USDA 

=a 

United States 

Department of 

Agriculture 

Forest Service 

Rocky Mountain 

Research Station 

General Technical 

Report RMRS-GTR-51 

May 2000 

Juniper Seed Sources 

in the Great Plains 

Richard A. Cunningham and Rudy M. King 

f 

RMRS-FILE COPY 
Cunningham, Richard A.; King, Rudy M. 2000. Juniper seed sources in the Great Plains. 

Gen. Tech. Rep. RMRS-GTR-51. Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station. 19 p. 

ABSTRACT 

At age 10, 100% of eastern redcedar ( L.) and Rocky Mountain juniper ( Sarg.) trees 

from several seed sources throughout the Great Plains had survived. Seed sources 

from southeastern Texas had the poorest survival. Eastern redcedar trees from Kansas 

seed sources grew tallest, and trees from Montana and southeastern Texas seed 

sources were the shortest. Rocky Mountain juniper trees survived better, were shorter, 

had smaller crowns, exhibited greater damage from Cercospora blight ( (Ellis and 

Everh.) Sutton and Hodges, formerly var. ) and Kabatina tip blight ( Schneider and V. 

Am) and less damage from cedar-apple rust than did eastern redcedar trees. Eastern 

redcedar trees were larger, had more horizontal branching, and exhibited a greater 

incidence of cedar-apple rust and bagworm. In the northern Great Plains, trees from 

seed sources 1 to 4 " latitude south of each test site survived best and grew the tallest. In the central Great Plains, trees from seed sources from 1 to 4 " latitude north of 

each test site survived best, while those from 4 " latitude north and 6 " longitude east 

grew the tallest. Cluster analyses differentiated groups of seed sources that performed 

in a similar manner in terms of survival, height, and crown characteristics. Although 

most clusters were composed of seed sources from the same species and geographic 

area, one cluster contained seed sources of both species and ranked near the median 

for most traits. Agelage correlations indicated that seed sources may be successfully 

selected for good survival and fast growth rate at age 5. 

Keywords: Rocky Mountain juniper, eastern redcedar, planting stock, seed sources, 

Great Plains, survival, height, and crown characteristics 

You may order additional copies of this publication by sending your 

mailing information in label form through one of the following media. 

Please send the publication title and number. 

Telephone (970) 498-1 392 

E-mail rschneider @ fs.fed.us 

FAX (970) 498-1 396 

Mailing Address Publications Distribution 

Rocky Mountain Research Station 

240 West Prospect Road 

Fort Collins, CO 80526-2098 
Juniper Seed Sources in 

The Great Plains 

Richard A. Cunninghaml 

Research Geneticist (retired) 

USDA Agricultural Research Service 

Rudy M. King2 

Biometrician 

USDA Forest Service 

Contents 

Application ......................................... 1 

Introduction.. ...................................... 1 

Materials and Methods ............................... 1 

Results ............................................ 3 

Discussion and Interpretation .......................... 18 

Acknowledgments ................................... 18 

Literature Cited ..................................... 18 

Cunningham is a collaborator at the Northern Great Plains Research Laboratory, 

Mandan, North Dakota. 

King is a biometrician at the Rocky Mountain Research Station, Fort Collins, GO. 

Application Materials and Methods 

Rocky Mountain juniper (Juniperus scopulorum Sarg.) 

and eastern redcedar (J. virginiana L.) were extensively 

planted in windbreaks and other conservation tree plantings throughout the Great Plains. The seed sources used 

by nurseries growing seedlings for these plantings were 

generally chosen on the basis of availability, rather than 

adaptability or performance potential. Determining and 

choosing the best seed source to produce planting stock 

for specific planting sites would improve the survival and 

performance of these species in windbreak and conservation plantings throughout the Great Plains. 

Seed Collection 

Field personnel of the USDA Soil Conservation Service 

(National Resource Conservation Service), USDA Forest 

Service, state universities, and state forestry agencies collected cones from 275 eastern redcedar and Rocky Mountain 

juniper trees in natural stands and windbreaks throughout the Great Plains (figure 1). Seed trees that had favorable windbreak characteristics were selected. The sampling 

scheme was based on seed zones proposed by Cunningham (1975). Cones were packaged in single-tree lots and 

shipped to the Forestry Sciences Laboratory, Lincoln, NE 

where seed was extracted and placed in -16 "C storage. 

Introduction 

Test Site Establishment 

Information about the genetic variation among populations of Rocky Mountain juniper or eastern redcedar 

has been limited to taxonomic studies of the distribution 

of each species, and their hybrids, within the Great Plains. 

Studies by Fassett (1944), Hall (1952), Van Haverbeke 

(1968), Schurtz (1971), von Rudloff (1975), Flake et. al. 

(1978), Comer et. al. (1982), and Adams (1983) characterized junipers in the Great Plains as a variable population, 

probably due to introgressive hybridihation. Variation 

was expressed in a gradual transition throughout the 

region. Van Haverbeke (1968) identified a zone of transition from southwestern South Dakota to northeastern 

Colorado. Trees to the north and west of this zone were 

typical of Rocky Mountain juniper, in i most traits. For 

most traits, trees to the south and east of this zone were 

typical of eastern redcedar in most traits. Very little was 

known about the adaptability of Juniperus seed sources to 

specific planting sites throughout the Great Plains. In particular, whether nonlocal seed sources would perform as 

well or better than local seed sources. 

