Variation in early growth of western white pine in north Idaho

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VARIATION IN EARLY GROWTH
OF WESTERN WHITE PINE
IN NORTH IDAHO

R. J. STEINHOFF

USDA FOREST SERVICE
RESEARCH PAPER INT-222
INTERMOUNTAIN FOREST AND RANGE EXPERIMENT STATION
FOREST SERVICE, U.S. DEPARTMENT OF AGRICULTURE

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USDA Forest Service
Research Paper INT-222
June 1979

VARIATION IN EARLY GROWTH
OF WESTERN WHITE PINE
IN NORTH IDAHO

R. J. Steinhoff

INTERMOUNTAIN FOREST AND RANGE EXPERIMENT STATION
Forest Service
U.S. Department of Agriculture
Ogden, Utah 84401

THE AUTHOR

R. J. (RAY) STEINHOFF received his formal education at the University of
Idaho, North Carolina State University, and Michigan State University.
From 1963 to 1965, he worked at the Institute of Forest Genetics at
Placerville, Calif., a part of the Pacific Southwest Forest and Range
Experiment Station. Since 1965, he has been at the Moscow Forestry
Sciences Laboratory where he is responsible for research on the genetics
of growth and adaptation of western white pine and grand fir.

ACKNOWLEDGMENTS

Special thanks are due to Tony Squillace and Burton Barnes, formerly
with this Station, who initiated the older studies reported on here and re-
viewed the manuscript. Bob Campbell, geneticist with the Pacific North-
west Station in Corvallis, also reviewed the manuscript and provided many
helpful suggestions. My colleagues Ray Hoff, Geral McDonald, and Jerry
Rehfeldt have also contributed materials, assistance, and encouragement.

RESEARCH SUMMARY

In nursery and field tests representing five different studies, most of the
variation in height growth or other traits in any one test area was among se-
edlings within family plots. Of tne useable genetic variation in those traits,
most (60-90 percent) was related to differences among the parent trees within
stands. The remaining genetic variation was spread among geographic areas,
elevational zones, or stands within these categories. When differences among
the latter catagories were significant, usually no pattern in the variation was
apparent. Thus, although seedlings within families and family means within
stands vary considerably, the lack of differences among stand means, or those
for broader geographic areas, and the absence of distinct variation patterns
suggests that inland western white pine is fairly uniform. However, seedlings
from high-elevation parents (generally 1,375 m or higher) were significantly
shorter than those of low- and mid-elevation parents in some low- or mid-
elevation tests. Even when elevational effects were not significant very few
high elevation families were among the tallest. Growth results to date indicate
that only two seed zones are needed in north Idaho--a low zone and a high zone
generally separated at 1,375 m. Within the lower zone, trees with good
growth potential can be found in nearly all stands.

CONTENTS

INTRODUCTION .....
PRIOR RESEARCH .
MATERIALS AND METHODS
RESULTS .

DISCUSSION AND CONCLUSIONS.
APPLICATIONS

PUBLICATIONS CITED.

APPENDIX

Page

20

21

INTRODUCTION

Western white pine (Pinus monttcola Dougl.) has long been an important component of
the forests of north Idaho, but the introduction of blister rust (Cronartium rtbicola
J.C. Fisch. ex Rabenh.) and the ensuing losses to the disease has dissuaded land mana-
gers from attempting to manage the species. Now, as efforts to produce trees resistant
to the disease show some promise, land managers are again beginning to include white
pine in their regeneration programs. In the long run, the natural selection process
should produce trees with sufficient blister rust tolerance or resistance to allow natu-
ral regeneration. For the present and near future, however, planting of resistant seed-
lings will be needed.

The production and utilization of resistant planting stock involves much more than
just securing an adequate level of resistance in a sufficient proportion of the seedlings
to be planted. The manager wants the seedlings to survive and grow on the sites he wants
to plant and he wants them to grow at a rate which fully utilizes the potential of the
Site. Attempting to achieve such an ideal match of seedling and site is not a realistic
goal at present because of our lack of knowledge of both trees and sites. Also, the
costs to approximate such a match would likely exceed the returns from greater yields
as the limits of the site are approached. The realistic goal for the present is to
attempt to achieve a balance that incorporates our present biological information and
approximate costs with expected growth rates and economic returns.

This paper will discuss past and current investigations into variation in growth
of western white pine in nursery trials and several experimental plantations. The
conclusions of the research will then be translated into guidelines for distributing
seed from orchards that have already been established and to evaluate alternative
designs for future seed orchards.

PRIOR RESEARCH

During the past 20 years several reports on variation in western white pine have
appeared. Squillace and Bingham (1958) opened the series with a report of "localized
ecotypic variation" based on germination and early growth of seedlings from quite dif-
ferent sites within a small area. They concluded that seedlings from trees growing on
poorer sites were shorter in nursery trials than those from trees growing on better
Sites. Germination of seeds in a sucrose solution was higher for poorer site trees
than for better site trees. In a low-elevation nursery, seedlings from low-elevation
trees were taller than those from high-elevation trees, but when transplanted to a
high-elevation test site the seedlings from the high-elevation trees grew faster. On
the basis of those findings, and other considerations, the applied program to produce
blister rust resistant western white pine was subdivided into three elevational zones
or breeding units with a seed orchard planned for each. These zones were: (1) Low-
elevation--areas below 1,000 m; (2) mid-elevation--areas between 1,000 m and 1,250 m;
and (3) high-elevation--areas above 1,250 m.

Progeny of many of the same parents used in the above studies were outplanted to
two additional sites for long-term evaluation. Rehfeldt and Steinhoff (1970) found that
at age 14 average growth at the two sites differed significantly and that the growth of
individual progenies differed significantly but the average growth of progenies from
different localities did not differ significantly.

In order to obtain materials representative of a broader range of sites, Barnes
(1967) systematically collected seed and recorded data from trees in a series of eleva-
tional plots along several north Idaho streams. He reported finding significant dif-
ferences in periodic annual growth rate and branch angle that were related to eleva-
tional differences along one transect. Trees in plots from 760 m elevation to 1,220 m
grew at essentially the same rate, but those from 1,400 m and above were slower growing.
Branch angle changed progressively from low to high plots, with trees in the lower plots
having more ascending branches and those in the higher plots having more nearly hori-
zontal branches. Differences among plots were also found in needle length and cone
scale length and width, but the variation did not follow a pattern related to elevation.

