An ecological study of creeping juniper (Juniperus horizontalis Moench.) in Montana by John Gage Miller A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Fish and Wildlife Management Montana State University © Copyright by John Gage Miller (1978) Abstract: Various ecological characteristics and relationships for creeping juniper (Juniperue horizontalis Moench.) in Montana were determined through intensive studies in 30 widely separated juniper stands. Supplementary data were obtained from an additional 21 study sites. Creeping juniper was associated primarily with the mountain foothills of north-central, central and southcentral Montana, and the open prairies of eastern Montana. Ridges, rimrocks, escarpments and hilltops adjacent to a river or mountain range were typical areas of occurrence. Stands occurred more frequently on northerly exposures, slopes with gradients less than 40 percent, and at elevations below 1500 meters (4900 ft). Creeping juniper was the most, important plant species in all stands, with mean canopy coverage of 32 percent. Graminoids were the most prominent plant associates, with Idaho fescue being the most common species. Trees typically were lacking. Eight creeping juniper "associations" were defined through simple ordination. Soils associated with creeping juniper stands appeared to be poorly developed with thin topsoils, large quantities of stone and exposed rock parent material, and some degree of surface erosion. They tended to be clay loam in texture, slightly basic (pH = 7.2), low to medium in organic matter content (4.6%), very low in phosphorus (18 ppm), medium to high in potassium (280 ppm) and low in salt hazard (.08 meq/100 gms soil). Creeping juniper plants averaged 6.4 cm in height, with 3.5 percent of the crown area dead. Most growth occurred horizontally on the peripheral branches or leaders. Annual twig growth commenced in early April, coincident with a crown color change from brown to green, and continued into September and possibly early October. The overall average terminal and lateral twig growth was 34.5 and 9.0 cm, respectively, during 1978. Longest twigs were produced in stands with northerly exposures and 13-14 inches of annual precipitation. Shortest twigs were produced in stands having a tree overstory. The "greening" of plants in early April coincided with staminate cone maturation. Once pistillate buds formed, berries generally formed and matured. Berry production peaked in mid summer (late June to early July). Ripening apparently requires 1-2 years after berries turn purple in late summer. Germination of seeds in the laboratory was very low (approximately 0.1%). Reproduction from seed also appeared to be a rare occurrence in the field where plants apparently are maintained vegetatively by rooting along branches. Plants producing the highest numbers of reproductive parts were associated with soils having relatively high phosphorus, moderate calcium and sodium levels and low salt hazard. Wildlife usage of creeping juniper included mule deer, small mammals and birds. Utilization by deer occurred mainly during the late winter and appeared to be influenced more by the distribution of deer, the relative availability and/or abundance of creeping juniper and the occurrence of other more preferred forage plants than by protein content. Data indicated that creeping juniper is subject to some damage and destruction by fire, though these effects may be less severe than reported for other species of juniper.  STATEMENT OF PERMISSION TO COPY In presenting this thesis in partial fulfillment of the requirements for an advanced degree at Montana State University, I agree that the Library shall make it freely, available for inspection. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by my major professor, or, in his absence, by the Director of Libraries. It is understood that any copying or publication of this thesis for financial gain shall not be allowed without my written permission. AN ECOLOGICAL STUDY OF CREEPING JUNIPER (Juniperus hovizontalis Moench.) IN MONTANA by JOHN GAGE MILLER A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Fish and Wildlife Management Approved: Gym Comm:mittee ead, Major Department -M raduate DeanG MONTANA .STATE UNIVERSITY Bozeman, Montana December, 1978 iii ACKNOWLEDGMENT The author wishes to express his appreciation and gratitude to Dr. Richard J. Mackie, Montana State University, for his guidance and constructive criticism in the preparation of the manuscript; Dr-. Theodore W. Weaver and Dr. Robert Eng, Montana State University, and Dr. Henry Jorgensen, Montana Department of Fish and Game, for critical reading of the manuscript and helpful suggestions for revisions; Dr. Richard Lund for assistance in preparation and interpretation of computer programs; Terry Lonner, Montana Department of Fish and Game, for assistance in setting up computer programs; Dr. John Rumely, Montana State University, for aid in identification of plant specimens; John McCarthy, Ken Hamlin and Bert Goodman, Montana Department of Fish and Game for assistance during the study; personnel of the Montana State University Seed Laboratory for aid during germination experiments; Forest Service personnel for assistance during distribution surveys; the private landowners who allowed me to establish study sites on their land; Mr. Larry Keown, Forest Service, for assistance during burn experiments; and to my wife, Connie, and mother, Edith, for assistance in typing and preparation of the manuscript. The study was supported by the Montana Department of Fish and Game under Federal Aid Projects W-120-R and W-130-R. I TABLE OF CONTENTS ACKNOWLEDGMENT .................................................. iii LIST OF TABLES.......................................... ; . . . v LIST OF FIGURES . . . ’ ........................................ vii ABSTRACT................................................. ' . . . x INTRODUCTION .................................................... ' I M E T H O D S .......................................... ........... 4 . RESULTS AND DISCUSSION................................. .. . . . 13 Distribution............................................ .. ' 13 Geographic .......................................... 13 Physiographic . .'.................................... 15 Phytosociological Characteristics .............. . . . . . . 24 Community Associations ............................... 24 Environmental Attributes of the Eight Associations . . 35 Vegetational Composition of Creeping Juniper Associations........................... '........... 39 Edaphic Characteristics and Relationships ................. 55 Growth Characteristics and Relationships ................ 65 Growth F o r m ............................... 65 Annual Chronology and Twig G r o w t h ........ ‘.......... 70 Reproductive Characteristics and Relationships . . . . . . 81 Forage Characteristics and Utilization by Wildlife . . . . 89 Nutritional Characteristics........... 89 Available Forage Biomass ................ 92 Utilization by W i l d l i f e ............................ . 93 Effects of Fire on Creeping Juniper....................... 99 Stanford B u r n ........................................ 100 Sun River B u r n ........................................ 103 Discussion of Fire Effects............ .......... > 104 CONCLUSIONS AND RECOMMENDATIONS.......... '...................... 107 LITERATURE C I T E D ................................. H O APPENDIX.......................................................... 117- Page V I T A ................................................................ ii V1. Distribution of 51 creeping juniper study sites among various exposures, slope gradients and elevations........ 23 2. Constancy, canopy coverage and frequency of low growing taxa for the eight creeping juniper associations as determined by examination of 2 X 5 decimenter plots on each a r e a ............................................ . 41- 3. Constancy, canopy coverage and density of three trees on each of the eight creeping juniper associations . . . . 54 4. Edaphic characteristics of creeping juniper associations ■ including pH, organic matter content, texture, salt hazard, soil depth, erosion and 5 important elements . . . 56 5. Comparison of six soil factors from samples taken from under and between creeping juniper shrubs................. 64 6. Height, decadence, biomass, and lateral and terminal growth of shrubs in eight creeping juniper associations, east of the Continental Divide ................. . . . . . 68 7. Reproductive characteristics of creeping juniper in Montana, and statistical comparisons among plants of eight juniper associations ............................... 85 8. Percentages of crude protein of creeping juniper plants fpt 12 study- sites during one year, 1976 to 1977 ........... 89 9. Comparison of spring (April, May, June) precipitation at 12 creeping juniper sites between 1976 and 1977 . . . . . 90 10. Percent of creeping juniper twigs utilized by small mammals during the summer of 1977 on the 30 study sites. . 95 11. Crown area and height of 21 creeping juniper plants within the Stanford experimental burn together with percent crown burned, post-fire fate of plants, temperatures of fire and fine fuel and soil moistures of the area LIST OF TABLES Table Page 102 Table Page 12. Comparison of the effects of fire on creeping juniper shrubs between the Sun River Game Range and Stanford experimental burns.......................................... 105 13. General and exact locations of the 51 Creeping juniper study sites................................................ 118 14. Exposure, slope, elevation and drainage of 51 creeping juniper sites in Montana. ........................ 120 15. Scientific and common names of the 24 graminoids, 104 forbs, 18 shrubs and 3 trees in this study................. 122 16. Percent canopy coverage, and percent frequency among 2 x 5, plot frames for the 30 intensively studied creeping juniper sites . ............................................ 129 17. Edaphic characteristics of 51 creeping juniper sites including pH, organic matter content, texture, salt hazard, soil depth, erosion and 5 important elements. * . 153 vi vii LIST OF FIGURES 1. Population trends of creeping juniper on 12 condition trend transects established by the Montana Department of Fish and Game along the east slope of the Continental Divide........ .. ................... . ............. . . 2 2. Study areas showing the approximate locations of 51 study sites............................. '................. 5 Figure - Page 3. General distribution of creeping juniper in Montana. . . . 14 4. Site I on an alluvial plain near Barr Creek, on the Sun River Game Range, Lewis and Clark County; JuniIpeTus- Festuoa important cover........................... .. 15 5. Site 3 near Haystack Butte southwest of Augusta, Lewis and Clark County; Juntpevus-Festuca important cover. . . . 16 6. Site 14 near Sage Creek east of Denton, Fergus County; Junipevus-Rhus-Sttpa important cover ..................... 17 7. Site 15 on edge of a Pinus pondevosa community south of Warhorse Reservoir, Petroleum County; Junipevus- Catamovitfa important cover.................................. 17 8. Site 16 on an open prairie south of Warhorse Reservoir, Petroleum County, showing high degree of erosion; Junipevus-Agvopyvon important cover......................... 18 9. . Site 19 in the foothills of the Big Snowy Mountains, Fergus County; Junipevus-Potentitta-Festuoa- important cover......................................"............... 18 10. Site 21 in the foothills of the Little Belt Mountains, Wheatland County; near a Pinus ftexitus— Pinus pondevosa community; Junipevus-Festuoa important cover ............ 19 11. Site 25 in an Absarokee mountain meadow near a community of Pinus ftexitus and Pseudotsuga mensiesii, Stillwater County, Junipevus and a variety of forbs important cover . 19 12. Site 26 on an eastern Montana prairie, northwest of Terry, Prairie County; Junipevus-Cavex important cover . . 20 viii • 13. Site 27 on an eastern Montana prairie/foothill area northwest of Terry, Prairie County; Junipevus- Andropogon important cover............................. .. 20 14. Site 32 on the west side of the Continental Divide, east of Lincoln, Lewis and Clark County; under a stand of Pinus contovta and Pseudotsuga menziesii; Avcto- staphylos-Agvopyvon-Junipevus important understory cover. 21 15. Site 37 under a stand of Pinus pondevosa north of Ryegate in the Little Snowy Mountains, Golden Valley County; Junipevus hovizontalis and scopulovum important under story cover.......................................... 21 16. Site 42 under a stand of Pinus pondevosa northwest of Broadus, Powder River County; Junipevus-Stipa important understory cover........................... " ............. 22 17. A simple ordination of 30 creeping juniper stands based on total vegetal composition, in two dimensions (X-Y axes) ..................................................... 29 18. The same ordination as Figure 17 only on a larger scale . 30 19. A simple ordination of the 30 creeping juniper stands showing the X-Z (top) and Y-Z (bottom) a x e s ............ 32 20. Soil depth (top) and percent frequency of stone (bottom) of soils in each of 30 creeping juniper stands plotted on the X-Y axes based upon the ordination in Figure 18. . 37 21. Percent soil surface eroded (top) and percent baregrourid cover (bottom) of soils of 30 creeping juniper stands plotted on the X-Y axes based upon the ordination in Figure 1 8 ...................• ....................... .. . 38 22. Elevations in meters of 30 creeping juniper stands plotted on the X-Y axes based upon the ordination in Figure 1 8 ................................................ 40 23. Annual chronology of creeping juniper in stands east of the Continental Divide as determined by spring, summer and fall observations............................. .. . . 71 Figure Page Ix 24. Growth curves of lateral (L) and terminal (T) twigs on creeping juniper shrubs in eight juniper associations as determined by the mean length of 15 terminal and 60 lateral twigs per site in 1977. a. Junipevus-PotentilZa- Festuoa association— sites 19, 20, 22, 25 . . . ......... 73 24. Continued, b. Junipevus-Agvopyvon association— sites 2, 3, 5, 17.................................................. ' 74 24. Continued, c. Pinus-Junipevus-Festuca association— sites 6, 7, 8, 3 1 ........................................ 75 24. Continued, d. Junipevus-Potentilta-Cavex association— sites 11, 12, 18.......................................... 76 24. Continued, e. Junipevus-Catamovitfa association— sites 15, 16, 26, 28, 29............................. 77 24. Continued, f. (top) Junipevus-Rhus-Stipa association— sites 13, 14 and 24g. (bottom) Junipevus-Hetiototviohon association— sites 10, 3 0 ............................... 78 24. Continued, h. Junipevus-Andvopogon-Festuoa association- sites I, 9, 27........................... 79 Continued, i. Non-clustered sites— sites 4, 21, 23 . . . . 80 Figure Page 24. ABSTRACT Various ecological characteristics and relationships for creeping juniper (Juniperus hovisontalis Moench.) in Montana were determined through intensive studies in 30 widely separated juniper stands. Supple­ mentary data were obtained from an additional 21 study sites. Creeping juniper was associated primarily with the mountain foothills of north- central, central and southcentral Montana, and the open prairies of east­ ern Montana. Ridges, rimrocks, escarpments and hilltops adjacent to a river or mountain range were typical areas of occurrence. Stands occur­ red more frequently on northerly exposures, slopes with gradients less than 40 percent, and at elevations below 1500 meters (4900 ft). Creeping juniper was the most, important plant species in all stands, with mean canopy coverage of 32 percent. Graminoids were the most prominent plant associates, with Idaho fescue being the most common species. Trees typ­ ically were lacking. Eight creeping juniper "associations" were defined through simple ordination. Soils associated with creeping juniper stands appeared to be poorly developed with thin topsoils, large quantities of stone and exposed rock parent material, and some degree of surface ero­ sion. They tended to be clay loam in texture, slightly basic (pH = 7.2), low to medium in organic matter content (4.6%), very low in phosphorus (18 ppm), medium to high in potassium (280 ppm) and low in salt hazard (.08 meq/100 gms soil). Creeping juniper plants averaged 6.4 cm in height, with 3.5 percent of the crown area dead. Most growth occurred horizontally on the peripheral branches or leaders. Annual twig growth commenced in early April, coincident with a crown color change from brown to green, and continued into September and possibly early October. The overall average terminal and lateral twig growth was 34.5 and 9.0 cm, respectively, during 1978. Longest twigs were produced in stands with northerly exposures and 13-14 inches of annual precipitation. Shortest twigs were produced in stands having a tree overstory. The "greening" of plants in early April coincided with staminate cone maturation. Once pistillate buds formed, berries generally formed and matured. Berry production peaked in mid summer (late June to early July). Ripening apparently requires 1-2 years after berries turn purple in late summer. Germination of seeds in the laboratory was very low (approximately 0.1%). Reproduction from seed also appeared to be a rare occurrence in the field where plants apparently are maintained vegetatively by rooting along branches. Plants producing the highest numbers of reproductive parts were associated with soils having relatively high phosphorus, mod­ erate calcium and sodium levels and low salt hazard. Wildlife usage of creeping juniper included mule deer, small mammals and birds. Utilization by deer occurred mainly during the late winter and appeared to be influ­ enced more by the distribution of deer, the relative availability and/or abundance of creeping juqiper and the occurrence of other more preferred forage plants than by protein content. Data indicated that creeping juniper is subject to some damage and destruction by fire, though these effects may be less severe than reported for other species of juniper. INTRODUCTION Creeping juniper, Junipevus hovizontalis Moench.',. occurs extensively on mountain foothill and prairie-plains habitat in Montana east of the Continental Divide. In many areas, it serves as an important source of forage for mule deer, especially during winter (Lovaas 1957, Kamps 1969, Eustace 1971,. Dusek 1971, Hamlin 1976). Coverage measurements for creeping juniper on 12 condition-trend transects, established by the Montana Department of Fish and Game on foothill winter ranges along the eastern fringe of the Rocky Mountains, show a general decline in the occurrence of mature juniper during the past 20 years (Fig. I). Little is known as yet about the ecology of creeping juniper. Previous studies have been concerned mainly with diseases (Brener et al. 1974, Nemec 1968), chromosomal biology (Evans 1971, Evans and Rasmussen 1972, 1974), ornamental value (Lamphear 1966), chemistry ■ (Couchman 1965), morphology (Bifoss 1947), associated insects (Bradley 1963, Nemec 1972), and hybridization (Fassett 1944a, 1944b, 1945a, 1945b, 1945c, Ross 1949). The lack of ecological information has hampered interpretation of possible interrelationships between the occurrence of creeping juniper and mule deer. This study was established in the spring of 1975 to obtain basic ecological data for creeping juniper in Montana. Specific objectives include: (I) to determine the natural distribution and associated . 2100- QC U-I CL. CD Z Q- U J U J QC 5 C-3 U J S 41 QC LAJ O CD U J CD QC UJ Q- r 1957 \ LEGEND — • —•Elk Creek --- -- ' T e t o n R i v e r - - - - - - Cobb Ranch - * * - Sun R i v e r Game Range 1959 1961 1963 \ \ \ \ 1965 1967 1969 1971 1973 Figure I. Population trends of creeping juniper on 12 condition trend transects established by the Montana Department of Fish and Game along the east slope of the Continental Divide. 3edaphic and climatological characteristics; (2) to determine community relationships, phytosociology, and animal use relationships; and (3) to determine plant characteristics such as reproduction, growth and development, forage biomass, production indices, fire influences, and nutritional aspects and, to relate these to environmental conditions. Field investigations were conducted primarily during the summers of 1975 and 1976 and in the spring and summer of 1977. Supplemental data were obtained in periodic field studies during the autumn and winter of 1976 and 1977. METHODS The natural distribution of creeping juniper in Montana was ascertained by ground reconnaissance throughout the State. Montana Department of Fish and Game range survey records and personal contacts with Department of Fish and Game and U. S . Forest Service • personnel along with general publications as Little (1971) provided additional information. Following the general distributional surveys, 30 sites represent­ ing an array of plant communities and physiographic situations (slopes, exposures, elevation, etc.) in which creeping juniper was observed to occur were selected for intensive investigation (Fig. 2). Additional basis for site selection were (I) a general and relative lack of major disturbance and (2) a history of importance for use by mule deer during winter. An additional 21 similarly selected sites were visited during the study to obtain supplementary data on edaphic and physiographic characteristics of creeping juniper stands. Locations of all sites are presented in Appendix Tables 13 and 14. Generally, all study sites were located in northcentral, east- central and southcentral Montana. A majority (39 or 76 percent) were in areas currently grazed by domestic livestock. Two (4 percent) were situated in game exclosures on previously grazed sites. Two others were located in recreation areas and three (6 percent) were on big game winter range from which livestock grazing had been removed for at least MONTANA ,Glasgow r e a l Fa l l s 26)(27 Hel ena l i es C i t y Big Timber B i l l i n g s Broadus LEGEND ----- CONT INENTAL D I V I D E O I NTENS IVE STUDY S I TES (no’s 1 - 3 1 except 24) Q SUPPLEMENTAL SITES (no' s 3 2 - SI plus 24 ) Figure 2. Study areas showing the approximate locations of 51 study sites. 