Range ecology and relations of mule deer, elk, and cattle in the Missouri River Breaks, Montana by Richard John Mackie A thesis submitted to the Graduate Faculty in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in Fish and Wildlife Management Montana State University © Copyright by Richard John Mackie (1965) Abstract: A study was conducted during the four-year period 1960-64 to obtain quantitative data on food habits, range use, and relations of mule deer (Odocoileus hemionus), elk (Cervus canadensis), and cattle on ponderosa pine-juniper (Pinus ponderosa/Juniperus scopulorum) and related vegetation types in the Missouri River "Breaks" in northcentral Montana. Eight vegetation types comprising 12 distinct communities were analyzed and described. Totals of 11,581 mule deer observations, 3,489 elk observations, and 25,125 cattle observations were recorded during morning and evening periods. Distributions, movements, and population trends during seasons and years were related to range use and forage conditions. Seasonal and yearly distributions of observations and animal activities were related to sunrise and sunset and environmental conditions, Group sizes during seasons and years were discussed in relation to social behavior and range use. Seasonal, yearlong, and year-to-year trends in relative use and importance of vegetation types were evaluated with respect to animal activity, forage utilization, plant growth, and weather conditions. Relative frequencies of occurrence of each of the three animals on slopes of various degrees of steepness and exposure were discussed. Relations between distributions of mule deer, elk, and cattle use on the area and distributions of water sources were described. Instances of plant use at 263 mule deer, 178 elk, and 184 cattle feeding sites were related to animal use of the various vegetation types to ascertain seasonal food habits. Analyses of rumen samples from 37 mule deer, 12 elk, and 18 cattle supplemented feeding site data. Yearlong trends in relative use and importance of forage classes and individual plant taxa were described. Year-to-year trends in food habits were related to precipitation and its influence on plant growth. Relations between elk and mule deer, elk and cattle, and mule deer and cattle were evaluated with respect to degrees of overlap in distribution of use on the area in time and space in relation to food habits, range use, animal behavior, and condition of "key" forage plants. Conclusions and recommendations relating to mule deer, elk, and "multiple-use" management programs on the area were presented.  RANGE ECOLOGY AND RELATIONS OF MtJLE DEER, ELK, AND CATTLE IN THE MISSOURI RIFER BREAKS, MONTANA by RICHARD JOHN MACKIE . A thesis submitted to the Graduate Faculty in partial fulfillment of the requirements for. the degree Of . DOCTOR OF PHILOSOPHY in ' Fish and Wildlife Management Approved: — - Hedqy Major Department ___ Chairman, Examining Committee Dean, Graduate Division MONTANA STATE COLLEGE Bozeman/ Montana June, 1965 iii ACKNOWLEDGMENT To the following among others, I wish to extend sincere appreciation for their contributions to this study: Dr. Don C. Quimby, Montana State College, who provided invaluable advice and assistance in preparation of the manuscript j Mr. G-Ien F. Cole, formerly of the Montana Fish and Game Department, for initial project planning and assistance during the first two years of the projectj Dr. Robert L. Eng, Montana Fish and Game Department, for advice, assistance, and critically reading the manuscript5. Mr. Thomas H. Leik, Montana Fish and Game Department, for assistance in preparing the field data for analyses by automatic data processing equip­ ment and with statistical analyses; Dr. WoE= Booth, Montana State College for verification of plant specimens; Dr; Richard J. Graham and Dr. P.D. Skaar, Montana State College, for constructive criticisms of the manuscript; Dr= John H. Rumely, Montana State College, for reading the vegetation analysis; Mr. R.G. Janson, Mr. Merle Rognrud, Mr. R.B. Knight, Mr. Arnold Foss, Mr. Kenneth Greer, Mr. Robert Varner, and other personnel of the Montana Fish and Game Department for assistance in various phases of the project; Personnel of the Bureau of Land Management Lewistown District Office and the Charles M. Russell Wildlife Range for their cooperation; and my wife, Barbara, for patience, encouragement, and assistance in preparation of the manuscript. The writer was employed by the Montana Fish and Game Department throughout the study under Federal ' Aid Projects W— 98—R-^ -I, W—9S—R—2, W —98—R=3# W—:98—R-A, and W—98—R—5° TT iv TABLE OF CONTENTS : * At o iT v A B L E A O 0 0 0 0 0 0 0 0 0 « INTRODUCTION . . . . . . . . . DESCRIPTION OF THE STUDY AREA Physiography . . . . . . . Clunah 6 . . . . . . . . . . Land-Use Characteristics ' . VEGETATION Artemisia/Agropyron Vegetation Type . . . . . . . . . Artemisia tridentata/Agropyron spicatum Association Artemisia tridentata/Agropyron smithii Associes -. , Artemisia tridentata/Agropyron smithii/Boutelona gra Assod3.*fciion 0 .0 . 0 0 0 ’ 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Agropyron/Poa Vegetation Type . . . . . . . . . . . . . Pinus/juniperus Vegetation Type . . . . . . . . . . . . Pinus ponderosa/Agropyron spicatum Association . . . Pinus ponderosa/Juniperus scopulorum Association . . Pinus ponderosa/Artemisia lohgifolia Association . „ Pseudot suga./Juniperus Vegetation Type . . . . . . . . Sarcobatus/Artemisia Vegetation" Type . . . . . . . . . Agropyron/Symphoricarpos Vegetation Type . . . . . . . . Xanthium strumarium Vegetation Type . . . . . . . . . . Artemisia longifolia Vegetation Type . . . . . . . . . Other■Vegetation Types . . . . 6 . . . . . . . . . . . POPULATIONS o o o o o o o o o o o e Mule Deer Distribution . . ,7. Movements and Home'Rahge' T A ' Population Trends Elk . Distribution Movements. . Population Trends Cattle Distribution Population Trends 6 ,0 6 0 0 6 6 0 0 0 0 6 0 0 6 0 0 0 0 0 0 6 0 0 0 0 0 0 0 0 0 6 0 0 0 0 0 O O O ■ O O 6 O O O O O 0 0 0 6 0 0 0 0 0 6 0 0 0 0 6 0 0 0 0 0 Page avi I . 3 14 14 .20 20 23 25 26 32 33 35 39 41 44 46 48 49 49 52 59 63 63 66 69 72 72 74 xQ O sV J VTABLE OF CONTENTS— Continued RANGE USE O O O O O O O O O O O O O O O O O O Niule Deer o o o o o o o o o o o o o o o o Activity Habits ......................... GrOUp SlZe O O O O O O O O O O O O O O Use of Vegetation T y p e s ............. . Summer (June, July, August) . . . . Fall (September, October, November),. Winter (December, January, February) Spring (March, April, May) . . . . ■ Yearlong and Year-to-Year Trends » . Use of Slopes » 0 0 0 0 0 0 0 0 0 0 0 0 Use of Exposures 0 0 0 0 0 0 0 0 0 0 0 Relations to Water » » , » 0 0 0. « 0 , Elk0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Activity Habits 0 0 0 0 0 0 0 0 0 0 0 Group Size 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Use of Vegetation Types 0 0 0 0 0 0 0 0 Summer (June, July, August) . . . . Fall (September, October, November) Winter (December, January, February) Spring (March, April, May) ......... Yearlong and Year-to-Year Trends „ „ Use of Slopes 0 0 0 0 0 0 0 0 0 0 0 0 0 Use of Exposures ......... 0 0 0 . 0 0 Relations to Viater 0 0 0 0 0 0 0 0 0 0 Cattle 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Activity Habits 0 0 0 0 0 0 0 0 0 0 0 » Group Size » 0 0 0 0 0 0 0 0 0 0 0 0 0 Use of Vegetation Types » '0 0 » . » , » Summer (June, JtSly, August) . 0 . 0 Fall (September, October, November) . Winter (December, January, February) Spring (March, April, May) 0 . 0 . Yearlong and Year-to-Year Trends . » U Se of Slopes 0 0 . » 0 0 0 0 0 0 0 0 0 0 Use of Exposures 0 0 0 0 . 0 0 . 0 0 0 Relations to Water . . . . . » . . . » FOOD HABITS 0 0 0 0 0 0 6 0 6 0 0 0 0 0 0 0 Mule Deer . . « . . » . . » . , Summer (June, July, August) , 0 O O O 6 O O Page 76 * 76 O 0 0 O O 0 0 77 O O O 0 O O O SI 0 • 0 0 0 0 83 O O O O 0 O 83 O O 0 O O O O 8 5 . O O 0 O 0 6 0 87 O 0 O O O O O SS O O 0 O O O O S9 O o' 0 O O 0 0 91 O O O O O O O 92 O O 0 0 O 0 0 92 O 0 0 O 0 0 O 95 O 0 0 6 0 0 0 95 O 0 0 O 0 0 0 99 O O O O O O 0 100 O O O O O 0 0 103 O 0 0 O O 0 0 104 O O O O O O 105 0 a a a 1 0 6 0 O O » O O O 106 » O O O O O O 108 O O 0 O O 0 0 109 O O O O O O O H O O O O O O O O 112 O 0 O O O O O 112 O O O O O O 0 115 O 0 0 0 0 0 0 115 O O O O 0 O O 117 0 O 0 O a O O 118 O O O O O O O 118 O O O 0 O O 0 • 119 0 O O O O O O 120 O O 0 O O O O 121 O O O O O O O 121 0 0 0 0 0 0 0 123 0 0 0 0 0 0 0 125 126 O 0 0 O O 0 O 126 TABLE OF CONTENTS— Contizmed Fall (Septembers Oetober5, November) , Winter (Decembers Januarys February) Spring (Marehs Aprils May), Yearlong Trends , . , . ............. Year-to-year Trends „ „ 0 » « » » Elk t .0 0 0 ■ 0 0 0 0 0 0 0 0 0 0 0 0 0 0 < Summer (Junes Julys August) . . . . , Fall (September,, Octobers November) , Winter (Deeembers Januarys February) Spring (Marehs Aprils .May) . . . . , Yearlong Trends . . . . . . . . 0 . < Year-to-Year Trends . . . . . . . . . Cattle 0 0 . 0 0 0 0 0 0 0 0 0 0 0 0 0 . Spring (Aprils May) . . . . . . . . . Summer (Junes Julys August) . . . . • , Fall (Septembers Octobers November) . Winter (December9 January) . . . . , Yearlong Trends . . . . . . . . . . . Year-to-Year Trends . . . . . . . . . 