Some of the seed collected by Barnes was sown in a nursery trial in Michigan.
After 4 years of growth, total height of seedlings representing the various elevational
and latitudinal collection areas did not differ significantly (Townsend and others 1972).
Seasonal growth rates differed, but they did not fit into a pattern related to elevation
or latitude. At a very low light level (130 lux) low-elevation seedlings (640 m) were
less efficient photosynthetically than higher elevation seedlings (1,065 m and 1,585 m),
but at higher light levels (425 and 615 lux) differences were not apparent.

MATERIALS AND METHODS

This report will present and integrate results from five studies, as follows:

I Elevational study.--This study was started to further investigate the effects
of elevation (as indicated in Squillace and Bingham 1958) and possibly latitude on
seedling growth. Wind-pollinated seed for the test was collected from individual trees
in plots (stands) at 120 m to 180 m (mostly 150 m) elevational intervals in several
stream drainages in north Idaho. Although the collection locations were often along
streams, they were kept out of the stream bottoms where cold air drainage effects are
common. Five trees from each of 1 to 3 plots per elevation and drainage were included,
for a total of 225 trees in 45 plots (fig. 1). In the Trestle Creek drainage, the
plots within an elevational zone were close together (mostly within 0.5 km), but in the
other areas the plots were often several kilometers apart. Further details of plot loca-
tion for the Trestle Creek drainage were presented by Barnes (1967).

A nursery test for early growth and seedling traits was conducted for 3 years at
Moscow, Idaho. The seed were sown at a spacing of 5x10 cm in 10-tree row plots in 4
replicates. The growing medium was a mix of forest soil, sand, and peat moss in equal
proportions. The beds were watered and fertilized as needed to maintain satisfactory
growth. Annual height measurements were taken after growth was completed.

Field plantations have been established at the Priest River Experimental Forest
(PREF) and near the North Fork of the Clearwater River on the Canyon Ranger District

(Canyon) (fig. 1). In each area, 3 test plantations were established with an eleva-
tional separation of 300 m (800 m, 1,100 m, and 1,400 m at PREF and 900 m, 1,200 mn,
and 1,500 m at Canyon). Aspect of the sites ranges from northeast to northwest. The

field tests contain representatives of nearly all the seed collection areas and eleva-
tion zones but do not contain representatives of all the plots within elevational zones.

LEGEND

© Elevational Study Collection Areas

Kaniksu © Natural Selection Study Collection Areas

Boulder Creek
VY Elevation-Aspect Study Collection Areas

Trestle yw Nursery Test Sites

Creek

SO

%& Elevational Study Plantations

© Vigor-Quality Plantations

Factorial Mating Design Study Plantation

St. Joe River

Crystal Creek A COCO
= A
\ x Fernwood ~— e VV Marble Creek
9 Neo Re Bae AY
Clarkia ‘| NG Gold Center Creek
6 | CJ White Rock
(@) SS sy oe a aa a SS 5555 eTcC5
Oneal

\ a)
| Elk Creek

Bertha Hill

N.F. Clearwater

Figure 1.--Seed collection areas and test sites for the vartous white pine studies in
north Idaho.

An additional plantation (Ida Creek) adjacent to the low PREF site contains representa-
tives of all the families included in the nursery test. The seedlings (3-0 stock) were
planted in 1971 at a spacing of 1.8 m in 4-tree row plots replicated 5 times. They were
all measured at age 4 (the end of the first field growing season). The Ida Creek plot
was measured again at age 6 and age 8 and the other plantations were measured at age 7.

Dis Elevation-aspect study. --An attempt was made to evaluate the effects of environ-
mental factors related to both elevation and aspect in this test. Study areas were
located in two drainages (Marble Creek and Gold Center Creek) near Clarkia, Idaho
(fig. 1). They consisted of a series of paired plots at 950 m, 1,175. m, and 1,400 m in
each drainage. One member of each elevational pair had a southern exposure and the
other a northern exposure. In each plot, four trees were chosen to be seed parents and
three to be pollen parents. Pollen from the three trees on a plot was mixed. Each seed
tree was pollinated with four pollens, the local pollen mix plus pollens from the low-,
mid-, and high-elevation north-aspect Marble Creek plots. For this portion of the study
seeds from the four trees on a plot were combined for each pollen type. Results of
crosses among individual trees within plots are being reported elsewhere (Rehfeldt, in
press).

A nursery test was conducted at Moscow for 3 years. Seed was sown in plots consist-
ing of 2 rows of 18 planting spots. Spacing was 5xl0 cm. There were two replicates.

The seedlings have been outplanted to forest test sites but have not yet been
measured.

Se Natural selectton study.--This test was originally designed for studying blis-
ter rust resistance, but its potential for studying growth variation within and between
stands was soon recognized. Ten stands were located and wind-pollinated seed collected
from 50 trees in each. Three of the stands are from the same general areas represented
by most of the trees in the vigor-quality study (No. 5 below) and one stand corresponds
to one of the elevational study collection areas (fig. 1). Six of the stands fall within
the low-elevation zone of the applied program (below 1,000 m) and four within the high
zone (above 1,250 m).

The seedlings were started and grown for one season in a greenhouse in 5x5x15 cm
tar paper plant bands in a soil, sand, peat moss mix. They were then transplanted, still
in the plant bands, in the PREF nursery site. Eight seedlings of each of the 497 fami-
lies were planted at random in each of 3 replicates. Spacing was 10xJ0 cm. The seed-
lings were given routine nursery care. They were inoculated with spores of the white
pine blister rust fungus (Cronartium rtbicola) at age 3 and measured at age 4, before
the disease had any noticeable effect on growth.

Other seedlings from the same families have been outplanted to a field test site
but have not been measured yet.

4. Factortal mating destgn study.--This is another test that can serve both blis-
ter rust resistance breeding and growth analysis. Three factorial mating designs, one
each for the low-, mid-, and high-elevation zones, were established. Four trees were
used as pollen parents and approximately 40 trees were used as seed parents in each zone.
However, the majority of the seedlings in the test plantation represent only 40 families
from each zone (4 pollen parents x 10 seed parents).

The seedlings were grown for 6 years in replicated row plots at the Moscow nursery.
They were inoculated for studying white pine blister rust resistance. No growth measure-
ments were taken in the nursery.