6twenty years. Five (10 percent) were within.areas fenced for crop­ lands which were ungrazed at the present time and may represent relict areas. Slope, exposure, elevation and exact location were determined for each study site. Slope was estimated to the nearest five percent using a pocket transit. Exposure was recorded from compass readings along the fall-line of the slope. Elevation was determined using a simple pocket altimeter. Within each of the 30 intensively studied stands, a 30.3 meters (100 ft) x 12.1 meters (40 ft) macro-plot was established. Sampling was conducted in the macro-plot along three permanent 30.3 meters (100 ft) transects spaced 6.1 meters (20 ft) apart along the contour of the. slope. Canopy cover of low-growing vegetation was recorded following the method of Daubenmire (1959) within 2 x 5 dm plot frames spaced 3.1 meters (10 ft) apart along the three, 30.3 meters (100 ft) lines. The cover of each plant species as well as bare ground, rock and litter was recorded visually by class as follows: (I) 0 - 1 percent; (2) I - 5 percent; (3) 5 - 2 5 percent; (4) 25 - 50 percent; (5) 50 - 75 percent; (6) 75 - 95 percent; and (7) 95 - 100 percent. Measurements were made in early June. Common and scientific names of plants follow Booth and Wright (1958), Hahn (1973), and Hitchcock and Cronquist (1973). When creeping juniper was encountered in the frames, the sex of the plant and an estimate of the crown decadence within the plot were also recorded. 7Where they occurred, trees and taller shrubs were sampled I separately using a line intercept method. Three 30.3 meters (100 ft) lines were established; one located 24.2 meters (80 ft) upslope from the upper permanent transect line, one along the center transect line, and one 24.2 meters (80 ft) downslope from the lower transect line. • Crown intercept was recorded to the nearest 0.3 meters (1.0 ft) along each line. On sites where few trees or tall shrubs occurred, all trees within a 60.6 meters (200 ft) x 60.6 meters (200 ft) plot at each site were measured as to species, crown diameter (mean of major and minor axes) to the nearest decimeter, percent of total crown dead, and percent live crown within an imaginary circle around the outer edge of the plant. For data analyses, study sites were grouped on the basis of vege- tational characteristics using a simple multidimensional ordination technique described by Bray and Curtis (1957) and modified by Beals (1960) and Orloci (1966). The ordination technique assumes that the compositional similarity between different stands can be used as compar­ ative distances within a spatial model. The phytosociologic distances are based on a similarity index with 0 percent being completely dissimi­ lar and 100 percent, theoretically, total similarity. The Spatz simi­ larity index (Mueller-Dombois and Ellenberg 1974) was used to compare sites on the basis of (I) the number of plant species common to both sites, and (2) the number of species present at each. site. Data used in the 8analyses were frequencies of occurrence of 84 low-growing species on study sites during June 1977. When indices were calculated for each two sites, the two most dissimilar sites became the reference points for the X-axis. A third site which was most distant from the X-axis (line A-B) became the third reference point for the X-Y plane. Finally a fourth site, chosen on its distance from (dissimilarity) the X-Y plane, became the fourth reference point for the X-Y-Z solid (third dimension). An X, Y, and Z coordinate was then calcu­ lated for each site using geometric formulae and the similarity indices. All sites were plotted first on X-axis, then on the X-Y plane, and finally on the X-Y-Z solid. Clusters of similar vegeta­ tion (associations) became apparent in this figure. Two soil samples were obtained from each of the 30 intensive study sites. One of these was a composite of 10 subsamples from soil to a depth of 10 cm beneath creeping juniper plants; the other a similar composite of 10 subsamples from interspaces. One composite soil sample was taken within each of the 21.supplemental sites. , Analyses were completed by the Montana State University Soils Testing Laboratory and included hand texture analysis, salt hazard, sodium, calcium, magnesium, potassium, phosphorus, and organic matter content, and pH. Depth to bedrock or parent material at each site was recorded as the mean of measurements with a sharply pointed (1.5 cm diameter) stake at 10 randomly located points at each site. An index of soil 9surface erosion at both intensive and secondary study sites was obtained by visually estimating the overall percentage of surface eroded at each site by class as follows: Class I = less than 5 percent; Class 2 = 5-25 percent; Class 3 = 25-50 percent; Class 4 = 50-75 percent; and Class 5 = 75-100 percent. Wind and water erosion were not differentiated. Edaphic characteristics were compared statistically among the eight creeping juniper associations identi­ fied by ordination. In addition, the possible influences of each of the soil factors on frequencies of occurrence and canopy coverage values for each of 17 graminoids (grasses and grass-like plants), 39 forbs, 9 shrubs and 2 trees among study sites were evaluated by multiple regression. Growth and reproductive characteristics of creeping juniper plants were measured during the spring and summer of 1977. Before the onset of growth, 15 major branches or. "leaders" were tagged on each of 3 randomly selected plants at each study site.. Leaders were marked with India ink on two year old or older growth, and the distances from the marks to the tips of leaders were measured to•the nearest millimeter. Remeasurements were made at two-week intervals until growth ceased. Four lateral twigs were similarly marked and measured on each of the 15 leaders. Standing crop of creeping juniper was estimated from the average air-dried weight of plant material, including needles, twigs 10 and new foliage growth clipped from ten, 2 x 5 dm frames per site. Average plant heights to the nearest 0.5 cm was determined as the mean of 10 measurements at each site. When growth had nearly ceased and "berries" were maturing, numbers of new pistillate (female) buds, old pistillate buds, new staminate cones, new "berries" and ripe "berries" were recorded at each site for each of 10 twigs on each marked leader for male and female plants respectively. Supplemental data were obtained by recording the same characteristics within 10 randomly placed 2 x 5 dm plot frames at each site. General phenological data were recorded for each study site at • approximately two-week intervals. These included browness-greeness of foliage .throughout the year, initiation and termination of growth, staminate cone formation, maturation and time of shedding,. pistillate bud and berry formation, berry maturation and pollen shedding. To evaluate reproductive potential from seeds, juniper berries were collected at 20 locations. Sufficient berries were collected to provide about 500 seeds per location (based on an average of 4 seeds per berry). Because juniper berries, may not ripen for at least 1 - 2 years after they turn purple and drop (U. S.. Dept. Agric. 1948), only berries found on the ground under plant crowns and which, by their pulpy exterior coat, were believed to be more than one year old were collected.. Germination procedures followed those outlined for Rocky 11 Mountain juniper (U. S. Dept. Agric. 1948). Minor modifications included stratification of seeds for 125 days at 4° C and a germi­ nation period of six weeks at 12° C. To determine nutritional value, samples of leaf-stem materials were collected at three month intervals from August 1976 through August 1977. Composite samples were clipped from a randomly . selected plant at each of 12 study sites. In August 1977, samples from two additional plants per site were obtained. Samples were returned to the laboratory, air dried and ground using a Wiley Mill with 20 mesh screen. Ground samples were oven dried at 80° C for 48 hours and analyzed for protein content by the Montana State University Chemistry Station. During the course of the study, incidental observations of twig use by small mammals and disease occurrence were made. Observations on the effects of fire on creeping juniper were made at two locations during the spring and summer of 1977. One area, located in the Blackball Hills area of the Little Belt Mountains south of Stanford, Montana, had been subjected to controlled burning under experiments conducted by Mr. Larry Keown of the U. S . Forest Service, Stanford Ranger District. Here, three adjacent plots, each 100 m^ were burned at different intensities (high, moderate and light) based upon temperature and rate of spread. Prior to the fire within each plot, seven juniper plants were randomly selected, and major and minor axes, and height of each were recorded to the nearest 1.0 cm. Soil and fuel moisture samples were taken and pyrometers were positioned adjacent to each shrub. Following the burn the fate and regrowth characteristics of each plant were monitored visually during the summer of 1977. Information recorded included percent foliage burned and evidences of regrowth. Total air- dried weight of regrowth was recorded at the end of the growing season. The second area, located on the Sun River Game Range west of 2 2 Augusta, included a single 30.3 m (100 ft ) burn plot and a control plot of equal size. Prior to burning, soil and fuel moisture samples were taken and pyrometers positioned at 6 locations within the burn-plot. Two 30.3 meters (100 ft) line transects were- established, one in the burn-plot and one in the control. Occurrence of litter, live and/or dead grass, forb or shrub, or bare ground was recorded at 0.3 meters (I ft) intervals along each transect. RESULTS AND DISCUSSION Distribution Geographic Juniperus horizontalis Moench. is one of about 60 species of evergreeen shrubs’ and trees included in the genus Juniperus. Members of this genus occur in the middle and lower latitudes of the northern hemisphere in North America, the West Indies, Canary Islands, northern and eastern Africa and southeast Asia. Fifteen species are native to the United States. Of these 13 species are found in the western United States, and four are native to Montana.■ These include common juniper (Juniperus communis) and Utah juniper (Juniperus osteosperma), both shrubs occurring primarily in mountainous regions, Rocky Mountain juniper {Juniperus scopulorum), a tree-like form, occurring throughout the state, and creeping juniper, a very low shrub found primarily in mountain foothills and open prairies. The center of distribution of creeping juniper in North America is in central Canada; the species occurs across the entire country in vast stands (Little 1971). In the northern United States it occurs from the east coast to the Rocky Mountains, primarily in scattered, local populations. Within Montana, creeping juniper can be found in nearly every county east of the Continental Divide (Fig. 3). The most extensive stands occur along the Missouri River upstream to Fort Benton; along MONTANAn G l a s g o w K r e a t f a l l s L e w i s t o w n ',Helena B ig T i m b e r B i l l i n g s Broadus LEGEND - - - CONTINENTAL DIVIDE Figure 3. General distribution of creeping juniper in Montana. 15 the Milk River upstream to the Canadian border; in foothills along the east face of the Continental Divide from Glacier Park south to Wolf Creek; and throughout the foothills of the Snowy Mountains, Judith Mountains, and eastern portions of the Little Belt Mountains. Scattered local populations are found in far eastern Montana; and in . the Beartooth-Absarokee Mountains south of Big Timber and west of Red Lodge. Areas where creeping juniper was common were generally .charac­ terized by rough "breaks" and ridges adjacent to a river or mountain range. Ridges, rimrocks, hilltops and interstream divides were common" landscapes supporting juniper communities; typically, however, juniper was restricted to the upper portions of these areas. Physiographic Creeping juniper is found in a wide variety of physiographic and physiognomic situations in Montana (Figs. 4 to 16). Study site eleva­ tions ranged from 2150 to 8200 feet; the average was 1326 ± 113 meters (4375 ± 373 feet). A majority of the sites (60 percent) were below 1500 meters (4900 feet) (Table I). • Slope gradients varied from 4.5 percent to 53.5 percent with a mean of 25.6 ± 4.0 percent; however, most of the sites were on slopes of less than 40 percent. All aspects or exposures were represented. Eight sites (15%) were ENE, 8 were SSE, 9 (18%) were NNE, 6 (12%) were WSW, 10 (20%) were NNW, 2 (4%) were WNW, 3 (6%) were SSW, 4 (8%) were ESE,, and I (2%) faced south. -16- Figure 4. Site I on an alluvial plain near Barr Creek on the Sun River Game Range, Lewis and Clark County. Juniperus-Festuaa dominant cover. Figure 5. Site 3 near Haystack Butte southwest of Augusta, Lewis and Clark County. Juniperus-Festuoa dominant cover. -17- Figure 7. Site 15 on edge of a Pinus ponderosa community south of Warhorse Reservoir, Petroleum County. Juniperus- CatamoviZfa dominant cover. -18- Figure 8. Site 16 on an open prairie south of Warhorse Reservoir, Petroleum County. Juniperus-Agropyron dominant cover. Figure 9. Site 19 in foothills of the Big Snowy Mountains, Northside, Fergus County. Potentilla-Festuoa dominant cover. -19- Figure 10. Site 21 in the foothills of the Little Belt Mountains, Southside, Wheatland County. Near a Pinus ftexilus — Pinus ponderosa community; Juniperus-Koeleria-Festuca dominant cover. Figure 11. Site 25 in an Absarokee mountain meadow, Stillwater County. Near a community of Pinus flexilus and Pseudotsuga menziesii; Juniperus-variety of forbs such as Zygodenus and Anemone dominant cover. -20- Figure 12. Site 26 on an eastern Montana prairie northwest of Terry, Prairie County. Juniper^us-Carex dominant cover. Figure 13. Site 27 on an eastern Montana prairie/foothill area north­ west of Terry, Prairie County. Juniperus-Andropogon dominant cover. —21 — Figure 14. Site 32 on the West side of the Continental Divide east of Lincoln, Lincoln County, under a stand of Pinus oontorta and Pseudotsuga menziesii. Arotostaphlos-Agropyron dominant understory cover. Figure 15. Site 37 under a stand of Pinus ponderosa north of Ryegate in the Little Snowy Mountains, Golden Valley County. Juniperus horizontalis and scopulorum dominant understory cover. -22- Figure 16. Site 42 under a stand of Pinus ponderosa northwest of Broadus, Powder River County. Juniperus-Stipa dominant understory cover. 23 Slope gradient, exposure, and elevation of individual study sites are given in Appendix Table 14. Table I. Distribution of 51 creeping juniper study sites among various exposures, slope gradients and elevations. Physiographic Categories Number of Stands Percent Exposure (Azimuth Degrees) NE 1-90 16 ' 31 NW 271-360 13 26 SE 91-180 15 • 29 SW 181-270 7 14 N 271-90 29 57 S 91-270 22 43 E 1-180 28 ■ 55 W 180-360 23 45 Slope Gradient (percent) Less than 40 42 82 . More than 40 9 18 Elevation (feet) Less than 4900 30 60 4900-5500 11 22 More than 5500 9 ■ 18 Creeping juniper appeared to be largely absent from three major regions of the state. These included all of Montana west of the Continental Divide with the exception of three areas (one near Rogers Pass, one near Red Mountain north of Lincoln, and one near Marias Pass); southwestern Montana; and a region along the Yellowstone River between 24 Columbus and Miles City (Fig. 3). These areas were typified either by mountains, in western and southwestern Montana, with dense forests and high precipitation, or flat grazing and farm-lands with deep and/or gravelly soils. • ' Phytosociological Characteristics Community Associations Because of its wide geographic range, creeping juniper occurs in numerous vegetational communities or associations along with a diverse array of plant species. However, most vegetational studies involving creeping juniper have been conducted along the southern border of its range. Breitung (1954) included creeping juniper as part of the shrub component of mixed-grass prairie vegetation (Agropyron-Stipa associa- tion) characteristic of the eastern edge of the Cypress Hills in southeastern Alberta and southwestern Saskatchewan. This vegetation type, which occurred at 900-1200 meters (3,000-4,000 ft) elevation, was dominated by mixed grass species characteristic of the Canadian prairie. He also noted creeping juniper occasionally occurring on dry exposed hillsides 1200-1500 meters (4,000 and 5,000 ft) elevation. In Montana, creeping juniper is important in two of the 22 vege­ tational types delineated by the Montana Agricultural Experiment . Station (1973) . These include the Badlands Grassland in eastern Montana and the Northeastern Grassland of northeast Montana. It also 25 occurs in the Foothill Grassland type, the Lodgepole Pine-Douglas Fir Forest type west of the Continental Divide, the Ponderosa Pine Savannah, the Missouri Breaks Scrub-Pine, the Northern Grasslands, and locally in the Teton River - Judith Basin Grasslands. The minor occurrence of creeping juniper in the Lodgepole Pine-Dpuglas Fir Forest type near Lincoln and near the Marias pass was its only observed occurrence west of the Divide. Ross and Hunter (1976) placed stands of creeping juniper in a single "riverbreaks" climax vegetation type characteristic of eastern and western sedimentary plants. . Major overstory associates included ponderosa pine (Finns ponderosa), Rocky Mountain juniper, and limber pine (Finns flexitns') . The most common grasses were bluebunch wheatgrass (Agropyron spiaatnm), green needlegrass (Stipa virdula), and prairie sandreed (Calamovilfa longifolia). I observed creeping juniper in this association mainly in eastern Montana, along the Marias, Milk and Missouri Rivers. In the Sweetgrass Hills of northcentral Montana, Thompson and Kuizt (1976) found creeping juniper in what they described as a Montana Grassland community covering most, dry, south-facing slopes and foothills between 1,500 meters (4,950 ft) and 2,000 meters (6,600 ft) as well as many non-forested areas of north-facing slopes. This community was dominated by rough fescue (Festuoa Soabretla), Idaho fescue (Festuoa idahoensis) and shrubby cinquefoil (Potentilla. fruitioosa). My study 26 sites along the eastern foothills of the Rocky Mountain front and in central Montana seemed to represent similar situations. Jorgensen (1976) described two tentative habitat types in eastern Montana which included creeping juniper, a Broom Snakeweed— Bluebunch Wheatgrass type, which occurred on very steep, unstable south­ facing slopes, and a Creeping Juniper— Bluebunch Wheatgrass type occur­ ring on north-facing slopes. The most abundant species in the former, which appeared to represent the vegetation of my study site number 26, were bluebunch wheatgrass, broom snakeweed and creeping juniper. Other plants of importance listed were rose, eriogonum, skunkbush sumac and prairie thermopsis. The latter type was dominated by creeping juniper with bluebunch wheatgrass, flowery anemone {Anemone ■ multifida) and Kelsey phlox as common associates, and was characteris­ tic of vegetation at study sites 27 and 28. In reviewing the foregoing studies on creeping juniper and its associates it should be kept in mind that each researcher used his own method of classification giving the impression that creeping juniper occurs in a wider variety of vegetational types than it actually does. In principle then, the various types described may only represent one or two types described by different men. Existing vegetational descriptions, together with my observations, suggest that several species are commonly associated with creeping juniper. Tree species, if present, include ponderosa pine, limber pine 27 and/or Douglas, fir (Pseudotsuga menZdessi). Principle shrubs include shrubby cinquefoil, common snowberry (Symphor-Learpos albus), skunk- bush sumac (Rhus tvi-lobata), prickly rose (Rosa ac-Lcular'is) and prairie rose (Rosa avkansana). Major grasses include bluebunch wheatgrass, Idaho fescue, rough fescue, spike oat (Eel1LctotT1Lchon hookeri), little bluestem (Andpopogon seopap-Lus) and prairie junegrass (KoeteT1La epistata). Two sedges, inflated sedge (Capex Ves1Leapa) and threadleaf sedge (Capex f 1Lt1Lfotta) , may also be prominent; while a variety of forbs commonly occur. The plant associates of creeping juniper in Montana presumably vary in their occurrence and relative importance with variation in environmental factors such as topography, soils, climate, fire, grazing and time; and "associations" (i.e., distinctive combinations of creeping juniper and other plant species which persist on the same area or on areas of similar site and environmental charac­ teristics) may not be readily apparent by examination of a few important plant species alone. As indicated earlier, I employed ordination of vegetational data for the 30 intensively studied stands to identify "associations" a s , well as to illustrate the nature of variation in creeping juniper communities and the relationship of this variability to the environment. The raw vegetative data for each site is presented in Appendix Table 16. An ordination of vegetational data for individual stands generates, a figure where stands with similar vegetational characteristics are ' i 28 I ! positioned adjacent to one another and dissimilar sites are widely I separated (Fig. 17). When environmental attributes of each site are plotted into this figure, those which are correlated with and possibly causal of the variation in vegetation will vary in parallel with the phytosociological structure while those which are not will vary randomly (Figs. 20, 21, 22). Specific ordination techniques were outlined under methods. Briefly, to identify "associations": (I) the phytosociological "distances" between stands or sites were calculated; (2) the most dissimilar sites were selected as reference points for the X-axis (Site Nos. 13 and 15), the X-Y plane (Site Nos. 13, 15 and 25) and the X-Y-Z solid (Site Nos. 13, 15, 25 and 16); and (3) each stand or site was plotted graphically, first on the X-axis, then on the X-Y plane and finally in the X-Y-Z solid. The principle associations in which creeping juniper occurs in Montana are illustrated in Figure 18, where total variability of sites is collapsed into one plane represented by the X and Y-axis (Figs. 17 and 18). Eight clusters, representing vegetational associations are recognizable. Individually, the associations were (I) Jimiipevus- Potentilla-Festuca3"{2) JunipeTus-Agropyron3 (3) Pinus-Juniperus- Festuea3 (4) Juniperus-Potentilla-Carex3 (5) Juniperus-Calamovilfa} (6) Juniperus-Helietotrichon3 (7) Juniperus-Rhus-Stipa3 and (8) Juniperus-Andropogon-Festuca.. Some clusters (associations) are riot clearly separated from the I 29 A simple ordination of 30 creeping juniper stands based on total vegetal composition, in two dimensions (X-Y axes). Figure 17. Axes represent compositional dissimilarity between stands. Juniperus- P o te n t i l la - Festuca Juniperus-Potentilla Pinus-Juniperus-Festuca. Jun ipe rus -A g ropyron Jun ip e ru s - Andropogon- Festuca JuniperusJuniperus- CaIamoviHa Jun iperus - H e lic to trichon Figure 18. The same ordination as Figure 17 only on a larger scale. 