0 6 0 ' -v 0 0 0 0 Page 133 135 136 138 141. 147 147 131 154 155 1$6 160 164 164 16? 170 171 172 174 RANGE RELATIONS . . . . . o . . . Elk and Mule Deer Relations . Elk and Cattle Relations . . . Mule Deer and Cattle Relations 178 178 182 184 CONCLUSIONS AND RECOMMENDATIONS FOR MANAGEMENT . o o o o o o o o o 18& SUMMARY O O O O ’ O O O O , 0 0 0 0 0 0 0 0 0 0 0 0 O 0 . 0 0 O O O 189 APPENDIX 198 LITERATURE CITED 227 vii TABLE LIST OF TABLES - Page Io • PRECIPITATION .DATA FOR MISSOURI RIVER BREAKS STUDY AREA, 1959-1964o UoSo DEPARTMENT OF COMMERCE, WEATHER BUREAU STATION, ROY 8 NE, MONTANA , 0 » . . . . . ........... 7 IIo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBU­ TION BY UNIONS OF TAXA ON THE ARTEMTSIA/AGROPIRON TYPE, » 15 !Ho GROUND-COVER CHARACTERISTICS FOR TWELVE RANGELAND COMMUNITIES0 DATA ARE FREQUENCIES OF OCCURRENCE OF BARE GROUND, LITTER, AND LIHNG PLANT MATERIAL AMONG 480 POINTS (160 IN EACH OF THREE STANDS) IN EACH COMMUNITY. I? IVo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA ON THE AGROPYRON SMITHIl/POA'SEGUNDA ASSOCIES O O O O O O O O O Q O O O O O O O O O O O O O O O 2^ Vo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXj'":ON ' THEi PINUS/JUNIPERUS TYPE . . / V ' . 27 Ho CONSTANCY, COVERAGE, AND FREQUENCY OF TREES AND SHRUBS GREATER THAN 12 INCHES TALL ON THE'PINUS/JUNIPERUS TYPE. . 30 HIo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY .AND DISTRIBUTION BY UNIONS OF TAXA ON THE PSEUDOTSUGA MENZIESIl/ JUNIPERUS SOOPULORUM ASSOCIATION . . . . . . . . . . . . . 37 HIIo CONSTANCY, COVERAGE, AND FREQUENCY OF TREES AND SHRUBS GREATER THAN 12 INCHES TALL ON THE PSEUDOTSUGA MENZIESIl/ ,TUNT PF-RUS SCOFUIORDM ASSOCIATION 39 IXo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA ON THE SARGOBATUS VERMICULATUS / ARTEMISIA TRIDENTATA ASSOCIATION. THE AGROPYRON SMITHIl/ . SYMPHORICARPOS OCCIDENTALIS ASSOCIATION, AND THE XANTHIUM STRIIMflRTUM ASSOCIATION . . . . . . . . . . . . . . . . . . . 42 Xo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA ON THE ARTEMISIA LONGI FOLIA ASSOCIES . . 47 Ho SUMMARY OF MOVEMENT DATA FOR MARKED AND RECOGNIZABLE MU RANGA DEER 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 53 viii TABLE LIST OF T A B m - Continued . HIo MOVEMENTS OF FAOT MULE DEER DURING FIRST SUMMER AID FAIL XTTT „ SEASONAL AND ANNUAL TREND IN NUMBERS OF MULE .DEER ON THE MISSOURI RIVER BREAKS STUDY AREA, 1960-64 . . =-„ . . XEVo • SEASONAL AND ANNUAL MOVEMENT OF RECOGNIZABLE ELK ON THE . MISSOURI RIVER BREAKS STUDY AREA. ; . . . . . . . . . . XV. WINTER ELK POPULATIONS ON THE MISSOURI RIVER BREAKS STUDY AREA, I960—64 o . . . . . . . . . . . . . . . . . . XVI. AVERAGE SIZE OF GROUPS FOR MULE DEER OBSERVED DURING SEASONS AND YEARS, 1960-64 . . . . . . . . . . . . . . . XVII. PERCENTAGES OF TOTAL MULE DEER OBSERVED ON SLOPE CLASSES, BY SEASON AND YEARS, I96O-64 . . . . . . . . . . . . . . . XVIII0 PERCENTAGES OF TOTAL MULE DEER OBSERVED ON VARIOUS EXPOSURES, BY SEASON AND YEARS, 1960-64 .. . . . . XIX. PERCENTAGES OF TOTAL MULE DEER OBSERVED AT ONE-FOURTH MILE DISTANCE INTERVALS FROM KNOWN WATER SOURCES, BY SEASON AND YEARS, 1960=64 . . . . . o . . . . . . . . . . XX. AVERAGE SIZE OF GROUPS FOR ELK OBSERVED DURING SEASONS AND YEARS, 1960=64. o . . . . . . . . . . . . . . . . . . XXIo PERCENTAGES OF TOTAL ELK'OBSERVED ON SLOPE CLASSES, BI SEASON AND YEARS, 1960=64 o . . . . . . . . . . . . . . . XXEI. PERCENTAGES OF TOTAL ELK OBSERVED ON VARIOUS EXPOSURES, BY SEASON AND YEARS, 1960=64 . . . . . . . . . . . . . . XXEII. PERCENTAGES OF TOTAL ELK OBSERVED AT ONE-FOURTH MILE DISTANCE INTERVALS FROM KNOWN WATER SOURCES, BI SEASON AND YEARS, 1960=64 . . . . . o . . . . . . . . . . . . . XXEVo AVERAGE SIZE OF GROUPS FOR CATTLE OBSERVED DURING SEASONS AND YEARS, 1960=64 . . o . . . . . . . . , . . . . . . . . XXVo PERCENTAGES OF TOTAL CATTLE OBSERVED ON SLOPE CLASSES, BI SEASON AND YEARS ,; 1960-64 . . . . . . . . . . . . . . . . . Page 61 70 71 82 91 93 94 100 108 109 111 H S 122 ix TABLE LIST OF TABIES=-Oontinued Page XXVI, PERCfflTAGES OF TOTAL CATTLE OBSERVED ON VARIOUS EXPOSURES, BI SEASON AND YEARS, 1960-64 . . . . . . . . . . . 122 XXVII, PERCENTAGES OF TOTAL CATTLE OBSERVED AT ONE=FdUR1TH MILE DISTANCE INTERVALS FROM KNOWN WATER SOURCES,' BI SEASON AND YEARS, I960—64. , . . , .... . , . . 1 2 4 XXVIIIo MULE DEER USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING.AT LEAST ONE SEASON, 1960-64 COMBINED . . . . . . . . . . . . . . . . 127 XXIX, FREQUENCY OF OCCURRENCE AND VOLUME PERCENTAGES OF PLANT TAXA AND FORAGE CLASSES AMONG 37 MULE DEER HJMEN SAMPLES 131 XXX. A COMPARISON OF PERCENTAGES OF MULE DEER USE OF GRASS, BROWSE, AND FORBS AT FEEDING SITES ON VARIOUS VEGETATION TYPES DURING SEASONS AND YEARS, 1960-64 . . . . . . . . . . 142 XXXI. A COMPARISON OF RELATIVE INTENSITIES OF MULE DEER USE OF SOME IMPORTANT FOOD.PLANTS DURING SEASONS AND YEARS, 1960-64 0 0 0 0 . 0 0 0 0 0 0 0 . 0 0 0 0 0 . o . . . . . . 143 XXXII, ' PERCENTAGES OF MULE DEER FEEDING SITES EXAMINED DURING SUMMER AND FALL SEASONS ON WHICH MELILOTUS OFFICINALIS WAS ABUNDANT, SCATTERED, AND ABSENT, 1960=63 . . . . . . 143 XXXIII, ELK USE OF TAXA RECEIVING AT LEAST TEN PER CENT OF THE TOTAL INSTANCES OF , PLANT USE AT FEEDING SITES ON AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1960—64 COMBINED . . . . . . . . . . . . . . . . . a . . . 14S XXXIV. FREQUENCY OF OCCURRENCE AND VOLUME PERCENTAGES OF PLANT TAXA AND FORAGE CLASSES AMONG 12 ELK RUMEN SAMPLES . . . 152 XXXV. A COMPARISON OF PERCfflTAGES OF ELK USE OF GRASS, BROWSE, AND FORBS AT FEEDING SITES ON VARIOUS VEGETATION TYPES DURING SEASONS AND YEARS, 1960=64 . . . . . . . . . . . . . 161 XXXVI. A COMPARISON OF RELATIVE INTENSITIES OF ELK USE OF SOME IMPORTANT FOOD PLANTS DURING SEASONS AND YEARS, 1960-64 162 TABLE LIST OF TABLES— Continued Page XXXVIIo CATTLE USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1961-63 COMBINED, , c o o , , , , , , , , , , , , , , , , 165 XXXVIII0" FREQUENCY OF OCCURRENCE AND VOLUME PERCENTAGES OF PLANT TAXA AND FORAGE CLASSES AMONG CATTLE RUMEN SAMPLES „ „ , ' l6S XXXIXo A COMPARISON OF PERCENTAGES OF CATTLE USE OF GRASS, BROWSE, AND FORBS AT FEEDING SITES ON VARIOUS VEGETATION TYPES DURING SEASONS AND YEARS, 1960-63 . « 176 .XL, A COMPARISON OF RELATIVE INTENSITIES OF CATTLE USE OF SOME IMPORTANT FOOD PLANTS DURING SEASONS AND YEARS, 1961-63 177 X U , CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION' BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6? PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT IN ANY COMMUNITY ON THE ARTEMISIA/AGROPYRON TYPE . , . . . . 1.99 XLIIo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING' CONSTANCY OF LESS THAN 6? PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT ON THE AGROPYRON SMITHIl/POA SECUNDA ASSOCIATION, , , , , 201 XLIIIo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6? PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT ON THE PINUS/JUNSPERUS TYPE . . 0 . . . o . , . 202 XLIV. CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6? PER ■ CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT ON, THE PSEUDOTSUGA‘MENZIESII/JUNIPERUS SCOFULORUM ASSOCIATION 203 XLVo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6? PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT ON THE SARCOBATUS VEEMICULATUS/ARTEMISIA TRIDENTATA ASSOCIATION, THE AGROPYRON SMITEIlTSYMPHORICARPOS OCCIDENTALS ASSOCIATION, OR THE XANTHIUM STRUMARlUM ASSOCIATION . . . ■ 204 - I Xl TABLE U S T OF TABLES— Continued Page XLVIo CONSTANCYs CANOPY-COVERAGE, AND FREQUENCY AND DISTRI­ BUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6? PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT ON THE ARTEMISIA LONGIFOLIA ASSOCIES, ... . . . . 205 XLVIIo PERCENTAGES OF TOTAL MULE DEER OBSERVED BI ACTIVITY CLASS DURING SEASONS AND YEARS, 1960=64 = 0 . . . . . . . . . 206 XLVIIIo AVERAGE NUMBERS OF MULE DEER OBSERVED BY ACTIVITY CLASS PER TRIP UNDER VARIOUS ENVIRONMENTAL CONDITIONS. . .. 20? XLIXo PERCENTAGES OF TOTAL MULE DEER OBSERVED ON VEGETATION TYPES BY ACTIVITY, SEASONS, AND YEARS, 1960-64 . . . . . . 208 Lo PERCENTAGES OF TOTAL MULE DEER OBSERVED ON VARIOUS VEGETATION TYPES IN RELATION TO WEATHER CONDITIONS . . . . . 209 Uo PERCENTAGES OF TOTAL, ELK OBSERVED BY ACTIVITY CLASS DURING SEASONS AND YEARS, 1960=64 . . . . . . . . . . . . . . . . 210 U I o AVERAGE NUMBERS OF ELK OBSERVED BY ACTIVITY CLASS PER TRIP UNDER VARIOUS ENVIRONMENTAL CONDITIONS . . . . . . . . . . 211 LIIIo PERCENTAGES OF TOTAL EIK OBSERVED ON VEGETATION TYPES BY ACTIVITY, SEASONS, AND YEARS, 1960=64 . . . . . . . . . . 212 UVo PERCENTAGES OF TOTAL ELK OBSERVED ON VARIOUS VEGETATION TYPES IN RELATION TO WEATHER CONDITIONS . . . . . . . . . 213 LVo PERCENTAGES OF TOTAL CATTLE OBSERVED BY ACTIVITY CLASS DURING SEASONS AND YEARS, 1960=64 . . . . . . . J ...' . 214 LVIo AVERAGE NUMBERS OF CATTLE OBSERVED BY ACTIVITY CLASS PER TRIP UNDER VARIpUS. ENVIRONMENTAL CONDITIONS . . . ...... . 215 LVIIo PERCENTAGES OP'TOTAL CATTLE OBSERVED ON VEGETATION TYPES BY ACTIVITY, SEASONS, ...AND. YEARS, 1960=64 . . . . . . . . . 