The survivors from the blister rust testing were outplanted to a low (750 m),
nearly flat site at PREF in 1971. A variable number (average 15) of seedlings per

family were planted as single tree plots at random throughout the plantation. Growth
during the 3-year period from 1973 to 1975 and height in 1976 have been measured.

5. Vtgor-quality study.--This is the oldest of the five studies. It was started
during the early stages of the program to produce blister rust resistant western white
pine (Squillace and Bingham 1954). Excess seed from the program was used for this study
to investigate the heritability of growth and quality traits. The parents of many of
the seedlings grow in the Crystal Creek drainage near Fernwood, Idaho, at an elevation
of about 870 m (fig. 1). A few others grow at three additional low sites within 45 km.
Four candidates from the White Rock area at an elevation of 1,525 m near Clarkia, Idaho,
were also included. Squillace and Bingham's (1958) report of "localized ecotypic varia-
tion in western white pine'' was based on other seedlings of many of these same families.

Seedlings (2-1) for the 1955 outplantings were grown in a nursery in Missoula, Mont.
All the others were grown in Spokane, Wash. The seedlings were not measured in the
nurseries.

Three plantations were installed during the years from 1955 through 1959. Data
from the 1955, 1956, and 1957 plantings at PREF and Deception Creek Experimental Forest
(DCEF) are included in this report. The PREF plantation is located on a moderate (10-
25 percent) north to northwest-facing slope at an elevation of 790 m. The DCEF planta-
tion is on a steep (30-50 percent) north-facing slope at 1,110 m (fig. 1). Seedlings
(mostly 2-1 transplants) were planted at a spacing of 2.4 m in 8-tree row plots in three
replications at each site. Competition from native vegetation has been reduced by weed-
ing and brush cutting at irregular intervals. The trees have been measured several
times--most recently when all had been in the field for 16 years.

RESULTS

Elevational study.--The most striking feature of the nursery results is the general
lack of patterns in the variation observed (table 1). Although the amount of variation
present was large, much of it was related to differences among the offspring of a single
tree or among the trees within a stand. This may be seen in the proportion of variance
attributable to the various sources and the common significance of the "'trees within
Stands' mean squares (table 2 and Appendix table 15). Because of the magnitude of the
within-stand variation, the differences among stands, elevational zones, or geographic
areas were seldom statistically significant.

Seed weight did not differ significantly from one area to another and only within
the Clearwater drainage were there significant differences among elevational zones.
There, seed from the higher elevation trees was lighter than that from the lower trees,
with a rather consistent gradation between.

Seeds from the Trestle Creek area germinated sooner than those from other areas,
but there was no pattern to the variation within areas. Whether the germination dif-
ferences are inherent or were related to cone collection time or subsequent handling is
unknown. Time of germination was closely related to first-year seedling height and
some of the effect was still detectable at the end of the third year. All data for
seedlings germinating more than 21 days after the start of germination were excluded
from these and the following analyses.

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Table 2.--Percentages of total vartatton attributable to vartous sources for several seedling traits in the elevattonal

study nursery test

Traits
Source Degrees :Germi-: lst :Adjusted: 2nd 2nd 3rd : Adjusted : Coty- : Coty-
of of : Seed :nation: year :lst year: year year year 3rd year :Needle ledon ledon
variation freedom :weight: time : height: height!: height:lammas: height: height? :length No. length

vartatton

Percent of tota

Trestle Creek vs.

Replications? 3 5.4 2.9 0.6 2.8 2.9 4.4 Ose) 4.0 0 (0)
Areas 1 0 24.4**4* 18. 6** Deals SOMA WGA MONG A (0) ily |* BED AIS 7
Elevation zones

within area 13 NZ. 6) eal 225) 0 eal 2.0 0.4 0.1 0.6 0.7 3
Stands within

elevation & area 18 0 So 8) 1.6 Zeal Ball Soe Bas Bs oer 0.6 0) 0.
Trees within stand,

elev., & area 130 87.4 DAO CA ey Arak ORS Se ORs aOR zis 2.4* SiS in Issaausese AL7/
Experimental error® 471 12.4 9.3 16.9 22 30.9 23.4 26.1 24.5 4.2 Wal
Within plot® 28.6 41.0 63.9 41.8 48.2 47.5 66.5 63.5 Voll 62

Clearwater, St. Joe, Trestle and Kaniksu but only 915 m, 1065 m, and 1220 m Elevation Zones

Replications? 3 Sad 1.9 0.4 Zea5) 4.5 25, ley 4.8 0.1 Le
Areas 3 0 Why Pett = ANA) Stes 1.9 HORSIX Seo Sr 0) aS OR 2 0
Elevation zones

within area 8 0 0.6 Dir 0 3.4 2is5 Si5) od 0 1.0 (0)
Stands within

elevation §& area 11 8) 0) 0) So/ LS 2.0 Pas) 2:14'* Soc 4.8* 6.
Trees within stand,

elev., §& area 92 98.2 ZOO R ee ZO MOR meplifio ee DNA Boek iskoxs S09 sa Ws ss AAI
Experimental error® 334 21.6 10.3 ikea 14.3 30.5 22.0 20.4 ONGd 5.3 9.
Within plot® 32.0 44.2 64.8 46.6 53.6 50.8 68.4 64.1 72.9 60.

TPirst year height adjusted for seed weight and germination time effects.

*Third year height adjusted for first year height differences.

30nly 2 replicates were used for cotyledon number and length analyses so degrees of
error were adjusted accordingly.

* and ** indicate significance at the 5 percent or 1 percent level, respectively.
variance see Appendix Table 15.

°Inciudes all interaction terms involving replication.

®Pooled estimate from 30 to 50 plots with 5 to 10 seedlings per plot.

Clearwater--All Elevation Zones from 455 m to 1585 m

S*

.5**

freedom for replicate and

For detailed analyses of

When the raw data for the first-year height were analyzed, the mean squares
associated with areas were highly significant. This was true for both comparisons,
for example, between Clearwater and Trestle where the broadest elevational range was
sampled, and among all areas but utilizing only the 915 m, 1,065 m, and 1,220 m eleva-
tion zones (those common to all areas). The early germinating Trestle seedlings were
taller than the later germinating seedlings from the Clearwater and Kaniksu areas.
However, when first-year heights were adjusted for seed weight and germination time
the variability related to areas was much reduced. The comparison of all areas using
the middle elévational zones showed no significant differences, but the Trestle Creek
and Clearwater areas differed at the 5 percent level when all elevation zones were
considered. These adjusted means differed by only 2 mm--approximately 4.5 percent of
the mean.