31 others on the X-Y plane (Fig. 17). However, these clusters clearly separate when the third dimension (X-Z and Y-Z plots) of the ordina­ tion is examined. For example, sites 19, 20, 21, 22, and 25 appear to represent a single association on the X-Y graph. However, examination of the X-Z and Y-Z plane plots (Fig. 19) readily separates site 21 as possessing a different plant composition from the other sites. Certain sites (4, 21 and 23) do not readily fit well into any of the associations listed. A larger number of samples would be required to understand their composition and environmental qualities. Since they appeared to be relatively unimportant, little further discussion will be devoted to them. Jun-Ipevus-PotentrHla-Festuoa This association occurs at elevations from about 1500 to 1800 meters (5000 - 6000 ft) in the foothill regions. It is typified by a high forb cover including death camas (Zygodenus venosus), Kelsey phlox {Phlox kelsey-l), kitten-tail {Besseya wyomlngens-ls), white point-loco (Oxytvop-is sevioeus), northern bedstraw {Galium boveale), and common yarrow {Achillea millefolium). Dominant graminoids include Idaho fescue, western.wheatgrass {Agvopyvon smithii) and inflated sedge; I while creeping juniper and shrubby cinquefoil were the major shrubs. This type is relatively open. 32 o ©7! 12,18O ? V 5 4,30 O 60 40- ? § 4 1 3.i i i? 2 » ! ? 10,30O 20 Figure 19. A simple ordination of the 30 creeping juniper stands showing the X-Z (top) and Y-Z (bottom) axes. 33 Jimiperus-Agroyyron This association occurs between 1200 - 1500 meters (4000 - 5000 ft) in the prairie-foothill regions. It is characterized by rocky, shallow soils with very little overstory. The cover is made up pri­ marily of graminoids including bluebunch wheatgrass, threadleaf sedge, Idaho fescue, and rough fescue. Common forbs are fringed sagewort (,Artemisia frigida) , northern bedstraw and alyssum-leaved phlox (Phlox alyssifolia). Creeping juniper and prickly rose make up the shrub complex. Pinus-Juniperus-Festuea This association is mainly found between 1500 - 1800 meters (5000 - 6000 ft). Rocky, shallow soils along with a limber pine overstory are characteristic. Bluebunch wheatgrass, threadleaf sedge, Idaho fescue and rough fescue are dominant graminoids. Prominent forbs are Kelsey phlox, fringed sagewort and Arizona hymenoxys (Hymenoxys acaulis). The only common shrub is creeping juniper. Juniperus-Potentilla-Carex Typically, this association occurs between 1200 and 1500 meters (4000 - 5000 ft) in the foothills. It commonly contains'a high forb complement including such prominent species as common yarrow, Arizona hymenoxys, alyssum-leaved phlox, Kelsey phlox and ball anemone (Anemone multifida). The major graminoids are threadleaf sedge, inflated sedge. 34 single-spike sedge (Carex sOirpoidea), spike oat, rough fescue and Idaho fescue. Shrubs present are creeping juniper and shrubby cinquefoil. Juniperus-Calamovilfa This association is characterized by a high amount of surface erosion and bareground. Also present is moderate tree cover mainly of ponderosa pine. It occurs at elevations between 900 - 1200 meters (3000 - 4000 ft). Dominant graminoids are prairie sandreed grass, threadleaf sedge, bluebunch wheatgrass and western wheatgrass. Forbs are pale, bastard toad flax (Comandra umbellate), fringed sagewort and prairie.thermopsis (Fhermopsis rhombifolia). Common shrubs include creeping juniper, prairie rose, and big sagebrush (Artemisia tridentata). Juniperus-Heliotrotriohon This association occurs at 1500 - 1800 meters (5000 - 6000 ft) in the foothills. Small trees, primarily limber pines, are common. Major graminoids are spike oat, bluebunch wheatgrass, threadleaf sedge, inflated sedge, Idaho fescue, rough fescue and prairie junegrass (Koelaria oristata). Common forbs are rose pussytoes (Antennaria rosea), buff fleabane (Erigeron oohroleuous) and Kelsey phlox. Dominant shrubs include creeping juniper, shrubby cinquefoil and prickly rose. 35 Junipevus-Rhus-Stipa This association is found between 900 - 1200 meters (3000 - 4000 ft), and is typically dominated by shrubs. No trees occur in this association. Prominent shrubs are creeping juniper, skunkbush sumac and common snowberry. Common graminoids are threadleaf sedge and needle and thread (,Stipa comata); narrowleaf poison vetch (,Astragalus peotinatus), kittentail, prairie trefoil {Lotus purshianus) and Kelsey phlox were major forbs. Juniperus-Andropogon-Festuoa This association occurs at elevations between 1200 - 1500 meters (4000 - 5000 ft) in the prairie regions. Prominent graminoids include bluebunch wheatgrass, Idaho fescue, rough fescue and threadleaf sedge, with prairie thermopsis and fringed sagewort as dominant forbs. Major shrubs are creeping juniper, shrubby cinquefoil and prickly rose. Environmental Attributes of the Eight Associations As noted above, environmental data from each site can be plotted into the X-Y ordination (Figs. 17 and 18). Factors apparently related to vegetational variation are plotted in Figures 20 to 22. Other environmental factors.studied, but which were not correlated with variation in vegetational characteristics, and/or were not plotted, include frost-free season, average annual temperature and precipitation, percent organic matter, calcium, magnesium, sodium, potassium, or 36 phosphorus content, salt hazard, soil texture, pH, north, south, east, west exposure and slope gradient. The Pinus-Jun-Lperus-Festuaa and Juniperus-Agropyran associations near the center of Figure 20 had the rockiest and shallowest soils. The soils averaged near 10.0 cm. deep and had frequencies of 90-100 percent stone occurrence. Soils near the top of the graphs were moderately stony (frequency 50%) and relatively deep (14 cm.) while those at the lower edges were deepest and relatively free of stones averaging near 25 cm. deep and 30-50 percent frequency of stone occurrence. Surface erosion increased toward the center and lower left corner of the graph (Fig. 21). The Juniperus-Catamovitfa association occurred on the most eroded sites and the Juniperus-Potentitta-Festuoa3 Juniperus-Andropogon-Festuaa and Juniperus-Potentitta associations were on the least eroded, averaging approximately 50, 3,, 3 and 3 per­ cent of the surface eroded respectively. Erosion probably is more rapid where vegetal cover is low; i.e., where total bare ground cover is high. • Bare ground is well correlated with erosion estimates (Fig. 21). •Associations, as the Juniperus-Catamovitfa and Juniperus-Rhus-Stipa3 at the lower edge of the graph, averaged around 30 percent of the sur­ face as bare ground. The associations toward the top, for example the two Juniperus-Potentitta associations (I and 4), averaged much less 37 t 13 H 10 1.5 ' I 3.3 y 2.1 « 16 » 10 'Yn 6 12 O 12,13 S ? G % 11,16 18 1.6 1.5 «-23 28 t 7 3.7 7.3 8.0 23 -n 3.3 8.7 33 7.3 8.7 3 50 13.70 ° 97 O 100,30 3 100 O 6-0 43,23 20- Figure 20. Soil depth in centimeters (top) and percent frequency of stone (bottom) of soils in each of 30 creeping juniper stands plotted at the locations of the respective sites within the ordination presented in Figure 18. 38 I ? 3.8 1.5 6.3 3? 1.5 3 1.5 ? I O V 15^5 I? 1.5 ? « 6 3 31>56 3-» Figure 21. Percent soil surface eroded (top) and percent bareground cover (bottom) of soils of 30 creeping juniper stands plotted at the locations of the respective sites within the ordination presented in Figure 18. 39 bare ground (approx. 5-10 percent). The association occurring at highest elevation, JunrLpevus- PotentLLLa-Festuea3 was near the top of the graph (Fig. 2.2). Ele­ vation appears to decrease toward the lower edge of the figure. Values ranged, from near 1500 meters (5500 ft) in the JunLpevus- PotentLLLa-Festuea3 JunLpevus-HeLLctotvLchon and PLnus-JunLpevus-- Festuea associations to less than 1200 meters (4000 ft) for JunLpevus- Rhus-StLpa and JunLpevus-CaLamovLLfa associations. Vegetational Composition of Creeping Juniper Associations Plant species associated with creeping juniper are listed in Table 2 together with their average constancy, percent canopy coverage and frequency of occurrence among 2 x 5.dm plot frames in each of the eight vegetation associations (Fig. 18). A total of 150 plant species was identified including 24 graminoids, 104 forbs, 18 Shrubs and 3 trees. Common and scientific names are presented in Appendix Table 15 The general vegetal composition of juniper stands may be obtained by averaging across all associations. Creeping juniper typically dominated the sites studied accounting for an overall average canopy coverage, of 32 percent of the total area sampled. Although some sites have high (60-70 percent) canopy coverage of vegetation, considerable bare ground and exposed surface stone were common, and together accounted for 30 percent of the total canopy coverage. Deciduous and t m s 1606 1606 15.76 1667 1030 1030 HSJi 14.24 148SJ364 6 fig. ISIS 1424 Q 1460273 1879 1485 1606.1636 G 1515 1030 1758 1576 \ *1000 3600^ 3 60 0- Figure 22. Elevations in meters of 30 creeping juniper stands plotted at the locations of the respective sites within the ordination presented in Figure 18. Table 2. Constancy, canopy coverage and frequency of low growing taxa for the eight creeping juniper associations as determined by examination of 2 X 5 decimeter plots on each area. ASSOCIATIONS Juniperua- Pinua- Jumperus- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Rhus- Andropogon- Taxa Festuca Agroppron Festuaa Carex Calovovilfa Heliototriahon Stipa Festuoa Totals Graminoids: Agropyron smithii* 100/ 6/45' Agropyron epicatwn * 25/ I/ 9 Andropogon seoparius ~ - Aristida longiseta - ~ ~ Bouteloua graoilis** ~ ~ Bromus inermis** — — — Bromus japoniaus* — — - Calamovilfa longifolia* ~ — Calamovilfa purpescens* - — — Carex elynoides* 25/tr/ 92 Carex fH i folia* 25/tr/ I Carex heliofila - “ - Carex soirpoidea* - ~ “ Carex vesioaria* 75/ 4/38 Canthonia intermedia ~ “ Festuca idakoensis* 100/ 8/64 Festuca saabrella* - - - Helictotriehon hookeri** - - - Juneus balticus - ~ ~ Koelaria eristata* “ - ~ Poa pattersoni* ~ ~ “ Poa spp. “ “ “ Stipa aornata - - “ Stipa viridula - - ~ Unknown Grass — — — Total Graminoids* 100/20/98 100/ 8/54 100/ 5/31 100/ 3/18 100/ 7/38 75/ 4/17 75/ I/ 5 100/ 8/53 75/ 5/27 75/ 2/12 75/ 6/37 75/ 6/32 25/tr/ 2 100/tr/12 33/ 1/12 66/ I/ 9 33/tr/ 3 66/ 1/12 33/ 3/16 66/ 2/22 100/ 3/22 66/ 4/22 100/ 5/24 33/tr/ 2 33/tr/ 3 40/ 4/22 60/ 4/26 20/tr/ I 60/ 5/33 20/tr/ I 60/ 3/22 20/ 1/10 40/ 2/10 40/tr/ 2 40/tr/ 5 20/tr/ 2 100/ 3/14 100/ 6/30 100/ 6/36 50/ 4/28 50/ 2/24 100/ 8/44 50/ 4/22 25/tr/ 4 25/tr/ 2 25/ I/ 6 100/29/97 100/27/99 33/tr/ 3 100/21/98 20/tr/ I - - - 40/ I/ 6 — — - 100/20/85++ 100/28/98 50/ I/ 6 100/ 4/24 50/ I/ 4 100/18/88 100/25/98 33/ 2/ 7 66/ 4/28 33/ 2/14 33/tr/ 2 100/ 5/28 100/ 2/12 33/ I/ 2 66/ 8/31 66/ 2/ 8 66/ 3/ 8 66/ 2/16 100/28/96 25/ I/ 9 63/ 3/28 4/tr/ 2 4/tr/tr 8/tr/tr 8/tr/ 2 4/tr/tr 11/ I/ 5 4/tr/tr 8/tr/ I 74/ 4/22 4/tr/ I 4/tr/ 2 74/ 2/19 4/tr/tr 74/ 5/30 44/ 3/14 33/ 2/ 8 4/tr/tr 47/ I/ 6 4/tr/tr 4/tr/tr 11/ 2/11 8/tr/ I 11/tr/ I 100/22/96 Table 2. (continued) ASSOCIATIONS Taxa Juniperus- Potentilla- Festuca Juniperus- Agropyron Pinus- Juniperus- Festuca Juniperus- Potentilla- Carex Juniperus- CalomoviIfa t/ uniperus- Heliatotriehon Juniperus- Rhus- Stipa Juniperus- Andropogon- Festuaa Totals Forbs: Achillea millefolium*1 75/ 3/29 50/ I/ 8 50/tr/ 2 66/ 2/ 4 20/tr/ I 100/ I/ 7 50/ I/ 7 66/ 1/11 55/ 1/10 Allium cernuum ~ ~ - - - - - - - 66/Cr/ 7 - ~ - 50/tr/ 4 — — — — — — - - - Allium textile*1 25/tr/ 6 75/ 1/25 75/ 1/26 33/tr/ 2 20/tr/ 2 50/tr/15 — — — 33/tr/ll 40/tr/ll Anemone multifida* 50/ 2/16 25/tr/ I - - - 100/ 3/18 20/ I/ 5 ~ - — — — 33/ 2/18 29/tr/ 7 Anemone patens** 75/ 5/41 25/tr/ 2 25/tr/ I 66/ 1/17 ~ ~ - 50/tr/ 2 — — — 66/ 2/ 9 37/ I/ 9 Antennaria rosea* 50/tr/ 7 75/ 1/11 100/ 1/21 66/tr/ 2 - - - 100/ 3/23 50/tr/ I 66/tr/ 5 59/ I/ 8 Arenaria conqesta 25/tr/ 7 25/tr/ I 25/tr/ 7 66/ 1/10 “ ~ “ - - ~ - - ~ 33/tr/ 2 22/tr/ 3 Arenaria hookeri ~ - 25/tr/ 9 25/tr/ 7 33/tr/ 3 - ~ - - - - ~ ~ ~ - - ~ 11/tr/ 2 Artemisia cana ~ ~ - ~ ~ ~ - - - - ~ ~ 20/tr/ 2 - “ “ 50/tr/ 2 - - — 8/tr/tr Artemisia friaida* - - — 100/ 2/20 100/ 4/33 100/ 2/25 60/ I/ 7 100/ 1/11 100/ 1/10 100/ 4/28 78/ 2/15 33/trZ 2 25/tr/ I 50/tr/ I 50/ I/ 4 50/tr/ 4 40/ I/ 6 50/ 1/12 33/tr/ I 33/tr/ I 50/ 4/19 4/tr/tr 18/tr/ 2 22/tr/ 2 4/tr/ 2 - " - - - ~ - - - 33/tr/ 3 - - - - - - - - - — — - 4/tr/tr Balsamorhiza saggitata* 25/ I/ 5 25/tr/ 2 25/tr/ I - - - 20/tr/ 2 - - - - - - 33/ I/ 4 18/tr/ 2 Besseua wuominaensis* 75/ 4/27 25/tr/ 3 25/tr/ I 66/tr/ 2 20/tr/ 3 50/tr/ 8 50/ 3/15 100/ 1/17 47/ I/ 9 Buvlerum amerieanum* - - - 75/ I/ 8 56/ I/ 3 33/tr/ I - - - 100/tr/ 7 - - - - - - 29/tr/ 3 Campanula rotundifolia* - “ - - - - - - - 33/tr/ I 20/tr/ I - - - - - - 33/tr/ 7 11/tr/tr Castelli.ia sessiliflora - - - - ~ “ - - 33/tr/ 2 _ _ 50/tr/ 2 8/tr/tr Cerastium arvense* 100/ 1/11 75/tr/ll 100/ 1/21 100/tr/24 20/tr/ 3 50/tr/10 50/tr/ I 66/tr/ 3 67/tr/ 3 Chrusopsis villosa - - - - - - - - - “ “ ~ 20/tr/ 2 — — — - - - — — - 4/tr/tr Cirsium undulatum - - - - - - - - - - - - 20/tr/ I “ - - - - - - - - 4/tr/tr Clematis pseudoalpina 25/ I/ 4 - - - - - - - - - - - - - — — - - - - “ - 4/tr/tr Comandra uhbellata* 75/ 2/16 50/tr/ll 25/tr/ 3 33/tr/ 2 80/ 2/15 50/ 1/11 100/ i/ 7 33/ I/ 9 55/ 1/10 *- ho T a b l e 2. (continued) ASSOCIATIONS Jumperus- Pirtus- Juniperus- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Rhus- Andropogon- Taxa Festuca Agropyron Festuaa Carex Calomovilfa Heliototrichon Stipa Festuca Totals Forbs: (continued) Crepis oeoidentalis Cryptanthe interrupta Delphinium bico lor Dodeaatheon eonjugens Eahinaceae pallida Erigeron eaespitosus Erigeron oehroleueusf Erigeron subtrinervis* * Erigeron spp. Eriogonum flavum Eriogonum umbellatum Eriogonum spp. Eritichum hovardii Erysium asperum 50/tr/ 5 25/tr/ 3 50/tr/10 25/tr/ 4 25/ 2/13 25/tr/ I 25/tr/ 2 75/tr/ 5 - - - 75/ 1/11 100/ 1/20 33/tr/ I 33/tr/ I 66/tr/10 25/tr/ 3 50/tr/ 4 25/tr/ 2 25/tr/ 3 Grindelia sguarrosa Guterrhiza sarothrae Haplopappus armeroides Haplopappus nuttali Haplopavpus spinulosus rum j 25/tr/ I 25/tr/ 2 25/tr/ 2 25/tr/ I 33/tr/15 20/tr/ 2 40/tr/ 2 20/tr/ I 20/tr/ 6 20/tr/ 3 50/ 3/ 5 100/ I/ 9 50/tr/ 2 Fvasera speciosa* 50/ I/ 6 25/tr/ I — — — 33/ I/ 9 — — - — — - Fritileria pudica — - - 25/tr/ I — — ~ — — — - ~ - - — - Gaillardia aristata* 50/ 1/12 25/tr/ 2 25/tr/ I 100/ 1/12 20/tr/ 2 50/tr/ 5 Galiwn boreale* 75/ 3/36 75/ 7/29 25/tr/ 2 66/ 1/13 ~ ~ 100/tr/ 9 Gewn tr if lorwn * 25/ I/ 4 25/ I/ 6 ~ - 33/tr/ I ~ ~ ~ 50/tr/ 4 50/tr/ 3 100/tr/ 3 50/ 1/12 50/tr/ 2 50/ I/ 7 50/tr/ 2 50/ I/ 5 50/tr/ 2 100/ 2/ 7 33/tr/ 4 33/tr/ I 33/tr/ I 33/tr/ I 100/tr/10 66/tr/ 4 4/tr/tr 4/tr/tr 8/tr/ I 8/tr/tr 8/tr/tr 4/tr/tr 29/tr/ 2 52/ I/ 9 11/tr/ I 11/tr/ I 4/tr/ I 22/tr/ 2 4/tr/ 2 4/tr/tr 18/tr/ 2 4/tr/tr 33/tr/ 4 55/ 1/14 25/tr/ 3 5/tr/tr 4/tr/tr 4/tr/tr 8/tr/ 2 4/tr/tr 18/tr/ 4 U> Table 2. (continued) ASSOCIATIONS JunipeitUS- Pinus- Juniperus- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Rhus- Andropogon- Taxa Festuca Agropyron Festuca Carex Calomovilfa Helictotrichon Stipa Festuca Totals Forbs: (continued) Hedysarwn sulfurescens* Leuarocrinum montanum 25/ 2/15 Lotus ourskianus Lupinus seriaeus Lupinus Dutheii Lupinus spp. Mertensia viridis Monarda fistulosa Musineon luteus 75/ 1/14 100/ 2/20 Linum verenne* 75/tr/ 2 75/tr/ 4 — ~ - Lithopermum ruderale — - — — — — 25/tr/ I Lomatium eous* 75/ 1/16 100/tr/12 75/tr/ 8 Lomatium marcoearpum — — — 25/tr/ I 25/tr/ 8 25/tr/ I 25/ I/ 5 25/ I/ 3 25/tr/ I 75/tr/ 6 25/tr/ 2 50/ 1/10 75/tr/ 3 Orthoearpus tenufclia - - 25/tr/ 2 Oxytropis besseyi - - - ~ ~ ~ Oxytropis seriaeus* 50/ 3/49 50/ 1/13 Oxytropis viscidia - - - 25/tr/ I Paroynahia sessiflora 25/tr/ 2 25/tr/ 4 Penstemon attenuates 25/ I/ 7 - - - Penstemon erianthus - - - 50/tr/ 2 Penstemon nitidus 66/ 1/11 33/ 2/ 3 33/tr/ 3 66/tr/ 3 33/ I/ I 33/tr/ 3 66/tr/ 7 33/ I/ 3 33/tr/ I 50/tr/ 4 100/ 1/48 - - - 33/tr/ I 20/tr/ I 40/tr/ I 60/tr/ 2 20/tr/ I 40/tr/ 3 40/ 1/16 50/ 1/18 50/tr/ 2 33/tr/ 3 8/tr/tr 37/tr/ 7 4/tr/tr 11/tr/ I 20/tr/ i 4 100/ I/ 8 - - - 47/tr/ 4 - - - - 33/tr/ I 11/tr/tr 5 100/tr/ 5 33/tr/ 2 67/tr/ 7 - - - - 33/tr/ I 11/tr/ I - 100/ 3/25 - - - 8/tr/ 3 ~ - - 33/tr/ 3 11/tr/ I - - - - 33/tr/ 2 15/tr/ I - - - - - - 8/tr/tr - - - - - - - 4/tr/tr - - - - - - - 4/tr/ I - 50/tr/ 3 66/ 1/15 37/tr/ 5 - - - - - - - 4/tr/tr - 50/tr/ I - - - 4/tr/tr - - - - - - - 4/tr/tr - - - - - - - 4/tr/tr - - - - - - - 8/tr/ 2 15 50/tr/ 3 33/tr/ 2 47/ 1/14 4 - ~ - - - - 8/tr/ 2 - - - - - - - 11/tr/ I - - - - - - - 4/tr/ I - - - - - - - 8/tr/tr - - - 4/tr/tr -O T able 2. (continued) ASSOCIATIONS Juniperus- Pinus- Juniperus- Potentilla- Juniperus- Juniperus- Potentilla- Juniperus- Taxa___________________Festuca________Agropuron_____Festuca______ Carex_________ Calomovilfa Forbs: (continued) Petaloatemon candidum — — - — — - - - - - - - - - Petaloatemon purpescena - - - - “ - 25/tr/ 2 - - * - - Phlox alyssifolia* - - - 100/ 4/26 - - " 100/ 5/32 20/tr/ Phlox hoodii* — — - 50/tr/ 4 - - - 33/tr/ 4 ~ - Phlox kelaeyi* 50/ 2/23 50/tr/ 6 100/ 3/26 33/ 6/26 40/tr/ Phlox multiflora* 25/tr/ I 75/ 1/14 75/ I/ 9 33/tr/ 4 Po In Q ala alba ~ - - - - - - - - - 20/tr/ Polygonum bistordis 50/ 1/14 - - - - - - - - - Potentilla gracilis 25/tr/ I - - - - - - - - - - - Potentilla hippana** ~ - - 25/tr/ I 75/tr/ 3 33/tr/ I - - Ranunculus glaberrimus ~ - - “ — - - - - - - 20/tr/ Sax-I fraaa rhomboidea 25/tr/ I - - - - - - - - Sisyrinchium sarmentosum - - - - - - - - - 33/tr/ 2 “ - 3 3 I I Sedum lanceolatum* 75/ 1/8 Senecio canus* 25/tr/ 2 Smilieia racemose Solidago oacidentalis 25/tr/ 2 - - - — — - 25/tr/ I 25/tr/ 2 33/tr/ 2 - - - - - - 33/tr/ 8 25/tr/ 2 - - - - - — — - — — — 20/tr/ I 20/tr/ I SiIISI - - - - * - - - - 33/tr/ 4 Thermopsis rhombifolia** 25/tr/ 3 25/ I/ 9 25/tr/ I 33/tr/ I Tragopogon dubius ~ - 25/tr/ I - - * - - - Vicia amerieana* 50/tr/ 2 75/tr/ 3 25/tr/ 2 66/tr/ 2 Viola nuttalli - - - - - - “ “ ~ - - - Yucca glauca** ~ - - - ~ ~ - - - Zyqodenus eleqans** 100/ 7/47 - - - - - - 100/ 1/13 Zyqodenus venosus* — — — 50/tr/ 4 50/tr/ 6 100/ 1/20 Total Forbs: 100/21/96 100/17/92 100/11/90 100/21/93 100/ 6/29 40/tr/ 5 20/tr/ I 100/13/76 Juniperus Juniperus- Juniperus- Rhus- Andropogon- Helietotrichon Stipa________ Festuea_______ Totals 50/ 1/14 100/ 5/43 100/ 2/20 50/tr/10 100/tr/ 2 100/tr/ 7 50/tr/ 2 50/tr/ 2 100/15/91 100/tr/ 5 100/ 3/45 50/tr/ 2 50/ 1/14 50/tr/ 4 50/ 2/11 100/ 2/13 50/ I/ 5 50/ I/ 4 100/ I/ 5 100/21/69 33/tr/ 3 33/tr/ 2 100/ 1/14 33/tr/ I 66/tr/ 3 66/ 7/58 33/tr/ I 33/tr/ 2 33/tr/ I 66/ 1/15 100/15/86 8/tr/ 2 4/tr/tr 40/ I/ 9 11/tr/ I 67/ 2/19 29/tr/ 4 4/tr/tr 8/tr/ 2 33/tr/ 2 4/tr/tr 4/tr/tr 4/tr/tr 4/tr/tr 18/tr/ 2 22/tr/ 2 4/Cr/ I 25/tr/ 2 8/tr/ I 4/tr/tr 47/ 2/13 4/tr/tr 47/tr/ 3 8/tr/ I 4/tr/tr 29/ I/ 9 47/tr/ 6 100/16/89 Table 2. (continued) ASSOCIATIONS Juniperus- Pinus- Juniperus- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Potentilla- JuniperrUs- Juniperus- Rhus- Andropogon- Taxa Festuca Agropyron Festuoa Carex Calomovilfa Heliatotrichon Stipa Festuaa Totals Shrubs: Apoaynum aannabium* Arctostauyhlos uva-ursi* Artemisia longifolia Artemisia tridentata Atriplex canesaens Berberis repens Juniperus horizontalisA Potentilla frutiaosa* Prunus virginiana 25/ I/ 8 25/tr/ I 100/32/75 75/ 7/27 25/tr/ 3 100/30/74 100/26/57 100/35/56 66/ 2/17 20/tr/ 2 20/tr/ I 20/ I/ 5 100/30/45 50/tr/ 3 100/27/47 100/ 4/28 100/35/50 33/tr/ I 100/47/61 66/ I/ 7 8/tr/ 2 4/tr/tr 4/tr/tr 4/tr/ I 4/tr/tr 4/tr/tr 100/32/57 29/ 2/ 9 4/tr/tr Rhus IrilobataitA Ribes spp. Rosa aciaularis* Rosa arhansana* Symphoriaarpos a IbusAA Symphoriearpos occidentalis* 25/tr/ 2 50/tr/ 2 50/ I/ 8 25/ I/ 7 25/tr/ 3 66/tr/ 2 20/tr/ I 20/tr/ 3 40/ I/ 7 100/ 2/ 8 100/ 4/12 100/ 2/20 50/ 7/37 33/ 2/13 33/tr/ I 11/tr/ I 4/tr/tr 37/ I/ 6 15/tr/ 2 8/ I/ 4 4/tr/tr TOTAL SHRUB 100/39/82 100/31/86 100/30/73 100/27/52 100/31/61 100/30/72 100/35/59 100/50/80 100/35/74 TOTAL LICHEN 100/ 3/23 50/ 2/14 25/ 2/22 100/ 4/18 100/ I/ 8 100/ 2/17 100/tr/ 2 100/tr/ I 82/ 2/15 TOTAL Sede* 13 100/ 5/29 25/tr/ I 75/11/29 66/ 6/26 100/tr/tr 100/ 7/25 100/tr/tr 100/ 4/11 82/ 4/15 TOTAL BAREGROUND ** 100/10/81 100/16/90 100/ 9/79 100/13/44 100/31/84+ 100/13/83 100/34/81+ 100/ 6/52 100/16/76 TOTAL ROCK ** 100/ 9/49 100/18/69 100/22/88' 100/ 3/21 100/18/62 100/21/79 100/ 2/15 100/ 9/42 100/14/57 TOTAL LITTER 100/13/95 100/26/98 100/18/96 100/29/97 100/17/94 100/24/97 100/21/99 100/26/96 100/21/96 * - specie involved in statistical analysis. ** = statistical difference of canopy coverage and frequency between the 8 associations at P=.05. + = statistical difference of canopy coverage between the 8 associations at P=.01. -H- = statistical difference of canopy coverage and frequency between the 8 associations at P=.01. // = statistical difference of frequency between the 8 associations at P=.01. 1 constancy (percent occurrence among sites)/canopy coverage (percent of area covered)/ frequency (percent occurrency among plots). 