216/, -s /* .' • I.;'1 ‘ ' ^ LVIIIo PERCENTAGES OF TOTAL CATTLE OBSERVED ON VARIOUS VEGETATION TYPES IN.RELATION TO WEATHER CONDITIONS ... . . . = ". . . 217 )TABLE III. LX. LXI. xli LIST OF TABLES— Continued MEfIE DEER USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST ONE PER CENT BUT LESS THAN TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETA- ' TION TYPE DURING AT LEAST ONE SEASON, 1960-64 COMBINED. . ELK USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST ONE PER CENT BUT LESS THAN TEN PER CfflT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1960-64 COMBINED. . . . . . . CATTLE USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST ONE PER CENT BUT LESS THAN TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1960-64'COMBINED. . . . . . . Page 218 221 224 xiii- LIST OF FIGUEES Figure I, Hap of the Missouri River Breaks Study Area. ■„ .. . . . . . 2 o Typical 8lBreaks11 of Uplands Adjacent to the Missouri River ■ in Nbrthcentral Montana. 81Piains68 Extend Southward in the Background . . ., . ■.. . . . . . . . . . . .... O 0 0 - 0 6 O 3. Artemisia/Agropyron Vegetation Type„ Ridgetop Occupied by Artemisia tridentata/Agropyron spicatum Association . . . 4. Artemisia/Agropyron Vegetation Type. Ridgetop Occupied by the Artemisia trldentata/Agbopyrbn smithii Assoeies 5o Artemisia/Aeropyron Vegetation Type. Ridgetop Occupied by V the.Artemisia tridentata/Agropyron smithii/Boutelbua gracilis Association, I o O O 0 . 0 O O O O O 0 0 0 0 . 0 O O O O 6° Agropyron/Poa Vegetation Type. Formerly Cultivated ■ Associes . . . . . '-i O ' O O O O O 4 O O O O O ! O O O 7. Pinus/juniperus Vegetation Type. Slight South-facing Slope Occupied by -Pinus ponderosa/Agropyron spicatum Association © o © © © © © © » © © ©. ©. ■ © © © © © © © © ©. 8. Pinus/juniperus Vegetation, Type. Moderate West-facing Shale Slope Occupied by-the'Pinus ponderosa/ 7 Juninerus Scopulorum Association O O O ' © O O O O O O O O O 9< Pinus/juniperus Vegetation Type. Steeps South-facing Shale Slope Occupied by the Pinus ponderosa/Artemisia L^onj||ifoli3^ Association © ©.© © © © © © © o © © © © © © - © © © Pseudotsuga/Juniperus Vegetation Type.. Steep North- . facing Slope Occupied by the Pseudotsuga menziesii/. Juniperus scopulorum Association . ... . . . . . » .... Sarcobatus/Artemisia Vegetation Type. Coulee-bottom Bench Occupied by the Sarcobatus Vermieulatus/Artemisia tridgSfttata Association . . . . . . . . . . . . . . . ■ ■. . . . Agropyron/Symphoriearpos Vegetation Type. Coulee-bottom Meadow Occupied by the Agropyron smithii/Symphoricarpos occidentalis Association . . . . . . . . . . . . . o . . . Page 4 .. 5 18 . 18 . 21 . 21 . 31 . 31 . ! . 34 . 34 . ,40 . . 40 Page xlv Figure LIST OF FIGURES— Gontintted 13o Xanthiium strumarium Vegetatios Type. Coulee-bottom Cut Occupied by the Xanthium strumarium Association . . . . . . 45 14. Artemisia longifolia Vegetation Type. Steeps Loose-Shale Slope Occupied by the Artemisia longifolia Assoeies . . . . 45 15. A Comparison-of Mean Numbers of Mule Deer Recorded Per Observation Trip on Extensive Open Ridgetops (Roadside)=, Predominantly Coulee Areas (Away from Main Roads), and the Entire Study Areas 1960-64° . . . . . . . . . . . . . . 51 16. Movements of Three Marked "Yearling** Males on the Missouri River Breaks Study Area. . . . . . . . . . . . . . '55 17° Movements of Two Marked Female Mule Deer on the Missouri River Breaks Study Area . . . . . . . . . . . . . . 56 18. Summer and Winter Distributions of Elk on the Missouri River Breaks Study Areas 1960-64 . . . . . . . . . . . . . 64 19. Movements of an Adult Female Elk from July 1962 to December 1964° The Animal Was Approximately F o w and One-half Years Old When Killed at Last Relocation . . . . . . . . . . . . 6? 20. Movements of a Male Elk Calf from June to December 1962 . ° 6? 21. Movements of a April 1961 . . "Yearling61 Male Elk from July i960 to 68 22. Movements of an Adult Male Elk from January 1.961 to December 1964. The Animal Was Approximately line and One- half Years Old When Killed at last Relocation' . . . . . . . 68 R i« .-,.a • • *' 1 23. Distribution of Cattle Denaities, "Generalised/1 for the April-November Gra^ing^8eas6m and Livestock Wintering Areas on the Missouri River Breaks Study Area . . . . . . . . . . 73 24. Percentages of Total Mule Deer Observed by Hour Relative to Sunrise and Sunset During Seasons . . . . . . . . . . . . . 78 - 25= Percentages of Total Mule Deer Observed by Hour Relative to Sunrise and Sunset During Years. . . . . . . o . . . . . . ... ?8 Page XVr Figure LIST OF FIGURES--Gontiraiaed 26o Percentages of Total Mxxle Deer Observed by Hour Relative to Sunrise/Sxmset in Relation to Temperature 0 . . BO 270 Percentages of Total Mule Deer Observed by Activity Class by Hour Relative to Sxanrise/Sxanset ........... 80 28. Percentages of Total Mule Deer Observed by Activity Class by Years . . . . . . . . . . . . . . . o . . . . . . . . . . . 80 290 Monthly Frequencies of Mule Deer Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the.Total Use During at Least One Month . . . . . . . . . . . . . . . . . .84 30. Percentages of Total Feeding Elk Observed by Hour Relative to Sunrise and Sunset During Seasons . . . . . . . . . . . 97 31. Percentages of Total Feeding Elk Observed by Hour Relative to Sunrise and Sunset Dxaring Seasons . . . . . . . . . . . 97 32. Percentages of Total Elk Observed by Activity Class5 During Normal Activity Periods5 Seasons5 and Years . . . . . . . . 98 33. Monthly Frequencies of Elk Occurrence on Vegetation l^rpes . Receiving More Than Ten Per. Cent of the Total Use During at Least One Month, (see Fig. 34 also) . . . . . . . . . . 101 34. Monthly Frequencies of Elk Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the Total Use During at Least One Month, (see Fig. 33 also) . . . . . . . . . . 102 .35. Percentages of Cattle Feeding=, Bedding5 and Traveling by Hoxxr Relative to Sxanrise/Sunset . . . . . . . . . . . . . . 114 36. Percentages of Cattle Feeding by Hoxxr Relative to Sunrise/ Sunset During Seasons . . . o . . . . . . . . . . . . . . . 114 37» Percentages of Cattle Feeding by Hoxar Relative to Sunrise/ Sunset During Years . . . . . o . . . . . . . . . . . . . . 114 38. Monthly Frequencies of Cattle Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the Total Use Dxxring at Least One Month . . . . . . . . . o . . . . . . . . . . 116 xvi ABSTRACT . A study was conducted during the four-year period I96O-64 to obtain quantitative data on food habits, range use, and relations of mule deer (Odocoileus hemionus), elk (Cervus canadensis), and cattle on ponderosa pine-juniper (Pinus ponderosa/Juniperus scopulorum) and related vegetation types in the Missouri River ‘‘Breaks” in northcentral Montana. , Eight vege­ tation types comprising 12 distinct communities were analyzed and described. Totals of 11,581 mule deer observations, 3*489 elk observations, and 25,12$ cattle observations were recorded during morning and evening periods. Distributions, movements, and population trends during seasons and years were related to range use and forage conditions. Seasonal and yearly dis­ tributions of observations and animal activities were related to sunrise and sunset and environmental conditions, Group sizes during seasons and years were discussed in relation to social behavior and range use. Seasonal, yearlong, and year-to-year trends in.relative use and importance of vegetation types were evaluated with respect to animal activity, forage utilization, plant growth, and weather conditions. Relative frequencies of occurrence of each of the three animals on slopes of various degrees o f . steepness and exposure were discussed. Relations between distributions of mule deer, elk, and cattle use on the area and distributions of water sources were described. Instances, of plant use at 263 mule deer, 178 elk, and 184 cattle feeding sites were related to animal use of the various veg­ etation types to ascertain seasonal food habits. .Analyses of rumen samples from 37 mule deer, 12 elk, arid 18 cattle supplemented feeding site data. Yearlong trends in relative use and importance of forage classes and indi­ vidual plant taxa were described. Year-to-year trends in food .habits were related to precipitation and its influence on plant growth. Relations between elk and mule deer, elk and cattle, and mule deer and cattle were evaluated with respect to degrees of overlap in distribution of use on the area in time and space in relation to food habits, range use, animal behavior, and condition of “key” forage plants. Conclusions and recommen­ dations relating to mule deer, elk, and “multiple-use” management programs on the area were presented. INTRODUCTION Vegetation characterized by ponderosa pine (Finns ponderosa) and rocky mountain juniper (Juniperus scopulorum) occurs extensively on rough uplands along the Missouri and Yellowstone River drainages and throughout central and eastern Montana, It represents the most important habitat type of mule deer (Odocoileus hemionus), is locally a superior habitat of elk (Cervus canadensis), and