Subsequent height measurements taken in the second and third years followed a pat-
tern similar to first-year height. Analyses of the raw data showed significant differ-
ences among trees within stands and among areas. There were few or no significant dif-
ferences among elevations within areas or stands within elevations. When the data were
adjusted to take into account first-year height differences, second- and third-year
height differences among areas were not significant. In other words, the Trestle Creek
seedlings, which germinated early and were tallest at age 1, were still tallest at age 3
but relative to their height at age 1 they had not grown as much as seedlings from the
Clearwater and Kaniksu areas.

Whether such adjustments should be made depends on the cause of the germination
time difference. If it is the result of genetic differences, no adjustment should be
made. If the difference resulted from cone collecting or seed processing timing or
techniques, then the adjustments would be appropriate. In the absence of information
to indicate genetic differentiation of germination time, I favor use of the adjusted
data.

If the range of 120- to 180-m elevational zones is combined across areas to form
three broader zones called low, mid, and high (<1,000 m, 1,000-1,250 m, >1,250 m) like
the classification in the following studies, then seedlings of the low zone were tallest
and those of high zone were shortest. At age 3, heights by zone were: low - 220 m,

mid - 211 mm, high - 207 mm. These means did not differ significantly.

Secondary leaf }ength (mature third-year leaves) was the only other trait in which
differences among areas were significant. Within the Trestle Creek and Clearwater
areas, differences between trees within stands were significant but for the other areas
they were not. Stand-to-stand differences within an elevational zone were sometimes
large but seemed to occur at random. Significant cotyledon differences--number and
length--were confined to trees within stands and stands within elevational zones.

Bud burst at the start of the third year did not differ significantly among stands,
elevations, or areas (table 3). The highest elevation source from Trestle Creek was
the first to break bud but the next to highest source was last. All buds broke within
a 2-week period and most within a 5-day period. The newly emerging leaves were rapidly
separated and in from 10 to 14 days the tip of the new bud was distinguishable. At
that time the seedlings were classified as having set buds even though stem elongation
and external bud development continued for nearly a month. No significant differences
in bud set time were observed except among trees within stands.

The seedlings were 7 years old and had been in the field plots for 4 years when
they were last measured. Overall survival of seedlings of Clearwater origin was about
10 percent less than that of Trestle seedlings with the differences greatest at the
Canyon series of test sites. However, survival differences so far do not clearly indi-
cate differences related to area or elevation of origin or their interactions with test
site conditions. Average height ranged from 54 cm at the best site, low PREF, to 34 cm
at the poorest, high PREF (table 4).

Table 3.--Analyses of vartance and percent of total variance for third-year bud burst
and bud set in the elevational study nursery test

Bud burst : Bud set
: ; Mean PNG Oie TCoOcel 3 ween Te oLatotall
Area : Source qide square ec) vwanialtaon : Square : variation

Trestle Replications 3 OSral 28 163.5 3.4
Creek
(640 to Elevation 6 26.4 OG LOE 12
1585 m plots)

Plots within

elevations 7 20.8 0.9 48.0 0
Trees within plots
within elevations 28 12.1 0 52. G36 8.1

Experimental error! 84 16.8 29.6 20.9 4.9

Within plot 672 Soe 66.1 Gall 82.4
1220 m plots Replications 3 14.2 Deal 104.4 555
from Clear-
WENEEIOS (Sed Areas 4 Sod Hoes 104.0 2.9
Joe, Trestle
Creek, Trees within areas 10 Woe 6.3 Og Pe aS)
Kaniksu

Experimental error! 42 6.5 9.0 20.2 5.0

Within plots 240 Aves 74.8 NS sal 74.6

lTIncludes all interaction terms involving replication.
2x and ** indicate significance at the 5 percent and 1 percent level, respectively.

At the individual test sites the results of the analyses of total height are very
similar to those from the nursery trial (table 5S presents the results for only the most
variable series of sites, i.e., PREF). Most of the variability is related to differences
among the families representing individual trees within stands. The variability within
plots (i.e., among seedlings of the same family) is also large--nearly equaling that
among plots (experimental error). This result indicates that within-family variation
is large or microsite differences exert strong influences or that both are important
contributors. The variation related to area of seed origin or to elevational zones
within areas was very small in comparison and usually not significant.

The Trestle Creek seedlings were taller than those from the Clearwater at the low
and high sites at both PREF and Canyon, but the Clearwater seedlings were as tall or
taller at the mid-elevation sites. No pattern could be seen in the variation in height
among seedlings related to the various elevational zones within areas. Neither was
there any correspondence between elevational zones in the different areas. For example,
seedlings from the highest Clearwater collections grew well at most test sites but those
from the highest Trestle Creek collection grew poorly. As a group, the seedlings from

Xe)

Table 4.--Height of seedlings at age 7 in the elevational study field tests

Seed collection : Test region and site

Canyon! PREF2 ‘Overall

Area : Elevation : Low : Mid : High : x : Low: Mid :High : x Ree
MOLT So Centimeters = = = ===> => = ===:

Clearwater 760 Sy, 40 38 38 55 39 30 43 41

915 42 40 37 40 53 38 34 42 41

1065 57] 4] 515 Si 51 35 32 40 38

1220 41 41 35 39 50 36 55) 40 40

1370 40 43 35 40 53 34 32 41 41

1585 44 43 38 42 54 36 34 43 42

x 40 42 36 39 53 S57/ 35 41 40

Trestle 760 42 42 36 40 53 Sy 35 42 41

915 40 42 42 41 5S 35 34 41 41

1065 45 41 38 41 Si/ 36 36 44 43

1220 39 41 35 39 55 39 35 44 42

1400 42 40 37 40 56 $5 34 43 41

1585 39 39 37 38 4 Si) 34 42 40

xX 41 4] 38 40 55 37 35 42 41

Overall x 41 4] 37 40 54 37 34 42 41

lCanyon Ranger District: low = 900 m, mid = 1,200 m, high = 1,500 m.
*Priest River Experimental Forest: low = 800 m, mid = 1,100 m, high = 1,400 m.