In 1973, the GP-13 Technical Committee of the Great 

Plains Agricultural Council initiated a cooperative seedsource study of Rocky Mountain juniper and eastern redcedar. The objectives of the study were to: 1) determine 

the genetic variation of major traits exhibited by both species; 2) identify the best sources of seed for the production of planting stock for use in each state; and 3) provide 

germplasm of known parentage for use in selection and 

breeding programs. 

In mid-August 1977, seed lots were sown in nursery 

beds at the USDA Forest Service Bessey Nursery near 

Halsey, NE. In March 1980, 2-year-old seedlings were 

slupped to 12 test-site cooperators throughout the Great 

Plains. Not all seed sources were planted at each site. Sites 

were stratified geographically into 3 regions (northern, 

central, and southern) and identical sets of seed sources 

were established at each site within the region. A randomized complete block with 5 replications of 4-tree, linear 

plots was established at each test plantation. 

After 5 growing seasons, survival and height data 

were collected and reported by Van Haverbeke and King 

(1990). At the Plattsmouth, NE site, a second test plantation including 131 eastern redcedar seed sources and 25 

Rocky Mountain juniper seed sources was established to 

study diseases (Zhang et al., 1997). In this planting, all 

the plots were inoculated for Cercospora blight in 1982, 

1984,1985, and 1986. Cercospora blight was recorded in 

1987; Kabitina tip blight occurred naturally, and was also 

recorded. 

Data Collection 

In 1989, tenth-year data was collected at 8 sites (Bridger, 

MT; Towner, ND; Mandan, ND; Brookings, SD; Plattsmouth, NE; Hastings, NE; Colby, KS; Sacville, MO) (figure 

2). Data collected included survival, height, crown width, 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Figure 1. Juniperus virginina (0) and J. scopulorum (A) seed sources sampled in the study. 

Figure 2. Locations of test plantations. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
crown density, branch angle, sex, strobili production, and 

pest incidence1 . Cedar-apple rust (Gymnosporangium juniperi-virginianae) and Cercospora blight were recorded in 

the South Dakota, Nebraska and Missouri sites. Kabatina 

tip blight occurred only in the Missouri and Nebraska 

sites. Bagworms (Thyridopteryx ephemeraeformis) were only 

recorded in the Missouri site. Schaefer (1995) has previously reported on the 10-year results of the test site in 

South Dakota. 

Statistical Analyses 

Statistical analyses were performed with SAS software 

(Statistical Analysis System, SAS Institute Inc., Cary, NC) 

and SPSS software (SPSS Inc., Chicago, IL). Analyses of 

site and species differences necessitated dropping 2 sites 

(Bridger, MT and Sacville, MO), which did not contain at 

least 121 seed sources as did the remaining 6 test sites. 

Mean responses for the 121 seed sources that were planted 

at all 6 test sites were initially analyzed in a 2-way analysis 

of variance of site and species. However, site-by-species 

interactions were strong in all analyses, and assumptions 

Provenance transfer models for each test plantation 

were estimated from least squares regression of differences in latitude, longitude, and elevation between source 

locations and plantation locations. 

Simple Pearson correlations were computed among 

seed source means for each trait at each test site. The correlation of each trait with species, latitude, and longitude 

was also examined with simple Pearson correlations. Categorical variables were assigned ordered, equally-spaced 

values. 

The value of the 5-year regression model in predicting 

the 10-year height at each test site was examined by 

adjusting the constant in the 5-year model to reflect the 

10-year mean height. Residuals for the difference between 

the calculated and the actual 10-year heights were calculated using the 5-year and the 10-year regression models. 

A univariate analysis was performed to characterize the 

mean residuals for each model at each test site. 

To determine the reliability of early seed source selection, age/age correlation was computed for survival and 

height of Ctree plots at ages 5 and 10. 

of normality and homogeneous variance were questionable. Therefore, the eastern redcedar and Rocky Mountain juniper were compared separately for each site using 

a Wilcoxon-Mann-Whitney rank test (Siegel and Castellan 1988). Sites were compared separately for each species using a Kruskal-Wallis rank test (Siegel and Castellan 

1988) followed by a Tukey modification of Dunn's procedure to specify pair-wise site differences (Hochburg and 

Tamhane 1987). Significance levels and multiple comparison procedures were adjusted using a Bonferroni correction to maintain Type I error at 0.05 for each set of 

analyses (Miller 1981). Categorical modeling was used to 

examine the pest scores (Rust, Cerc, Kab, Bag) for those 

sites where they were recorded. 

Cluster analyses (Ward's minimum variance) were used 

to identify seed sources with similar performance characteristics across the test plantations. Variables used in those 

analyses were survival, height, crown width, crown density, branch angle, and number of terminals. Only seed 

sources present in all 6 test sites were included in the 

analysis. Genotype x environment interaction was examined by a 2-factor analysis of variance with clusters and 

plantations as factors. 