is heavily utilized in grazing by livestock. Much of this important game habitat type is publicly owned, mostly by the Federal Government, and administered by the Bureau of Land Management. Paucity of quantitative data on big game and livestock food habits, range-use habits, and relations has hindered the establishment of sound effective management programs. A clarification of relationships is essen­ tial for development of cooperative multiple-use management programs. To aid in the formation of these, an intensive investigation was conducted on a representative 75,000 acre area in the Missouri River Breaks approxi­ mately 25 miles northeast of Roy, Montana. Full-time1 field work was con­ ducted from June i960 through September 1963» Additional data were obtained during several one-day to two-week periods from October 1963 to June 1964. The primary objectives were to obtain quantitative data on range-use and food habits of mule deer, elk, and cattle on the ponderosa pine-juniper and associated vegetational types and to develop methods and criteria for management. Habitat and population studies were conducted concomitantly to evaluate the effects of range condition, intraspecific and interspecific conflicts, and current game management practices on mule — 2 deer and elk populations„ These data also aided in interpretation of food and range-use relationships„ Seasonal and year-long food and/or range-use habits of mule deer and elk have been studied intensively on many parts of their ranges= Available data on range food and use habits of cattle on .western rangelands appear largely observational or derived from forage utilization and condition studies. Some quantitative} seasonal and year-long data for cattle are available for a few areas where big game-livestock relations have been studied, Few detailed studies of mule deer-cattle, elk-cattle, or mule deer-elk relationships have been made. Julander (1955) possibly presented the most comprehensive data on mule deer-cattle relationships in his report on the i Qak Creek Unit of Fishlake National Forest, Utah. Detailed studies involv­ ing mule deer-cattle forage relations have been reported by Julander (1937) and Kimball and Tfatkins (1951) for the Kaibab deer range, Arizona and Dasmann (1949) for the interstate deer winter range, California. Elk-mule deer relationships have been considered previously in comparative winter food habit studies in northern Idaho and Montana (DeNio 1938)s the Blue Mountains of Oregon ("Cliff 1939), and Jasper Park, Alberta (Cowan 1947). Morris and Schwartz (1957) made a year-long study on food habits of these two species on the National Bison Range, Montana. Elk-cattle relationships were studied by Stevens (1964) in the Elkhorn Mountains, Montana. The only study of mule deer, elk, and cattle relationships known to the writer was reported by Julander and Jeffery (1964) on summer range relations in Utah. DESCRIPTION OF THE STUDY AREA Physiography The study area (Figure I) extended approximately 20 miles as a 4 to 7-mile wide belt adjacent to the Missouri River in northeast Fergus Countys Montana. This area is part of a broad dissected plateau which slopes gently to the north and east, • The terrain is characterized by "breaks" (Figure 2), Ridges, sharply cut coulees, and creek bottoms resulting from erosion of the plateau are closely interspersed, . Elevations vary from about 3,100 feet along the southern boundary to.approximately 2,250 feet along the Missouri River, Rolling plains extend, with slightly increasing elevation, to the south and southwest» The soils, derived from the underlying Bearpaw Formation which con­ sists largely of clay shale containing moderate amounts of alkali and other salts, are predominantly heavy clay loams of the Lismas and Pierre series (Gieseker 1938). " Shallow layers of glacial lag may occur on- higher level ridgetops. Shale outcrops are common. Alluvial soils of the Havre silt loam and Lohmiller clay loam series are predominant on Missouri River bottomlands, (Mr, E=K, Hogan, Pers, Comm,, June 25, 1964)« Several other soil types may be found locally on uplands, terraces, footslopes, fans, and bottomlands, As a group, these "gumbo” soils are relatively imperm­ eable and runoff is high, especially from eroded and sparsely vegetated upland sites. They are considered too shallow, too heavy and plastic, and generally too steep for cultivation (Gieseker, op. cit.), — PAVED HIGHWAY ■ — —OBSERVATION ROUTE -------- JEEP TR AIL - = - DIKI ROAD ^% ^-CR EEK or COULEE A RESERVOm SCALE rMILES' NORTH Figure I. Map of the Missouri River Breaks Study Area. 1- 5 - Figure 2. Typical "Breaks" of Uplands Adjacent to the Missouri River in Northcentral Montana. "Plains" Extend Southward in the Background. - 6 - In addition to intermittent streams of the Sand Creek Coulee and Carroll Coulee drainages which run water only during spring runoff or heavy rains, several natural springs or seeps, and one well, occur on the area. About 30 of the 55 stock-water reservoirs developed by stockmen and/or the Bureau of Land Management hold water throughout the summer. These are widely distributed except on the extreme southeast "portion where no perm­ anent stock-water sources exist. Most of these upland water sources dried up during the drought of I960 and 1961. Access is provided by three graded dirt roads leading from US Highway 191 which forms the western boundary, and by numerous jeep or truck trails which cover all parts of the area (Figure I). As a rule, these roads and7trails are impassable when wet. Climate The semi-arid climate is characterized by moderately low rainfall, comparatively low relative humidity,- great extremes in summer and winter temperatures,..and many- sunny days.- Records, for the U .8, Department of Commerce weather station at Roy 8 NE,. located approximately 15 miles south­ west of the center of the area, show a 21-year (1943-1963) annual mean temperature: of 44°F . January, with aln average temperature of l6*F, is the coldest.month and July, with an ,average^of 71®F, is: the ,warmest. Monthly and annual precipitation data for the January. 1959-May 1964 period and 25-year (1939-1963) .means., .are. -shown in -Table -1 - Annual rainfall fluctuates widely between the lowest recorded value of 7.03 inches for 1952 and - 7 - TABLE Io PRECIPITATION DATA FOR MISSOURI RIVER BREAKS STUDY AREA, 1959-1964« UoSo DEPARTMENT GE COMMERCE, WEATHER BUREAU STATION, ROY 8 NE, MONTANA. , 25-Year _______ ______ Precipitation in Inches Month Mean 1959 I960 ■ 1961 1962 1963 1964 January 0.39 0.52 0 .8? 0.26 , 0,47 .0.85 0.23 February 0.40 0.72 0.43 0.20 • 0.48 0.92 0.08 March 0.54 0 .2 6 . 0.28 0.16 ' 0.33 T 1.00 April ' 0.88 1.01 1.78 0.15 0.30 1.18 1.53 May 2.26 2.84 1.54 1.25 . 7.58 2.65 June 3.42 2.94 1.58 1.41 5.03 3.10 July Io 50 . 1.21 0.23 2.77 2.08 1.25 = =™ = . August 1.29 0.41 1.33 0.21 ■ 3 .6? 1.62 September 1.14 2.46 0.34 1.77 . 0.37 ' 1.41 «=™=™ October 0.81 1.66 0.06 0.94 ' .1.89 0.23 November 0.47 1.13 O'. 12.,„ 0 .3& 0.78 0.37 December 0.33 0.03 0.33 G.ll 0.16 0.71 Totals \ 13.49 15.19 8.89 10.61 23.14 14.29 Deviation from mean 4-1.70 —4 060 -2.88 + 9.65 40.80 'f - 8 - highest of 23-14 recorded during 1962. Eleven inches or less total precip­ itation was recorded for 7 of the 25 years» These years are notable in that rainfall during the critical growing season months of May5 June5 and July was also severely reduced. This suggests that drought conditions might be expected about every three to four years„ '.Annual precipitation totals for the four-year study period represented nearly the complete range ■of expected conditions, The year prior to inception of the Study5 1959s was quite "normal"; however, I960 and 1961 were .marked by drought condi­ tions unmatched in the 25-year records. The total precipitation from January i960 through April 1962 was only 21.08 inches,. This includes only 8.42 inches during the most severe drought period which prevailed between I ‘ June i960 and July 1961» Exceptionally heavy rain fell during the late spring and summer of 1962, and near average amounts of precipitation were recorded for 1963- The mean annual precipitation recorded on a home rain gauge at the Tver Mathison ranch located on the south-central part of the area was 10.86 inches for the 1957-1962 period. This does not include water content of moderate snowfalls. A comparison with the 13 - 94-inch mean recorded for the same period at the Weather Station suggests slightly less annual rainfall on the study area. Total snowfall amounts to about 25 inches annually. Snow cover seldom persists for more than a few days except during the coldest periods. Warm southwesterly "Chinook" winds, characteristic of the region., temper the harshness of the winter and prevent accumulation of snow. - 9 - Land-Use Characteristics Widespread grazing began on the study area and vicinity during the early 1880?s (Gieseker 1938). According to Gieseker9 et al (1953)s large livestock companies grazed herds on range forage throughout the year until the early 1900?s when many of these organizations failed or were dissolved and were succeeded