0.74
80.91
AUG
2
TExc
1.41
1.07
1.25
79.67 18.91
AUG
2
Graze 76.14 19.03
4.84
1.32
0.53
AUG
2
1.26
CExc
85.98
11.22
2.80
0.41
AUG
2
1.21
TExc
17.54
3.65
0.72
78.81
AUG
thin
2
Graze 72.12
26.02
1.86
1.28
0.69
AUG
3
Cont
CExc
11.17
0.60
1.11
1.25
88.23
AUG
3
Cant
7.19
0.93
TExc
33.48
3.56
59.33
AUG
3
Cont
0.34
1.15
Graze 88.39
11.27
0.65
AUG
Thin
3
CExc
82.96
17.04
0.00
1.22
0.63
AUG
Thin
3
TExc
15.24
0.85
1.19
0.83
83.91
AUG
3
Thin
Graze 79.83
1.51
1.11
18.66
0.76
Timber harvest treatments: Thin rommerciai thinning; Cont control.
Herbivory treatments: Graze cattle and big game grazing; CExc cattle exciosure,
game grazing only; TExc total exciosure, exclusion of cattle and big game grazing.
2
2
CExc
TExc
83.65
87.08
67.95
95.33
95.25
75.31
92.52
88.72
94.85
Shrubs
0.17
0.09
0.62
0.08
0.04
0.96
0.08
0.07
0.07
0.22
0.14
0.35
0.04
0.09
0.03
0.02
0.07
0.23
0.46
0.14
0.74
0.63
0.34
0.33
0.05
0.27
0.06
0.00
0.08
0.43
big
205
Table 3. Nutritional quality of diets collected in June and August of 2001 and 2002 for
arand fir sites.
YEAR
TIME
Site
LOGTRT
GRAZTRT
NDF
ADF
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2901
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
1
Clear
Clear
Clear
Thin
Thin
Thin
Cont
Cont
Cont
Clear
Clear
Clear
Thin
Thin
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
0.55
0.55
0.50
0.56
0.55
0.53
0.58
0.58
0.55
0.57
0.57
0.56
0.57
0.56
0.55
0.33
0.34
0.34
0.37
0.35
0.35
0.41
0.37
0.40
0.34
0.41
0.35
0.37
0.39
0.39
0.40
0.42
0.39
0.35
0.34
0.32
0.39
0.36
0.40
0.43
0.38
0.43
0.38
0.39
0.37
0.40
0.37
0.44
0.45
0.42
0.44
0.37
0.42
0.39
0.42
0.41
0.41
0.44
0.43
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
3
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
2
Thin
Cont
Cont
Cont
Clear
Clear
Clear
Thin
Thin
Thin
Cont
Cont
Cont
Clear
Clear
Clear
Thin
Thin
Thin
Cont
Cont
Cont
Clear
Clear
Clear
Thin
Thin
Thin
Cont
Cont
055
0.60
0.58
0.52
0.53
0.53
0.59
055
0.57
056
0.55
0.58
0.60
0.62
0.60
059
0.57
0.60
0.56
0.66
0.65
0.59
0.66
0.60
0.59
0.60
0.52
0.61
0.65
IVOMD
0.82
0.78
0.74
0.77
0.78
0.70
0.75
0.74
0.75
0.76
0.75
0.77
0.76
0.76
0.72
0.74
0.77
0.75
0.77
0.76
6.73
0.76
0.76
0.72
0.75
0.74
0.72
0.74
0.75
0.78
0.69
0.76
0.61
0.75
0.74
0.72
0.75
0.70
0.70
0.70
0.72
0.70
0.72
0.71
CP
14.32
16.08
18.44
16.43
15.62
20.13
16.87
16.49
17.42
12.19
14.46
14.77
14.12
15.55
13.12
15.64
13.98
12.74
14.67
14.21
14.25
13.97
13.10
14.37
16.50
14.97
15.14
10.09
10.39
9.91
9.08
8.93
8.87
10.70
9.17
9.83
8.46
9.85
9.06
8.72
9.49
9.47
9.67
8.52
206
Table 3. continued,
YEAR
TIME
Site
LOCTRT
GRAZIRT
1'JDF
ADF
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
AUG
AUG
2
TExc
Graze
0.64
AUG
3
AUG
AUG
AUG
AUG
AUG
AUG
AUG
3
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
1
1
Cant
Clear
Clear
Clear
Thin
Thin
Thin
Cont
Cont
Cant
Clear
Clear
Clear
Thin
1
Thin
1
Thin
1
Cont
0.43
0.34
0.35
0.34
0.37
0.39
0.39
0.46
0.43
0.46
0.42
0.44
0.42
0.43
0.44
0.37
0.44
0.45
0.45
3
3
3
3
3
3
3
1
1
CEXc
TExc
Graze
CExc
rExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
0.57
0.58
0,59
0.56
0.61
0.58
0.61
0.64
0.58
0.56
0.59
0.59
0.53
0.60
0.48
0.51
0.57
1
Cant
JUNE
1
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
2
2
Cant
Clear
Clear
Clear
Thin
Thin
Thin
Cant
2
Cant
JUNE
2
JUNE
JUNE
JUNE
JUNE
JUNE
3
3
Cant
Clear
Clear
Clear
Thin
3
Thin
JUNE
3
Thin
CExc
TExc
JUNE
JUNE
JUNE
3
Cant
Graze
3
Cant
3
0.45
0.40
0.59
0.59
0.58
Graze
0.54
CExc
1
Cant
Clear
Clear
Clear
Thin
Thin
Thin
CEx
TExc
Graze
CExc
TExc
1
Cont
Graze
1
Cont
Cant
TExc
0.57
0.59
0.53
0.56
0.55
0.44
0.42
0.40
0.41
0.40
0.44
0.45
2
2
2
2
2
3
3
AUG
AUG
1
AUG
AUG
AUG
AUG
AUG
AUG
AUG
1
1
1
1
1
Graze
0.57
0.58
0.59
0.58
0.53
0.55
0.57
0.52
0.52
0.60
0.56
CExc
0.55
TExc
0.58
0.48
0.60
Graze
TExc.