2tr. = trace; canopy coverage less than 0.5 percent< 3Sede = seleginella densa. ON 47 coniferous litter together covered 21 percent of the area and except for grass cover, was the most frequently encountered ground cover. Graminoids covered 67 percent as much surface area as juniper with a mean 22 percent canopy coverage. Forb cover was locally high but overall averaged only 16 percent. Aside from creeping juniper, total shrub cover accounted for 3 percent of the total canopy coverage. In addition to the other live plants, club moss (Setag-LneZZa densa) and lichen represented 6 percent of the ,surface cover. Although club moss had an overall cover value of 2 percent, in one association, Pinus- JunipeTus-Festuoa3 the value was 11 percent as a result of a high canopy cover recorded at site 8. In this site, northwest of Choteau, club moss occurred in 93 percent of the plots and covered 42 percent of the area. Cover classes and the frequency of occurrence of individual plant species varied within and between the eight associations (Table 2). As noted previously, bare ground and stone averaged 30 percent of the sur­ face covered. Values ranged from 15 percent in the Junipevus- Andvopogon-Festuoa association to 49 percent in the Junipevus- CaZamoviZfa association. The low value in the former association was, a result of the high shrub canopy cover, whereas the high value in the latter association may have been a result of the relatively high surface erosion. The bare ground cover was significantly higher (P=-OS) in the Junipevus-CaZamoviZfa (31%) and the Junipevus-Rhus-Stipa, (34%) 48 associations than in the other associations. Litter cover ranged from 13 percent in the Junipevus-Potentilla- Festuea association to 29 percent in the Junipevus-Potentilla-Cavex association with an average of 21 percent. No significant difference in the canopy coverage or the frequency of occurrence of litter between the eight associations was noted. Of. the 24 species of graminoids encountered, 8 had an overall canopy coverage of more than I percent (Table 2), The most important grass was Idaho fescue occurring in 74 percent of the sites; canopy coverage (C) 5 percent, frequency (F) 30 percent. Its importance ranged from its low in the Junipevus-Calamovilfa association (C < 0.5%, F = 28%)to its highest values in the Junipevus-Potentilla-Festuea association (C = 8%, F = 64%). Threadleaf sedge was second in overall importance (C = 4%, F = 22%), occurring'at 74 percent of the sites I Its importance ranged from low values in the Junipevus-Potentilla- Cavex association (C < 0.5%, F = 1%) to highest in the Junipevus- Helietotviehon (C = 6%, F = 30%) and Junipevus-Agvopyvon (C = 5%, F = 31%) associations. Other graminoids of importance included blue-, bunch wheatgrass, inflated sedge, rough fescue, spike.oat, needle and thread grass and prairie junegrass. Most associations contained several grasses in approximately equal proportions with no single species dominating the others. How­ ever, four exceptions were noted. Needle and thread grass was the most v, 49 important grass within the Junipevus-Rhus-Stipa association (C = 18%, F = 88%), with a significantly higher cover and more frequent occur­ rence than in any other association (P=.05). The occurrence of this species may be inversely related to that of Idaho fescue as these species did not occur in the same locations. Spike oat was signifi­ cantly more frequent and had a higher canopy coverage in the Junipevus- Helietotviahon and Junipevus-Potentilla-Cavex association than in the other six associations (P=.05). Blue grama (Bouteloua gvaeilis) and smooth brome (Bvomus inevmis) occured more frequently with a higher canopy cover in the Junipevus-Rhus-Stipa and Junipevus-Potentilla- Cavex associations respectively (P=.05). Total grass occurrence and canopy coverage in the Junipevus- Calamovilfa association (C = 20%, F = 85%) was significantly less than the other associations (P=.05). The three sites, which did not cluster in the ordination (4, 21, and 23), contained total grass cover similar to the eight associations with the exception of No. 23 (C = 32%, F = 100%). The higher grass cover may have been a result of no grazing by livestock for at least 20 years. ■ - The number of forb species recorded per site varied from I to 34 (Table 2). Among forbs, Kelsey phlox, fringed sagewort and prairie thermopsis were most important, (C = 2 , 2, 2%, and F = 19, 15, 13% respectively). Fringed sagewort was the most widespread forb occurring 50 in 78 percent of the sites. Other important forbs included common yarrow, pale bastard toad flax, three-veined fleabane (Eri-geron subtrinervis), kittentail and alyssum-leafed phlox. Although contrib­ uting little to the total canopy cover, mountain lomatium (Lomatium oous) and field chick weed (Cevastium avvense) were widespread, occurring in 67 percent of the sites. Forb cover appealed to be unevenly distributed within each associa­ tion. Several dominant forbs provided most of the cover, while the remainder was comprised of around 30 species of low individual impor­ tance. Additionally, the importance of several species was variable between the associations. The cover and frequency of pasqueflower varied significantly (P=.05) among the eight associations; it occurred mainly in the Jun-Lpevus-Potentilta-Festuaa (C = 5%, F = 41%) and Junipevus-Andvopogon-Festuaa (C = 2%, F = 90%) associations. Death camas and Arizona hymenoxys were most important in the Junipevus- Potentilla-Festuca (C = 7%, F = 47%) and Pinus-Junipevus-Festuaa (C = 2%, F T= 20%) associations respectively (P=.05). Prairie thermopsis, Misscmrigpldenrod (Solidago missouvensis), horse cinquefoil (Potentilla hippiana) and three-veined fleabane were other forbs whose importance varied statistically among the eight associations (P=.05). No differ­ ence in general forb cover between the eight associations and the three unclustered sites (4, 21 and 23) were noted. Site 21 had a relatively high forb complex similar to the Junipevus-Potentilla-Festuaa association 51 Total shrub importance ranged from a low in the Jun-Lpevus- Potent-Llla-Cavex association (C = 27%, F = 50%) to a high in the Ptnus-Jun-Lpevus-Festuoa association (C = 50%, F = 80%). Overall shrub canopy coverage and frequency of occurrence did not vary signif­ icantly among the eight associations. However, three shrubs, common snowberry, shrubby cinquefoil and skunkbush sumac individually showed a variation between associations (P=.05). Both snowberry (C = 7%, F = 37%) and skunkbush sumac (C = 4%, F = 12%) were statistically more important in the Juntpevus-Ehus-St-Lpa association than in the other seven associations (P=.05). Shrubby cinquefoil varied signif­ icantly between the eight associations occurring most abundantly in the Juntpevus-Potenttlla-Festuca (C = 7%, F = 27%), Juntpevus- Heltototvtohon (C = 4%, F = 28%) and Juntpevus-Potenttlla-Cavex associations (P=.05). Creeping juniper was by far the most important shrub as well as the most important species (C = 32%, F = 57%). Coverage estimates ranged from 26 percent in the Pinus-Juntpevus-Festuea association to 47 percent in the Junipevus-Andvopogon-Festuoa association. At site 27 within the latter association, creeping juniper covered 83 percent of the area demonstrating the high degree of community dominance attainable by this shrub. A common associate and the second most important shrub was shrubby cinquefoil (C =2%, F = 9%) occurring at 29 percent of the sites and 50 percent of the associations. Other 52 important shrubs included prickly rose (C = 1%, F = 6%) and common snowberry (C = 1%, F = 4%). Shrubs encountered but not sampled were Rocky Mountain juniper,. silver buffalo berry (Shepherdia argentea) and common juniper. Of the three unclustered sites, No. 21 was the only one to show any peculiar shrub characteristics. The total shrub cover was near 64 percent, mainly as a result of creeping juniper (C = 63%, F = 80%). Both canopy coverage and frequency of creeping juniper were nega­ tively and highly correlated with the southern azimuths (C, r = -.53; F, r =-.59). Generally those associations with the most creeping juniper had a north to northwest exposure. The slope gradient had little direct effect on the presence of juniper. It occurred on slopes ranging from 5 to 42 percent, although 67 percent of the sites were on slopes greater than 20 percent. In areas where slope was apparently a major factor as to the establishment of vegetation and/or providing a mesic environment, it may have indirectly limited the occurrence of creeping juniper. Although possibly not a major influence, elevation was negatively correlated with canopy coverage of juniper (r = -.37). Maximum cover values were attained in areas between 1300 meters (4000 ft) and 1500 meters (5000 ft). In areas less than 1300 meters or greater than 1500 meters percent of ground covered was considerably Typically, creeping juniper sites lacked a definite tree overstory. less. 53 Only limber pine, ponderosa pine and Douglas fir were identified as being associates of creeping juniper within the eight associations (Table 3). Although deciduous trees were encountered within several hundred meters of several study sites, they invaribly were associated with a different community. At one supplemental location (Site 24) high in the mountains south of Big Timber, whitebark pine (Pinus albicaulis) and lodgepole pine (Pinus oontorta) were also encountered. Total tree densities ranged from zero in the Juniperus-Rhus-Stipa association to 159 plants/ha in the Juniperus-Helietotriehon associa­ tion with canopy coverage estimates varying from less than 0.5 percent in the Juniperus-Ehus-Stipa and Juniperus-Agropyron associations to 24 percent in the Pinus-Juniperus-Festuoa association. Limber pine, the major tree associate, occurred at 48% of the study sites with an overall mean canopy coverage of 4% and density of 43 plants/ha. This species was also the most important tree among the eight associations, ranging from less than 0.5% coverage in the Juniperus-Agropyron and Juniperus-Rhus-Stipa associations to 18 percent in the Pinus-Juniperus-Festuoa association and, from 0 to 158 plants/ha. Although the canopy coverage of trees was higher in the Pinus-Juniperus- Festuoa association than in the Juniperus-Eeliototriohon association, the tree density was about one half. This may indicate the difference in age structure of limber pines between, these two associations. The other two tree species were ponderosa pine, which occurred only in the 54 Table 3. Constancy, canopy coverage, and density of three trees on each of the eight creeping juniper associations. Associations Pinus flexilus TAXA Pinus pondevosa Pseudotsuga menziesii Total Trees/ha Total Tree Cover (percent) Junvpevus- Totent-IVla- Festuea 25/ I/ 141 25/tr/ I. 15 I Jun-Lp evus- Agvopyvon 50/tr/ 9 — — — — — — 9 tr2 Pinus Junipevus. Festuea • 75/18/ 73 25/ 6/12 85 24 Junipevus PotentilLa Cavex 66/ 3/ 34 34 3 Junipevus ■ Cdlamovilfa 20/ I/ 41 20/20/32 — — — 71 21 Junipevus Helieto- tviehon 100/ 4/158 50/tr/ I 159 5 Junipevus- Rhus- Stipa Junipevus- Andvopogon- Festuea 66/ I/ 13 — — — 33/tr/ 2 15 I TOTAL 48/ 4/ 43 4/tr/ 4 15/ I/ 2 49 • 5 1Constancy (percent occurrence among sites)/percent canopy coverage/ density per Ha. ■ 2Tr. = canopy coverage less than 0.5%. r 55 Junipevus-Catamovi-Zfa association in central and eastern Montana, and Douglas fir, which occurred in several associations at low densities. . Edaphic Characteristics and Relationships Soils associated with creeping juniper stands appeared to be poorly developed, and were often characterized by thin topsoils, large quantities of stone, and exposed rock parent material. Also, because of their moderately steep slopes, relative openness, and sparse vege­ tation, some degree of erosion was typical on most sites. Analyses of soil samples from study sites (Table 4, Appendix Table 17) further indicated that soils tended to be clay loam in texture, slightly basic, low to medium in organic matter content, very low in phosphorus, medium to high in potassium, and low in salt. Average apparent soil depth (to parent material) was approximately 16 cm, with a range from 9 cm to 33 cm for 51 sites (Appendix Table 17). A majority (73%) of the sites had soil less than 20 cm deep. Soil depth differed significantly between the eight associations (P=.05). Deepest soils (ave. 25.3 cm) supported the Juniperus-Rhus-Stipa associ­ ation. Among individual plant species tested, soil depth was positively correlated with frequency of occurrence and coverage of purple reedgrass (Calamovilfa purpescens) (r = .52 and r = .52,' respectively), and Hood's phlox (Phlox hoodii) (r = .54 and r = .53) and the frequency of pasque flower (Anemone patens) (r = .51). A negative relationship between soil depth and coverage of limber pine (r = -.54) was noted. Table 4. Edaphic characteristics of creeping juniper associations including pH, organic matter content, texture, salt hazard, soil depth, erosion, and five important elements. Associations pH Organic Matter Percent Tex- ture^ Phos­ phorus 2ppm Potas­ sium ppm Magne­ sium rX meq- Sod­ ium meq Cal­ cium meq Salt Hazard mmhos^ Soil Depth cm Surface Erosion Percent Juniipevus- PotentiHa- Festuaa 7.3 6.5(H) SiL 23(VL) 373(H) 2.5 0.05 30.0 0.95 12.9 3.0 Juniperus- Agropyron 7.4 4.2 (M) L ' 19(VL) 198 (M) 1.7* 0.03 22.6 1.2 13.7 12.0 Pinus- Juniperus- Festuca 7.i 4.9 (M) CL 17(VL) 251(H) 1.8 0.05 24.6 0.83 10.8 12.0 Juniperus- Potentilla- Carex 7.4 6.2(H) SiL 14(VL) 375(H) 3.6 0.13 29.6 1.4 17.4 3.0 Juniperus- Calamovilfa 6.8 2.7(L) CL 23-(VL) 230 (M) 3.1 0.13 28.0 0.58 21.1 43. O+ Juniperus- 7.0 Heliatotriahon 4.I(M) SCL 8 (VL) 262(H) 4.4 0.10 22.2 0.45 15.6 9.0 JuniperuS- Rhus- Stipa ■7.5 2.8(L) C 17(VL) 246(M) 2.4 0.10 70. O+ 3.4** 25.3* 15.0 Table 4 (Continued). Associations pH Organic Matter Percent Tex­ ture^ " Phos­ phorus 2ppmr Potas­ sium ppm Magne­ sium meq Sod­ ium meq Cal­ cium meq Salt Hazard mmhos^ Soil Depth cm Surface Erosion Percent Juniipevus- Andropdgon- Festuca 7.5 5.2 (M) CL 12(VL) 344(H) 5.2 0.03 27.5 0.87 16.8 3.0 • Average 7.2 4.6 (M) CL 18(VL) 280(H) 2.9 0.08 29.5 1.1 15.9 12.5 Site 4^ Site 21 Site 23. 7.6 7.3 8.0 2.7(L) 6.8(H) 3.8(L) LS L • CL 18(VL) 21(VL) 9 (VL) 71(VL) 399(H) 312(H) 1.2 3.0 1.9 Tr.5 Tr. Tr. 11.2 34.0 21.0 0.7 1.7 0.9 17.9 17.5 20.8 15.0 3.0 15.0 SiL=Siltloam; L=Loam; CL=Clayloam; SCL=Sandyclay loam; C=Clay; LS=Loamy Sand. 2 ppm=pounds/one million pounds of soil. ^TTieq=Millequivalents/100 grams of soil. 4 mmho s =miIlimho s Imho=1/Ohm. ^Ratings from Montana Soils Testing Laboratory Report, ST-Form 2; VL=Very Low; L=Low; M= Medium; H=High. Tr.=Trace; value less than 0.1%. 6 . Individual sites; not clustered. * and + mean significant difference at P=.05 and .001, respectively. **=Slightly salty. 58 The proportion of soil surface eroded averaged 15 percent, but var­ ied from 3 percent to over 60 percent at five sites. Seven sites (14%) showed 38 percent surface erosion, while the remaining 39 were judged to be 15 percent or less eroded. The eight associations differed significantly in surface erosion (P=.001). The JunrLpepus-CaUmovtLtfa association occurred on more heavily eroded sites (ave. 43%). The other associations averaged only eight percent erosion. The frequency, and/or coverage of four plant species appeared to be negatively correlated with soil erosion. These included frequency and coverage of rough fescue (r = -.54 and -.57, respectively), field chickweed (r - -.57 and r = -.66) and flowery phlox (r = -.64 and r = -.51), and frequency of Idaho fescue (r = -.49). Creeping juniper with its ability to root along its branches and form large mats may be able to tolerate the high soil surface erosion. Clay loams, loams, silt loams, clays, sandy clay loams, sandy loams, sandy clays, silt clays and loamy sands comprised 31, 18, 14, 12, 10, 8, 4, and 2 percent of the sites respectively. The average of all sites was a clay loam (35% silt, 34% clay and 35% sand). The finest textured of these soils may impose difficulty for some plants to become established (Russell 1973); creeping juniper with its shallow root system has apparently been able to cope with the soils. Soil pH values ranged from strongly acid (4,4) to moderately alkaline (8.5) among the 51 sites. The overall average was 7.39 or a 59 slightly alkaline condition. A pH value such as this provides an environment in which most of the common elements are available. The eight associations were not significantly different, with all ranging near 7.2. The Jurvtperus-Catamov-ilfa association was an exception with a pH of 6.8. Positive correlations between soil pH and the frequency and coverage of western snowberry (Symphorioarpos oac-identalis) (r = .59), frequency of flowery phlox (r = .49) and the coverage of field chickweed (r = .51) were noted. A single negative correlation between pH and the frequency and coverage of prairie rose (r = -.62 and -.59, respectively) was also noted. The organic matter content of a soil influences such physical and chemical properties as cation-exchange capacity, soil stability and nutrient supply and availability (Buckman and Brady 1969). Twelve sites (24%) were considered very low, 15 (29%) low, 10 (20%) moderage and 14 • (27%) high. Average overall organic matter content was 4.1 percent (high) with a range from 0.8 to 7.4 percent. The areas of highest organic matter content were those having a relatively large percent of. the ground covered by forbs and grasses. The converse was true at sites with low soil organic matter. The organic matter content in the eight associations, which ranged from 2.7 percent (low) in the Juniperus- Calamovilfa association to 6.5 percent (high) in the Juniperus- Potentilla-Festuaa association, did not differ significantly. Only two associations were rated low, the others were, either medium (4) or I 60 high (2). Plant species positively correlated with organic matter content included frequency and cover of Hood's phlox (r = .59 and r = .74, respectively), and cover of Idaho fescue (r = .61) and textile onion (Attium textile)- (r = .63). Frequency of flowery phlox (r = -.51) and coverage of Arizona hymenoxys (r = -.53) were negatively correlated with organic matter. Phosphorus is a critical element in the growth of plants influ­ encing the processes of reproduction, development and the intake of r the other important elements (Buckman and Brady 1969). Three, sites (6%) were rated low (45.0, 49.0 and 43.0 ppm), I (2%) was high (103.0 ppm) and the rest, 47 (92%), were very low. (less than 30.0 ppm) in this element. .The site near Lincoln, west of the Continental Divide, was found to be very high in comparison to all others, with a value of 103.0 ppm. The mean value for all sites was 15.6 ppm (very low); ex­ cluding the site near Lincoln, it drops to 13.5 ppm. In either case, the Lincoln value was approximately seven times greater than the average for the other sites. A combination of precipitation, location under conifers and relatively low levels of calcium may have attributed to this condition (Buckman and Brady 1969). Significant differences were not detected between the eight associations where the average was 16.6 ppm and the range was from 8 ppm (very low) in the Junipevus- Helictotvichon association to 23 ppm (very low) in the Junipevus- Calamovilfa and Junipevus-Potentilla-Festuca associations. Positive 61 correlations between phosphorus and plant parameters included coverage of western wheatgrass (r = .63) and field chickweed (r = .62), fre­ quency of flowery phlox (r = .53) and frequency and cover of arrow- leaf balsamroot (Balsamorhiza sagg-ltata) (r = .64 and .66, respectively). Potassium is an element essential to the photosynthetic process and transportation of its products (Russell and Russelll974). Additionally, it is important in the synthesis' of amino acids and proteins. Twenty- one sites (41%) were found to have high levels, 24 (47%) medium, 4 (8%) low and 2 (4%) very low. The content of samples varied from 69.0 ppm (very low) to 580.0 ppm (high), with a mean of 260.1 ppm (high). The eight associations were not statistically different at the .05 level. Potas­ sium content varied from 198 ppm (medium) in the Juniperus-Agvopyron association to 375 ppm (high) in the Juniperus-Potentitla-Festuoa asso­ ciation with a mean of 284.9 ppm (high). This element was positively . correlated with more plant species than any other soil factor. Positive correlations between potassium and plant occurrence included frequency and cover of mountain lomatium (r = .53 and .51, respectively), Kelsey phlox (r = .51 and .60, respectively), pasque flower (r = .50 and .51), blue flax (Linum perenne) (r = .57 and .55)., common yarrow (r = .73 and .55), common dandelion (Taraxicum officinale) (r = .59 and .59), and blanket flower (Gaillardia aristatd)' (r = .55 and .49), frequency of northern bedstraw (r = .51), and coverage of meadow death camas (r = .55) and spotted frasera (Frasera speoiosd) (r .= .67). Frequency and r 62 coverage of inflated sedge (r = -.57 and -.58) and three-veined fleabane (r = -.51 and -.51), and frequency of flowery phlox (f = -.55) were negatively correlated with this element. j ; Calcium is essential for growth of meristems and particularly for the proper growth and function of root tips ("Russell and Russell 1974). Calcium contents of samples from the 51 sites varied between 9.6 and 98.0 meq/ 100 grams of soil with a mean of 29.3 meq/100 grams of soil. The eight associations ranged from 22.2 meq for the Jun-Lipevus-Hel-Ietotr-Lchon association to 70.0 meq in the Jun-Lpevus-Rhus-St-Lpa association with an average of 31.9 meq. One association, Jun-Lpevus-Rhus-St-Lpa, differed significantly (P=.001) from the other seven. This association had a calcium level of approximately 2^ times the others and may have account­ ed for the relatively high average height of creeping juniper (ave. = 9.6 cm). Positive correlations of plant performance with calcium levels included cover of inflated sedge (r = .53) and frequency of pale bastard toad flax (r = .54). Frequency and cover of flowery phlox (r = -.66 and -.65, respectively) and frequency of mountain lomatium (r = -.53) were negatively correlated with calcium content. Magnesium is a necessary element as a constituent of chlorophyll (Russell and Russell 1974) , Overall, the element ranged from 0.7 to 7.3 meq with a mean of 2,8 meq. Variation within the eight associations was from 1.7 meq in the Jun-Lpevus-Agvopyvon association to 5.2 meq in the Jun-Lpevus- Andvdpogon-Festuea association. The average content was 30.9 meq. The 63 Junipevus-AgropyTon association soil possessed significantly lower levels of magnesium (1.7 meq) from the other seven associations. Plant species positively correlated with this element comprised frequency and cover of inflated sedge (r = .49 and .54, respectively),, kinnikinik {Avotostayphtos uva-uvsi) (r = .58 and .58), Japanese brome {Bvomus japonious) (r = .58. and .58), pale bastard toadflax (r = .61 and .54), and frequency of leafy musineon Qiusineon divavieatum) (r = .49). Nega­ tive correlations included frequency and cover of mountain lomatium (r = -.77 and -.53, respectively), flowery phlox (r = -.56 and -.54), and cover of Arizona hymenoxys (r = -.54). Sodium levels at all sites were less than 0.40 meq with the average at less than 0.1 meq. Within the eight associations it ranged from 0.03 to 0.13 meq and an average of 0.08. No significant difference was noted. A majority of the correlations with plant species were negative. These included frequency and cover of common yarrow (r = -.68 and -.54, respectively), common dandelion (r = -.51 and -.51), cover of spotted frasera (r = -.63), and frequency of blue flax (r = -.49). The only positive correlation was frequency of western wheatgrass (r = .65). Generally the salt concentration at all sites was low enough to have little effect on vegetation. Only two sites (Nos. 14 and 32) gave any . indication of saltiness. Only the Junipevus-Ehus-Stipa association showed signs of saltiness and this was due to site 14. Frequency and cover of blue grama (r = .75 and r = .75), soapwell (Yucca glauca) 64 (r = .75 and .75), flowery phlox (r = .65 and .68) and cover of skunk- bush sumac (r = .50) were positively correlated with salt hazard. Negative relationships included frequency and cover of prairie rose (r = -.53 and -.50) and inflated sedge (r = -.54). No significant difference was noted in. comparing six soil factors of samples taken from under creeping juniper shrubs with those between shrubs (Table 5). This appears to indicate that creeping juniper shrubs do not modify the soil in their immediate vicinity. A slightly higher organic matter content was noted under the plants, but not enough to be of any importance. , Table 5. Comparison of six soil factors from samples taken under and between creeping juniper shrubs. Location pH Phosphorus ppm Potassium ppm Salt Hazard mmhos Sodium meq/lOOgm Organic Matter percent under 7.18 21.1 380.1 1.2 .09 5.8. between 7.20 . 