by local stockmen and smaller herds. Much of the area was taken up under the Homestead Acts and settled between 1906 and 1915 when attempts were made to cultivate upland sites. Recurrent drought conditions, especially during the 1918-1921 and 1930-37 periods, and other factors led eventually to abandonment of most of these lands and a gradual depopulation of, the area. At present, only three headquarter ranches persist. Formerly cultivated tracts have reverted to grasslands currently utilized for grazing or occasionally cut for hay. ' Publicly owned lands now comprise 51,210 acres or 68.3 per cent of the total area. Of this, 47,850 acres are owned by the Federal Government9 3,320 acres by the State9 and 40 acres by Fergus County. Approximately 44 per cent of the area lies within the Charles M. Russell Wildlife Range established in 1936 and administered by the U.S. Fish and Wildlife Service.. Most of the area is open range and grazed by domestic livestock— primarily cattle but a few horses are included. Occasionally, domestic sheep are grazed into the "breaks" on the south-central portion. Livestock grazing is administered by the Bureau of Land Management and the Indian Buttes State Grazing District. The grazing season extends from April I through Movaaber 30. - ID - Winter livestock use is restricted largely to Missouri River bottomlands and two privately owned upland pastures» Fourteen individual livestock operators hold permits to graze Ij, 646 cattle and 13 horses on the study area and/or adjacent prairie« Approximately 1,500 acres (2 per cent of the total) are cultivated. Except for one.small dryland grain field on the western edge of the area, these lands are restricted to bottomlands along the Missouri River' where alfalfa and a limited amount of barley are grown. In addition to mule deer and elk which range throughout the area, white-tailed deer (Odocoileus virginianus) are common,.on the Missouri River- bottomlands-. Pronghorn antelope (Antilocapra americana) and an occasional bighorn sheep (Qvis canadensis) range onto the area from adjacent range­ lands . VEGETATION Ecological studies of rangelands in eastern Montana are few. Wright and Wright (1948) studied grassland communities in southcentral Montanajl and Jonas (1964) described plant communities in the Long Pines of south­ eastern Montana. Neither these, nor studies in the adjacent Great Plains of Canada (Coupland 1950, 1961) and western North Dakota (Hanson and Whitman 1938), seem entirely applicable to the vegetation of the study area. The present analyses were undertaken to ascertain the vegetal char­ acteristics for determining intrinsic relationships between the animal, species studied and their habitat. , Reconnaisance permitted tentative classification of eivegetation types." Refinements followed. Beginning during the winter of 1961-62, records were made of the presence and abundance of plant species at sites used for feeding by mule deer, elk, and/or cattle. Examination of 287 sites on eight vegetation types during all seasons of 1962 and 1963, aided in determining phenologies of the various taxa, recognition of unions, and delineation of communities. Statistical studies were conducted during July 1963 when taxa which characterized communities throughout most of the year had attained maximal development, and those of spring aspects had com­ pleted their, annual cycles. For each of the 12 rangeland communities, three representative stands with one each in the eastern, central, and western portions of the study area were subjectively selected for analysis. ■The degree and direction of slope, soil type and condition, and major influent hoofed mammals for each stand were recorded. Sites which appeared - 12 - .-to have received recent significant prior animal use by grazing were avoided. The primary method of vegetation analysis'followed Daubemnire (1959)° ,,Forty 2 x 5 dm plots were systematically placed within a, 20 x 50 ft sample unit located within a relatively homogeneous and undisturbed portion of , ' each vegetation stand. The per cent canopy-cover' of each taxon and per­ centages of bare ground and rock were estimated by class for each plot. Classes were: -.Class I = 0-5%5 Class 2 = 5-25%; Class 3 — 25=50%; Class 4 = 50-75%; Class 5 — 75-95%; and Class 6 = 95-100%. The midpoint of each class was the value used in data tabulations» Taxa occurring within each stand but not in any of the plots were also recorded. Ground cover was further evaluated by recording whether each of the four ^ sharply pointed legs of the plot frame contacted bare ground=, rock=, Iitters or living plant material. Tree and taller shrub composition as well as canopy-coverage were, studied in the four timbered communities by superimposing four (two in each of the three stands of the Pseudotsuga/Juniperus type) line transects at 50 ft intervals across the ^ established sample unit. The 100 ft tape was suspended approximately 30 inches above the ground.between two steel stakes. Crown intercept of each taxon was measured by using, the densio- meter (Lemon 1956) in a method described by Dealy (I960). While straddling the Iines points of beginning and end of intercept were located with the densiometer. A plumb-line attached to the densiomet er/;fell:r;against the tape at the point of recorded measurement. T - /)■ - 13 - Phytosbciologic terms used in describing the vegetation are as follows: "vegetation typb" denotes the collective area occupied by a single association or a group of associations characterized by the same taxa, e.g.,the Artemisia/Agropyron Type includes the Artemisia tridentata/ Agropyron spicatum Association^ the Artemisia tridentata/Agfbpyron smithii/ Bouteloua gracilis Association, and the A. tridentata/A. smithii Associes« Serai stands are also included; i.e=, the Pseudotsuga/luniperus Type includes stands in which Pseudotsuga menziesii is sparse or absent but unions characteristic of the Pseudotsuga menziesii/juniperus seopulorum Association are well represented. The terms "union" and "association" are used in the sense of Daubenmire (1952). The union represents the smallest structural unit of organization. It is comprised of a single species or of several species that are closely similar in ecology as indicated by similarity of local environmental amplitude and phenology. Association is \ applied only to what are believed to be climax (either climatic, edaphic, or topographic) communities, and indicates the distinctive combination of unions superimposed on the same area. "Associes" are developmental equiv­ alents of associations (Weaver and Clements 1938). As used herein, the ' term denotes the unions or distinctive combination of unions which persist on the same area under disclimax conditions resulting from animal activity (including man’s) and/or peculiar topoedaphic conditions which appear to be somewhat less than "permanent." Artemisia/Agropyron Vegetation Type This type occurs extensively on level to gently rolling1' ridgetops where it is largely restricted to less-than-15-degree slopes and clay sub­ soils more than 15 inches deep over shale» The. type •'occupies approximately 23>000 acres or 31 per cent of the total area= Recognition is afforded by its physiographic location and prominence of the Artemisia tridentata union. The type includes three distinct communities> the distribution and organization of which seems related to livestock distribution^, intensity of grazing,'and edaphic (soil) conditions. Comparative statistical data for these communities and the type as a whole are.presented in Table II. Ground-cover characteristics of these and the other rangeland communities are listed in Table III. Artemisia tridentata/Agropyron spicatum Association.— This association (Figure 3) occurs where a "light," one-to-three inch layer, of "topsoil" persists and livestock use is much restricted as a result of inadequate water supplies or physiographic barriers. It attains widest distribution in areas greater than one mile from a permanent water source, though remnant stands or vestiges may be found at lesser distances. The assoc­ iation is most extensive on the very lightly grazed southeastern portion of the study area, but isolated stands occur throughout. The presence of "topsoil" seems important. Observations indicate that A. spicatum occasionally occurs in the A. tridentata/A. smithii associesj but on iso­ lated or marginal areas of both communities where severe disturbance and TABLE II. CONSTANCY, CAEOP^eOVERA^'^IB' ■ FREQUENCY Aim DIE3TRI3UTIOK: BY. UEIONS 01' TAXA OK ■ THE ARTEMISIA/AGROFYROH- TYPE. 