Table 5.--Analyses of variance and percent of total vartatton for hetght of 7-year-old
seedlings itn the elevattonal study sites at PREF

Low site : Mid site : High site
0 me

Source : Ghgate : Mean : % of total : Mean % of total : Mean : % of total
: :square: variation :square: variation :square: variation

Replicates 4 134 0.0 114 0.0 269 0.4
Area 1 1,013*! 0.8 5 0.0 793% WO
Elevation

within Area 10 255 0.0 155 0.4 68 0.0
Rep * Area

G. Eley) (Exon 21) 44 240 2.4 Si 6.0 192 6.4
Trees within

Elev § Area 47 BO 6.6 140** Holl 220 8.2
Repey aree

(Exro7 92) 170 Wes, S38) 79 SoZ 95 2.4
Within Plot 565 149 84.9 70 81.3 88 81.6

lx and ** indicate significance at the 5 percent and 1 percent levels,
respectively.

10

the 760 m Clearwater collection grew poorly at the low- and mid-elevation sites at Canyon
but well at the high site. In contrast, at PREF they grew well at the low- and mid-
elevation sites but poorly at the high site. However, these fluctuations are rather
small--only 2 or 3 cm (5S percent) above or below the site mean--and, so far at least,
appear more accidental than the result of specific tree-site interactions.

When the overall variability across sites within test regions was analyzed (table 6),
the variation associated with trees within elevational zones was still predominant over
that for elevational zones or areas. The site x tree interaction effect was significant
at PREF. When the tree mean squares were tested against this interaction term rather
than the error mean squares, the tree effects were significant at only the 5 to 10
percent level.

The average height of the seedlings over the three sites at PREF was essentially
the same as that for the Canyon sites but the range at PREF was much greater. Neither
group of sites shows a typical spread of conditions for the area. At PREF, growth and
survival at the mid-elevation site has been reduced due to poor site quality and resid-
ual competition resulting from site preparation problems. At Canyon, the low site
trees suffered from similar site preparation problems.

Table 6.--Analyses of vartance and percent of total variation for hetght of 7-year-old

seedlings of the elevational study combined over sttes at Canyon and PREF

Canyon : PREF
Meantem iotmtotale i aMean:) sof Ttotal
Source : Gl site : square: variation :square: variation
Test sites! 2 3,829 4.9 106, 038 SDP
Replicates within sites 2 302 sil 72 0
Area 1 W27/ 0 493 0
Site * Area 2 Wag 0 658 0.4
Elevation within Area 10 290 OZ 278 0
Seen ELevation 20 168 0 101 0
RepramAweaG Elev, (Error, 1) NZ 149 DS 188 250,
Trees within Elevations
and Areas 48 DADE 1.9 283 Os
SES FSS 94 142 Drool DN 28
REO ~ eee ((lsieie@ie 2) 500 111 0 119 Wes)
Within Plot 116 87.0 109 40.4

lhLow, mid, and high elevation.

2* and ** indicate significance at the 5 percent and 1 percent level,

respectively.

At Canyon d.f. = 1,001, at PREF dof.

1]

1,694.

In the Ida Creek plantation adjacent to the low PREF plot, where all the families
included in the nursery test were planted, area differences were significant at age 4
but became less important and nonsignificant by age 6. Actually, the difference between
area means remained nearly constant (2 cm between Clearwater and Trestle Creek), but as
a percentage of total height that difference dropped from 7 percent at age 4 to only
2 percent by age 8.

When all the families were grouped as to low, mid, or high origin, as with the
nursery data, then height at age 8 decreased from low to high elevation (101, 100, and
98 cm, respectively). These differences were not significant and that result is con-
trary to the findings in the adjacent low plot where the high-elevation seedlings were
slightly taller than the low- and mid-elevation seedlings.

Elevation-aspect study.--Height at the end of the third year in the nursery
(table 7) did not differ significantly among drainages, elevations, aspects, or pollen
types. Rehfeldt (in press) also found that at age 6 differences among drainages, eleva-
tions, or aspects were not significant when tested by individual control-pollinated
families within plots. Seedlings from south aspect plots in this study were generally
taller than those from north aspect plots in the same drainage and at the same elevation.
This was also the result for the Gold Center Creek drainage in Rehfeldt's study, but
seedlings from north and south aspect Marble Creek plots were essentially equal in height.

Table 7.--Third year hetght of seedlings in the elevation-aspect study

Seed parent location : Pollen source
: : Marble Creek
Elevation : > north aspect
Drainage : zone! : Aspect : Low Mid High =: Local, <a) gx. 3 Seabees

------ Hetght (millimeters) - - - - - -

Gold Center Low North Wal Vall US 64 70 75
Creek South 67 84 80 87 79
Mid North 75 73 83 -- 80 83 78
South 76 69 87 108 85
High North WS 85 65 72 74 76
South Wil 73 84 84 78
Marble Low North -- -- 88 85 86 85
Creek South 84 87 V7 87 84
Mid North 74 eo 75 79 76 78 81
South 84 78 74 84 80
High North -- -- -- 75 75 78
South 64 84 87 84 80
5d a 7380 83
x x
fs 79 N 76
M 80 S 8l
H 78

ILow = 950 m, mid = 1,175 m, high = 1,400 m.

Overall, the seedlings produced by local pollen were taller than those produced
with pollen from the north aspect plots but the differences were not significant. For
the south aspect plots, local pollens produced seedlings significantly taller than those
produced by the north aspect pollens. On the north aspect plots, pollen from one of the
other north plots usualiy produced taller seedlings than did local pollen.

Among the north aspect Marble Creek plots, seedlings from the low plot were tallest
but pollen from the plot produced the shortest seedlings when used on trees in other
plots. For the high plot, the results were the opposite (seedlings from the plot trees
were short but pollen from the plot produced tall seedlings when used elsewhere). Such
reversals were probably the result of large amounts of within-plot variability coupled
with the small numbers of trees chosen to be pollen or seed parents in any one plot
rather than resulting from plot interactions.

Natural selection study.--In this study only growth during the fourth growing season
was measured and analyzed (tables 8 and 9). The mean growth of the 4 high-elevation
stands was significantly less than that of the 6 low-elevation stands (10.8 cm vs.

11.8 cm). However, the best high-elevation stands grew more than the poorest low-
elevation stand. Among the 6 low-elevation stands, the 4 fastest growing ones differed
significantly from the slowest growing one.