- - 

Data were measured or scored as: survival (%); height (cm); 

crown width (cm); crown density (very sparse = 1, sparse = 2, 

moderate = 3, dense = 4, very dense = 5); branch angle (acute 

< 30°, moderate = 30" - 60°, horizontal > 60"); sex (male, 

female, both); strobili production (none = I, few = 2, moderate 

= 3, many = 4); and pest incidence ( absent = 1, light = 2, moderate = 3, heavy = 4, very heavy = 5). 

Results 

Site - Species Analysis of Variance 

Survival 

Mean performance at all 8 test sites is shown in table 1. 

Table 2 lists the means by site and species for the 6 sites 

included in the site-species analysis of variance. The 2 species did not differ sipficantly in survival except at Plattsmouth, NE where eastern red cedar survived better than 

Rocky Mountain juniper (table 2). Within respective species, Rocky Mountain juniper survived best at Hasting, 

and eastern redcedar survived best at both Nebraska sites. 

Height 

Eastern redcedar grew taller than Rocky Mountain 

juniper at each site except Towner, ND where no significant difference was apparent. Within species, eastern redcedar grew tallest in Nebraska and shortest at Towner, 

ND. 

Crown Width 

Eastern redcedar had a significantly greater crown 

width than Rocky Mountain juniper at each test site. 

Within each species, eastern redcedar was widest at Plattsmouth, NE and Colby, KS. Rocky Mountain juniper was 

widest at Colby, KS and narrowest at Towner, ND. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Table 1. Mean performance by site averaged over all seed sources. 

Site Surva Htb CwC Termd Cde Anglef Strobg Rusth Cerd Kabj Bagk 

--- 

% cm cm no score 

Bridger, MT 97 231 

Towner, NDI 77 151 

Mandan, ND2 77 274 

Brookings, SD 73 239 

Plattsmouth, NE1 92 369 

Hastings, NE2 95 305 

Colby, KS 79 286 

Sacville, MO 69 289 

All 82 275 

S. E. 0.8 2.6 

N 1094 1055 

asurvival; bheight; Ccrown width; dterminals; ecrown density; 'branch angle; gstrobili; hcedar-apple rust; iCercospora blight; JKabatina 

blight; kbagworm. 

Table 2. Means by site and species for the 6 sites in the site-species analysis. 

Site Species Surv H t Cw Term Cd Angle St ro b 

% cm cm no score 

Towner, ND ERCa 81 154 187 1.8 2.6 1.9 3.0 

RMJb 87 147 146 1.7 3.4 1.3 2.0 

Mandan, ND ERC 83 288 21 5 2.5 3.5 1.9 3.5 

RMJ 86 262 174 2.5 4.5 1.5 3.5 

Brookings, SD ERC 

RMJ 

Plattsmouth, NE ERC 

RMJ 

Hastings, NE ERC 

RMJ 

Colby, KS ERC 

RMJ 

All 88 280 244 1.6 3.4 1.9 2.6 

S.E. 0.7 3.3 2.7 0.03 0.02 0.01 0.03 

N 726 726 726 726 726 726 726 

a eastern redcedar. 

Rocky Mountain juniper. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Crown Density 

Rocky Mountain juniper had significantly denser 

crowns than eastern redcedar at each test site, except at 

Plattsmouth and Hastings, NE. Comparing sites with 

each species, eastern redcedar was densest at both 

Nebraska sites and least dense at Towner, ND and Brookings, SD. Rocky Mountain juniper was densest at Mandan, 

ND and Colby, KS. 

Terminals 

The species differed significantly in the number of terminals only at the Nebraska sites where Rocky Mountain 

juniper had the greater number of terminals. Comparing 

sites, within species, Rocky Mountain juniper had the 

most terminals in Mandan, ND and the least in Brookings, SD and Colby, KS. Eastern redcedar had the greatest number of terminals at the North Dakota sites and the 

least at the Nebraska and Kansas sites. 

Branch Angle 

At each test site, eastern redcedar had a significantly 

greater branch angle than Rocky Mountain juniper. Comparing sites, within species, eastern redcedar at Plattsmouth, NE and Hastings, NE had greater branch angles 

than Mandan, ND and Colby, KS. Brookings, SD and 

Towner, ND had the least branch angles. Rocky Mountain 

juniper at Plattsmouth, NE and Hastings, NE had greater 

branch angle than at any other sites. 

Strobili 

The species differed significantly in strobili production 

at each site, except Hastings, NE. Comparing sites, within 

species, Rocky Mountain juniper had the most strobili at 

Mandan, ND and the least at the Nebraska sites. Eastern 

redcedar had more strobili at Towner, ND and Mandan, 

ND and the least at both Nebraska sites. 

Pest Scores 

Cedar-apple rust and Cercospora blight were most 

prevalent in the Missouri plantation, while Kabatina blight 

was most severe in the Plattsmouth, NE plantation. Bagworms (T. ephemaraeformis) were recorded only in the Missouri plantation. 

Seed sources from Texas exhibited the poorest survival (< 

30%), with a source from Smith County, TX surviving at 

only 11%. Overall survival was 82% (table 1). 

Four of the 5 tallest seed sources were from Kansas. The 

fifth-tallest was from Fourd county, in northwest Texas. 