'l/ . . . . : . . A. tridentata/ A. tridentata/ A. tridentata/ Combined Community A. spicatum A. smithii A. smithii/ Type Association Associes B. gracilis Totals Association Union: Taxa 2/ Cy/Cv/Fr 3/ Cy / Cv/Fr C^/Cv/Fr Cy/Cv/Fr Agropyron spicatum: . Agropyron spicatum 1 0 0 / 4 3 / 8 9 3 3 / + / + 44/14/30 Agropyron smithii: . Agropyron smithii IOO/18/56 1 0 0 / 5 7 / 9 8 I O O / 3 0 / 9 8 1 0 0 / 3 5 / 8 4 Bouteloua gracilis 3 3 / 2 / 1 0 6 7 / + / 2 I O Q / 1 0 / 5 2 67/ 4 /2 1 Koleria crista,ta 1 0 0 / 3 / 2 3 1 0 0 / 6 / 2 8 1 0 0 / 1 0 / 7 5 100/ 6 /4 2 Stipa comata 3 3 / 2 / 1 3 1 0 0 / 5 / 2 7 3 3 / + / 3 2 2 / + / 6 Stipa viridula, 1 0 0 / 5 / 2 6 6 7 / I / 5 ' 8 9 / 4 / 1 9 Carex spp. 6 7 / 4 / 2 6 3 3 / + / ' - 4 3 3 / 3 / 1 9 44/ 3/16 Artemisia tridentata: ~ Artemisia tridentata 100/15 /50 1 0 0 / 1 5 / 5 2 1 0 0 / 1 9 / 6 7 100/16/56 Chrysothamnus nauseosus 3 3 / + / 2 - ' 3 3 / + / 5 ' Sr/ +/> - '' 44/ +/ 4 Gutierrezia sarothrae 3 3 / + / 2 3 3 / + / + ' 1 0 0 / t / l 3 5 6 / + / 5 Artemisia, frigida 3 3 / l / 6 '67/ l/ll 3 3 / + / 6 Hymenoxys richardsonii 3 3 / 1 0 0 / + / 3 3 3 / . + / I ' Petalostemum purpureum - 3 3 / + / 5 6 7 / +-/ 8 3 3 / + / 3 Phlox hoodii 3 3 / + / + 1 0 0 / 2 / 3 7 56/ +/l4 Tragopogon dubius: Comandra, umbellatum 1 0 0 / l / 2 0 1 0 0 / 1 / 2 8 6 7 / + / 1 3 8 9 / + / 2 0 Melilotus officinalis 3 3 / + / 3 100/ 2/12 6 7 / + / + 6 7 / l / 5 Opuntia. polyacantha 100/ 1/10 100/ 1/10 100/ 1/10 100/ 2/13 . Selaginella densa 100/ + / 1 2 . 3 3 / I / 5 6 7 / 1 / 1 3 - 33/ +/ 6 Sphaeralcea coccinea, 100/ +/19 6 7 / + / 8 8 9 / + / 1 3 Tragopogon dubius 100/ +/ 3 1 0 0 / +/ 3 33/ +/ + 7 8 / + / 2 Vicia americana, 6 7 / + / 1 3 1 0 0 / +/ 3 100/ 4/i4 8 9 / 2 / l 4 TABLE TI. Continuedo Community A. tridentata/ A. spicatum Association- A. tridentata/ A. smithii Associes A. tridentata/ A. smithii/ B. gracilis Association Combined Type Totals Union: Taxa 2/ Cy/Cv/Fr 3/ Cy/Cv/Fr Cy/Cv/Fr Cy/Cv/Fr Poa sgcunda: Poa secunda 6 7 / t / 5 1 0 0 / 1 5 / 5 8 100 / 10/67 8 9 / 9 / 4 3 Descuriana pinnata 6 7 / + / 3 22/ +/ I Oenothera caespitosa 6 7 / + / 3 33 / +/ + 3 3 / + / I Plantago spinulosa 3 3 / + / + 6 7 / +/ 8 3 3 / + / 3 Sarcobatus vermiculatus: Schedonnardus paniculatus 3 3 / +/ 8 6 7 / +/ 7 . 3 3 / + / 5 Bare ground 1 0 0 / 4 6 / 9 0 1 00 /5V 100 - ' l O O / 5 0 / l O O 100 /50 /100 Rock 3 3 / . + / 1 9 1 0 0 / 2 / 5 0 100 / 2 / 4 0 ' ' . 1 0 0 / 1 /36 l/ Data are mean percentages for 120 two x. five dm plots (40 in each of three stands) in each community. 2/ Includes only those species which occurred in at least two stands, i.e., constancy of 67 per cent.or more, or had mean canopy-coverage of at least one per cent in any community. Other taxa are considered in Appendix Table XLI. 3/ Constancy/Canopy-coverage/Frequency. +/+ indicates canopy-coverage or frequency values less than one per cent; a single + in place of these two values denotes occurrence in the stands but not in the plots. - 17 - TABLE III. GROUND-COVER CHARACTERISTICS FOR 12 RANGELAND COMMUNITIESo DATA ARE FREQUENCIES OF OCCURRENCE OF BARE GROUND, LITTER, AND LIVING PLANT MATERIAL AMONG 480 POINTS (160 IN EACH OF THREE ■ STANDS) IN EACH COMMUNITY. Conmmnity Bare Ground . Litter Live Plant ' - Pet. Pet. Artemisia tridentata/Agropyron spicatum ' 45.2 39 .1 15.6 Artemisia tridentata/Asropyron smithii 54.0 . 34.6 11.2 Artemisia tridentata/Agropyron smithii/ : Bontelpna gracilis 55T0": 28.1 16.8 Agropyron smithii/Poa secunda 57.7. 33.9 8.3 Pinns ponderosa/Agropyron spicatum 13.7 82.0 4.1 Finns ponderosa/Juniperus scopulorw 19.3 ■ 71.8 8.7 Finns ponderosa/Artemisia, longifolia 77.0 22.5 0.4 Pseudotsuga menziessii/Juniperus scopulorum 0.2 ■61.4 38.3 Sarcobatus vermiculatus/Artemisia tridentata 47.7 38.5 13.7 Agropyron smithii/Symphoricarpos occidentalis 30.4 58.3 11.2 •Xaiitfium strumarium 93.7 . - 5.6 0.6 \ Artemisia longifolia 95.6 . 2 .7 1.6 - 18 - Figure 3* Artemisia/Agropyron Vegetation Type. Ridgetop Occupied by Artemisia tridentata/Agropyron spicatum Association. LI % * 'Ti I 1"" ■ ■ ■ ■ ■ ■ . ' .?/!(' '' M ■ # ' » a>.. f Figure 4 . Artemisia/Agropyron Vegetation Type. Ridgetop Occupied by Artemisia tridentata/Agropyron smithii Associes. ■ ; . ' ’ ra' 19 ' ■ subsequent erosion., partially or- ■ completely removed, the topsoil leaving either a shallow^ gravelly ©r el&y-p&n Soilx, A. spieatma is either absent or persists only on ’’pads68 (amali elevated, mounds of soil) around the base of Artemislao ■■■-'. ■ • The Ao spicatum lmlom is. dominant (.Table II)» Artemisia trldentata and the. Ao smithii union, comprised of the principal grasses of the mixed prairie (Coupland 1950) s. are - well represented -. but to a lesser degree than . in the- other two Communities of ...the Artemisia/Agropyron type-o-- The fact t;that, the A» smithii union i s relatively well developed here may indicate some disturbance in the stands o The ..Poa secunda union, which. includes the ' " ■ ■ • •. ' ■ ■ earliest spring herbs, is .scarcely -represented! although Alliira textile and Lomatium foenieulatum are common during the springo The Tragopogoncdubius union, comprised of.species of relatively wide ecological amplitude and which attain maximum development in- early summer, is best ■ represented by Comandra umbellatum. Qpuntia polycantha, Sphaeralcea coccinsa, Tragopogon dubius. and Vieia americana.' 1 ' ■. ■ This.association appears to have somewhat greater total ground cover­ age than any of the other-communities- which .occur on open ridgetops • (Tables II and ill) „ These data ..support general observations ■ of less run­ off during•rainstorms- a n d .suggest greater soil- stability on sites where it occurs. In view of its distribution on deep, undulating, upland clay-loam . soils throughout the-relatively undisturbed,portions' of the area, this association may represent the- climatic climax type in the central Missouri Breaks. — 20 — Artemisia tridentata/Agropyron smith!! Associes,— On sites accessible to hoofed mammals throughout t h e . study area where topoedaphic conditions seem similar to those supporting the A. tridentata/A. sulcatum association, Agropyron spicatum appears to have been greatly reduced or eliminated and dominance assumed by the A. smith!! union (Figure.4)! "The distribution of this community seems to correspond with areas where livestock use is periodic, fluctuates as. a result of the lack of permanent, water, dr is otherwise restricted by man or grazing habits, ' Agropyron smith!! is the usual dominant (Table II); however, either Koleria•cristata or Stipa viridula may dominate local stands. The A, tridentata union co­ dominates with little change in importance from the Artemisia tridentata/ Agropyron spicatum association. The Poa secunda union is well represented, especially by P. secunda and during the spring, by Allium textile, Lomatium foeniculatum, Mieroseris nutans, Musineon divaricatum, and Oenothera caespitosa. The Tragopogon dubius union is conspicuous in both the number and abundance of representative species, . The distribution of this community, the paucity of A, spicatum, the greatly increased importance of the A, smithii union, and the increased representation of forbs all suggest a disclimax condition resulting from disturbance of the Artemisia tridentata/Agropyron spicatum association. Artemisia tridentata/Agropyron smit This community. (Figure 5) is especially prevalent on the western and central portions and along the southern boundary of the study area. I — 21 — Figure 5- Artemisia/Agropyron Vegetation Type. Ridgetop Occupied by the Artemisia tridentata/Agropyron smithii/Bouteloua gracilis Association. Figure 6. Agropyron/Poa Vegetation Type. Formerly Cultivated Ridgetop Now Occupied by the Agropyron smithii/Poa secunda Associes. - 22 - Its distribution seems to correspond with that of shallow* gravelly* or clay-pan surface soils and generally heavy grazing pressures„ Normally*, severe spot or sheet erosion is evident in that islands of higher-lying soil and vegetation continue to existj A. tridentata frequently is elevated o n ,small mounds of soil which are anchored by litter* Selaginella demsa, and roots of both A. tridentata and herbs which grow through or beneath it. This suggests that degradation of sites normally supporting either of the other two communities has been at least partially responsible for its developmenti Vegetation cover is typically sparse and depauperate. Much . of the total canopy-coverage is often concentrated on small islands where species and unions are superimposed. The A. tridentata and A. smithii unions are dominant (Table II). - Artemisia tridentata is somewhat more abundant than in either of the * , I previously described communities and its associates are better represented. Chrysothamnus nauseosus and Gutierrezia sarothrae are conspicuous. Agropyron smithii, although less dense than in the Artemisia tridentata/ Agropyron smithii associes* remains dominant; and Bouteloua gracilis and Koleria eristata become quite prominent. In contrast to the other commun­ ities of the Artemisia/Agropyron type* Stipa viridula is only poorly represented. The Poa secunda union is also prominent and on some sites may dominate the community. In local areas* particularly on the western part of the study area* the Sareobatus vermiculatus union is frequently I associated with this community. “ 23 - Observations suggest that this association represents an edaphic climax on sites where it occurs. Agropyron/Poa Vegetation Type This type (Figure 6) occurs entirely on sites usually characterized by the Artemisia/Agropyron type.' I t 'is largely restricted to approximately Ij200 acres on ridgetops in the central and