Even though the differences between elevational groups and among stands were signif-
icant, the bulk of the genetic variation (61 percent) was associated with differences
among trees within stands. Of the remaining genetic variation, 28 percent was due to
elevational differences and 11 percent to differences between stands within elevational
zones. Overall, 62 percent of the variation was related to variation among seedlings
from the same tree, 2 percent to block interactions, and 36 percent to genetic variation.

Table 8.--Fourth year height growth of seedlings tn the natural selection study

Number of

: Elevational : : Range of : families
Stand : zone! : Growth : family means : in top 10%
- - - - Centimeters - - - - -

Beaver Creek pole Low eayaa 8.8-15.4 12
Crystal Creek Low N52 9.3-14.9 9
Beaver Creek mature Low Ao, &\ 8.3-16.1 12
Montford Creek Low FOF alb 9.4-15.2 7
Fighting Creek Low 11.7 abc 9.3-16.4 53
Rescue Creek High 14) bed 8.2-14.8 3
Bertha Hill High Iota cd 7.8-14.7 1
Elk Creek Low 10.9 cd 85 2=15 57 1
Boulder Creek High HO Siaca 8.0-14.9 2
White Rock High 10.6 d PeSoN GeO 0)

lLow = below 1,000 m, high = above 1,250 m.
Means not having a letter in common differ at the 5 percent level of significance.

13

Table 9.--Analysts of fourth year hetght growth of seedlings in the natural selection

study
Percentage
: : : Variance : of total

Source 5 Glaiee : Sao a= WIbS> : F : component : variance
Block 2 622 Sill 0.09 0.005
Stand 9 3260 362 7.9%! JOS) .02

Elevation 1 1728 22 9. 0* .24 .014

Stand (Elev) 8 1538 192 AO mull . 006
Block*Stand 18 825 46 6 dil . 006
Tree (Stand) 487 15468 32 Dro 495 . 06
Block*

Tree(Stand) 969 14727 WS .00 .00
Within family 7054 108648 5 15.40 Sil

l* and ** indicate Significance at the 5 percent or 1 percent level, respectively.

Factortal mating destgn study.--Seedlings from crosses among mid-elevation (1,000-
1,250 m) trees grew the most between ages 8 and 11 and were tallest at age 12 (table 10).
As a group they differed significantly from both the low- and high-elevation families in
growth and from the high-elevation families in height (table 11). The low- and high-
elevation means did not differ significantly.

Approximately 90 percent of the total variability was associated with differences
among seedlings within families and environmental effects. Of the remainder, 67 percent
of the variability in growth and 80 percent of the variability in height was associated
with differences among families within elevation zones and only 33 percent or 20 percent
associated with differences among elevation zones.

Although the mid-elevation group was tallest as a whole, the tallest family was
from the low-elevation group. The low group also had a higher ratio of better families
than either the mid- or high-elevation groups (1/5 vs. 1/8 and 1/36 families in the top
10 percent, respectively).

Table 10.--Growth and hetght of seedlings tn the factorial mating destgn study

Elevational : Growth between : Height
zone! : ages 9 and 11 : age 12
--------- Centimeters - - ----- -
Low 44 101
Mid 48 106
High 45 99

lLow = below 1,000 m, mid = 1,000 to 1,250 m, high = above 1,250 m.

14

Table 11.--Analysts of growth and hetght in the factorial mating design study

Variance
: : : Percentage
Source : Gl538 : M.S. : Value : of “totall
Growth between ages 9 and 11
Elevational zone 2 2206**! 4.3 iy
Family (Elev. ) 95 ASD 8.6 56 Hf
Within Family 1382 301 301.4 95.29

Height age 12

Elevational zone yD. PIMOS 14.8 1.9
Family (Elev. ) 95 GDI 59.9 LO
Within Family 1382 716 GAS 7 90.5

1* and ** indicate Significance at the 5 percent and 1 percent level,
respectively.

Vigor-quality study.--The main reasons for including this study are that the trees
are old enough to have reached a stable growth pattern and to show how the results have
changed over the years. The mean height of the low elevation families has been consis-
tently greater than that of the high elevation families (table 12). Significant dif-
ferences during the early years, ages 4 to 8 (table 13), probably resulted from the
combination of a carryover of nursery differences (Squillace and Bingham 1958) and
limited competition on the recently cleared sites. By age 8 competition from the native
vegetation was increasing and until age 12 or 14 when most trees were taller than the
brush all families probably grew less than their potential. That competition probably
was also responsible for keeping the differences between elevational zones relatively
constant while between-family and within-plot variation increased more rapidly. After
the trees overtopped the brush, their growth potential could be realized more fully.
Then height and growth rate differences between the low and high zones again became
Significant.

During the last 3 years, differences between the low and high families continued to
increase but at a slower rate. At the same time, differences among families within
elevational zones became larger, resulting in a decrease in the significance of the
elevational difference. The elevational difference exhibited in this study is probably
larger than would be found if more stands, and trees, had been sampled. The results from
the natural selection study reported earlier indicate that trees of the high elevation
stand (White Rock) used in this study produce the slowest growing seedlings of the high
elevation stands sampled. Also, most of the low elevation trees are from a stand (Crystal
Creek) that produces some of the fastest growing seedlings. Thus, by chance, these
results and those of Squillace and Bingham (1958) contrast the poorest high trees we have
Studied with some of the best low trees. Even then, the best high family has been taller
than several of the low families.

Height growth of the young trees at the two sites was essentially equal until age
10 and did not differ significantly until age 16. Interactions between sites and eleva-
tional origin of the seedlings were essentially nonexistent even after the height dif-
ference between the sites became highly significant.