CExc
0.57
0.57
0.58
O.41
0.41
0.41
0.41
0.40
0.45
0.43
0.42
0.48
0.38
0.37
0.37
0.39
0.43
0.44
0.44
0.44
0.41
0.46
IVOMD
0.68
0.76
0.77
0.74
0.75
0.68
0.69
0.69
0.63
0.67
0.84
0.81
0.81
0.80
0.82
0.85
0.79
0.79
0.86
0.79
0.81
0.79
0.84
0.83
0.81
0.79
0.86
0.82
0.83
0.85
0.83
0.84
0.84
0.83
0.79
0.80
0.79
0.75
0.70
0.75
0.71
0.76
0.71
0.72
0.76
0.74
CP
8.52
9.13
10.32
9.42
9.34
10.00
8.62
9.37
9.40
9.07
14.92
12.97
14.95
14.34
14.51
14.75
14.48
14,57
14.49
14.88
14.02
14.29
14.52
16.37
13.98
15.79
13.80
12.68
14.29
15.76
15.20
14.16
14.26
14.48
15.38
14.50
14.31
10.81
10.37
11.67
10.80
9.16
9.92
10.17
10.80
11.16
207
Table 3. continued.
YEAR
TIME
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
AUG
2
AUG
2
Site
LOGTRT
GRAZTRT
NDF
ADF
JVOMD
CP
0.57
0.46
0.71
9.96
0.58
0.40
0.73
9.80
AUG
0.72
2
0.60
0.45
12.20
AUG
0.38
0.75
2
0.54
11.02
AUG
0.37
0.78
2
0.50
11.76
AUG
Thin
0.59
0.43
0.72
2
10.62
AUG
0.37
0.76
2
Cont
0.55
11.05
AUG
0.43
0.71
2
Cont
CExc
0.53
11.52
AUG
0.77
2
Cont
TExc
0.62
0.46
10.61
AUG
0.38
0.77
3
Clear
Graze
0.56
11.87
AUG
Clear
0.54
0.41
0.76
12,09
3
CExc
AUG
0.75
13.44
3
Clear
TExc
0.56
0.37
AUG
Thin
Graze
0.54
0.40
0.72
12.97
3
AUG
0,53
0.42
0.72
11.60
3
Thin
CExc
AUG
Thin
0.54
0.41
0.73
12.53
3
TExc
AUG
3
0.59
0.44
0.74
10.37
Cont
Graze
AUG
3
Cont
CExc
0.58
0.44
0.68
10.96
AUG
3
Cont
TExc
0.53
0.44
0.63
12.01
Timber harvest treatments: Clear clearcut, Thin crownthinning; Cont contrOl.
Herbivorv treatments: Graze cattle and big game grazing; CExc cattle exciosure, big
game grazing only; TExc total exclosure, exclusion of cattle and big game grazing.
Clear
Clear
Clear
Thin
Thin
Graze
CExc
TExc
Graze
CExc
TExc
raze
Table 4. Botanical composition and relative preference index for steer diets collected in
June and August of 2001 and 2002 for thgd fir sites.
otanicalCQrnpJtiQfl
YEAR
TIME
Site
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
JUNE
JUNE
JUNE
1
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
1
Timber
1
Clear
Clear
Clear
Cont
Cont
1
Con
1
Thin
Thin
Thin
Clear
1
1
1
1
2
2
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
3
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
Herb.