southwestern portions of the study area which were formerly cultivated and/or artifieally reseeded with exotic grasses. Differences in times of abandonment or reseeding and past treatment have resulted in diverse vegetal characteristics. The three stands selected for analysis include an area grazed primarily during the winter and infrequently cut for hay, a.stand grazed only during the grazing season and occasionally cut for hay, and a recently reseeded stand which is also grazed during the grazing season. All but the latter are now dominated by native species. Because of the minor extent of the type (1.5 per cent of area) and some indication that the successions! trend on all reseeded areas is toward dominance by the same native grasses, these communities are recognized as a single associes in which the A. smithii and P. secunda unions dominate (Table IV). Although the Artemisia tridentata union appears to be regenerating slowly, it is only poorly represented by scattered young plants and a few depauperate older plants which persist despite occasional cutting, browsing, and trampling. Agropyron cristatum. an exotic, joins the A. smithii union in this community where deliberate or accidental seeding has occurredj however, it dominates only the recently reseeded stand. Bromus japonicus, another exotic, also joins the - 24 ™ TABLE IVo CONSTANCYj CANOPY-COVERAGE, AND FREQUENCY AND.DISTRIBUTION BY UNIONS OF TAXA ON THE AGROPYRON SMITHIl/POA SECUNDA ASSOCIES, l/ Union: Taxa 2/ Cy/Cv/Fr 2/ Agropyron smithii: Agropyron cristatum Agropyron smithii Bromus .Iaponicus Stipa viridula 67/20/33 100/46/82’ 67/ 3/13 100/ 5/22 Artemisia tridentata: Artemisia frigida\ Aster occidentalis 33/ 2/21 100/ -V -s- Poa secunda: - Poa secunda 100/16/48 Tragopogon dubius: Melilotus officinalis Opuntia polycantha Polygonum erectum Sphaeralcea coccinea Vicia americana 100/13/42 100/ -V 3 ■ 67/ +/19 100/ -V 9 100/ -V 7 Bare Ground 100/61/100 Rock 67/ +/ 6 . / ' y Data are mean percentages for 120 two x five dm plots distributed as 40 in each of three stands„ 2/ Includes only those species which occurred in at least two stands, i.e., constancy of 6? per cent or greater or had mean canopy-coverage at least one per cent in the three stands« Other taxa are considered in • in Appendix Table XLD&;-+:- 2/ Constancy/canopy-coverage/frequency. +/+ indicates canopy-coverage or - frequency values less than one per cent. )A. smlthii union on local areas where disturbance seemingly has provided an avenue for invasion. At the time of analysis in late Julyjl character^ . istic :'forbs were those of the Tragopogon dubius union.. During the spring, however, observations indicate the P. secunda union is well represented by Allium textile. Lomatium foeniculatum. and Microseris nutans. The total ground cover provided by this type is somewhat less than that afforded by any community of the Artemisia/Agropyron type (c.f. Tables II, III, arid IV). Pinus/Juniperus Vegetation Type This is a major type comprising approximately 45 per cent of the vegetation. It occurs extensively on slight to moderately steep slopes and soils which vary from clay-loam less than 15 inches deep over shale to raw shale. It is prevalent on south or west-facing slopes, but also occurs on northerly or easterly exposures where the degree of slope is slight. The type frequently acts as an ecotone between the Artemisia/Agropyron and Pseudotsuga/juniperus types. Stands of pine are seldom dense. A near- complete overhead canopy may be found in only a few scattered stands on slight, north-facing slopes. Frequently, only a few scattered pines occur to give a savannah-like appearance, especially on old burn sites and along the upper limits of the type where it contacts the Artemisia/Agropyron type. The characteristically slow growth of P. ponderosa is attested by data obtained by the Bureau of Land Management for a "site" tree in each of 14 pine stands on and adjacent to the study area. These trees, which had a mean approximate age of 83.2 (51 to 141) years, averaged only 32.6 (22 to 46) feet in height and 8.5 (6.4 to 11.4) inches dbh (diameter 4.5 — 25 \ 1 ~ 26 — feet above the ground). The mean growth during the last 10 years was Oo29 inches for 36 trees, (Unpubl, Sample Plot Datai, U=So Bureau of Land Management, Lewistown, Montana, 1964)» The type is .comprised of three more or less distinct communities» The distribution and organization of these appears relatedato, topographic and edaphic factors. Statistical data for the three communities and the.type as a whole are presented in Tables V and V I . • Pinus ponderosa/Agropyron spieatum Association»^Wherever the Finns/ Juniperus type abuts upon ridgetops ..dominated by the Artemisia/Agropyron type or the Agropyron/Poa-type, it is characteristically,.represented-by • the Finns ponderosa/Agropyron spieatum association (Figure 7)« . At least a shallow layer of clay-loam soil seems essential to support the usually well- developed grass-understory. Theumore exposed,. .xeric upper slopes are also characteristic = Qn less: .exposed-,. -more.-.mesic footslopes, and draws this community is replaced by the Plnus ponderosa/Juniperus scopulorum ■ _ . association. - Pinus ponderosa is dominant and better represented than in either of the other communities (Table V). Typical openness of stands is attested to by the 27 per cent coverage afforded by P. ponderosa and the total tree-shrub intercept of only-32 per .cent. The. principal^undergrowth is herbaceous and. usually dominated by the.Agropyron spieatum union. . .. Frequently, however, the Agropyron smithii .union.assumes this position. This may be correlated with the fact that the association characteristically occurs on gentle slopes.susceptible to -moderate or heavy- use- by hoofed TABLE V. CONSTANCY, .iCAEOEY'^OVEKAGEy. -MB FREftyMCY AM) DISTRIBUTION BY.; UNIONS Oi TAXA ON ECO PiNUs/ALYDTOILS E F T OM .;i/ :. .. ;' , Association . Union: Taxa 2/ Agrppyron. spicatum: Agropyron spicatum MuhlenBergia cuspjdata. . Agropyron smithii: Agropyron smithii Bouteloua gracilis Koleria cristata Stipa viridula -Carex spp. Artemisia tridentata: Artemisia, tridentata Artemisia frigida Poa secunda: - Descuriana pinnata, - Microseris nutans Juniperus scopulorum: . .. Juniperus scopulorum : Rhus trilohata Rosa arkansana i Symphoricarpos alba ; Carex geyeri : Antennaria dimorpha Tragopogon dubius: Achillea millifolium .•Artemisia ludoviciana . Aster commutatus . Bahia oppositifolia 'Pinus ponderosa/ Pinus ponderosa/ Pinus ponderosa/ Combined Agropyron spicatum J. ^copuborum Artemisia longifolia Type total Association Association . Association Cy/Cv/Fr 3/ Cy/Cv/Fr Cy/Cv/Fr Cy/Cv/Fr 1 0 0 / 2 0 / 5 3 6 7 / + / 5 I O O / 1 5 / 3 9 6 7 / + / 3 6 7 / + / + 6 7 / + / + ' 8 9 / 1 2 / 3 1 6 7 / + / 3 100 /1 7 /6 2 ‘' . 3 3 / . + 6 7 / 3 / 1 3 1 00 /1 2 /3 4 .6 7 / 9 / 4 8 ' ' _ 6 7 / 5 / 1 9 6 7 / + / 3 6 7 / + / 8 1 0 0 / 4 / i i 6 7 /1 0 /4 0 1 0 0 / 4 / 2 9 3 3 / + / + 3 3 / + / 2 8 9 / 9 / 3 7 .3 3 / + / + 44 / 1 / 7 7 8 / 5 / 1 5 5 6 / 6 / 3 0 1 0 0 / + / + 3 3 / + 6 7 / + 6 7 / + / 4 3 3 / + / + 3 3 / + 6 7 / + / + 44 / + / 1 .3 3 / + 6 7 / + / 4 6 7 / + / 3 6 7 / + / 2 5 6 / + / 2 2 2 / + / 1 1 0 0 / 3 / 4 1 0 0 / 2 / 5 6 7 / + / 6 1 0 0 / 3 /22 . 3 3 / 9 / l 8 3 3 / + / 2 1 0 0 / 1 9 / 3 8 . 1 0 0 / 2 / 1 0 1 0 0 / 2 / 9 ■ 3 3 / + / + U % \ 1 0 0 / 2 / 3 1 0 0 / + / 2 3 3 / + / 6 1 0 0 / 8 /1 5 1 0 0 / 1 / 6 6 7 / l / 7 . 44 / 1 / 7 2 2 / 3 / 6 3 3 / + / 2 6 7 / 2 / 1 5 6 7 / + / 3 6 7 / + / 3 .3 3 / 1 / 1 8 1 0 0 / 3 / 2 5 6 7 / + / 5 1 0 0 / + / 8 3 3 / V 2 6 7 / + / 3 3 3 / + / 2 5 6 / 2 / 1 3 44 / + / 3 7 8 / + / 6 3 3 / + / 7 TABLE V, Continueda Pirns ponderosa/ Pinus ponderosa/ Pinus ponderosa/ Combined Association Agropyron spicatum J. scopulorum Artemisia longifolia Type Total Association . Association Association Union: Taxa 2/ Cy/Cv/Fr 3/ Cy/Cv/Er Cy/Cv/Fr .Cy/Cv/Fr Chenopodium album 6 7 / + / 2 6 7 / + / 8 1 0 0 / + / 5 7 8 / + / 5 . Comandra umbellatum 6 7 / + / 8 2 2 / + / 3 Gaura coccinea .6 7 / + / 3 6 7 / + / + 3 3 / + / 4 3 3 / + / 2 _ Grindelia . squarrosa . 3 3 / + ' 3 3 / + / + Lactuca pulchells, 3 3 / + / + 6 7 / + / 6 6 7 / + 3 3 / + / 2 Melilotus officinalis . 3 3 / + / + 6 7 / + / 4 5 5 / + / 2 Opuntia polycantha 6 7 / + / + 6 7 / + / 7 1 0 0 / + / + 7 8 / t / 2 Phacelia linearis 6 7 / + / 3 6 7 / + / 1 0 4 4 / + / 4 . ,Sblaginella densa 1 0 0 / + / 8. 6 7 / - + / 1 3 2 2 / + / 4 ", .,Solldago mias.ouriensis 1 0 0 / 2 / 1 7 6 7 / + / 8 8 9 / l / l l Stephanomeria r'uncincata 6 7 / + / 8 .3 3 / + / 3 1 0 0 / 2 / 1 7 § 7 / + / 9 Thermopsis rombifolia 1 0 0 / 1 / 1 8 6 7 / . + / 4 , $ 6 / + / 7 Tragopogon dubius 1 0 0 / 2 / 1 2 6 7 / + / 3 6 7 / +. 