19.1 351.3 1.1 .06 . 5.0 t-value1 .06 .43 .51 .64 .75 1.3 1C-Value from table in Dixon and Massey, 1969, is 2.1 for P=.05. The three sites (4, 21 and 23) which did not "fit" into any associ­ ation had soil characteristics generally comparable to those of the eight associations with the following exceptions; site 4 had lower levels of magnesium, calcium and potassium possibly contributing to 65 poor plant conditions; and site 21 had relatively high organic matter probably caused by high creeping juniper ground cover (63%). Growth Characteristics and Relationships Growth Form ' Creeping juniper typically occurred as a very low shrub, in dense mats less than one dm tall, growing horizontally or "creeping" along the ground. Occasionally, taller or small, bush-shaped plants occurred where browsing on peripheral branches reduced spreading or where disease caused abnormal growth. In some areas or stands, with histories of heavy use by deer, mats were often broken up, and individual plants were impossible to differentiate. Mats consisted of a central section and peripheral, spreading branches or leaders on which most of the plant growth occurs. Little if any net growth appeared to occur within the central mat. Excavations showed that this central portion has a major root system consisting of several major and numerous minor roots. As the shrub enlarges, roots sprout along the peripheral branches anchoring them to the ground. Rooting generally started 10-20 cm from the tips of leaders. Depths of major roots of excavated plants ranged from 13.0 to 35.0 cm, with a mean of 22.4 ± 1.3 cm. On relatively flat sites, plants appeared to grow laterally in all directions; on steeper slopes, lateral enlargement occurred mainly on the downhill side of the plant. As a rule, growing leaders adhered to 66 the ground, only rarely extending their tips off the ground, such that the low plant profile was maintained. Annual growth occurred both laterally and terminally on indivi­ dual branches or leaders less than 4-5 years old; and most new twigs and leaves originated from one and two-year-old stems. However,, some new leaves and twigs were also noted to occur on older portions of the plant. Current annual growth of creeping juniper is essentially impossible to differentiate from previous years' growth by visual examination; though close inspection of known-age materials showed several possible distinguishing features. Current year's growth on twigs was bright green, with short, awl-like leaves closely appressed to the stem, and with very short interaxial leaf spaces. Second year growth generally also had bright green leaves, but they were less appressed to the stem and interspaces were approximately 5-10 times longer than on new growth. After the first two years, the flaring of leaves and interspaces remained uniform and similar to the second year growth, though the leaves turned a reddish brown and appeared to be dying. When annual growth was nearly complete, both staminate and pistil­ late "flowers" developed from the tips of lateral and terminal twigs located both within the plant and peripherally. The flowers occurred on both first and second year twigs. The height of creeping juniper plants varied significantly (P=.005) 67 between sites occupied by the eight associations in which- this species occurred (Table 6). Mean plant heights ranged from 4.8 cm in the Junvperus-Potentilla-Festuoa association to 9.6 cm in the Juniperus- Rhus-Stipa association. The overall mean plant height was 6.4 cm for all associations. Plants were significantly taller (P=-OS) in the Juniperus Calamovilfa (8.5 cm) and Juniperus-Rhus-Stipa associations than in other associations, while plants in the Juniperus-Potentilla- Festuca association were significantly shorter (P=.OS) than in the others. The two associations having the tallest juniper plants occupied sites which were fairly dry, at relatively low elevation, and had relatively low grass and forb coverage; whereas the sites occupied by Juniperus-Potentilla-Festuoa were quite mesic, at high elevations, and had high grass-forb coverage. Heights of creeping juniper plants appeared to be highly influenced by weather or climatic factors, little influenced by topo-geographic factors, and not influenced by soil factors. Among weather, factors, maximum and average annual temperatures for 1976, and mean annual precipitation were postively correlated with height (r = .55, .74, .51, respectively). Minimum 1976 temperatures and normal annual temperatures were negatively correlated (r = -.57 and -.63, respectively). Overall, weather factors were highly correlated (r = .76) with plant heights. Although the effects of individual weather factors were difficult to separate within this correlation, it generally appeared that 68 Table 6. Height, decadence, biomass, and lateral and terminal growth of shrubs in the eight creeping juniper associations east of the Continental Divide. Associations Height (cm) Decadence (percent) Biomass Mean Crown Standing Area1 Crop2 (Ibs/f.t'2) (Ibs/acre) Lateral Growth (mm) 3 Terminal Growth (mm) Juniperus- Potentilla- Festuca 4.8* 3.3 .027 386.1 8.7 30.8 Juniperus- Agropyron 5.7 5,8 .032 416.0 7.2 30.7 Pinus- Juniperus- Fsstuca 5,9 3.3 .036 436.2 . 6.5 17.8 Juniperus- Potentilla- Carex 5.3 4.0 .026 417.9 12.4 69.6 Junipsrus- Calamovilfa 8.5* 1.4 .032 376.7 11.7 46.0 Junipsrus- Eslietotriehon 5.4 1.5 .031 354.9 6.2 21.3 Junipsrus- Rhus- Stipa 9.6* 5.0 .038 548.6 11.5 36.9 Juniperus- Andropogon- Festuea 6.2 8.3 .029 589.2 7.9 19.8 Total 6,5 3.1 .031 444.9 9.0 34.5 Site 23 6.6 2.0 .039 290.5 13.7 57.9 Site 21 . 9.9 4.0 .035 908.2 5.5 21.1 Site 4 4.3 3.0 .038 485.2 4.0 9.3 ^^statistical difference between associations at P=.05 ^pounds of creeping juniper per foot2 of creeping juniper crown area 2Pounds of creeping juniper per acre of total vegetation ^growth during 1977 growing season 69 temperature and precipitation had greatest influence on the height of creeping juniper plants throughout its range in Montana, overshadowing the effects of other factors such as length of the growing season which showed no significant correlation with height. Elevation was the only topo-geographic factor which appeared to be correlated with plant height (r *= -.65), with plants at lower elevations tending to have a higher mean stature. Among edaphic factors, neither the pH, organic matter content, various mineral contents, nor texture of soils at study sites were correlated with the height of juniper plants. The proportion of dead crown was relatively low among creeping juniper plants on all study areas. The mean ranged from 1.4 percent in the JunipeTus^Catamovitfa association to 8.3 percent in the Junipevus- Andvopogonr-Festuca association with no significant difference between any of the eight associations. The average,decadency for all plants and sites was 3.5 percent. This was much lower than reported for two other browse species in Montana, including skunkbush sumac -29 percent (Martin 19.71) and curl-leaf mountain mahogany -26 percent (Duncan 1975). Soil characteristics apparently had no significant influence on the.occur­ rence of dead crown; and elevation and slope had little effect. How­ ever, aspect, the north-south exposure of a site, was significantly negatively correlated with decadency (r = -.50). Those associations generally having a northern exposure, as Junipevus-Andvopogon-Festuaa, contained highest levels of dead crown while those typically found on 70 southerly exposures, as Jumipevus-CdtamovjIlfa had the least amount of dead crown. The high proportion of dead crown may also be a reflection of high amounts of deer utilization as noted by Kamps (1968) and in this study. Annual Chronology and Twig Growth A chronology of phonological events in the annual cycle of creeping juniper, based on observations and measurements during 1977, is presented in Figure 23. The annual cycle usually began in early April when, plants almost simultaneously began to "green-up" and initiate twig growth. Among individual sites, dates of "green-up" and initial twig growth ranged from early April to early May. The "green-up" or change in leaf color from a reddish-brown to a deep green coincided with the breaking of dormancy near the beginning of April and was completed in all plants by the first of May. Twig growth was first noticed about the second week in April on plants of sites in the vicinity of Lewistown and continued into Septem­ ber. Generally, growth of all terminal and lateral twigs was completed by early September; however, plants on several sites continued to show growth well into September. Observations at site 29, on the Sun River Game Range, west of Augusta, over a three-year period indicated that a small amount of growth occurred after mid-September and possibly into . early October. The period of most rapid growth was usually from the , first of June to the end of July, where plants at all sites put on Pistillate bud formation Pistillate berry formation Pistillate berry ripening “ Browness” “ Greeness" Twig growth C > < > > - Staminate cone formation Staminate cone ripening Pollen shedding O Staminate cone shedding I I I I I I I I I I I I I I I I I 15 30 15 30 15 30 15 30 15 30 15 30 15 30 15 30 April May June July August Sept Oct Nov Figure 23. Annual chronology of creeping juniper in stands east of the Continental Divide as determined by spring, summer and fall observations. 72 between 10 and 100 mm growth. Patterns of current annual growth of terminal and lateral twigs of creeping juniper plants in the eight associations plus the three stands which did not cluster are shown in Figures 24a through 24i. Seasonal growth, particularly of terminal twigs followed the "S"-shaped curve commonly found for most vegetative growth. Lateral and terminal twig growth differed considerably, as might be expected. Lateral twig growth was slow but consistent whereas the terminal twigs put on a relatively greater proportion of their growth between June and July. Overall, the ratio of terminal to lateral growth was consistent within each association; that is, relatively long termi­ nal growth on plants was accompanied by relatively long lateral growth and vice versa; however this ratio differed among plants in the different associations. Terminal and lateral twig growth varied between sites of the. same association as well as among those of different associations, though differences were not statistically significant (Table 6, Figs. 24a to 24i). For example, a comparison of the growth of plants among sites of the Junipevus-TPotent-LtZa-Capex association (Fig. 24d) shows that annual growth on plants at site 18 was much longer, averaging more than 130 irmi terminally and nearly 20 mm laterally, than that of plants at sites 11 and 12. Among sites of the Junipevus-CalconoviZfa association (Fig. 24e), plants at site 26 produced longer terminal twigs (ave. 60 mm) than the M IL LI M ET ER S . - - - - - - 2 2 - T - - - - - - • 2 2 - L APR I L M A Y JUNE J U L Y AUGUST SEPT Figure 24. Growth curves of lateral (L) and terminal (T) twigs on creeping juniper shrubs in eight juniper associations as determined by the mean length of 15 terminal and 60 lateral twigs per site in 1977. a. Juniperus- Potentilla-Festuea association— sites 19, 20, 22, 25. MI LL IM ET ER S 1 7 - T • 1 7 - L 3 0 15 A U G U S T Figure 24. Continued, b. Juniperus-Agropyron association— sites 2, 3, 4, 17. M IL LI M ET ER S 1 5 3 0 AUGUSTAPR I L Figure 24. Continued, c. Pinus-Juniperus-Festuca association— sites 6, 7, 8, 31. MI LL IM ET ER S 1 4 0 - i 120- 100- - • 1 8 - L 12 - T ----- 12- L AUGUST Figure 24. Continued, d. Juniperus-Potentilla-Carex association— sites 11, 12, 18. LI ME Tt RS - • 2 8 - 1 - --- . jg- j - - - - - - - - 15 -1 15 30 AUGUST Figure 24. Continued, e. Juniperus-CalamoviLfa association— sites 15, 16, 26, 28, 29. 20- APR I L MAY JUNE JULY AUGUST SEPT Figure 24. Continued, f. (top) Juniperus-Rhus-Stipa association— sites 13, 14 and 24 g. (bottom) Juniperus-Helictotrichon association— sites 10, 30. ± 20— A P R I L A U G U S T Figure 24. Continued, h. Juniperus-Andropogon-Festuaa association— sites I, 9, 27. 15 AUGUS TA P R I L Figure 24. Continued, i. Non-clustered sites— sites 4, 21, 23 81 other four sites, with site 15 a close second averaging 50 mm. Similarly, most other associations also included at least one site on which terminal twig growth was somewhat higher than at other sites. Overall, the longest twigs generally occurred in stands on open sites of northerly exposures and in areas of moderate precipitation (13-14 inches). However, there appeared to be little obvious correlation overall between twig growth and environmental factors. Among associations, plants of the Juwilpevus-PotentrLtla-CaTex association produced longer mean terminal and lateral twig growth than those of the other associations (Table 6). Stands of this association all occurred on northerly-facing slopes and in areas receiving rela­ tively high amounts of precipitation continually throughout the growing season (18-19 inches annually). Plants of the Ptnus-JunrLpevus-Festuoa association produced the shortest mean terminal and lateral twig growth. This may reflect suppression of growth under an overstory. The three sites (4, 21 and 23) which did not "fit" into any association possessed growth characteristics generally consistent with the eight associations (Fig. 24i). Reproductive Characteristics and Relationships Creeping juniper plants are dioecious, the unisexual flowers being borne on separate plants. In this study, the sex ratio was found to be 2:1 in favor of males. The staminate cones are cylindri­ cal, pale brown, 4-5 mm long and 2 mm in diameter. They are composed 82 of scales occurring in rows of 4-5 each having 4 scales (Stephens 1973). Scales are approximately circular and 1-2 mm in diameter, each with 4 yellow pollen sacs attached near the base (Stephens 1973). At maturity, the scales spread and the pollen is shed. Pistillate buds consist of 2-6 oval ovules at the tip of a short, curved twig or branchlet (peduncle) approximately 2-3 mm long. The ovules are modified scales which fuse and enlarge to later form the berries. When first formed, "berries" or "fruits" are green, turning chalky purple when maturing. The peduncles of each are curved, leafy and about 3-5 mm long. The berries are nearly round, 8-10 mm across, and contain an average of 4 seeds (Fassett 1945). The exterior coat of each berry contains large resinous glands. The seeds are oval to rounded, acute, slightly grooved and ridged, 4-5 mm long, 2-3 mm wide, and red-brown with a hard, bony exterior and a large white scar on the base. The "greening" of plants during the first two weeks of April coincided with initiation of maturation and opening of staminate cones. Pollen was shed two to three weeks later, with a peak during the last ten days of April and the first week of May. The staminate cones persisted for approximately two to three weeks after pollen was re­ leased before being shed. New staminate cones first appeared June 23 at site 2. Among individual plants, the new cones typically formed within about four weeks of the date on which old cones were shed. They 83 usually developed on twigs which had not formed cones during the previous year. Cone formation extended over a period of nearly three months after which cones became dormant until maturation the following spring. Berries appeared about one to two weeks after the pollen was shed, in mid to late June, on the ends of twigs. Berry formation was first noted at site No. 3 on June 25. The light green berries continued to be produced for approximately two months, until mid-July. They remained green for about two more months before turning a chalky, purple color in the process of ripening. By mid-November, nearly all berries had become purple in color. These berries apparently remained on the plant, in the process of maturation, similar to Rocky Mountain Juniper (U. S. Dept. Agric. 1948), for one to two years before dropping to the ground. Germination experiments conducted during this study indicated that berries (seeds) do not ripen until at least the second year after formation. About one month after berries of the. current year began to form, in mid to late June, the curved penunclate twigs characteristic of the new pistillate buds began to form. Observations indicated that creeping juniper rarely reproduces by seed. Rather, plants and/or populations appear to be maintained primarily by vegetative means spreading horizontally and sprouting roots along branches or leaders. Observations indicate that new plants apparently are then formed by the break-up of old shrubs. Among the 84 study sites, only six plants were found which were believed to have been less than 10 years old and originated from seeds. Lonner (1971) found 52 creeping juniper plants from various sites in eastern Montana to range in age from 20 to 140 years. The mean age of 56.7 ± 3.7 years ranked creeping juniper second in longevity among 11 browse species studied. Only one of the 52 shrubs was 20 years old or less and only six were less than 30 years of age, which would also indicate a lack of recent reproduction from seed. The low incidence of seedlings of any plant species may be attributed to a number of factors, including low seed production, maturation- and/or germination, high seedling mortality, vegetative reproduction instead of seed reproduction, or species longevity. The following discussion of reproduction characteristics of creeping juniper seems to suggest that several of these factors are involved with the lack of creeping juniper seedlings. Overall, the number of ripe berries per 100 twigs was 32.5 and 2 the number per .1 m plant crown area was 68.8 berries (Table 7). The number of ripe berries ranged from 5.5 per 100 twigs examined in the Pinus-Junrlpevus-Festuoa association to 101.4 per 100 twigs in the 2 Jun'ilperus-CaZamov-iZfa association, and from 37.2/.I m of juniper crown 2 area in the Juni-Tpevus-AgvoipyTon-Festuoa association to 107.4/.I m in the Junipevus-CaZamoviZfa association. These data are difficult to interpret in the absence of comparative studies for creeping juniper. T a b l e 7. R e p r o d u c t i v e c h a r a c t e r i s t i c s of c r e e p i n g j u n i p e r in M o n t a n a , a nd s t a t i s t i c a l c o m p a r i s o n s a m o n g p l ants of eight juni p e r ass o c i a t i o n s . _______________________________________________ ASSOCIATIONS_________________________________________________________ Juniperus- Finus- Juniperus- Juniperus- Juniperus Potentilla- Juniperus- Juniperus- Potentilla- Juniperus- Juniperus- Rhus- Andropogon- Characteristics Festuca Agropyron Festuca Carex Calamovilfa Helietotriehon Stipa Festuca Average % twigs with: buds 15 9 7 16 32* 40* 15 7 18 berries 15 7 4 10 25 18 15 4 12 ripe berries 14 6 4 6 24 18 11 4 12 new buds 21 16 2* 36 36 56* 9 8 23 male cones 52 49 74 77 49 70 78 51 60 Number per 100 twigs: buds 51.3 22.3 16.5 33.0 132.8 99.5 39.5 9.0 52.8 berries 32.2 16.6 6.1 20.3 111.6 33.5 38.5 4.0 36.8 ripe berries 29.5 14.1 5.5 19.9 101.4 32.5 20.5 3.7 32.5 new buds 81.3 73.5 2.5* 193.3 250.0 230.0 28.5 66.7 115.0 male cones 810.0 490.0 870.0 1150.0 490.0 910.0 880.0 520.0 750.0 Number per .1 m2 crown area buds 93.1 177.7 170.8 93.3 179.9 129.8 153.6 78.1 138.8 berries 63.3 112.0 62.8 73.7 118.9 70.2 97.4 46.2 83.0 ripe berries 49.1 89.3 51.7 64.0 107.4 55.9 72.6 37.2 68.8 new buds 88.8 64.7 51.2 38.2 85.8 56.4 66.6 80.1 68.5 male cones 1018.0 726.6 432.8 888.0 1862.0 939.2 520.0 498.1 929.5 % Success: buds forming 68/621 63/74 37/37 78/61 66/84 54/34 63/97 60/44 60/70 berries buds forming 77/91 mature berries 80/85 82/90 87/98 90/90 77/94 74/50 80/92 84/88 berries 53/57 50/63 30/34 68/60 60/76 42/32 47/50 48/41 49/62 maturing Sex Ratios 42/582 28/72 41/59 63/37 20/80 29/71 31/69 51/49 36/64 ^First number is success rate for ' per m" crown area" data; second number is success rate for "per twig" data. ^First number is * percent females; second number is percent male plants. = represents a significant difference between associations at P=.05. 86 however, they indicate that this level of berry (seed) production probably is sufficient to maintain the species under natural conditions. The percentage of twigs bearing new pistillate buds was signifi­ cantly higher (P=.05) in the Juntpevus-HetyLototv-Iahon association and significantly lower (P=.05) in the Pinus-Junipevus-Festuoa. association than in the other associations. The percentage of twigs holding old buds was significantly higher in the Junipevus-Hetiototvichon and Junipevus-Catcmovitfa associations. Numbers of old buds, berries, ripe berries and new buds per 100 twigs all differed significantly (P=.05) between the eight associations, with the Junipevus-Catamovitfa associations being highest in all categories, while the Junipevus- Andvopogon-Festuoa association was lowest in the first three categories and the Pinus-Junipevus-Festuoa association was lowest in the fourth. 