HarvestTrt.
Clear
Clear
Cont
Cont
Cont
Thin
Thin
Thin
Clear
Clear
Clear
Cont
Cont
Cont
Thin
Thin
Thin
Clear
Clear
Clear
Cont
Cant
Cant
Thin
Thin
Thin
Clear
Clear
Clear
Cant
Cant
Cant
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
Relative Pref.
Grass
Forhs
Shrubs
Grass
Forbs
Shrubs
87.92
83.04
83.05
77.94
59.46
72.01
81.71
81.82
82.56
86.43
88.25
89.61
77.88
73.75
80.35
92.72
69.55
82.91
90.05
93.81
88.04
77.54
64.89
71.82
87.00
89.85
75.31
92.71
12.08
16.96
16.95
22.06
38.65
27.52
18.29
18.18
17.44
13.57
11.12
10.39
22.12
23.52
19.65
7.28
30.01
15.01
9.95
6.19
11.96
21.15
31.73
28.18
11.83
9.43
19.18
0.00
0.00
0.00
0.00
1.55
1.44
0.66
0.61
0.52
0.84
9102
8.98
6.93
14.13
35.90
45.50
11.87
22.56
15.74
7.51
10.25
16.31
14.75
17.54
18.30
0.00
0.00
0.00
0.00
0.06
0.02
0.00
0.00
0.00
0.00
0.03
0.00
0.00
0.13
0.00
0.00
0.02
0.06
0.00
0.00
0.00
0.06
0.08
0.00
0.06
0.04
0.30
0.00
0.00
0.00
0.05
0.02
0.00
0.00
0.05
0.00
0.00
0.00
0.02
0.02
0.08
0.06
93.07
84.62
63.57
54.50
88.13
76.17
84.26
92A9
89.75
82.95
84.59
80.G7
80.60
/.29
1.89
0.48
0.00
0.00
0.00
0.00
0.63
0.00
0.00
2.73
0.00
0.00
0,44
2.08
0.00
0.00
0.00
1.31
3.37
0.00
1.17
0.72
5.51
0.00
0.00
0.00
1.26
0.53
0.00
0.00
1.26
0.00
0.00
0.00
0.74
0.67
1.79
1.09
147
1,55
1.57
211
1.08
1.24
1.23
1.64
1.21
1.63
1.49
1.00
1.11
1.92
1.05
151
1.33
1.24
1.02
1.34
2.15
1.31
1.17
1.27
1.06
1.64
1.58
1.65
1.68
1.68
1.59
1.16
1.15
1.25
1.75
1.23
1.51
1.62
1.10
1.12
1.21
0.67
1.14
1.79
0.80
1.39
1.22
1.08
3.56
4.36
1.89
0.19
3.60
1.27
0.37
0.79
1.45
1.03
1.11
1.47
1.93
0.76
1.81
0.40
0.32
0.21
0.54
1.13
1.11
0.74
2.22
0.72
0.77
1.13
1.69
2.37
3.25
1.76
209
Table 4. continued.
Bo1inic&rnposition
YEAR
TIME
Site
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2001
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
AUG
AUG
2
AUG
AUG
2
2
3
Timber
Harvest
Thin
Thin
Thin
Clear
Clear
Clear
Cont
Cont
Cont
AUG
AUG
AUG
AUG
AUG
AUG
AUG
AUG
3
JUNE
JUNE
JUNE
JUNE
1
JUNE
1
JUNE
JUNE
1
Clear
Clear
Clear
Cont
Cont
Cont
1
Thin
JUNE
1
JUNE
JUNE
JUNE
JUNE
JUNE
1
JUNE
JUNE
2
Thin
Thin
Clear
Clear
Clear
Cont
Cont
Cont
Thin
Thin
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
JUNE
AUG
AUG
AUG
AUG
AUG
3
3
3
3
3
3
3
1
1
1
2
2
2
2
2
2
2
2
3
3
3
3
3
3
3
3
3
1
1
1
1
1
Thin
Thin
Thin
Thin
Clear
Clear
Clear
Cant
Cont
Cont
Thin
Thin
Thin
Clear
Clear
Clear
Cont
Cont
Herb.
Trt.