15

Table 12.--Hetght and growth of young trees tn the vigor-quality study at vartous ages

Height DCEF PREF Combined
at : Low High Low High Low High
age elevation:elevation elevation:elevation elevation:elevation
families!: families? families: families families: families
----------------- Meters ---------------
A ae 0.11 0.09 Oeil 0.09 0.11 0.09
Range 0.08-0.15 0.09-0.10 0.08-0.13 0.05-0.13
Qo se 0.61 0.56 0.62 0.51 0.62 0.54
Range 0.51-0.72 0.46-0.62 Oe4AnsSOo72l Wo SlsO Se
10” 0.97 0.91 0.94 0.85 0.95 0.88
Range 0276=0. 12 0576-1200 0.67-1.10 0.58-1.00
12) ae 1.6 1.5 LBS 1.6 a) 16
Range WS B=11© Ie Saal, 7 Wess,0) dene ss
AP Ded Dis 2.9 Des 5 DG 2.4
Range 2.0-3.0 2.0-2.5 Ze Shel Lo MaZn T
WG Se 3.5 B50) Ave! 3.6 BiB o 553
Range 3.0-4.1 2.8-3.3 Seana 2.8-4.1
197 xX 5.4 AT, 6.3 5.8 5.9 522
Range 4.6-6.1 4.0-5.1 5.2-6.8 4.7-6.5
Growth
between
ages
--------------- Meters/year - - -------------
12 &
16 x 0.47 0.38 0.58 0.50 0.52 0.44
Range 0.38-0.53 0.32-0.42 0.51-0.60 0.42-0.57
ORG ae
19 x 0.63 0.56 575 0.75 0.69 0.64
Range 0.57-0.69 0.46-0.61 0.60-0.85 0.68-0.79

laverage elevation 925 m.
2average elevation 1,525 m.

16

Table 13.--Analysts of the effect of elevation of seed origin and test stte on height
and growth of trees in the vigor-qualtty study

Test site and source of variation

DCEF : PREF : Combined
Trait : Elevation : Elevation . Test : Elevation: Gaiteux
measured : zone! : zone! : site? : zone! __:elevation?
Height at age: '
4 Oo SO? eV <l Pel? <l
8 9S 4.84* <] Oia 1.86
10 <l 1.48 <1 2.06 ela
12 1.96 BESS) SS 3.46 <]
14 4.12 Opaliox Wo ita! 8.84* <All
16 So sie To OS hee Sie SA <]
19 8S. 4.98 Ska NORE <]
Growth between
ages:
IZ & kG WiOne Nabe Aes Sona SOMOLA <l
16 & 19 5.04* <] WO ows 4.04 <1

Elevation zone mean square

inp value resulting from the comparison ———————————__————————
Trees within elevation zone mean square

Test site mean square

2"Et™ value resulting from the comparison ———————__+——______
Rep within site mean square

Site X elevation mean square

3"P" value resulting from the comparison
8 P Rep within site X elevation mean square ~

** and ** indicate significance at the 5 percent and 1 percent level, respectively.

Although the differences among families within elevational zones were large (the
range between tallest and shortest varied from 10 percent to 35 percent of the mean at
different ages), the differences were seldom significant because of the small numbers
of families tested and large amounts of within-family variability. The variance attrib-
utable to family-by-site interactions was small and nonsignificant except at age 8.

The question of family-by-site interactions is even more confusing if one also includes
year of planting. A few families were planted in each of 2 or 3 years. For some of
these families the seedlings planted in 1955 may have grown faster at PREF than DCEF
but those planted in 1957 grew faster at DCEF or vice versa.

DISCUSSION AND CONCLUSIONS

To date all of our studies with western white pine indicate that the majority of
the variation which might be utilized in a tree improvement program is related to the
differences among individual trees within stands rather than to differences among stands,
elevational zones, or geographic areas. When stand or area differences were significant,
there did not appear to be a pattern to the variation. In most of the tests, seedlings
from high elevation parent trees (generally 1,375 m or above) grew slower than those
from lower elevation parents. The differences were significant in only about half of
the tests and the average growth of seedlings from some high stands was good. However,

W/,

if one were selecting only those families that were in the top 10 percent group for
height in any one test, very few of the high-elevation families would be selected
(table 14). Among the lower elevation zones there again appeared to be no pattern to
the variation.

The general finding from these studies that seedlings from high elevation trees are
slower growing than those from lower trees agrees with the results reported earlier by
Squillace and Bingham (1958). However, these data do not agree with their findings that
high-elevation families grew slower than low-elevation families in a low nursery and low
plots but faster at a high plot. In the vigor-quality study, low-elevation families were
Significantly taller than high-elevation families at both low- and mid-test sites. While
the differences in sites were not as great as in Squillace and Bingham's study, the site
means did differ significantly and there was no indication of the high-elevation families
growing faster at the higher site. In fact, the difference between low- and high-eleva-
tion families was greater at the higher site even though total height was less. Simi-
larly, as mentioned above, in the elevational study high-elevation families were as tall
or slightly taller than low- or mid-elevation families at the low sites but slightly
shorter at the high sites. Thus, up to this time the results of the present studies

suggest that groups of families representing various elevational. zones perform nearly
the same whether tested at low-, mid-, or high-elevation sites.

Table 14.--Ratio of trees tn top 10 percent group in vartous studies

Study : Seed collection elevational zone
Low : Mid i High
Elevational
Nursery 5/7 Iya 1/39
Ida Creek WT 1/10 L/SS
PREF
Low site 1/20 e/a LALO
Mid site 1/4 0 Wf 2O
High site 1/20 Wd 1/10
x 1/9 1/10 Wale
Canyon
Low site 0 1/10 YS
Mid site 1/20 1/20 1/5
High site 1/5 0 1/6
x LPl2 1/20 1/6
Natural selection WW WP LY WSS
Factorial mating design 1/5 1/8 1/36
Overall La 1/10 1/26

18

Chance selection and sample size appear to contribute much to the conflicting
results of the various studies. The vigor-quality study generally shows a marked con-
trast between low- and high-elevation stands, but it mainly samples only two stands and
only four trees in the high one. Subsequent results from the natural selection study
where 50 trees were sampled per stand confirm the differences among the low and high
stands represented in the vigor-quality study, but also show that other choices of single
low and high stands could lead to the conclusion that low and high stands do not differ.

On the other hand, the elevational study sampled a much broader range of stands but
again with few trees per stand. Consequently, the lack of differences among low and high
stands may be too conservative. For example, two 5-tree collections were obtained from
a stand at Bertha Hill at 1,585 m in the Clearwater area for the elevational study.
Seedlings from one of these collections were the tallest of the Clearwater collections
but those from the other were about average. The mean height of the two was only
slightly less than that for the best elevational zone. However, the results from the
50-tree collection made for the natural selection study indicate that although seedlings
from this stand are among the fastest growing of the high-elevation stands they are
slower growing than those of five of the six low-elevation stands. Among the low-
elevation stands in the natural selection study, even a 50-tree sample indicated only
that seedlings of the poorest stand differed significantly from the best ones.