2 The number of reproductive parts per .1 m crown area was not signifi­ cantly different between associations. The dioecious nature of creeping juniper may also contribute to the low incidence of seedlings. Twice as many male plants as female plants were recorded during this study. Only one plant in this study was noted as not bearing either staminate or pistillate flowers. Although the viability of seeds was not evaluated directly, only 9 of the approximately 9,500 new and two year old creeping juniper seeds tested in the laboratory germinated. None of the approximately 500 seeds of the current year germinated. Similar finds have been,reported 87 for juniper species in general (Heit 1966, 1967). Once pistillate buds form, berries also generally form and mature. Overall, 60 percent and 70 percent of the buds fruited (formed berries) in the interior (frame data) and along the periphery (twig data) of the shrubs, respectively (Table 7). About 84 and 88 percent of the buds fruited with the fruits continuing on to maturity within and at the edges of the plants, respectively; and 49 and 62 percent of the berries, once formed, matured within and at the edges, respectively. These data indicate generally higher success in fruiting and maturation along the edges of the plant, where a relatively greater amount of plant activity probably occurs, than within the interior of the plants. In contrast to vegetative growth, the reproductive performance of creeping juniper varied in relation to soil characteristics of study sites with less,influence from climatic and geographic factors. Normally, numbers of reproductive parts per 100 twigs on plants were negatively correlated with salt hazard (r = -.62) and positively correlated with calcium (r = .49) and sodium content (r =. .78) of the 2soil. Numbers per 0.1 m crown area showed positive correlations with phosphorus (r = .60) and sodium (r = .61) content. Percentages of individual twigs bearing reproductive parts were negatively correlated with organic matter content (r = -.68), potassium content (r = -.62) and salt hazard (r = -.63), and positively correlated with sodium (r = .57)-and calcium (r = .50) content of soils. Overall, salt hazard. sodium, calcium and phosphorus levels in the soil appeared to be most influential with respect to numbers of reproductive parts on plants. Success rates within and at the periphery of plants were . generally negatively correlated with pH of the soil. The associations with highest production estimates generally were those such as Juni-Ipevus-CatamovLtfa and Junipevus-Hetietotviohon .with soils higher in phosphorus than the other associations, but still very low, moderately high in calcium and sodium, and having a low salt hazard rating. Associations as Pinus-Junipevus-Festuea and Junipevus- Andvopogon-Festuea with low seed production had opposite soil factor values. Additionally, associations with slightly acid or alkaline soils seemed to have higher reproductive success than either of those in more acid or alkaline soils. Overall climatic factors examined by multiple regression appeared 2 to obviously influence only the number of reproductive parts per .1 m crown area; but single factors were not well correlated. Positive correlations were rioted among the combined influence of all weather factors (temperature, precipitation and frost-free periods) and number 2 2 of buds per .1 m crown area (r = .59), numbers of berries per .1 m crown area (r = .71) and numbers of ripe berries (r = .72). . Generally associations occurring on sites with higher annual maximum temperatures, and lower mean annual temperatures as the Junipevus-Catamovitfa3 Juni- pevus-Hetietotviehon and Junipevus-Rhus-Stipa association had plants with higher production. 88 Forage Characteristics and Utilization by Wildlife Nutritional Characteristics Average crude protein content of composite samples of creeping juniper leaves and twigs (Table 8), was 6.3 percent for summer (August 1976), 5.7 percent in autumn (November 1976), 5.7 percent in winter (February 1977), and 5.5 percent during spring (May 1977). Seasonal variations, from summer to autumn, autumn to winter and winter to spring, were not significant; while the difference between summer and spring was highly significant (P=.01). Table 8. Percentages crude protein of creeping juniper plants for ___________ 12 study sites during one year, 1976 to 1977.______________ 89 August 1977 Site August 1976 November 1976 February 1977 May 1977 (Ave.) Plant■ I Plant 2 Plant 3 4 6.5 5.2 4.9 5.3 (5.5) 6.5 4.8 5.2 7 5.5 5.0 5.2 4.7 (5.1) 5.1 5.8 4.4 " 11 7.6 7.4 6.7 5.9 (6.9) 5.9 5.9 6.1 13 7.2 6.6 6.1 6.2 (6.3) 6.0 6.5 ' 5.7 15 5.8 4.4 5.1 ■ 4.7 (5,0) 5.1 4.3 4.6 17 7.4 5.5 — 5.6 (6.0) 6.0 4.9 6.0 19 5.7 5.6 6.1 6.1 (5.9) 5.2 6.1 5.7 21 5.3 5.5 5.5 4.8 (5.3) 5.3 5.7 5.9 22 6.8 5.4 5.8 6.3 (6.1) 5.2 5.1 5.9 23 4.8 4.7 5.7 5.6 (5.2) 5.3 5.8 5.7 ■ 27 5.5 6.1 5.3 5.6 (5.6) 6.0 5,7 6.1 31 7.0 7.4 6.3 6.0 (6.7) 6.3 6.5 6.1 Average 6.3 . 5.7 5.7 5.5 (5.8) 5.7 5.6 5.6 90 The average protein'content of samples collected from one plant, at each of 12 study sites during August 1976 (6.3%) was higher than that of the same plants in August 1977 (5.6%), though the difference was not significant. Comparison of precipitation at the 12 study sites during spring (April, May and June), between 1976 and 1977, indicated that these differences may be due to moisture differences (Table 9). At nearly every site the precipitation was higher in 1976. than in 1977 and the protein levels varied similarly. Table 9. Comparison of spring (April, May, 12 creeping juniper sites between June) precipitation at 1976 and 1977. Spring 1976 Spring 1977 Precipitation Precipitation Sites (inches) (inches) 4 5.8 3.1 7 6.3 3.9 11 4.3 2.7 13 8.0 5.7 15 8.3 4.2 17 5.5 5.3 19 7.1 5.3 21 6.9 3.5 ■ 22 6.9 3.5 23 7.9 3.6 27 6.0 5.1 . 31 3.8 3.3 91 Crude protein content of individual plants varied somewhat within sites (Table 8), but differences were not significant. Differences among plants within given sites ranged from about 0.2 percent between plants at site 11 to 1.7 percent between plants at site 4. There were significant differences between the means for various sites (P=-OOl). For example, the annual (August 1976-May 1977) average protein content of plants at site 11 was 6.9 percent as com­ pared with the low average of 5.0 percent among plants at site 15 and the overall average of 5.8 percent for all sites combined. Generally those sites with highest protein content were more mesic sites at elevations above 1400 meters (4500 ft). Overall, the crude protein content of creeping juniper was lower than that reported for many other important deer browse plants in Montana (Eustace 1971, Duncan 1975, Morton 1976). The occurrence of maximum protein content in the late summer sample contrasts with other shrub species such as big sagebrush, silver sagebrush,.skunkbush sumac curl-leaf mountain-mahogany, rabbitbrush, and chokecherry in which highest protein levels typically occur during early spring (Eustace 1971). The nutritional value of creeping juniper, at least in terms of crude protein, appears to be relatively low throughout the year in relation to the minimal protein requirements of deer. Further, the time of maximum utilization, mid-to-late winter, corresponds with 92 minimum annual protein levels in the plants. ,This would further suggest that mule deer select and use creeping juniper on some basis other than its nutritional value or palatability, such as availability. Available Forage Biomass Creeping juniper plants are capable of supplying an abundance of forage throughout the year. Biomass availability measurements, obtained from clippings of old and new leaves, and twigs less than 5 mm diameter from plants in stands of the eight associations are presented in Table 6. Total average biomass of creeping juniper crown area in pounds per ft ranged from .026 in the JUnvpevus-Potenti,Ita-Caeex association to .038 in the Juniperus-Rhus-Stipa association, with no significant differences between any of the associations. The overall mean biomass 2 for all stands and associations was .031 pounds per ft . .A figure of possibly greater value in evaluating availability of creeping juniper as a forage source is the mean pounds per acre or standing crop in re­ lation to the total vegetation resource. This value ranged from 376.7 pounds in stands of the Junipevus-Calamovilfa association to 589.2 pounds in the Junipevus-Andvopogon-Festuea association. The mean for all stands and associations was 432.0 pounds per acre with no difference noted between the associations. Overall the associations with higher standing crops of creeping juniper were those associated with northerly slopes, areas of moderate precipitation (13-15 inches annually), relative­ ly higher annual temperatures (45° F) and occurring at lower elevations. 93 Utilization by Wildlife Studies of the food habits of wild ungulates in Montana indicate that creeping juniper is utilized primarily by mule deer. Reports of important consumption by other species include pronghorn'antelope (Anti-locapra amevicana) during winter in northern Montana (Martinka 1967) and white-tailed deer (JDdooo-IZeus v-irg-ln-Lana) during January in eastern Montana (Hamlin 1976). Martinka (1967) did not believe that creeping juniper was a suitable forage for pronghorns and attributed its usage to the scarcity of other forage during, the severe 1964-65 winter when antelope experienced heavy mortality. Cole (1956) found minor use of this species by antelope during winter and spring in central Montana, while Dirchl (1963) reported minor antelope use of creeping juniper during fall and important usage during winter on one of his study areas in Saskatchewan, just north of the Montana border. Martinka (1968) reported light use of creeping juniper by white-tailed deer during winter in the Bear Paw Mountains of northcentral Montana. Minor utilization of the species by elk during the winter has been reported for the Sun River area by Casagranda and Janson (1957), Knight (1970) and Schallenberger (1965). Schallenberger (1965) also reported trace usage of creeping juniper by bighorn sheep and white-tailed deer in the Sun River Canyon during winter. ■Creeping juniper is also apparently important to other mammals and birds in Montana. Yde (1977) recorded use of creeping juniper • berries and vegetation by juvenile and adult sharptailed grouse in Phillips County, Montana. Berries occurred in approximately 70 percent of the grouse, crops collected in September 1976, comprising 40 percent of the total contents by volume. Vegetation and seeds occurred in about 20 percent of the crops and made up one percent of the volume. Nielsen (1977), working on the same area, recorded juniper berries in 46 percent of the sharptail crops comprising 25 percent of the total volume. Juniper was the second food item utilized. During my study the incidence of small mammal use on 75 twigs exam­ ined on each of the intensive study sites throughout the summer of 1977 (Table 10) suggested that in some areas substantial small mammal use may • occur. Percentage of twigs used ranged from 0 at several sites to 15 ' percent at site 20. The average for all study sites was 2.6 percent. At least some twigs were used at 80 percent of the sites. A species of Marmota was sighted several times in the vicinity of site 20 and the high use of twigs at that site might be attributed to that mammal. In addition to foliage use, gnawing of berries by small mammals was also noted. The importance of creeping juniper in the diet of mule deer has been documented by several studies in central and eastern Montana (Cada 1971, Compton 1966, Dusek 1971, Eustace 1971, Hamlin 1975, Kamps 1969, Coop 1.975, Lovass 1958, Martinka 1968, Schallenberger 1965, Stoneberg 95 Table 10. Percent of during the creeping juniper twigs utilized by small mammals summer of 1977 on the 30 study sites. Twigs Utilized by Small Mammals Site (percent) I 4 2 - 3 — 4 I 5 4 6 I 7 - 8 I 9 3 10 I 11 3 . 12 - 13 4 14 I 15 - 16 3 . '■ 17 - 18 7 19 I 20 15 21 - 22 4 23 4 • 25 -12 26 3 27 - 28 4 29 I 30 I 31 — Average 2.6 96 1975 and McCarthy et al. 1977) as well as western North Dakota (McKean 1954). However, utilization apparently is highly variable between areas on which creeping juniper occurs. . In areas of important usage, greatest consumption occurs during the winter months-(January, February and March). Use generally begins in late December or early January and gradually increases to March. Thereafter, consumption generally has decreased dramatically, apparently due to the availability of other forages with spring green-up. Some exceptions to this pattern have been reported by Dusek (1971), who observed maximum, usage during April, and Eustace (1971), who compared food habits of mule deer in Garfield and Rosebud Counties of eastern Montana. On the Garfield area, consumption of creeping juniper peaked between December and February, with moderate use occurring from Septem­ ber through November. In contrast, maximum use in Rosebud County occurred between March and May, with no apparent use during the fall months. In general, however, reports of utilization during spring and fall are rare. No summer usage has been reported. Although I did not specifically attempt to quantify utilization patterns on the study sites, general observations of mule deer.activity and utilization supported the conclusion that if creeping juniper is utilized, the majority of use occurs in late winter. Spatial and seasonal variations in use of creeping juniper may be related to corresponding variations in the distribution of mule deer 97 and/or the relative availability of other, more desirable forage plants. This may be particularly important on mountain-foothill ranges where seasonal movements and use of cover types varies in relation to weather conditions. In the Snowy Mountains of central Montana, Kamps (1968) reported a progressive shift in mule deer use from cover types dominated by Douglas Fir and common juniper in late fall and early winter, to ■wind-blown ridges dominated by creeping juniper later, in the winter when this species comprised 45 percent of the plant use at feeding sites. A similar shift in mule deer habitat usage leading to use of creeping juniper has since been observed in other areas (Coop, McCarthy pers. comm.). This shift may be dependent upon weather conditions, and the extent to which it occurs will vary between ranges and years. In the Little Belt Mountains, Lovaas (1958) observed major differences between the food habits of mule deer wintering in Dry Pole. Canyon and those using an adjacent prairie type. Common juniper was the most important forage plant on the former, a predominantly forested area, while creeping juniper was relatively unimportant in spite of moderate abundance. On the prairie area creeping juniper was the most . important forage, while the rare common juniper was found only as a trace item in one rumen .sample. Other plant species associated with creeping juniper oh the prairie type were predominantly less desirable winter forages such as shrubby cinquefoil, rose, snowberry, fringed sagewort, and kinnikinick. 98 Highly palatable browse plants such as antelope bitterbrush (PuVsiiia tvidentata), serviceberry (Amelanchier OlrvifodZda), choke- • ! I cherry (PTunus vdvgdndana) and curl—leaf mountain mahogany, were only rarely found in creeping juniper stands examined during this study. Thus, when deer used these areas, creeping juniper typically was the most abundant, available and, relatively, the most palatable browse species present, particularly during the winter. Coop (1975) felt variations in creeping juniper use was due to availability. Measurements and general observations of plant height and cover suggested that moderate browsing had minimal effect on these parameters of creeping juniper. Heavy utilization, however, left plants either totally destroyed or with large sections of stripped and dead foliage. The latter was especially evident in stands in the foothills of the Snowy Mountains in central Montana and, in eastern and southcentral Montana. The trend in live crown coverage of creeping juniper on transects established and measured by personnel of the Montana Department of Fish and Game in northcentral Montana (Fig. I) was generally downward from the late 1950* s, when measurements were first obtained, to 1973. This trend is similar to that found south of Big Timber in the Picket Pin Canyon and Horseman Flat areas. In the view of Montana Fish and Game and U. S . Forest Service personnel (pers. comm.) familiar with the area, the downward trend may have been the result of two major factors: 99 livestock grazing and/or high deer numbers. In the past, wintering deer numbers were reportedly high in areas, such as Picket Pin Canyon, where creeping juniper showed major declines. In other areas, such as that immediately south of the Sun River Game Range where livestock grazing was and is being controlled, the lack of grazing appears to have allowed the forb and grass complex to increase to such a level as to possibly be competitive with creeping juniper. Competition between grasses and other species, including one-seed juniper (Smith et al. 1975), may be sufficient to inhibit root growth and actually limit many junipers to certain topographies as escarpments and steep rocky ridges (Wells 1970). Effects of Fire on Creeping Juniper Fire or the lack of it has been considered one of the major factors influencing the distribution and abundance of junipers (Foster 1917, Johnson 1962, Leopold 1924), and considerable research has been directed to its effects on various species (Beuhring et al. 1971, Burkhardt 1976, Dalrymple 1969, Dwyer 1967, Foster 1917, Jameson 1962, Johnson et al. 1962, Leopold 1924, Martin et al. 1955, Parker 1945, Wink et al. 1973). Most of this work has focused on the use of burning to control juniper on rangeland. While none of these studies was concerned with the effects of fire on creeping juniper, findings for other, species gener­ ally have indicated that burning results in nearly 100 percent mortality . 100 of plants less than 0.3-0.45 meters (1-1.5 ft) in height. Mortality rates of taller plants vary by species, height of the shrubs or trees, and fuel and fire characteristics. Observations in one stand of creeping juniper, burned by a lightning-caused wildfire, during the reconnaissance phase of this study suggested that burning may be less damaging to this species than reported for other junipers. Subsequently, observations and measurements of the response of creeping juniper to fire were made on two experimental burns. One of these was located in the Blacktail Hills, approximately 10 miles southwest of Stanford in central Montana; the other on the Sun River Game Range, 22 miles west of Augusta in westcentral Montana. Stanford Burn - The three hectare burn site was in an area of slightly rolling topography, with a slope of 5 percent and a northerly exposure. The average elevation was 1785 meters (5890 ft). The vege­ tation was characterized by a dense stand of shrubby cinquefoil, a wide variety of grasses and forbs, and a sparse overstory of Douglas fir and lodgepole pine. Creeping juniper plants occurred exclusively beneath 2 the shrubby cinquefoil plants at a density of 0.034 plants/m . This area is heavily grazed by livestock which apparently avoid use of both the cinquefoil and juniper plants. A total of 21 individual creeping juniper plants was tagged before the burn. Each was measured as to crown dimensions, amount burned and 101 I . • killed and the amount of regrowth, if any (Table 11). Fire tempera­ tures, fuel moisture and soil moisture at the time of burning were also recorded near each of these shrubs (Table 11). The plants were all 2 relatively small, averaging 0.08 m in crown area and 7.8 cm tall. At the time of burning, the average soil moisture beneath these pIdmtS ■ was 49.6 percent with a range of from 45 percent to 65 percent. Aver­ age fire fuel moisture content varied from 4.0 to 23.0 percent, with a mean of 15.0 percent. Although an effort was made to obtain three 2 intensities of fire in separate 100 m plots, burning actually resulted in a mosaic of varying intensities in each plot as evidenced by the wide range of temperatures at tagged plants: 78°C (109°F) to more than '406.9°C (700°F). The mean temperature was 205°C (337.6°F). Post-burn measurements indicated that 53 percent of the plants were burned and died, 43 percent were burned to some extent but re­ mained alive, and 4 percent (I plant) were totally unharmed (Table 11). Approximately 48 percent of the total canopy coverage of creeping juniper on the plots were totally consumed, while 41 percent were burned to a lesser extent and 11 percent were unburned. The percentage of crown area burned on individual plants ranged from 0 (I plant) to 100 percent (10 plants); the average was 85. percent. All plants with crowns totally consumed died. Only one of the plants was killed where less than 100 percent of the crown was burned. Plants not killed tolerated fire temperatures ranging from 78°C ■(109°F) to 267°C (450°F) 102 Table 11. Crown area and height of 21 creeping juniper plants within the Stanford experimental burn together with percent crown burned £ post-fire fate of plant, temperature of fire, and fine fuel and soil moistures of the area. Crown Plant Crown Area (m* 2) Height (cm) Area Burned1 (percent) Regrowth (gms/ plant) Temper­ ature (°c) Fine Fuel Moisture (percent) Soils Moisture (percent) I .08 11.0 100 2 407 23 3 45 3 2 .09 7.5 37.5 — 43 23 45 3 .15 8.0 97.5 1.32 204 23 45 4 .01 6.5 100 — 343 . 23 45 5 .10 3.0 100 — 267 23. 45 6 .06 12.0 62.5 1.35 78 23 45 7 .09 11.0 100 . ---- 118 23 45 8 .11 5.0 100 — 343 23 45 9 .16 4.5 100 — 407 16 47 10 .13 8.0 100 — 204 8 51 11 .23 7.0 97.5 3.95 118 . 5 52 12 .05 8.5 85.0 2.10 78 5 46 13 .04 3.5 97.5 2.00 78 18 65 14 .02 4.0 85.0 0.21 78 8 .65 15 .06 9.0 0.0 — 78 ' 13 46 16 .05 6.0 100 — — 343 5. 58 17 ’ .03 11.0 100 ---- ■ 62 14 .48 18 .10 . 9.0 100 — 204 4 • 57 19 .02 7.0 62.5 0.86 78 9 ' 51 20 .04 8.0 62.5 0.37 118 4 : 45 . 21 .05 5.0 97.5 0.14 267 22 . 52 Average .08 7.8 85.0 1.40 205 . 