Ironically, 5 of the 7 shortest seed sources were from counties in southeast Texas. The other 2 of the 7 shortest seed 

sources were from Montana. 

Means for crown width mirrored those for heights with 

4 Kansas seed sources among the 5 seed sources averaging 

the widest crown width. Seed sources with the narrowest 

crowns included those from both Rocky Mountain juniper (WY) and eastern redcedar (TX). Seed sources with 

the greatest crown density were eastern redcedar sources 

from North Dakota, Oklahoma, and Kansas. An exception 

was a Rocky Mountain juniper source from South Dakota. 

The least dense crowns were typical of seed sources from 

Texas and Oklahoma. Seed sources with the fewest terminal shoots were from Kansas, Nebraska, and South 

Dakota. Seed sources from Texas and Oklahoma had the 

greatest number of terminal shoots, probably as a result 

of winter damage in the more northern test sites. 

Most of the seed sources with the most acute branch 

angle were Rocky Mountain juniper from Wyoming, 

Montana, and North Dakota. More horizontal branching 

habit was typical of seed sources from Texas, Oklahoma, 

Kansas, and South Dakota. 

Rocky Mountain juniper seed sources from Wyoming, 

Montana, and North Dakota had the least incidence of 

cedar-apple rust, whereas seed sources of eastern redcedar from Kansas, Nebraska, and South Dakota had the 

greatest incidence. Cercospora and Kabatina blights were 

most severe on Rocky Mountain juniper seed sources 

from Montana, North Dakota, South Dakota, and Wyoming. In a separate inoculation study at the Plattsmouth, 

NE site, this same trend was reported (Zhang et al., 1997). 

Seed sources from southeastern sites at lower elevations 

exhibited higher resistance to Cercospora and Kabatina 

blight than seed sources from northwestern sites at higher 

elevation. 

Incidence of bagworm was greatest on some eastern redcedar seed sources from Nebraska, while other Nebraska 

seed sources had very little damage. Rocky Mountain juniper seed sources from Wyoming, Montana, and North 

Dakota had no incidence of bagworm damage. 

Seed Source Performance 

at Individual Test Sites 

Seed Source Means Averaged Over All Sites 

Averaged over 8 test sites, several seed sources had a 

survival rate of 100%. Seed sources with perfect survival 

were scattered throughout most of the Great Plains states. 

Significant differences among seed sources existed at 

each test site for survival, height, crown width, crown 

density, and branch angle. Number of terminals differed 

significantly among seed sources at all sites except South 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 5 
Figure 3. Juniperus seed-source clusters as determined by cluster analysis. 

Dakota. At sites where they were recorded, significant 

differences among seed sources existed for Cercospora 

blight, Kabatina tip blight, cedar-apple rust, and bagworm. Instead of focusing on site-specific results, we 

chose to look for more general patterns of variation using 

cluster analysis. 

Cluster Analysis 

The cluster analysis included 121 seed sources that 

were present in all 6 test sites. The analysis differentiated 

5 clusters composed of seed sources that performed similarly in terms of survival, height, crown width, crown density, branch angle, and number of terminals. These clusters 

were not geographically distinct in ail cases (figure 3). 

Cluster #1 was a disjunctive combination of eastern 

redcedar from Kansas, Oklahoma, and Texas, and both 

species from Montana, North Dakota, and South Dakota 

(figure 4). Survival of Cluster #1 seed sources averaged 

80% over all plantations (figure 5), with highest survival 

occurring in the central plains (figure 6). Height growth 

of cluster #1 seed sources was greatest in Nebraska and 

poorest in Mandan, ND (figure 7). Among the 5 clusters, 

cluster #1 seed sources ranked second for mean height 

growth and mean crown width, and fourth for mean 

crown density (figures 5, 8). Mean number of terminals 

ranked second highest and branch angle was intermediate (figure 9). 

Cluster #2 seed sources were all eastern redcedar and 

were scattered over the Great Plains from South Dakota 

to Texas (figure 10). Survival, height, and crown width 

of these seed sources ranked highest among the 5 clusters (figure 8). Crown density tied for second place (figure 

8). Mean number of terminals ranked lowest and branch 

angle tied for highest (most horizontal) among the 5 clusters (figure 9). Survival and height ranked highest at all 

sites and was best in Nebraska (figures 6,7). 

Cluster #3 consisted of only 2 seed sources and both 

were eastern redcedar from Oklahoma and northern Texas 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Figure 4. Juniperus seed sources in cluster #I (A = J. scopulorum; 

= J. virginiana). 

survival Height 

Figure 5. Mean survival and height of Juniperus seed-source clusters. 

1 2 3 4 5 

Cluster 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Test Sites 

Cluster 

Figure 6. Genotype x environment interactions for survival (seed-source clusters vs. test 

sites) of Juniperus in the Great Plains. 

- ND1 ND2 NEI NE2 SD KS - 

Test Sites 

Figure 7. Genotype x environment interactions for height (seed-source clusters vs, test sites) of 

Juniperus in the Great Plains. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
1 2 3 4 5 

Cluster 

Crawnwid(h Crwn density 

Cluster Means 

terminals branch agk 

300 

275 

250 

225 

200 

175 

150 

Figure 9. Mean number of terminals and branch angle of Juniperus seed-source clusters. 