The results of this group of studies also differ from those found in other western
conifers. Callaham and Liddicoet (1961) found that mid-elevation ponderosa pine families
were tallest at low-, mid-, and high-elevation sites during the first 20 years after
outplanting. However, although the high-elevation trees were doing very poorly at the
low- and mid-elevation sites at age 20, at the high site they were taller than the low
trees and nearly as tall as the mid trees. This pattern continued through age 29
(Conkle 1973). Conkle also found that the variance component related to elevation zones
was much larger than that for families within zones at the low and mid sites but not at
the high site. In our studies the family component was always much larger.

In working with Douglas-fir, Rehfeldt (1974) found that differences between low,
or warm, habitat type trees and high, or cool, habitat type trees could be detected with
seedlings from as few as 7 trees per stand. He also found growth differences between
stands on the east and west sides of the mountain ranges along the border between Idaho
and Montana.

In summary, the results from this series of tests indicate that, at least within
the north Idaho portion of its interior range, western white pine has a different varia-
tion pattern than that of most other conifers that have been intensively studied.
Western white pine is highly variable, but most of the variation is related to differ-
ences among the offspring of a single tree or among the trees in a stand. Although
differences among stands, elevation zones, or geographic areas were sometimes signifi-
cant, the proportion of the variance attributable to these sources was usually smaller
than that for trees within stands.

All of the seedlings involved in the studies reported here were started under pro-
tected and irrigated nursery conditions. They escaped the variety of environmental
conditions that might lead to ecotypic differentiation during the critical germination
and early establishment growth phases often found when reproduction occurs naturally.
Squillace and Bingham's (1958) germination test results may indicate such a differenti-
ation, but the results reported here do not show ecotype formation for other traits.

19

APPLICATIONS

The results to date indicate that only two seed zones are necessary for western
white pine in north Idaho, and probably for the adjacent white pine areas in north-
eastern Washington and northwestern Montana as well. The main, or low, zone would
include all the area below 1,375 m, with the exception of the most severe sites where
the upper boundary might be lowered to 1,250 m. The second, or high, zone would include
all the area above 1,375 m and a limited area related to severe sites between 1,250 m
ands lSy7- Same

Two seed orchards, with selections related to the above seed zones, should be suf-
ficient for the next phase of the blister rust resistance breeding program. Siblings,
or vegetative propagules, of seedlings intended for either orchard should be tested at
several sites to facilitate selection of those individuals or families that are both
broadly adapted and fast growing.

Seed from the present low- and mid-elevation orchards can be combined and used on
low- and mid-elevation sites throughout north Idaho. It can probably be used at eleva-
tions up to 1,375 m on most sites. Seed from the high orchard should be used for sites
above 1,375 m.. This is a conservative approach, as our results so far show little or
no difference in survival or growth among seedlings of low-, mid-, or high-elevation
origin at the high sites. However, long-term effects have shown up in other species.

PUBLICATIONS CITED

BarnespaBie. Wie
1967. Phenotypic variation associated with elevation in western white pine. For. Sci.
13(4):357-364.
Calilaham,; Ra Zon and AS Re haddvcoet:
1961. Altitudinal variation at 20 years in ponderosa and Jeffrey pines. J. For.
59(11):814-820.
Conkle, M. T.
1973. Growth data for 29 years from the California elevational transect study of
ponderosa pine. For. Sci. 19(1):31-39.
Rehfeldt, G. E.
1974. Genetic variation of Douglas-fir in the Northern Rocky Mountains. USDA For.
Serv. Res. Note INT-184, 6 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Rehikeiidtay Gree
[In press.] Ecotypic differentiation in populations of Pinus monttcola in north
Idaho--myth or reality? Am. Nat.
Rehkelidte (Gleb a wand ahead me soteaminostetee
1970. Height growth in western white pine progenies. USDA For. Serv. Res. Note
INT-123, 4 p. Intermt. For. and Range Exp. Stn., Ogden, Utah.
Squillace, A. E., and R. T. Bingham.
1954. Breeding for improved growth rate and timber quality in western white pine.
Je Eon. 152)(9) 2656-662
Squillace, A. E., and R. T. Bingham.
1958. Localized ecotypic variation in western white pine. For. Sci. 4(1):20-34.

Townsend, A. M., J. W. Hanover, and B. V. Barnes.
1972. Altitudinal variation in photosynthesis, growth, and monoterpene composition of
western white pine (Pinus montzcola Dougl.) seedlings. Silvae Genet. 21(3-4):
133-139.

20

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Steinhoff, R. J.
1979. Variation in early growth of western white pine in north Idaho.
USDA For. Serv. Res. Pap. INT-222, 22 p. Intermt. For. and
Range Exp. Stn., Ogden, Utah 84401.

In several nursery and field tests most of the useable variation among

western white pine seedlings or saplings was related to differences among
the parent trees within stands. When differences among geographic areas,
elevational zones, or stands within those categories were significant us-
ually no pattern to the variation was apparent. However, seedlings from
high elevation parents (generally 1,375 mor higher) were often shorter than
those from lower elevations.

KEYWORDS: geographic variation, elevational variation, Pinus monticola,
seed zones

Steinhoff, R. J.
1979. Variation in early growth of western white pine in north Idaho.
USDA For. Serv. Res. Pap. INT-222, 22 p. Intermt. For. and

Range Exp. Stn., Ogden, Utah 84401.

In several nursery and field tests most of the useable variation among
western white pine seedlings or saplings was related to differences among
the parent trees within stands. When differences among geographic areas,
elevational zones, or stands within those categories were significant us-
ually no pattern to the variation was apparent. However, seedlings from
high elevation parents (generally 1,375 mor higher) were often shorter than
those from lower elevations.

KEYWORDS: geographic variation, elevational variation, Pinus monticola,
seed zones

yx U.S. GOVERNMENT PRINTING OFFICE: 1979—677-019/92 REG. 8

L LIBR:

i

102296617

Headquarters for the Intermountain Forest and
Range Experiment Station are in Ogden, Utah.
Field programs and research work units are
maintained in:

Billings, Montana

Boise, Idaho

Bozeman, Montana (in cooperation with
Montana State University)

Logan, Utah (in cooperation with Utah State
University)

Missoula, Montana (in cooperation with
University of Montana)

Moscow, Idaho (in cooperation with the
University of Idaho)

Provo, Utah (in cooperation with Brigham
Young University)

Reno, Nevada (in-cooperation with the
University of Nevada)

2
3. =
“MENT OF AGRIOS