15 49.6 !percent of plant burned and killed. 2IOO percent = total consumption. 3Samples I to 8 averaged and value shown is the average. 103 or an average of 134°C (209.2°F). However, the fact that several plants which experienced fire temperatures below 267°C (450°F) also died, indicated that under certain conditions, fires of even light intensity may be damaging to creeping juniper Where plants were totally consumed and killed, fire damage usually extended into the root system, below ground level. Plants which remained alive or resprouted did not suffer this severe damage. These plants typically produced new leaf material and showed good growth during the first growing season; the regrowth averaging 1.4 grams per plant (0.14-3.95). Sun River Burn - This site was located on a five percent slope of northeast exposure at an elevation of 1500 meters (4950 ft).. The vegetation was characterized by a dominance of creeping Juniper, grasses, including bluebunch wheat grass and rough fescue, and. forks, of which the major species were prairie thermopsis, silky lupine (Lupinus 'seviceus) and miIkvetch (,AstragaZus spp.). The fire was of low intensity and moved slowly across the plot at a rate of about 0.61 meters (2 ft) per minute. Flames never exceeded one meter in height, moving at an angle of 60 - 90° from the horizontal. The aver­ age fire temperature was 91°C (132.3°F). Average soil and fire fuel moisture were 11 percent and 32 percent, respectively. These temper­ atures averaged only about half as hot as those experienced by plants on the Stanford burn, while soils were drier and fuels contained more 104 moisture as compared with the Stanford site. Effects of fire on individual plants were not measured as on the Stanford burn. Rather, only an estimate of effects on canopy coverage was obtained. Comparison of 2 x 5 plot frame data from the burn plot and the adjacent control plot suggest that approximately 31 percent of the total canopy coverage of creeping juniper were consumed, 29 percent were burned but not destroyed, and 40 percent were not burned (Table 12). This and supplemental data combined indicate that about 60 percent of the total crown area of creeping juniper were burned to some extent and 40 percent were unburned. The terminal portions of all of the 10 major branches dr leaders tagged prior to burning died. However, none of the plants was totally ■killed; and regrowth was detected on all tagged leaders during the first growing season (the summer of 1977). Discussion of Fire Effects The general effects of the two controlled burns on creeping juni­ per plants are summarized in Table 12. The result's indicated that creeping juniper is subject to damage and destruction by fire, though these effects may vary by area and may be less severe than reported for other species of juniper. Several factors may account for these differences. The very low mat-like stature and extensive root.system, together with the size of individual creeping juniper plants seem to be especially important. The low, dense mats may result both in poor 105 Table 12. Comparison between the burns. of the effects of fire on Sun River Game Range and ■ creeping Stanford juniper experimental Effect Characteristic Sun River Game Range Burn Blackball Hills Burn - Stanford Canopy Coverage Consumed (percent) 31 48 Canopy Coverage Burned and/or Killed (percent) 29 41 Total Canopy Coverage Burned (percent) ■ 60 89 Canopy Coverage Unharmed (percent) 40 11 Plants Burned and Not Killed (percent) N.M.1 43 Plants Burned and Killed (percent) N.M. 53 Plants Unharmed (percent) N.M. 4 Plants Consumed When Burned (percent) N.M. 48 Plants Partially Defoliated (percent N.M. 48 Plants Unharmed (percent) N.M. 4 1N-M. = Not Measured. 106 air circulation around the plant and in low fire fuel supplies in and about the plant. Also, when the shrub occurs in larger, dense mats as on the Sun River burn area, fire may not carry completely over or across the plant because of low fuel and/or high moisture or relative air humidity in and around the crown area. With smaller, definitive plants as on the Stanford burn, greater amount of fuel may occur in, around and over the plants resulting in more complete burning and more destructive damage. Where more extensive, rooted mats occur, unburned portions of plants may survive. CONCLUSIONS AND RECOMMENDATIONS The results of this study provide baseline information concern­ ing various biological and ecological attributes of creeping juniper in Montana. Generally, this information may be useful in evaluating wildlife usage and management needs and opportunities - in rangeland habitats. In some cases, the results also serve to clarify existing knowledge and concepts about the plant, its use by wildlife, espe­ cially mule deer, and its importance in rangeland ecosystems. In many instances they point to the need for additional study. The study clearly indicates that creeping juniper is an impor­ tant component of rangeland vegetation. It occurs extensively on foothill and prairie ranges throughout most of Montana east of the Continental Divide, and was the dominant shrub or plant in at least eight different plant associations occurring on ridges, rimrocks, escarpments and hilltops. On these sites, which typically were very open and had only poorly developed soils, creeping juniper may be extremely important in both maintaining and building soils and also tends to be available for use by wildlife most of the time. Creeping juniper appears to be capable of producing and provid­ ing large amounts of forage for wildlife, especially mule deer on sites where it occurs, although annual production varies between sites. Generally, sites with higher annual precipitation (13-15 inches) produced greater amounts of forage. Utilization by deer normally did not appear to adversely affect annual growth and forage production, 108 while light to moderate usage of twigs seemed to stimulate lateral growth of twigs. Concentrated utilization on local sites for long ■ periods of time, however, may result in destruction of plant crowns. Domestic livestock grazing, which reduces competition from grasses and other plants, may be an important factor in maintaining viable stands of creeping juniper at least on some sites. Management of creeping juniper stands should be directed to maintaining existing stands and plant productivity. Because of the two-dimensionality of plants, this species lends itself to measure­ ment from aerial photographs to estimate trends in plant size and density, available biomass and reproduction. The apparent lack of natural reproduction from seed would appear to preclude attempts to stimulate increased plant densities by this means, however, additional seed germination experiments are important to fully understand this idea. Similarly, controlled burning appears to have little applica­ tion in increasing plant densities and productivity, and may actually be detrimental to the species. Range fertilization appears to have little effect on productivity but may' be a viable management tool when increased fruit success and production is desired. Additional research would be desirable to determine factors associated with the distribution of creeping juniper in Montana, especially those responsible for its general absence from western and southwest Montana. Further definition and study of the plant 109 associations in which the species, occurs and the age structure of creeping juniper stands with possible successional relations would be valuable. Further study into the digestibility for deer would be important in assessing the value as a forage species. Relationships between site factors and total nutritional characteristics require further study. Additional research into the effects of various site and environmental factors including grazing, burning, utiliza­ tion by deer, climate, microclimate, and usage by wildlife other than deer on productivity, nutritional aspects and dominance of creeping juniper would also be useful. LITERATURE CITED Ill Beals, E. 1960. Forest bird communities in the Apostle Islands of Wisconsin. Wilson.Bull. 72:156-181. Bifoss, C. G. 1947. The water conducting capacity and growth habits of Jurvipevus Tiovizontali-S Moench and Junipevus vivginiana L. Ecology 28 (3):281-289. Booth, W.' E. and J. C. Wright. 1959. Flora of Montana, part II, Dicotyledons. Mont. State Coll., Bozeman. 280.pp. Bradley, G. A. 1963. Two new species of Cinava cuvtis (Homoptera: Aphididae) from Junipevus hovizontalis Moench. 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Chromosome counts in three cultivars of Junipevus. L. Bot. Gazzette. 132(4):259-262. _____ and H. P . Rasmussen. 1972. Anatomical changes in developing graft unions of Junipevus L . J . Am. Soc. Hort. Sci. 97(2):228-232. 113 _____ and ._____ . 1974. Serological determination of tissue contri­ bution from the graft union - of"Juniperus :L. grafts. Bot.: Gaszette. 135(3):163-169. Fassett, N. C. 1944a. Juniperus Virginiana3 J. horizontalis and j. scopulorum - I. The specific characters. Tor. Bot. Club. Bull. 71(4):410-418. ' 1944b. Juniperus Virginiana3 J. horizontalis and J. scopulorum, II. Hybrids of J. virginiana and J. scopulorum. Tor. Bot. Club. Bull. 71(5):475-483. _____ . 1945a. Juniperus Virginiana3 J. horizontalis and J. scopulorum III. Possible hybridization of J. horizontalis and J. scopulorum. Tor. Bot. Club. Bull.. 72(1):42-46. _____ . 1945b. Juniperus Virginiana3 J. horizontalis and J. scopulorum IV. Hybrids of J. virginiana and J. horizontalis. Tor. Bot. Club. 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One-seed juniper invasion of northern Arizona grasslands. Ecol. Monogr. 32(3):187-207. Jorgenson, H. E. 1976. Classification, description and ecology of the vegetation of eastern Montana, with special reference to the prairie region. Job Prog. Rept., Mont. Dept. Fish and Game, Fed. Aid Proj. W-120-R-7. 7 pp. Kamps, G. F. 1969. Whitetail and mule deer relationships in the Snowy Mountains of central Montana. M.S. Thesis. Mont. State Univ., Bozeman. 59 pp. ' • Knight, R. R. 1970. The Sun River elk herd. Wildl. Monogr. No. 23. 66 pp. Lanphear, F. 0. 1966. Influence of the stock plant environment of the rooting of Junvpevus hov-tzontdlis plumosa. Prbc. Am. Soc. Hort. Sci. 89:666-671. Leopold, A. 1924. Grass, brush, timber and fire in southern Arizona. J.For. 24 (6):1-10. Little, E. L. 1971. Conifers and Important Hardwoods .- Atlas of U.S. Trees. USDA For. Serv., Washington, D.C. 1:221-N, 221-E. Lonner, T. N. 1972. 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State Univ., Bozeman. 70 pp. / APPENDIX 118 Table 13. General and exact locations of the 51 creeping juniper. __________ study sites. __ ■ Site County Nearest Post Office Distance/ Direction from P. 0. Location I Lewis & Clark (kilometers) Augusta 25 NW ' T21N R 8W S16- SEh 2 Lewis & Clark Augusta 17 SW . T19N R 7W S17 NE% 3 Lewis & Clark Augusta 17 SW T20N R 7W S16 NWk 4 Lewis & Clark Augusta 17 SW T19N R 7W S 9 -SWk- 5 Lewis & Clark Augusta 17 SW T19N R 7W S16 SEk 6 Lewis & Clark Augusta 25 NW T21N R 8W S 4 SWk 7 Lewis & Clark Augusta 25 NW T21N R 8W S22 SEk 8 Teton Choteau 42 W T25N R 8W Sl6 NWk 9 Lewis & Clark Augusta 20 W T 2 ON R 8W S 8 SEk 10 Lewis & Clark Augusta 20 W T20N R 8W S 9 NWk 11 Pondera Dupuyer 33 W T28N R 8W S31 NWk 12 Pondera Dupuyer 33 W T28N R 9W ■S25 SEk 13 Fergus Denton 8 E T18N R15E S35 SEk 14 Fergus Denton 8 E ' T18N RISE S35 SEk 15 Petroleum Teigan 8 N T15N R25E S 6 SEk 16 Petroleum Teigan 8 N T15N R25E S 7 .NWk 17 Fergus Hilger 8 E T19N Rl 7E S16 NEk 18 Fergus Hilger 8 E T19N R17E S. 9 SEk 19 Fergus Moore 17 S T13N R16E S35 NWk 20 Fergus Moore 17 S T13N Rl 7E S30 SEk 21 Wheatland Judith Gap 33 W TllN R14E S23 SWk ■ 22 Wheatland Garneil 25 E TllN R17E S16 SWk 23 Sweetgrass Nye 17 NW T 5S R14E S I NWk 25 Sweetgrass Nye 25 W ' T 5S R14E s. 9 NWk 26 Prairie Terry 42 NW T15N R4-8E S19 NWk 27 Prairie Terry 42 NW T15N R48E S29 SEk 28 Prairie Terry 42 NW T15N R47E S36 NWk 29 Lewis & Clark Augusta 25 NW T21N R 8W S21 SEk 30 Pondera Dupuyer 33 SW T27N R 8W S17 NWk 31 Teton Choteau . 42 W T25N R 8W S 9-NWk 32. Lewis & Clark Lincoln 20 E T15N R 7W S23 NWk' 33 Toole Shelby 20 SE T31N R IW S28 NWk 34 Choteau Ft. Benton 8 E ■ T24N R 9E si9 swk 35 Hill Box Elder 12 E T29N R14E S 2' NEk 36 Phillips Dodson 7 E T30N R27E s I swk 37 Fergus Ryegate 58 N T12N . R20E • S27 SEk 119 Table 13. (continued) Site County Nearest Post 'Office Distance/ Direction from P. 0. Location (kilometers) 38 Meagher Checkerboard 25 S T 9N R 9E S30 SW% 39 Meagher White Sulfur 42 W TllN R 3E . S25 NW% 40 Rosebud . Ingomar 8 W TlON R34E S20 SW^ 41 Bighorn Lame Deer 17 W • T 3N R39E S13 NW^ 42 Powder River Broadus 8 NW T 4S R50E S. 2 NW% 43 Carter Ekalaka 25 SE T 2S _R60E S24 NEk 44 Prairie Ismay 5 N T 9N R55E S 8 SWk 45 Wibaux Wibaux 17 SE T13N • R60E -S34 NWk 46 Roosevelt Culbertson 5 N T28N .R56E S16 SWk 47 Daniels Scobey 7 E- T35N R49E XS 8 SWk 48 Valley Ft. Peck 3 S T26N R40E S20 NWk 49 Garfield Jordan 25 N T20N R38E S18 NWk 50 . Cascade Monarch 3 N T16N R 6E S23 NWk 51 Sweetgrass Melville 17 SW T 3N R13E S13 SEk 24 Sweetgrass Ney 42 NW T 4S R14E S29 SWk c. 120 Table 14. Exposure, slope, elevation and drainage of 51 creeping juniper sites in Montana. Site Slope (percent) Elevation (meters) Exposure Direction/Degrees Drainage I 5 1515 ENE • 65 Picture Basin 2 14 1439 SSE 178 Elk Creek 3 33 1417 NNE 13 Elk Creek 4 5 1326 ENE 72 Elk Creek 5 14 1270 WSW 268 Elk Creek 6 36- 1412 ENE 50 Dickinson Reservoir 7 41 1485 WSW 238 Barr'Greek 8 28 1885 ENE 60 Chicken Coulee 9 26 1594 ENE 55 Cutrock Creek 10 21 1564 WSW 233 Cutrock Creek 11 27 1448 NNW 342 Dupuyer Creek 12 9 1479 WNW 280 Dupuyer Creek 13 38 1094. NNE 30 Sage Creek 14 35 1070 SSW 190 Sage Creek 15 5 1012 . ENE 80 Warhorse Reservoir 16 11 1030 NNE 340 Warhorse Reservoir 17 40 1406 SSE 15.8 Deer Creek 18 15 1364 WNW 310 Deer Creek 19 19 1600 . NNE 38 Rock Creek 20 24 1606 ESE ■ 99 Rock Creek 21 12 1670 SSE 178 Roberts Gulch 22 40 1560 NNW 350 Dry Creek 23 . 30 ■ 1776 S 180 Picket Pin Creek 25 10 1794 NNE 25 Stillwater River 26 35 1040 ESE 123 Agate Creek 27 31 1048 NNE 10 . Agate Creek 28 36 1030 SSW 195 Cherry Creek 29 42 1542 SSE 157 ■ Rose Creek 30 19 1776 SSE 165 Dupuyer Creek 31 21 1845 SSE 165 Bynum Creek 32 . 34 1500 ESE 117 Blackfoot River 33 54 909 NNE 17 Marias River 34 38 848 NNW 347 Missouri River 35 40 936 NNW 330 Big Sandy Creek 36 24 . 758 WSW 240 Milk River 37 5 . 1621 ESE 117 North Willow Creek 38 7 2091 NNW 347 Musselshell River >■ 121 Table 14. (continued) Exposure Site. Slope (percent) Elevation (meters) Direction/Degrees Drainage 39 46 1561 NNW 353. Thomas Creek 40 42 . 939 NNW 357 Big Porcupine Cr. 41 50 1000 NNE 8 Rosebud Creek 42 33 969 NNW 317 Mizpah Creek 43 ■ 31 984 NNW 357 Spillman Creek 44 15 772 ENE 55 0'Fallon Creek 45 28 772 ENE 73 Beaver Creek 46 24 651 WSW 257 . Missouri River 47 37 848 SSE ' 153 ■ Poplar Creek 48 26 788 WSW 263 Missouri River 49 30 939 NNE 18 Hell Creek 50 5 1788 NNW 330 Belt Greek 51 19 — SSE 175 Big Timber Creek 24 6 2485 SSW 200 Picket Pin Creek 122 Table 15. Scientific and common names of the 24 graminoids, 104 forbs, 18 shrubs and 3' trees .in this study. GRAMINOIDS Agvopyron Sm-LthyL-I Western wheatgrass Agvopyron sp-Lcatim Bluebunch wheatgrass Andvopogon soopavius Little bluestem Av-Lst-Lda LongyLseta Red threeawh BouteLoua gvae-LL-Ls Blue grama Bvomus -Lnevm-Ls' Smooth brome Bvomus japonieus Japanese brome CgLamovLLfa LongLfoLLa Prairie sandreed CaLamovLLfa puvpeseens Purple reedgrass Cavex eLynoLdes Kobresia-Iike sedge Cavex fLLLfoLLa Threadleaf sedge Cavex heLLofLLa Sun sedge Cavex scLvopoLdea Single-spike sedge Cavex VesLoavLa Inflated sedge DanthonLa LntevmedLa Timber danthonia Festuea LdahoensLs Idaho fescue Festuea scabveLLa Rough fescue HeLLetotvLehon hookevL Spike oat Juneus DaLtLcus Wire rush KoeLavLa evLstata Prairie junegrass Poa pattevsonL Patterson’s bluegrass Poa spp. Bluegrass species StLpa eomata Needle and thread StLpa vLvLduLa Green needle grass Table 15. (centinued) FORES kdhiVlea WiVLefolyLim Allivm eemuvm Allivm. textile Anemone rmltifida Anemone patens Antennavia vosea Avenaria eongesta Avenccvia hookevi Avtemisia eana Artemisia fvigida Astvagalus bisulatus Astvagalus euaosmus Astvagalus gilviflovus Astvagalus peetinatus Astvagalus tenellus Bdlsamovhiza saggitata Besseya wyomingensis Buplevum cmevioanum Campanula votundifolia Castellija sessiliflova Cevastium arvense Chvysopsis villosa Civsium undulatum Clematis pseudoalpina Comandva umbellata Cvepis oeeidentalis Cvyptanthe intevvupta Western yarrow Nodding onion Textile onion Ball anemone Pasque flower Rose pussytoes Ballhead sandwort Hooker sandwort Silver sagebrush Fringed sagewort Two-grooved milkvetch Elegant milkvetch Three-leaved milkvetch Narrow leaf pdisonvetch Loose flower milkvetch Arrowleaf balsamroot Kittentail American thorway Roundleaf harebell Downy-painted cup Field chickweed Golden aster Wavy leaf thistle Climbing-purple clematis Pale bastard toad flax . Western hawksbeard Bristly cryptantha 124 Table 15. (continued) Forbs DeZphini-Um bieotov Dodecatheon conjugens Eehinaceae paZZida Evigevon eaespitosus Evigevon oehvoZeueus Evigevon subtvinemis Evigevon spp.' Eviogonum fZavum Eviogonum umbeZZatum Eviogomm spp. Evitiehum howardii Evysium aspevum Fvaseva speeiosa FvitiZevia pudiea GaiZZavdia avistata GaZium boveaZe Geim tvifZovum GvindeZia squavvosa Gutevvhiza savothvae HapZopappus avmevoides HapZopappus nattaZi HapZopappus spinuZosus Hedysavum boveaZe Hedysavim suZfuveseens Hymenoxys aeauZis Leucvocvinum montamm Liatvis punetata Linum pevenne Low larkspur Shooting star Purple coneflower Tufted fleabane Buff erigeron Three-veined, fleabane Erigeron species • Yellow eriogonum Sulfur eriogonum Eriogonum species Howard’s alpine forget me not Plains wallflowers Spotted frasera Yellowbell Blanket flower Northern bedstraw Prairie smoke Curlcup gumweed Broom snakeweed Thrift goldenweed Nuttall goldenweed Spring goldenweed Northern sweetvetch White sweetvetch Arizona hymenoxys Sand lily Dotted blazingstar Blueflax ■ 125 Table 15. (continued) Forbs L-Lthopemrum vudevale Lomat-Lwn oous Lomatiwn macvooavpwn Lotus purshianus Lupinus serioeus' Lupinus wytheii Lupinus spp. Mevtensia vividis . Monavda fistulosa Musineon divarication Myosotis sytvatioa ■ Opuntia potycantha Orthoeavpus Luteus Orthoearpus tenufolia Oxytropis besseyi Oxytropis serieeus Oxytropis viseidia Paroynchia sessiflora Penstemon attenuates Penstemon erianthus Penstemon nitidus Petalostemon earididwn Petalostemon purpeseens Phlox alyssifolia . Phlox hoodii Phlox kelseyi Phlox multiflora Western gromwell Mountain lomatium Large-fruit lomatium Prairie trefoil Silky lupine Wyethe lupine Lupine species Greenleaf bluebell Horse mint Leaf musineon Alpine forget me not' Plains prickly pear Yellow owl clover Thin-Ieaf orthocarpus Bessey pointvetch White pointloco Sticky pointloco Stemless nailwort Sulfur penstemon Fuzzytongue penstemon Waxleaf penstemon White prairie clover Purple prairie clover Alyssum-Ieaved phlox Hoods phlox Kelsey phlox Flowery phlox 126 Table 15. (continued) Forbs Polygala alba Polygonum b-istovdis Potentilla gvacilis Potentilla hippiana Ranuneulus glabevrimus Saifraga rhomboidea Sisyrinehium sarmentosum Sedum laneeolatum Seneeio eanus Smilieia raeemosa Solidago missourensis Solidago oeeidehtalis Taraxieum officinale Thermopsis rhombifolia . Tragopogon dubius Vicia 'americana Viola nuttalli Yueea glauea Zygondenus elegans Zygodenus venosus White milkwort Western bistort Northwest cinquefoil Horse cinquefoil Sagebrush buttercup Diamondleaf saxifrage Blue-eyed grass Lance-leaf stonecrop Wooly groundsel Falsq spikenard Missouri goldenrod Western goldenrod Common dandelion Prairie thermopsis Common salsify American vetch Nuttall violet Soapwell Glaucos zygodenus Meadow death camas 127 Table 15. (continued) SHRUBS Apocynwn oannabiwn Avetostayphtos uva-wvst Avtemista tongifotia Avtemisia tvidentata Atviptex caneseens Bevbevis .vepens Junipevus hovizontatis Potentitta fvutieosa Pvunus vivginiana Rhus tvitobata Ribes spp. Rosa aeicutaxis Rosa avkansana Symphovieavpos atbus Symphovicavpos oecidentalis Hemp dogbane Kinnikinick Longleaf sagebrush Big sagebrush Fourwing saltbrush Oregon grape Creeping juniper Shrubby cinquefoil Chokecherry Skunkbush sumac Currant species Prickly rose Prairie rose Common snowberry Western snowberry 128 Table 15. (continued) TREES Limber pine Ponderosa pine Douglas fir TPinus ftexitus Pinus ponderosa ' Pseudotsuga menziesii Table 16. Percent canopy coverage and percent frequency among 2x5 plot frames for the 30 intensively studied creeping juniper sites. Sites Taxa I 2 3 4 5 6 7 8 9 GRAMINOIDS: Agropyron smithii (Agsm) - - - - - - - - - - - - - - - - - - Agropyron spieatum (Agsp) 5/471 13/77 3/30 7/77 5/79 8/90 16/87 1/13 11/63 Andropogon soopar-tus (Ansc) - - - - - - - - - - - - - - - - - - Ar-Lst-Lda tong-Lseta (Arlo) - - - - - - - - - - - - - - - - - - Bouteloua grao-Ll-Ls (Bogr) _ _ _ _ - - - - - - • - - Bromus -LnemrLs (Brin) Bromus japonieus (Brja) Calamov-Llfa Iong-Lfolia I/ 7 (Calo) Calamovilfa purpesoens (Capu) Carex elynoides (Gael) Carex fH i folia (Cafi) 3/23 11/50 3/23 10/57 3/27 6/40 2/20 8/40 Carex heliofila (Cahe) Carex soirpoidea (Case) Carex vesioaria (Cave) 1/13 1/10 1/10 — — I/ 7 4/27 I/ . 7 1/13 tr/10 Danthonid intermedia (Dain) - - Festuoa idahoensis (Feid) 9/53 tr/ 3 10/60 4/47 17/77 8/47 — — 11/70 22/73 Festuoa. soabrella (Fesc) 6/33 4/23 9/37 — — 2/10 • 2/13 — —- 11/73 — — 129 Table 16. (continued) Sites Taxa 10 11 12 13 14 15 16 17 ■ 18 19 20 GBAMINOIDS: (4.gsm) - — - - — - - - - - - - 10/63 - - - - 5/63 16/27 (Agsp) I/ 7 3/47 - - - - - - 1/ 3 - - . 