- 

- 

- 

- 

- 

- 

- 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 

Figure 8. Mean crown width and crown density of Juniperus seed-source clusters. 
Figure 10. Juniperus seed sources in cluster #2 (A = J. scopulorum). 

Figure 11. Juniperus seed sources in cluster #3 (a = J. virginiana). 

(figure 11). These seed sources were intermediate in survival, height, and crown width (figures 5, 8), but were 

above average in crown density, number of terminals, and 

branch angle (figures 8,9). Survival of these seed sources 

was best in Nebraska, but very poor in North Dakota and 

South Dakota (figure 6). 

Cluster #4 seed sources were all eastern redcedar from 

Oklahoma and Texas (figure 12). Survival, height, crown 

width, and crown density (figures 5, 8) ranked lowest 

among the 5 clusters, while the number of terminals was 

intermediate and the branch angle was tied for the most 

horizontal (figure 9). Survival was highest in Nebraska, 

and poor at all other sites (figure 6). Height growth of 

these clusters ranked lowest at most sites and was second 

lowest at one North Dakota site (ND2). 

Cluster #5 was entirely Rocky Mountain juniper seed 

sources from the northern Great Plains, and one from New 

Mexico (figure 13). Overall, these seed sources ranked 

second best in survival and intermediate in height (figure 

5). Crown density ranked highest, number of terminals 

intermediate, and branch angle most acute among the 5 

clusters (figures 5,8). Survival of these seed sources was 

excellent at all of the sites. Height growth was intermediate at most sites. (figure 7). 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Figure 12. Juniperus seed sources in cluster #4 (0 = J. virginiana). 

Figure 13. Juniperus seed sources in cluster #5 (A = J. scopulorum). 

Averaged over 6 test sites, seed sources from cluster #2 

ranked best in survival, height, and crown width. Cluster 

#2 seed sources tied for second in crown density. Cluster 

#4 seed sources ranked lowest in these same traits. 

Based on performance at each test site, cluster #2 seed 

sources performed best. Cluster #4 seed sources survived 

the poorest at test sites in North Dakota, Nebraska, and 

Kansas; however, their survival improved to second poorest in South Dakota. Average heights of cluster #4 seed 

sources were shortest at all sites except Mandan, ND. 

Genotype by environment interaction was evident in 

the survival and height of seed sources in clusters #3 and 

#5 (figs 6,7). Cluster #3 seed sources survived poorly in 

the northern plains and Kansas, but well in Nebraska. 

Height growth of Cluster #3 seed sources was moderate 

in Nebraska, but poor in North Dakota and Kansas. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
The frequency of seed sources classified within each 

cluster that occurred within cluster #2, which was the 

best cluster, are shown in table 3. Of all the seed zones 

in cluster #2, about 89% were classified as belonging to 

cluster #2. About 5% of the seed sources from the seed 

zones in cluster #2 were classified as belonging to cluster 

#1. Only 4% of the seed sources from the seed zones in 

cluster #2 were classified as belonging to cluster #4. Less 

than 1% of the seed sources from the seed zones in cluster 

#2 were classified as belonging to cluster #3. None of the 

seed sources from the seed zones in cluster #2 were classified as belonging to cluster #5. The high probability that a 

seed source in cluster #2 belongs to the highest performing cluster indicates the practical utility of choosing seed 

sources from the cluster #2 territory. 

Provenance Transfer Models 

Survival was significantly related to latitude at all test 

sites except Bridger, MT (table 4). Seed sources from up 

Table 3. Frequency of seed sources classified within 

each cluster that occurred within cluster #2. 

Cumulative 

Cluster No. No. YO No. % 

to 4" latitude south of the test sites survived best in the 

northern Great Plains (table 5, figures 14-16). In the central 

Great Plains and Missouri, seed sources from the same 

latitude, to 4 " north of the test sites survived best (figures 17-21). Longitude was weakly related to survival 

among seed sources tested in Montana, South Dakota and 

Nebraska test sites where seed sources from the same latitude as the test sites survived best (table 5, figures 16-18). 

Elevation of the seed source was very weakly related to 

survival at Bridger, MT (3,656 ft), Mandan, ND (1,740 ft), 

Brookings, SD (1,650 ft) and Sacville, MO (1,300 ft). Seed 

sources from the same elevation as the test site survived 

best at these sites (table 5). 

The influence of latitude on height of the seed sources 

was similar to its effect on survival, although the distance 

from the test site to the origin of the best seed source was 

often greater than for survival (tables 5,6). At some northern test sites, such as Mandan, heights peaked for seed 

sources about 4 " latitude south of the test site (table 5, 

figure 15). In South Dakota and Nebraska, seed sources 

from the same latitude as the test sites grew tallest (figures 16-18). At Colby, KS, heights peaked for seed sources 

about 4 " latitude north of the test site (figure 19). In Missouri, seed sources from 1 degree latitude north of the 

test sites grew tallest (table 5). In Montana, seed sources 

from the farthest latitude north or south of the test site 

grew tallest (figure 20). Longitude of the seed source was 

related to the heights of the seed sources at all test sites 

except Bridger, MT and Sacville, MO (table 5). At several 

sites, seed sources from 1 to 6 " longitude east of the test 

sites were tallest. At Mandan, ND and Plattsmouth, NE, 

seed sources from the same longitude as the test sites were 

tallest. Elevation of the seed source had a minor effect 

on height at South Dakota, Missouri, and Montana sites 

(table 6). At these sites, seed sources from the same elevaTable 4. Provenance transfer model relating survival to source location relative to plantation location. 