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Graze
CExc
TExc
Relative Pref. Index
Grass
Forbs
Shrubs
Grass
Forbs
Shrubs
85.54
82.86
81.97
95.57
95.08
93.09
77.36
57.60
79.78
95.67
73.64
93.59
95.50
12.84
16.50
17.47
1.61
1,77
1.26
1.49
1.41
1.25
1.08
1.33
1.91
1.45
1.28
1.04
1.32
1.69
1,51
1.67
1.53
2.00
2.01
1.16
1.26
1.21
1.81
1.29
1.75
1.57
1.15
1.23
1.95
1.35
1.64
1.41
1.28
1.04
1.28
1.87
1.64
1.25
1.06
1.18
1.62
1.42
1.55
1.19
1.17
0.33
1.98
1.48
0.16
0.63
0.84
0.99
1.17
0.93
0.63
1.92
0.60
0.25
0.46
0.18
0.83
0.76
0.74
0.75
1.59
0.85
0.44
0.47
0.30
2.82
2.91
1.08
0.15
1.24
0.82
0.17
0.32
0.13
0.02
0.02
0.00
0.00
0.00
0.11
0.22
0.10
0.02
0.14
0.00
0.00
0.00
0.00
0,06
0.00
0.04
0.00
0.02
0.00
0.00
0.08
0.03
0.00
0.00
0.00
0.00
0.02
0.00
0.00
0.00
0.00
0.07
0.13
0.00
0.00
0.03
0.10
0.00
0.00
0.00
0.03
0.24
443
0.64
0.56
0.00
0.00
0.00
2.27
9.19
2.53
0.46
2.41
0.00
0.00
0.00
0.00
94.20
76.76
4.92
6.91
20.37
33.20
17.69
3.88
23.94
6.41
4.50
12.87
5.80
21.94
75.81
24.19
0.00
68.83
87.91
30.23
0.94
12.09
0.00
8343
8162
16.09
18.38
4.30
0.48
0.00
0.00
4.29
2.90
1.54
1.17
0.00
0.00
0.00
0.00
0.49
0.00
0.00
0.00
0.00
1.52
5.52
0.00
0,00
0.58
1.92
0.00
0.00
0.00
0.79
7.18
87.13
95.70
94.17
95.93
82.48
84.32
88.73
94.14
89.:19
90.29
95.27
97.3
89.68
74.40
56.44
90.04
93.59
74.62
83.33
91.58
82.02
87.49
59.66
44.28
1752
15.68
11.27
5.86
10.32
9.71
4.73
2.47
10.32
24.08
38.05
9.96
641
24.80
14.75
8.42
17.98
12.51
39.54
48.55
1.31
1.25
1.17
1.34
0.52
1.04
1.99
1.39
0.46
0.64
0.39
1.50
1.52
210
Table 4. continued.
Baicijiposition
YEAR
TIME
Site
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
2002
AUG
1
AUG
1
AUG
1
AUG
1
Timber
Harvest
HerD.
Cont
Thin
Thin
Thin
Clear
Clear
Clear
Cant
Cant
Cont
Thin
Thin
Thin
Clear
Clear
Clear
Cant
Cont
Cant
Thin
Graze
Trt.
CExc
TExc
Gra
Relative Pref. Index
Grass
Forbs
Shrubs
Grass
Forbs
Shrubs
62.07
88.48
75.69
79.60
88.53
83.54
65.35
70.08
80.52
84.63
83.32
84.30
78.75
37)2
0.90
0.00
0.00
1.81
1.17
1.16
1.18
1.68
1.15
1.19
1.34
1.09
1.17
1.72
1.28
1.43
1.27
1.29
1.13
1.14
1.46
1.46
1.04
1.00
1.11
0.90
0.72
2.30
0.87
0.04
0.00
0.00
0.15
0.04
0.15
0.25
0.50
0.00
0.06
0.19
0.00
0.06
0.17
0.00
0.09
0.38
0.22
0.34
0.52
0.27
0.35
1152
31
I89
1.61
9.79
1.68
1.00
CExc
AUG
2
TExc
13.82
2.64
1.52
AUG
2
Graze
25.85
8.80
2.68
AUG
2
CExc
20.57
1.51
9.36
AUG
2
19.48
0.00
TExc
3.61
AUG
2
Graze
14.35
1.03
1.38
AUG
2
CExc
14.29
2,38
0.36
AUG
2
15.70
0.00
TExc
1.88
AUG
2
Graze
19.22
2.03
1.62
AUG
3
0.85
CExc 8582 13.33
0.49
AUG
3
0.00
TExc 98.13
1.87
0.24
AUG
3
1.49
0.48
Graze 98.03
0.18
AUG
3
8.06
1.26
CExc 66.13 25.82
AUG
9.25
3
TExc 43.99 46.76
1.64
AUG
3
Graze 80.24 10.83
8.93
0.57
AUG
10.05
3
CExc
7732 12.63
2.06
AUG
3
Thin
4.60
TExc 70.64 24.76
1.99
AUG
Thin
6.43
3
Graze 78.57 15.00
1.41
Timber harvest treatments: Clear clearcut, Thin crownthinning; Cant control.
Herbivory treatments: Graze cattle and bg game grazing; CExc cattle exclosure, big
game grazing only; TExc total exclosure, exclusion of cattle and big game grazing.
AUG
2