2/30 - - (Ansc) - - - - - - - - - - - - - - - - - - - - - - (Arlo) - - - - - - - - - - - - - - - - - - - - - - (Borg) - - --- ---. --- 2/13 - - - - - - --- - - --- (Brin) — — 2/20 2/17 — — — — — — — — — — — — tr/ 7 — — (Brja) --- --- --- --- --- --- --- - - --- --- --- (Calo) - - - - - - - - - - 12/80 (Capu) - - 2/13 (Gael) - - - - - - - - - - (Cafi) 4/20 3/23 3/23 2/17 6/30 - - - - 2/23 - - - - 1 / 3 (Cahe) --- - - --- - - - - - - - - - - - - - - (Case) — — — — 14/63 — — — — — ' — — — — — — — — — — — (Cave) 1/50 5/53 3/23 2/ 7 - - - - - - 6/30 - - 1/37 3/33 (Dain) - - - - - - - - - - - - - - - - - - (Feid) 9/57 3/13 7/63 - - - - - - tr/ 3 tr/ 3 3/13 14/97 6/40 (Fesc) - - 15/67 2/20 - - - - - - - - - - - - - - 130 Sites Taxa 21 22 23 25 26 27 28 29 30 31 GRAMINOIDS: (Agsm) 5/37 I/ 7 25/93 2/13 4/17 7/27 - - (Agsp) . - - - - 1 / 7 2/37 3/33 - - - - 12/67 1/20 8/80 (Ansc) - - - - - - - - - - 3/43 - - - - - - - - (Arlo) - - - - - --- 1/ 3 - - - - - - - - - - (B°rg) - - ----- --- --- --- --- - - - - --- (Brin) - - - - - - - - - - - - - - - - - - --- (Brja) - - - - - - - - - - - - - - - - --- - - (Calo) x- - - - - - - - 2/23 - - 4/27 - - (Capu) - - - - - - - - - - - - - - - - - - - - (Gael) - - - - - - 1/37 tr/3 (Cafi) - - - - - - - - 8/40 7/47 5/40 1/10 7/40 12/50 (Cahe) --- --- - - ---t r / 3 ------ --- (Case) - - - - - - - - - - - - - - - - - - - - (Cave) 1/20 10/83 - - - - 6/23 3/27 1/10 - - 3/23 (Dain) - - - - - . - - - - - - - - - - - - - - - - - (Feid) 3/33 5/63 - - 8/57 - - - - - - 1/ 3 - - 1/30 (Fesc) _ _ _ _ _ _ _._ _ _ - - - - - - 6/47 11/63 Table 16.. (continued) 131 Table 16. (continued) Sites Taxa 1 2 3 4 5 6 7 8 9 GRAMINOIDS: Heliototviohon liookevi (J&eho) 7/27 - - - - - - - - 1 / 7 - - - - 1 / 3 Junous baltious (Juba) ■ - - - - - - - - - - - - - - - - - - Koelavia ovistata (Kocr) 4/13 I/ 7 2/10 - - - - 1/10 I/ 3 tr/10 - Poa pattevsoni (Popa) - - - - - - - - - - - - - - - - - - Poa spp.. . - - - - - - - - - - - - - - - - - - Stipa oomata (Stco) - - - - - - - - 1 / 7 - - - - - - - - Stipa vividula (Stvi) - - - - - - - - - - - - - - - - - - Unknown grasses FORES: Aohillea millefolium (Acmi) — — - - 2/27 1/17 — — tr/ 3 — — — — 1/23 Allium oevnuum (Alee) Allium textile (Alte) Anemone multifida (Anmu) 1/33 1/30 1/23 1/40 3/47 1/23 3/40 1/40 ' — — Anemone patens (Anpa) 2/10 — — 1/10 tr/ 3 — — tr/ 3 Antennajoia vosea (Anro) tr/ 7 tr/ 3 1/13 tr/ 3 1/10 tr/ 7 5/60 I/ 7 Avenavia oongesta (Arco) I/ 7 — — I/ 3 1/23 — — Avenxzvia hookevi (Arho) Avtemisia oana (Area) — — - — — 1/37 Avtefnisia fvigida (Arfr) 8/13 1/17 2/23 1/23 3/27 1/20 8/50 1/20 2/54 132 Table 16. (continued) Sites Taxa 10 11 12 13 14 15 16 17 18 19 20 GRAMINOIDS: (Heho) (Juba) (Kocr) (Popa) Poa ssp-. (Stco) (Stvi) Unknown grasses 13/60 tr/ 3 10/40 - - - - - - - - - - 9/53 — — — — tr/ 6 — — — — — — — — — — — — — — — — 1/13 - - - - - - - - 2/17 tr/ 3 I/ 7 ---- - - - - - - - - 1/10 13/87 22/90 _ _ _ _ _ _ _ _ _ _ _ _ 4/23 - - - - - — — I/ 6 — — — — FORBS: (Acmi) (Alee) (Alte) (Anmu) (Anpa) (Anro) (Arco) (Arho) (Area) (Arfr) tr/10 — — tr/ 3 2/13 - - - - ■ 1 / 3 5/57 4/47 6/50 I/ 7 tr/10 tr/10 tr/ 7 1/23 — —' 4/33 1/10 - - tr/ 3 2/13 — — — — tr/ 3 4/47 tr/ 3 11/77 3/27 4/30 — — I/ 3 1 / 3 - - - - 5/30 tr/ 3 — — — — — — 2/17 tr/10 3/tr — — 1/10 1/10 2/30 tr/ 3 1/10 1/10 - - 1/10 1/13 3/13 - - 133 Table 16. (continued) Sites Taxa 21 22 23 25 26 27 28 29 30 31 GRAMINOIDS: (Heho) tr/ 3 - - - - - - - - - - - - - - - - --- (Juba) - - - - - - - - - - - - - - - - - - - - (Kocr) 3/20 - - 3/20 - - - - 1 / 3 - - 1 / 3 7/43 1/23 (Popa) --- --- --- --- 1 / 7 --- --- --- --- --- Paa ssp. - - - - - - - - - - - - - - - - - - - - (Stco) - - - - - - - - - - - - - - - - - - - - (Stvi) - - - - - - 2 / 3 Unknown - - - - - - 3/17 - - 2/13 - - - - - - - - grasses FORES: (Acmi) 1/10 - - 1/ 7 tr/17 tr/ 3 1/10 - - - - I/ 3 tr/ 3 (Alee) - - - - tr/ 3 - - - - - - - - - - - - - - (Alte) --- - - - - --- --- --- ---tr/10 1/30 (Anmu) - - 2/23 - - 4/40 ---- 5/53 3/23 (Anpa) I/ 7 — — — — 7/60 — — — — — — — — — — — — (Anro) 1/10 1/13 - - 2/13 - - - - - - - - 2/17 tr/ 7 ■ (Arco) - - - - - - - - - - - - - - - - - - - - (Arho) --- --- - - - --- --- --- - - - - - - 1/20 (Area) - - - - - - - - 1/10 - - - - - - - - - - - (Arfr) - - - - 3/23 - 2/13 I/ 3 - 2/13 1/13 1/50 134 Table 16. (continued) Sites Taxa I 2 3 4 5 6 7 8 9 FORES: Astragalus btsulatus (Asbi) - - . - - - - - - - - - - - - - - - - Astragalus euoosmus (Aseu) - - - - - - - - 2 / 3 - - 1 / 3 - - - - Astragalus g-ilwiflorus (Asgi) I/ 3 • 1 / 3 Astragalus IpeotyLnatus (Aspe) - - - - - - - - - - - - - - _ _ Astragalus tenellus (Aste) - - - - - - - - - - - - - - _ _ Balsamorhiza saggitata (Basa) - - - - • - - ■ - - tr/ 7 - - - - - - _ _ Besseyawyomingensis (Bewy) tr/ 3 - - 2/13 - - - - - - - - - - _ _ Buplerum ameriaanum (Buam) - - 1/ 7 2/20 - - 1/ 3 3/23 - - - - _ _ Campanula rotundifolid.Caxo') _ _ - - - - - - - - ' - - - - Castellioa sessiliflora^3-3^) - _ - - - - _ _ _ _ _ _ _ _ _ _ _ _ Cerastium aaroense (Gear) 1/27 tr/ 3 tr/23 tr/17 tr/17 1/23 tr/ 3 tr/20 1/13 Chrysopsis villosa (Chvi) - - - - - - - - - - - - - - _ _ Cirsium undulatum (Ciun) - - - - - - - - - - - - - - - - - - Clematis pseudoalpina (Clps) - - - - - - - - - - - - - - - - - - Comandra umbellata (Coum) - - 1/17 - - .1/17 1/27 tr/10 - - - - - - Crepis oooidentalis (Croc) - - - - - - - - - - - - - - Cryptanthe ■ interrupia (Crin) - - . - - - - - - - - 1 / 3 - - - - - - Delphinium bieolor (Debi). - - - - - - -- - - - - - - - - - - Dodeeatheon eonjugens (Doco) - - - - - - 1 / 3 - - - - - - - - - - Eehindeeae pallida (Ecpa) - - - - - - - - - - - - - - - - - - - 135 Table 16. (continued) Taxa 10 . 11 12 13 Sites 14 • 15 16 17 18 19 20 FORBS: (Asbi) tr/ 7 — — _ __ (Aseu) (Asgi) I/ 3 — — — — — — (Aspe) 7/37 (Aste) — — — — 1/10 (Basa) 4/20 (Bewy) 1/17 tr/ 3 — — 5/30 — — — — ' 3/17 — — tr/ 3 10/80 3/20 .(Buam) I/ 7 tr/ 3 (Caro) — — — — tr/ 3 I/ 3 (Case) — — — — I/ 7 (Gear) tr/20 tr/37 tr/17 tr/ 3 tr/17 1/17 2/16 (Chvi) (Ciun) (Clps) 3/17 - - (Coum) — — tr/ 7 — — I/ 3 1/10 - - 5/40 — — — — 2/13 tr/ 7 (Croc) (Grin) (Debi) 1/13 tr/ 7 (Doco) — — I/ 7 1/13 — - (Ecpa) 136 Table 16. (continued) Sites Taxa 21 22 23 25 26 27 28 29 30 31 FORES: (Asbi) - - - - - - - - - - - - - - - - - - - - (Aseu) - - - - - - 4/27 I/ 7 tr/ 3 - - - - 2/13 (Asgi) - - - - - - - - - - 1 / 3 - - - - 2/23 tr/ 3 (Aspe) - - - - 5/53 - - - - - - - - - - _ _ (Aste) - - - - - - - - - - - - - - - - - - - _ (Basa) - - - - 2/17 - - - - - - - - 1/10 - - 1/3 (Bewy) - - - - - - - - - - 1/13 - - - - - - 1/ 3 (Buam) - - - - - - - - - - - - - - - - tr/ 7 _ _ (Caro) - - - - - - - - 1/20 (Case) - - - - - - - - - - - - - - - - 1 / 3 - - (Gear) 1/23 tr/10 2/13 tr/ 3 - - - - - - 1/13 - - 2/37 (Chvi) - - - - - - 1/10 (Ciun) - - - - - - - - tr/ 3 - - - - - - - - - - (Clps) - - - - (Coum) 2/27' 4/43 1/40 - - 1/ 3 - - 3/27 2/ 7 2/33 - - (Croc) - - - - - - - - .--- - - 1/10 (Crin) - - - - - - - - - - (Debi) • - - - - - - - - - - - - - - - - (Doco) " - - - - - - - - - - - - - - - - (Ecpa) - - - - - - - - 1/7 - - 1/ 3 - - 137 Table 16. (continued) Taxa ■ . I 2 3 Sites 4 5 6 7 8 9 FORBS: Evigeron oaespitosus(Erca)- - Evigeron ochroleueus'(Eroc')- - . tr/10 tr/ 7 I/ 3 1/13 Evigevon subtvinevvis (Ersu)- - Evigevon spp. - - . 1/10 2/13 3/37 2/20 2/37 tr/ 3 2/37 — — Evigevon■faivum (Erfl) ------- • Eviogoniun Z^mbeZZatw(Erum)- - I/ 3 tr/ 3 I/ 3 Eviogonum spp. - - Evitiohwn howavdii (Erhq) - - 1/13 — — tr/ 3 I/ 3 Evysiwn aspevwn ■ (Eras) - - Fvaseva speoiosa (Frsp) I/ 3 'I/ 3 Fvitilevia pudioa (Frpu) - - Gaitlardia aristata (Gaar)- - I/ 3 I/ 7 I/ 3 Galiwn boveale (Gabo) tr/ 7 — 1/10 tr/ 3 • 2/30 tr/10 — — — — 1/20 Gewn tviflovum (Getr)' - - 2/23 — — tr/ 7 I/ 3 tr/ 7 Gvindelia sqnavvosa(Grsq) - - — — Gutevvhiza savqthvae(Gusa) - - Haplopappus avmevoides(Haar)— Haplopappus nuttali (Hanu)- - Haplopappus spinulosus(Hasp)— Hedysarwn boveale (Hebo) - - T- — I/ 3 - - — — 138 Table 16. (continued) • Sites Taxa 10 11 12 .13 14 15 16 17 18 19 - 20 FORES: (Erca) (Eroc) (Ersu) ETigevon spp. (Erf I). (Erum) Eviogonum spp. (Erho) (Eras) (Frsp) (Frpu) (Gaar) (Gabo) (Getr) (Grsq) (Gusa) (Haar) (Hanu) (Hasp) (Hebo) — — — — tr/ 3 tr/ 3 I/ 3 — — I/ 7 — —- — — —■— — — 1/10 1/13 tr/17 _ _ 1/4 _ _ _ _ _ _ _ _ tr/13 I/ 7 - - 'tr/ 7 , 1/17 tr/ 7 - - 1/33 I/ 3 tr/ 7 - - 1/33 2/10 4/27 2/17 I/ 7 tr/ 3 2/13 :: 2/10 9/83 I/ 7 4/67 3/27 3/17 — — ■ I/ 3 1 / 3 I/ 7 6/47 139 Table 16. (continued) Sites Taxa 21 22 23 25 26 27 . 28 29 30 31 FORES: (Erca) - - - - - - - - - - - - - - - - - - - - (Eroc) 1/3 - - - - - - - - - - - - - - 1/10 (Ersu) - - 1/27 tr/13 - - - - - - - - 1/ 7 tr/ 3 Er-Lgevon spp. - - - - I/ 7 - - - - - - - - - - - - - - (Erfl) - - - - - - - - - - - - 1/31 (Erum) - - - - - - - - - - - - 2/33 - - - - - - Er-Logonum spp. - - - - 1 / 7 - - - - - - - - - - tr/ 3 tr/ 7 (Erho) - - 8/53 - - - - - - - - - - - - - - - - (Eras) . - - - - - - tr/ 3 - - - - - - - - - - - - (Frsp) --- --- --- --- --- --- --- --- - - -- (Frpu) --- --- --- --- ---. ---- --- --- --- -- (Gaar) tr/ 3 — — — — 1/40 — — — — 1/10 — — tr/10 — — (Gabo) • 1/13 - - 1/40 3/60 - - 1 / 3 - - - - tr/10 (Getr) - - - - - - - - - - tr/ 3 - - - - - - - - (Frsq) --- --- --- --- --- --- -- ---.------ (Gusa) - - - - - - - - - - - - 2/17 - - - - (Haar) - - =-- ---- - - - --- - - - - (Hanu) - - - - - - - - - - - - - - - - - - (Hasp) - - _ - - -- - - - - - - - - - - - - 1/7 (Hebo) - - - - - - - - - - - - - - - - - - - - 140 Table 16. (continued) Taxa Sites I 2 3 4 5 6 7 8 9 FORES: Hedysarion sulfuresaens (Resn) - - - - - - - - - - - - - - - - - - Hymenoxys acaul-is (Ryac) 2/27 2/20 1/13 tr/7 1/3 2/17 2/27 - - LeucroarjLnwn montanumCLemo) - - - - - - - - - - - - - - - - LjLatris punctata (Lipu) - - - - - - - - - - - - - - - - - - Linum perenne(Lipe) - - tr/3 - - tr/13 tr/10 - - - - - - - - Lithopermuin ruderaie (Liru) - - - - 1/3 - - - - - - - - - - - - Lomatium eous (Loco) tr/7 1/7 tr/20 tr/7 tr/7 - - - tr/7 tr/17 - - Lomatium maerocarpum(Loma) ■ . - - - - 1/3 - - - - - - - - 1/30 - - Lotus purshianus (Lopu) - - - - - - ^ - - - - - - - - - - - - Lupinus sericeus (Luse) - - - - - - 3/17 - - - - - - - - 1/10 Lupinus wytheii(Luwy) - - - - - - - - - - - - Lupinus spp. - - - - - - - - - - 1/7 - - - - - - Mertensia viridis (Mevi) - - - - - - - - - - - - - - - - - - Monarda fistulosa (Mofl) - - - - - - - - - - - - - - - - - - Musineon divarication(Mudi) tr/3 1/17 - - - - - - tr/3 1/7 • tr/3 3/43 I4yosotis SyLvatica(Mysy) - - - - - - - - - - - - - - - - - - Opuntia polycantha(O-ppo) - - - - - - - - - - - - - - - - Orthocarpus Leuteus(Orle) - - - - - - - - - - - - - - - - - - 141 Table 16. (continued) Sites Taxa 10 11 12 13 14 15 16 17 18 19 20 FORES: (Hesu) - - tr/7 3/27 1/7 3/7 - - - - 7/60 (Hyac) - - 1/10 ' - - - — - - - - - - - - - - - - - (Lemo) - - 1/7 - - - - - - - - - - - - - - - - - - (Lipu) — — — — 1/7 — — — — — — tr/3 — — 1/3 — — — — (Lipe) - - - - - - 1/10 1/7 - - tr/3 1/3 3/53 - - tr/3 (Liru) - - 10/7 - - - - - - - - - - - T - - - - - - (Loco) tr/17 t r / 7 ---tr/7 tr/3 - - ---tr/13 1/13 2/23 2/17 (Loma) - - - - - - - - - - - - - - - - - - - - (Lopu) — — — — — — tr/3 7/4.7 — — — S T T T T T T T T (Luse) - - 2/10 (Luwy) - - - - - - - - - - - - - - - T - - Lup-inus spp. - - — 1/3 - - - - - - - - - - - - - - - - (Mevi) - - - - - - - - - - - - - - - - - - - - - - (Mofi) --- --------- --- --- --- -- --- --- ---6/20 (Mudi) - - - - - - - - 1/7 - - 1/7 3/23 (Mysy) --- --------- --- --- --- --- - - --- --- -- (Oppo) - - - - - - tr/3 - - - - - - - - - - - - (Orle) --- ---- T ------ --- --- --- --- --- --- --- 142 Table 16. (continued) Sites Taxa 21 22 23 25 26 27 28 . 29 30 31 FORBS: (Hesu) --- ---.------ --- ---•------ ' ---- 1 / 7 --- (Hyac) — - - - - - - - - - - - - tr/3 1/37 2/40 (Lemo) - - --- - - - - - - - - - - - - - - - - - (Lipu) - - - - - - - - tr/3 - - — — ’ — - — — - - (Lipe) tr/3 tr/3 1/17 tr/3 ' tr/3 ' 1/3 tr/7' (Liru) - - - - tr/3 - - - - - - - - - - - - . - - (Loco) - - --- - - 1/23 --- --- ---tr/7 tr/3 .tr/10 (Loma) - - - - - - - - - - - - - - - - - - - (Lopu) - - - - - - - - - - - - - - - - (Luse) - - --- - - - - - - - - - - - - I/7 ~ - (Luwy) . - - - - - - - - - - - - - - - - - - tr/20 Lupinus spp. ■ - - - - - - - - - - - - - - - - - - - - (Mevi) - - - - - - 1/3 - - - - - - - - . (Mofi) - - - - - - - - - - (Mudi) ' tr/17 - - 1/10 - - - - - - 1/10 (Mysy) - - ~ - - 2/13 - - - - - - - - - - - - (Oppo) • --- ~ ~ (Orle) ---- - - - - 143 Table 16. (continued)____________________________________________ .______________ ____ Sites Taxa I 2 3 4 . 5 6 7 8 9 FORBS: Orthocappus PenufoZia(Orte) Oxytvopis Toesseyi(Oxbe) Oxytvopis sevieeus(Oxse) Oxytvopis visoida(Oxoj±) Pavoynohia sessifZova(Vase) Pensiemon attenuates(Peat) Penstemon evianthus(Peer) Penstemon nitidus(Peni) PetaZostemon oandidum(Peca) Petdlostemon puvpescens(Pepu) Phlox dlyssifolia(Phal) Phlox hoodii(Pbbo) Phlox kelseyi(Pbbe) Phlox multiflova(Phmu) Polygala alba(Poal) Polygonum bistovdis(Pobi) Potentilla gvaoillis (Pogr) Potentilla hippiana(Pohi) tr/7 1/20 4/33 3/37 - - 1/3 - - 2/13 _ _ _ _ — — — — — — — — — — 1/3 — — — — - - tr/3 tr/3 - - - - - - - - tr/10 - - 8/33 1/10 1/13 1/13 - - 1/3 — — tr/13 4/40 — — 3/30 - - - - - - 1/7 1/10 - - - - - - - - 2/27 4/43 1/23 1/13 1/3 1/7 - - 1/23 - - tr/3 - - - - - - - - - - - - - - - - — — — — — — — — — — tr/3 — - tr/3 — — 144 Table 16. (continued) Sites Taxa 10 11 12 13 14 15 . 16 17 18 19 20 FORES: (Orte) - - - - ~ - ~ ------- - ~ - ~ ~ - ~ - - - - - (Oxbe) - - - - - - - - - - - - - - - - - - - - - - (Oxse) 2/17 1/87 - - 1/7 - - - - - - - - 1/47 . 2/17 (Oxvi) - - - - - - - - - - - - - - - - - - - - - - (Pase) — — tr/3 — — — — — — — — — — 1/17 — — — — 1/7 (Peat) - - - - - - - - - - - - - - - - - - - - - - (Peer) --- ---. --- --------- --- --- --- --- --- --- (Peni) ' - - - - - - - - - - 1/3 ' (Peca) - - - - - - - - - - - - - - - - - - - - - - (Pepu) --- --- --- ---'----- ---■ ---- - - --- -- --- (Phal) - - 3/37 2/20 1/3 1/7 - - 2/17 3/23 10/43 (Phho) - - 1/13 - - - - - - - - - - - - - - - - (Phke) . 2/23 - - 1/17 3/23 2/27 - - - - (Phmu) - - 1/13 - - - - - - - — - -- - - 1/3 (Poal) - - - - ~ ~ - - - - - - (Pobi) - - - - - - - - - - - - 1/53 (Pogr) 1/1.7 - - - - 1/3 - - -• - - - - - - - tr/3 (Pohi) - - 1/3 - - - - - - - - 1/3 145 Table 16. (continued) Sites Taxa 21 22 23 25 26 27 28 29 30 31 FORES: (Orte) - - - - - - - - - - - - . - ~ 1/7 (Oxbe) - - - - - - - - - - - 4/43 3/33 (Oxse) - - ' 12/90 - - - - - - - - - - - - 1/13 - - (Oxvi) - - - - - - - - - - - - - - - - - 1/27 - - (Pase) - - - - - - - - - - - - - - - - - - - - (Peat) - - - - _ _ _ _ - - - - - - - - - - - - (Peer) - - - - - - - - - '-- - - . - - - - - - - - (Peni) - - - - - - - - --- - — - - - - - - - - (Peca) 1/20 - - - - - - --- 1/10 - - - - 2/27 (Pepu) - - - - - - . - - -- - - - - - 1/10 (Phal) 3/37 - -. 4/43 - - - - - - - - - - - - (Phho) - - - - - - - - - - (Phke) . - - - - - - - - 1/3 1/17 - - 1/13 7/63 2-/10 (Phmu) - - - - - - - - - - - - 1/7 (Poal) - - - - - - - - tr/3 - - - - - - - - , '- - (Pobi) - - - - ---tr/3 - - (Pogr) - - - - - - - - - - - - - - 2/23 (Pohi) . 1/3 - - . - - - - - - - - - - , tr/3 146 Table 16. (continued) Taxa Sites I 2 3 4 5 6 7 3 9 FORBS: Ranunculus glabevrimus(Ragl) Saxifraga rhomboidea(Sarh) Sisyrinchium sarmentoswn(Sisa) Sedum IanceoVatum(Sels) tr/7 tr/3 Senecio canus (Seca) — — tr/3 - - - - tr/7 — — Smilieia raeemosa(Smra) Solidago missourensis(Somi) tr/17 tr/7 1/3 Solidago. ocaidentalis (Sooc.) ■ Tarasieum offieianle(Taof) tr/10 Thermoipsis rhombifolia(Thrh) 3/47 3/37 1/3 — — — 11/70 Tragopogon dubius(Trdu) — — tr/3 — — 1/3 Vicia Omericana(Vram) tr/3 — — tr/3 - - 1/7 Viola nuttalli (Vrwr) Tueoa glauca(Yugl) Zygodenus elegans(Zyel) tr/3. 1/10 Zygodenus venosus(Zyve) tr/3 — — tr/3 - - - 1/20 3/43 147 Table 16. (continued) Taxa Sites 10 11 12 13 14 15 ' 16 17 18 19 20 FORBS: (Ragl) (Sarh) (Sisa) (Sela) (Seca) (Smra) (Somi) (Sooc) (Taof) (Thrh) (Trdu) (Viam) (Vinu) (Yugl) (Zyel) (Zyve) - - - - - - - - - - - - - - - - 1/7 - - - - ■ tr/20 - - - - - - - - - - - - - - - - - --' 1/10 2/17 1/20 1/7 - - --- - - - - - - - - - - 1/7 — — — — 1/20 — — — — — — — — — — - — — — — — — 1/7 - - - - 2/27 - - - - - - - - - - - - - - 1/7 - - tr/3 - - - - • T - - - '-- - - - - 1/13 - - - - 1/10 - - - - - - - 3/23 18/57 5/30 - - 1/3 - - - - - - - tr/3 — — 3/23 1/3 2/20 1/3 1/3 — — — — 2/10 2/7 - - tr/7 tr/3 2/27 2/10 tr/3 7/57 1/7 1/3 1/20 tr/3 1/7 1/3 - - - - - 1/13 3/37 — — — — 148 Table 16. (continued)___________ ;___________________________________ ;__________________ Sites Taxa 21 22 23 25 26 27 28 29 30 31 FORBS (Ragl) _ 1/3 _ _ _ _ _ _ (Sarh) - - - - 1/3 - - - - - - - - - - - - (Sisa) - - - - - - - - - - . (Sela) - - - - T- - - tr/3 - - - - _ _ ■ _ _ (Seca) - - - - - - - - - - - - tr/3 - - (Smra) - - - - - - - - - - - - - - - - (Somi) - - - - tr/3 - - - - . - - tr/7 - - (Sooe) - - - - - - - - - - - - - - (Taof) — — — — 2/10 — - — — - - — - — - — — — — (Thrh) - - 1/10 - - - - 1/3 - - 1/3 4/57 (Trdu) - - - _ - - - - - - J - - - - (Viam) tr/10 - - 3/53 - - - - - - - tr/3, ■ - - tr/10 (Vinu) - - - - - - - - 1/7 (Yugl) . - - - - . - - - - - - - - - - . - - (Zyel) , 1/3 ‘ 6/57 ' - - 10/70 - - - - - - (Zyve) 1/7 - - ' - - - - - - - ' - - 1/3' - - tr/3 149 Table 16. (continued) Taxa Sites I 2 3 4 ' 5 6 7 8 9- SHRUBS Apooynum cannabium (Apca) ■ Arctostayphlos uva-ursi(Aruv) Artemisla longifolia(Artlo) Artemisia tridentata(Artr) Atriplex canescens(Atca) Berberis. repens (Bere) Juniperus horizontalis(Jubo) Potentilla frutieosa(Vofr) Prunus VirginiarUa (Prvi) tr1?* Iobata (Rhtr) Ribes spp. Rosa acieularis(Voae) Rosa arkansana(Roar) Symphoriearpos albus(Syal) Symphoriearpos occidentalis(Syoe) TREES 'Pinus flexilus(Vifl) Pinus ponderosa(Vipo) Pseudotsuga mensiesii'(Vsme) 1/3 43/60 19/67 33/73 31/77 35/83 36/33 34/67 16/43 16/23 1/3 - - - - - - - - - - - - - - 4/17 1/13 1/40 / 1/20 1/15 - - 1/22 22/91 48/138 1/13 2/21 — - 23/47 - — - - tr/5 150 Table 16. (continued) Taxa Sites 10 11 12 13 14 15 16 17 18. 19 20 SHRUBS: (Apca) (Aruv) (Artlo) (Artr) (Atca) (Bere) (Juho) (Pofr) (Prvi) (Rhtr) Ribes spp. (Roac) (Roar) (Syal) (Syoc) TREES (Pifl) (Pipo) (Psme) 5/33 — — — — — — 3/13 5/23 - - - - - - - - - - - - - - - - - - - - t r / 3 27/53 47/77 30/47 53/73 16/27 17/33 35/50 32/73 28/43 40/73 9/37 4/20 3/20 4/30 - - - - - - - - - - - - - 13/50 9/30 - - - - - - - - - - - - - - - - 1/13 2/10 5/13 - - - - - - 1/3 - - tr/3 1/13 2/27 - - - - 4/27 tr/3 - - - - --- - - - - 3/23 1/13 — — — — — — 14/73 — — — — — — — — — — 1/7 1/3 3/23 2/10 2/93 1/93 1/7 - - - - - - tr/2 - - . --- - - - - - - 4/160 ' 151 Table 1.6. (continued) Taxa Sites21 22 23 25 26 27 28 29 30 31 FORBS: (Apca) - - - - - - - - - - - - - - 2/10 - - - - (Aruv) - - - - - - - - - - - - - - - - - - - - - (Artlo) - - - - - - - - - - - - - - - - - - - - (Artr) - - --' - - - - - - --■ - - - - - - - - - (Atca) - - - - - - - - - - - - - - - - 1/10 - - (Bere) - - - - - - - - - - _ _ _ _ - - - - - - (Juho) 63/80 26/63 11/60 57/90 56/63 83/100 25/53 10/27 27/40 20/33 (Pofr) - - 5/27 - - - - - - — - - - - - - - - - (Prvi) - - - - - - - - - - - - - - - - - - (Rhtr) ~ — — ~ — — ~ — ~ ~ — - ~ ~ 1/3 — — — — R-Lbes spp. - - - - - - - - - - - - - - - - - - - - (Roac) - - - - - - - - - - --- 2/13 .- - 2/13 - - (Roar) - - 1/3 - - - - - - - - - - - -' - - - - ■. (Syal) ’ -- - " , ~ ------------- ---. ---- -- (Syoc) . - - - - - - - - tr/3 . - - - -' - - - - TREES. . ' (Pifl) 10/70 - - 4/48 5/58 - - - - - -. 3/200 5/220 - - (Pipb) - - - - .. - - . tr/2 - - — — - - - - - - - - (Psme) . 1/23 - - '- - - - - - - - -. - - - -- 152 Table 17. Edaphic characteristics of 51 creeping juniper sites including pH, organic matter content, texture, salt hazard, soil depth, erosion, and five important elements. Sites PH Organic Matter Percent Tex­ ture1 Phos­ phorus ppm2 Potas­ sium ppm Magne­ sium meq3 Sod­ ium meq Cal­ cium meq Salt Hazard mmhos1* ■ Soil Depth cm Surface Erosion Percent I 7.8 4.5(M)5 CL 2.(VL) 199 (M) 2.1 Tr.6 36.0 0.6 23.6 3.0 2 7.7 3.2(L) SL 17(VL) 162 (M) 1.3 Tr. 28.4 . 1.0 12.2 15.0 3 7.0 3.9(1) L 18(VL) 107(L) 1.5 Tr. 12.8 1.3 16.8 3.0 4 7.6 2.7(VL) LS 18 (VL) 71(VL) 1.2 Tr. 11.2 0.7 17.9 15.0 5 7.5 3. 3(L) L 16(VL) 171 (M) 1.5 0.1 20.0 0.9 13.4 15.0 6 6.9 3.7(L) CL 9 (VL) 208(M) 3.6 Tr. 15.6 0.2 . 10.1 15.0 7 7.8 3.8 (L) CL 14(VL) 180 (M) 0.7 Tr. 34.0 • 0.6 9,2 38.0 8 6.0 6.2(H) L 26(VL) 199 (M) 2.3 0.1 15.2 1.9. 9.4 3.0 9 6.6 6.3(H) L 28(VL) 539(H) 6.2 0.1 19.2 1.2 10.8' 3.0 10 6.0 5.2 (M) CL IO(VL) 284(H) 4.1 0.2 12.8 0.5 14.7 • 3.0 11 7.6 6.0(H) SiL 13(VL) 255(H) 5.0 0.2 28.0 ■ 1.0 30.7 3.0 12 7.3 6.2(H) SiL 13(VL) 312 (H) 3.6 0.2 24.4 • 1.2 10.6 3.0 13 7.5 • 3.9(L) SiC 13(VL) 293(H) 4.0 0.1 42.0 0.6 27.8 3.0 14 7.5 1.6(VL) C 20(VL) 199 (M) 0.7 0.1 98.0 6.2 22.7 38.0 • 15 4.4 1.8(VL) C 49 (L) 236(M) 2.2 0.2 33.0 1.1 22.5 63.0 16 6.2 2.2(VL) CL 45 (L) 190(M) 3.3 Tr. 22.0. . 0.4 20.5 63.0 17 7.2 6.2(H) SiL 23(VL) 351(H) 2.4 0.2 29.2 1.4 12.2 15.0 18 7.4 6.3(H) L 17(VL) . 519(H) 2.3 Tr. 36.4 2.0 11.0 3.0 19 7.2 7.4(H) SiL 23(VL) 419(H). 2.5 0.1 32.0 1.4 13.8 3.0 20 7.6 6.8(H) SiL 43 (L) 439(H) 2.0 Tr. 37.6 0.8 10.3 3.0 21 ■7.3 6.8(H) L . 21(VL) 399 (H) 3.0 Tr. ■ 39.0 1.7 17.5 3.0 22 7.9 4.8 (M) L 6 (VL) 144 (M) 2.9 Tr. 34.0 0.8 14.8 3.0 • 23 8.0 3.8(L) CL 9 (VL) 312(H) 1.9 Tr. 21.0 0.9 20.8 15.0 25 6.4 6.8(H) L 20(VL) 488(H) 2.4 0.1 16.4 0.5 12.8 3.0 26 7.9 ■ 3.9(L) SiL . 7 (VL) . 190(M) 3.9 0.1 30,4 0.6 15.5 . 15.0 27 8.0 4.8 (M) SC ■ 7(VL) 295(H) 7.3 Tr. 27.2 0.7 16.1 3.0 . 28 8.5 I-O(VL) CL 2 (VL) 69(VL) 3.7 Tr. 23.2 0.4 32.6 38.0 29 7.1 4.5(M) ■ CL 14(VL) 463(H) 2.4 .33 24,0 0.4 14.3 38.0 153 Table 17. (continued) Sites pH • Organic Matter Percent Tex­ ture1 Phos­ phorus ppm2 Potas­ sium ppm ■ Magne­ sium • meq3 Sod­ ium meq Cal­ cium meq Salt Hazard Imnhoslf . Soil Depth cm Surface Erosion Percent 30 8.0 2.9(VL) SCL 6 (VL) 23 9 (M) ■ 4.6 Tr. 31.6 0.4 16.4 15.0 31 . 7.7 5.8(H) CL 17(VL) 415(H) ■ 0.7 0.1 ■ 33.6 0.6 14.4 ' 3.0 32 6.7 3.8(L) CL 103(H) 200 (M) 1.3 Tr. ■ . 9.6 6 . 4 N.M.7 . 3.0 33 7.5 1.5(VL) C . H(VL) 184 (M) 1.1 0.3 35.4 3.7 11 15.0 34 7.9 2.1(VL) c .• 4 (VL) . 236 (M) 3.4 0.3 37.1 0.4 . Il . 3.0 . 35 7.5 5.9(H) CL 6 (VL) 333(H) 5.3 0.4 28.0 ' 0.7 3.0 36 8.0 I.I(VL) SCL 8 (VL) 120(L) 3.3 Tr. 31.2 0.4 Il 15.0 37 7.8 4.5(H) CL ■ 9(VL) 390(H) 2.6 Tr. 35.2 6.8. Il 3.0 38 7.7 3.9(L) C .7 (VL) 290 (M) 1.4 0.1 40.0 0.6 . ' n 3.0 39 6.5 6.6(H) SCL 14(VL) 333(H) 2.8 0.4 17.5 0.3 Il 3.0 • 40 8.3 0.8(VL) SL 7 (VL) IlO(L) 3.3 Tr. . 25.2 0.4 II 15.0 41 8.2 0.8(VL) CL . 6(VL) 160(H). 3.2 0.1 31.6 0.7 Il 38.0 42 6.9 3.5 (L) SCL 9 (VL) 201 (M) ■ 2.5 0.3 10.0 0.2 Il 3.0 43 7.4 4.8 (M) CL 2 (VL) . 342(H) . 4.4 0.3 36.6 0.5 11 3.0 44 8.1 3.4(L) C 4 (VL) 229(M) 4.3 0.3 28.4 0.4 . If 63.0 45 8.0 6.4(H) SL . 6 (VL) 218 (M) . 3.0 0; 3 28.0 0.6 If 3.0 • 46 8.1 2.9(L) CL 6 (VL) 210 (M) 3.5 Tr. 37.2 0..5 11 38.0 47 8.0 5.9(H) SCL 6 (VL) 184 (M) 3.7 0.3 37.1 0.6 Il 38.0 48 8.1 2.7(L) CL 6 (VL) 218 (M) 4.3 0.3 36.6 0.5 II. 15.0 49 8.2 . 1.8(VL) CL 15(VL) IlO(L) 2.0 Tr; 24.4 0.4 11 63.0 50 7.0 6.3(H) SiL 19(VL) 340(H) 1.9 Tr. • 37.2 0.8 Il 3.0 51 8 . 0 I.8(VL) SC 6 (VL) 218 (M) 1.0. 0.4 34.6 .0.4 If 63.0 24 7.0 7.0(H) L . . 28(VL) 580(H) 1 . 6 Tr. 32.4 1.5 11.2 ■' 3.0 ^Sii1=Silt Loam; L=Loam; CL=ClayLoam; SCL=Sandyclay Loam; C=Clay; LS=Loamy Sand; SC=Sandy Clay; SiG=Silty Clay. . 2ppm=poundsybne million pounds of soil. 3meq=Millequivalents/100 grams of soil. ^mmhos=Tnillimhos Imho=I/ Ohm 5Ratings from Montana Soils Testing Laboratory Report, ST-Form 2; VL=Very Low; M=Medium; H=High. 6Tr.=Trace; less than 0.1 percent ?N.M.=hot measured. 154 MONTANA 3 1762 UNIVERSITY LIBRARIES 001 4939 O