Regression coeff icientsa 

Site R SEb Constant A Lat. A Lat. A Long. A Long. A Elev. A Elev. 

Towner 0.68 14.9 

Mandan 0.77 11.5 

Brookings 0.74 14.1 

Plattsmouth 0.54 6.4 

Hastings 0.24 4.3 

Col by 0.52 10.7 

Sacville 0.75 13.2 

Bridger 0.56 1.4 

a Differences between sources and plantations in latitude (negative values represent sources south of plantations); in longitude 

(negative values represent sources east of plantations); in elevation (negative values represent sources lower than plantations). 

Units of latitude and longitude in degrees; elevation in 1000 feet. 

Standard error of estimate (p< 0.01 ). 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
tion as the test site grew slightly taller than seed sources 

from higher or lower elevations (table 5). 

After adjusting the constant for the 10-year mean, prediction of 10-year values by the 5-year models did not 

indicate bias in the 5-year models (table 7). Only Towner, 

ND provided any indication of possible bias, but it was 

not significant (p = 0.87). 

Correlation 

Among Traits 

Correlation analyses of seed-source means at each test 

site revealed significant correlations among many traits 

Table 5. Location of best seed sources for each test site. 

(tables 8-10). Cedar apple rust was more highly correlated with seed sources from southern latitudes at all sites 

except Mandan, ND. Trees from easterly longitudes had 

more rust at the Missouri site, but the effect was non-significant elsewhere. More rust occurred on Rocky Mountain juniper at Mandan, ND, but it occurred more on 

eastern redcedar at Nebraska and South Dakota sites. Rust 

was more prevalent on taller trees at Missouri, Nebraska, 

and South Dakota, but not at Mandan, ND. Better surviving trees had more rust at Missouri and Nebraska, but 

the effect was non-significant at North Dakota and South 

Dakota. 

Cedar apple rust and Cercospora blight were negatively 

correlated at the Nebraska sites and were non-significant 

Location 

Survival Height 

Lat.a Long.b Elev.= Lat.a Long.b Elev.= 

Bridger 

Towner 

Mandan 

Brookings 

Plattsmouth 

Hastings 

Colby 

Sacville 

a Degrees of latitude from the test site to the origin of best seed sources. 

Degrees of longitude from the test site to the origin of best seed sources. 

Difference in elevation of seed sources in relation to test site. 

Minimal influence. 

Equal to that of the test site. 

Table 6. Provenance transfer model relating height to source location relative to plantation location. 

Regression coeff icientsa 

Site R SEb Constant Lat. Lat. Long. Long. Elev. E lev. 

Towner 

Towner 

Mandan 

Brookings 

Plattsmouth 

Hastings 

Colby 

Sacville 

Bridger 

a Differences between sources and plantations in latitude (negative values represent sources south of plantations); in longitude 

(negative values represent sources east of plantations); in elevation (negative values represent sources lower than plantations). 

Units of latitude and longitude in degrees; elevation in 1000 feet. 

Standard error of estimate (p< 0.01). 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
MOT 

Figure 14. Provenance transfer model for survival and height at Towner, ND. The vertical line at the origin represents 

the test site location. See table 4a for definitions of latitude and longitude. 

Figure 15. Provenance transfer model for survival and height at Mandan, ND.The vertical line at the origin represents 

the test site location. See table 4a for definitions of latitude and longitude. 

Figure 16. Provenance transfer model for survival and height at Brookings, SD. The vertical line at the origin 

represents the test site location. See table 4a for definitions of latitude and longitude. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.200C 
Figure 17. Provenance transfer model for survival and height at Plattsmouth, NE. The vertical line at the origin represents the test site location. See table 4a for definitions of latitude and longitude. 

Figure 18. Provenance transfer model for survival and height at Hastings, NE. The vertical line at the origin 

represents the test site location. See table 4a for definitions of latitude and longitude. 

Figure 19. Provenance transfer model for survival and height at Colby, KS. The vertical line at the origin represents the 

test site location. See table 4a for definitions of latitude and longitude. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Figure 20. Provenance transfer model for survival and height at Bridger, MT. The vertical line at the origin 

represents the test site location. See table 4a for definitions of latitude and elevation. 

Figure 21. Provenance transfer model for survival and height at Sacville, MO. The vertical line at the origin 

represents the test site location. See table 4a for definitions of latitude and elevation. 

Table 7. Comparison of regression models. 

5-year model 10-year model 

Site xa pb S. E.~ x a pb S. E.= 

Towner, ND -0.420 0.871 23.76 0.001 1 .OOO 20.71 

Mandan, ND -0.001 0.999 35.98 0.001 0.999 26.46 

Brookings, SD 0.001 1 .OOO 65.04 -0.001 1 .OOO 35.22 

Plattsmouth, NE 0.001 0.999 60.66 -0.001 0.999 36.59 

Hastings, NE 0.001 1 .OOO 42.37 0.001 0.999 26.76 

Colby, KS 0.001 0.999 25.75 -0.001 0.999 20.77 

a Mean residual. 

Probability there was no bias in applying regression model to 10-year data. 

Standard error of estimate. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
elsewhere. Kabatina and Cercospora were positively correlated at the Nebraska sites, but negatively correlated 

at the Missouri site. More Cercospora blight occurred on 

shorter trees at the Nebraska sites, but the effect was nonsignificant at the Missouri and South Dakota sites. 

Kabitina blight and cedar apple rust were negatively correlated at the Missouri site. Kabitina blight was more prevalent on better surviving trees at the Nebraska sites, but 

at the Missouri site the effect was non-significant. Shorter 

trees had less Kabitina blight in Nebraska. More Kabitina 

blight occurred on Rocky Mountain juniper in Nebraska, 

but the correlation in Missouri was non-significant. 

Bagworm, recorded at the Missouri test site only, was 

positively correlated with cedar apple rust, survival, and 

height. Bagworm was more prevalent on eastern redcedar than on Rocky Mountain juniper. 

Survival and height at age 10 was significantly correlated with survival and height at age 5 (table 11). Correlation of 5-year height with 10-year height was particularly 

high (0.96). 

Table 8. Correlation of diseases and bagworm with 

selected variables at each test site.a 

Trait 1 Trait 2 MO ND2 NE1 NE2 SD 

SURV 

HT 

CER 

KAB 

SURV 

HT 

KAB 

SURV 

HT 

SURV 

HT 

RUST 

CERC 

RUST 

RUST 

RUST 

RUST 

CER 

CER 

CER 

KAB 

KAB 

BAG 

BAG 

BAG 

BAG 

a All values are significant (p<0.01). 

Table 9. Correlation of species with selected variables at each test ~ite.~g -- 

Trait 1 Trait 2 KS MO MT ND1 ND2 NE1 NE2 SD -- - -- - - - -- 

SURV SP NS -0.85 0.36 NS NS NS NS 0.23 

HT SP NS -0.78 NS NS NS -0.6 -0.55 NS 

RUST SP - -0.73 - - 0.47 -0.62 -0.56 -0.72 

CERC SP - 0.32 - - - 0.84 0.8 NS 

KAB SP - NS - - 0.51 0.55 - 

BAG SP - -0.43 - - - - - 

a All values are significant (p<O.Ol). 

Species were coded: J. virginiana = 1 ; J. scopulorum = 2. 

Table 10. Correlation of latitude and longitude with selected variables at each test site.a - - 

Trait 1 Trait 2 KS MO MT ND1 ND2 NE1 NE2 SD - - - - - - - 

SURV LAT 0.60 -0.52 0.62 0.52 0.59 NS 0.41 0.56 

HT LAT 0.24 -0.53 0.44 0.33 0.33 -0.32 NS 0.28 

RUST LAT - -0.47 - - 0.30 -0.31 -0.35 -0.27 

SURV LONG NS -0.65 NS NS NS NS NS NS 

HT LONG NS -0.62 NS NS NS NS NS NS 

RUST LONG - -0.50 - - NS NS NS NS 

a All values are significant (p<0.01). 

USDA Forest Service Gem Tech. Rep. RMRS-GTR-51.2000 
Table 11. Correlation among survival and height the best and grew the tallest. Seed sources from the same 

measured at 5 and 10 years.a latitude and longitude as the test site survived best and 

from the same latitude and 1 " longitude east, grew the Variables tallest in South Dakota. In Nebraska, seed sources from 

the same to 4 " latitude north of the planting site survived 

Surv-I 0 Ht-5 Ht-lo best and those from the same latitude and from the same 

Su rv-5 0.68 0.39 0.37 to 4 " east of the Hastings planting site, grew the tallest. 

0.57 Seed sources from 4 " north of the Kansas site survived Surv-10 0'54 best, and those from 4 " latitude north and 6 " longitude Ht-5 east grew the tallest. In Missouri, seed sources from 2 " 

a All values are significant (pc0.001). north survived best and those from 1 " latitude north grew 

the tallest. Seed sources from the same longitude and elevation as the Bridger test site survived best, while those 

from 4 " latitude north or 6 " latitude south of the test 

site grew the tallest. The unusual effect of latitude of the 

Discussion and Interpretation 

The wide geographic distribution of seed sources sampled in this study and the inclusion of Rocky Mountain 

juniper and eastern redcedar, resulted in significant differences among the seed sources for most of the traits 

measured or recorded. Differences between the species 

are not unexpected for many traits, particularly crown 

characteristics and susceptibility to pests. When the performance of the seed sources was averaged over all 6 test 

sites and clusters of those performing in a similar manner 

were derived, most of the resulting clusters were composed of seed sources of the same species and geographic 

region. However, seed sources in cluster #I included both 

species and were from opposite ends of the Great Plains. 

Mean survival, height, and crown characteristics of cluster #I seed sources were .intermediate in rank among the 5 

clusters. This may be evidence of hybridization and introgression between the 2 species in the areas where these 

seed sources originated. Schaefer (1995), however, suggests 

that results from the South Dakota test site and molecular 

research (Stephen Ernst, Univ. Nebr., pers. Comm.) indicate 

that such introgressive hybridization is highly unlikely. 

Of the clusters comprised primarily of eastern redcedar, seed sources in cluster #2 performed best in survival, 

height, and crown width. Seed sources in cluster #5 were 

exclusively Rocky Mountain juniper. These seed sources 

survived well and had compact, conical crowns. 

The strong correlation of 5-year height with 10-year 

height and the univariate analysis reveals that the 5-year 

model could predict the 10-year height with little probability of bias. This suggests that seed sources that perform well at age 10 can be successfully identified at age 5, 

thus reducing the time necessary to choose the best seed 

source for a particular planting site. 

The results of this study provide a guide for selecting 

seed sources for seedlings to plant at specific sites in the 

Great Plains. In North Dakota, seed sources from 4 " latitude south (1 " = 70 miles) of the planting site survived 

seed source on height at Bridger, MT was unexpected and 

difficult to explain, because the results at other test sites 

in the northern Plains generally show that seed sources 

from north of the test site grow more slowly. 

The use of a seed source from within 70 miles of the 

planting site would be a conservative approach to seed 

collection. In most cases, this approach would produce 

planting stock that would survive well and grow to an 

acceptable height. However, using the above stated guidelines could result in significant increases in growth rate 

while maintaining adequate survival. 

Acknowledgments 

The authors acknowledge the Great Plains Tree Improvement Committee for coordinating this research and 

thank the following cooperators: R. Laframboise, N. D. 

For. Serv., Towner, ND; P. R. Schaefer, SDSU, Brookings, 

SD; W. R. Lovett, Univ. Neb., Lincoln, NE; M. E. Dix, 

USDA-FS, Washington, DC; W. A. Geyer, KSU, Manhattan, KS; T. R. Robison, MO Dept. Conserv.; J. G. Scheetz, 

USDA-NRCS, Bridger, MT. 

Literature Cited 

Adams, R.P. 1983. Intraspecific terpenoid variation in Juniperus scopulorum: evidence for Pleistocene refugia and recolonization in western 

North America. Taxon. 32: 30-46. 

Comer, C.; Adams, R.P.; Van Haverbeke, D.F. 1982. Intra- and interspecific variation of Juniperus virginiana and J. scopulorum seedlings 

based on volatile oil composition. Biochemical Systematics and Ecology. 10: 179-306. 

Cunningham, R.A. 1975. Provisional tree and shrub seed zones for the 

Great Plains. Res. Pap. RM-150. Fort Collins, CO: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Forest and Range 

Experiment Station. 15 p. 

18 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 
Fassett, N.C. 1944. Juniperus virginiana, J. horizontalis and J. scopulorum - 11. Hybrid swarms of J. virginiana and J. scopulorum. Bull Torrey Bot. 

Club. 71: 475-483. 

Flake, R.H.; Urbatsch, L.; Turner, B.L. 1978. Chemical documentation of 

allopatric introgression in Juniperus. Systematic Botany. 3: 129-144. 

Hall, M.T. 1952. A hybrid swarm in Juniperus. Evolution. 6: 347-366. 

Hochburg, Y.; Tarnhane, A.C. 1987. Multiple Comparison Procedures. 

New York: John Wiley and Sons. 450 p. 

Miller, R.G., Jr. 1981. Simultaneous Statistical Inference, 2"* ed. New 

York: Springer-Verlag. 299p. 

Schaefer, P. 1995. Ten-year results of an eastern redcedar and Rocky 

Mountain juniper provenance test in eastern South Dakota. Northern J. Applied Forestry 12 (1): 30 - 35. 

Schurtz, R.H. 1971. A taxonomic analysis of a triparental hybrid swarm 

in Juniperus. Lincoln, NE: University of Nebraska. 90 p. Ph.D. 

dissertation. 

Siegel, S.; Castellan, N.J., Jr. 1988. Nonparametric Statistics for the Behavioral Sciences, 2nd ed. New York: McGraw-Hill. 399 p. 

Van Haverbeke, D.F. 1968. A population analysis of Juniperus in the 

Missouri River Basin. New Series 38. Lincoln, NE: University of 

Nebraska Studies. 82 p. 

Van Haverbeke, D.F.; R.M. King. 1990. Genetic variation in Great Plains 

Juniperus. Res. Pap. RM-292. Fort Collins, CO: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Forest and Range 

Experiment Station. 8 p. 

von Rudloff, E. 1975. Volatile oil analysis in chemostytematic studies of 

North American conifers. Biochemical Systematics and Ecology. 1: 

131-167. 

Zhang, J.W.; Klopfenstein, N.B.; Peterson, G.W. 1997. Genetic variation 

in disease resistance of Juniperus virginiana and J. scopulorum grown 

in eastern Nebraska. Silvae Genetica 46 (1): 11-16. 

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-51.2000 

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