7 8 / + / 5 . .Vicia americana 1 0 0 / 1 / 2 0 1 0 0 / i / 1 2 6 7 / + / l l Symph or icarpos. Occident alls: 6 7 / 6 / 2 4Symphoricarpos- occidentalis 2 2 / 2 / 1 5 . Agropyron trachycaulum 6 7 / 2 / 9 2 2 / + / 3 Prums * Tirginiana: Prunus Tirginia ns. 3 3 / I / 6 1 1 / + / 2 Ribes cereum ' 6 7 / + / + 22/ +/ + Agrostis scabra 1 0 0 / 1 9 / 4 3 33/ 6/l4 . Smilaoina''. stellata Artemisia longifolia: .33/ l/ 3 11/ +/ + Artemisia longifolia 33/ + 33/ + 100/ 2/ 7 56/ +/ 2 Calomovilfa, longifolia 33/ + 100/12/29 4 4 / 4 / 1 0 . Oryzopsis hymenoides . Mentzelia laevicaulis Mirabilis linearis % % I 67/ +/ 2 22/ +/ + 22/ +/ + 22/ + / + TABLE V. Continued Association .Pinus ponder0sa/ Agropyron spicatum Association Pinus ponderosa/ J. scopulorum Association Pinus ponderosa/ Combined Artemisia longifolia Type Association Total Union: Taxa, 2/ Cy/Cv/Fr/ 3/ Cy/Cv/Fr Cy / Cv/Fr Cy/o/Pr Xanthium strumarium: Conringa orientalis . 67/ +/ + 22/ +/ + Bare Ground 100/20 /6 4 100/56/21 IOO/80/97 100/52/61 Rock 33/ +/ + 33/ +/ + - 67/ +/ 3 ' 44/ +/ + l/ Data are mean percentages for 120 two x fiv.e dm plots (kO in each'of three stands) in each « association. ■ ' " 2/ Includes only those species which occurred in at least two stands, i.e.y.constancy of 67 per cent or more, or had mean canopy-coverage at'.least one per cent in any association. . Other taxa are considered in Appendix Table XLIII. 3/ Constancy/canopy-coverage/frequency. -+/+.indicates canopy-coverage or frequency values less than one per cent: a single + in place of these two values denotes occurrence in the stands but not in the plots. TABLE VI. CONSTANCY, COVERAGE, AZD FREQUENCY OF TREES AND SHRUBS GREATER THAN 12 INCHES’ TALL ON THE PINUS/JUNIPERUS TYPE, l/ Association Pirns ponderosa/ . Pinus.ponderosa/ Pinus ponderosa/ Combined Agropyron spicatum Juniperus scopulorum Artemisia longifolia %rpe Total Taxa Cy/Cv/Fr 2/ Cy / Cv/Fr Cy/Cv/Fr Cy/Cv/Fr Pinus -Donderosa lOO/27/lOO 100/14 /75 100/18/73 IOO/20/83 Juninerus sconulorum 100/ 4/58 lOO/53/lOO 1 0 0/ 8/88 100/21/83 Rhus trilobata 100/ +/50 100/ 7/88 100/ +/36 100/ 3/54 Artemisia tridentata 33/ +/ 8 67/ 1/50 33/ 1/2 7 44/ +/26 Chrvsothamnus nauseosUs 67/ + /17 22/ +/ 6 Prunus virginiana 33/ 6/50 11/ 2/13 Ribes cereum U) CO C O 6 7/ +/25 33/ +/10 Artemisia cana 33/ +/ 9 11/ +/ 3 Artemisia longifolia 67/ 3/73 22/ 1/26 Sarcohatus vermiculatus 33/ +/ 9 33/ +/ 3 Total Intercept (per cent) 32.2 80.9 30.7 / . 4 4 .3 l/ Data are mean percentages for twelve 100 ft line-intercept transects (four' in each of ~ three stands) in each association. . . " 2,/ Constancy/coverage/frequency. Frequency is the percentage occurrence artoAg the twelve 100 ft lines = + indicates canopy-coverage values less than one per cent.;' - 31 - Figure 7. Pinus/Juniperus Vegetation Type. Slight South-facing Slope Occupied by Pinus ponderosa/Agropyron spicatum Association. Figure 8. Pinus/Juniperus Vegetation Type. Moderate West-facing Slope Occupied by Pinus ponderosa/Juniperus scopulorum Association. - 32 - mammals o The A 0- smithii union seems almost identical in. composition and abundance to its occurrence in the Artemisia tridentata/Agropyron spicati; association; however# the A. spicatum .union appears less abundant but Muhlenbergia cuspidata is somewhat better represented here than in that associationo Principal forbs are those of the Tragopogon■dubius union / V - V v : . ' . - y . which differs only in the relative abundance of member species from its occurrence on the Artemisia/Agropyron type, . The Poa secunda union is very poorly representBd5, although Allium textile, Fritillaria pudica, Lomatium foeniculatum, and Microseris nutans are common during spring. The shrub layer is comprised mainly of a sparsely represented Juniperus scopulorum union (Tables. V and VI) in which either J 0 scopulorum, Rhus trilobata, or Symphoricarpos alba may dominate, in individual stands, Occasionally, shrubs of the Artemisia tridentata, Prunus virginiana, Sarcobatus vermieulatus and Symphoricarpos occidehtalis unions occur as scattered individuals, Pinus ponderosa/Juniperus scopulorum Association, — On less exposed, more mesie sites along shallow drainageways and south or west-facing slopes, the J, scopulorum union supplants the Agropyron unions in under­ story dominance (Table V) to give this association a distinctive physiog­ nomy characterized by P. ponderosa and dense growths of J, scopulorum (Figure 8), Although P, ponderosa is Usually prominent, it appears less abundant than in the Pinus.ponderosa/Agropyron spicatum association (Table VI), ' This may be due to curtailed reproduction by the shade- intolerant pine under the dense canopy of J, scopulorum, - 33 - Symphoricarpos alba, which frequently is a very prominent member of the tJ, seopulorum union in open stands of the Pinus ponderosa/ Agropyron spicatum association^ is suppressed and the Prunus virginiana union, comprised of species typical of north-facing slopes, may develop under the shaded mesic conditions, Pseudotsuga menziesii may also occur. The herbaceous layer is usually dominated by A, spicatum (Table V); but on especially protected sites Agrostis scabra, a member of the P= virginiana. union, may dominate. The Agropyron smitfaii union is of secondary importance, occurring primarily on alluvium along bottoms of smaller drainageways'. The Symphoricarpos occidentalis union usually is associated with the A, smi-thii union on these sites, Porbs are largely those of. the Tragopogon dubius union with only minor changes in species representation occurring as compared to the Pinus ponderosa/Agropyron spicatum associa­ tion. Distributional limitations suggest that this association represents a topographic climax on the sites where it occurs,. Pinus ponderosa/Artemisia longifolia Association,— This association (Figure 9) is restricted to shale outcroppings on steep slopes where it may represent a topoedaphie climax. These sites are generally unstable, and individual stands characterizing them exhibit considerable variation with local conditions. The community typically is characterized by a • sparse pine overstory, a variable understory which usually is dominated by the A, longifolia union, comprised of species more or less restricted to shale outcrops, and a bare shale soil surface (Tables III, V, and VI), - 34 - Figure 9. Pinus/Junipenis Vegetation Type. Steep, South-facing Shale Slope Occupied by the Pinus ponderosa/Artemisia longlfolia Association. Figure 10. Pseudotauga/junlperua Vegetation Type. Steep North-facing Slope Occupied by the Pseudotsuga mengiesii/ Junlperus scopulorum Association. - 35 - The Juniperus scopulorum union is quite prominent and may dominate local standso Although scarcely represented in the stands analyzed5 the Sarcobatus Termiculatus union often occurs in this association; on Steeps, open slopes along deeply cut coulees in the northeastern part of the study area5 it occasionally dominates the shrub layer. The poorly developed herbaceous layer is usually dominated by Calamovilfa longifolia g howevera Agropyron spicatum and/or A. smithii may dominate on more stable sites. Forbs typically are poorly represented. Chenopodium album and Stephanomeria runcincata of the Tragopogon dubius'union seem most commons but other species of the T. dubius, Artemisia longifolia. and Xanthium strumarium unions may be locally abundant. Pseudotsuga/juniperus Vegetation Type - This type predominates on approximately IOjlOOO acres .(13 per cent of the total) of moderate to steep northerly exposures where cool mesic condi­ tions prevail (Figure 10). Occasionally5 where well®developed stands of the Pinus ponderosa/Agropyron spicatum association occur on slight north- • 1 facing slopes and provide protection^ it extends upward onto more level sites. It seems to become slightly more extensive in the Missouri River Breaks west of the study area. .To the' east, it gradually becomes more restricted. Recognition is afforded by the dominance of Pseudotsuga menziesii or on serai or burned sites, the presence of conspicuous Prunus virginiana and Symphoricarpos occidentalis unions. Mature and undisturbed sites are characteristically occupied by the Pseudotsuga menziesii/juniperus scopulorum association. Stands of this — 36 “ association are usually characterized by a well-developed tree layer dominated by,?, menziesii and dense shrub undergrowth dominated by the . J. scopulorum and P. virginiana unions (Tables H I and H I I ). Finus ponderosa typically is present but poorly represented in the over­ story. Although this species may dominate, open, serai or burned stands, tree reproduction in mature stands, except for open disturbed spots, is limited to P. menziesii. This suggests that pine is serai on these sites, as it is in the Pseudotsuga menziesii zone of the northern Rocky Mountains (Daubenmire 1952). The Symphoricarpos occidentalis union, is best repre--. sented by S. occidentalis and Agropyron trachyeaulum. but Artemisia cana and Chrysothamnus viscidiflorus frequently occur in more open or burned stands. Both P. virginiana and G. viscidiflorus are susceptible and sensitive to heavy browsing by mule deer and elk. Dead and depauperate remnant plants attest to greater abundance of these species in former years. The herbaceous layer typically is dominated by Agrostis scabra.and/ or Agropyron spicatum. The latter is predominant in more open, serai', or burned stands. As a group,'forbs appear less.abundant than on any of the other vegetation typesj however, Selaginella densa often forms extensive mats on the organic matter which tends to accumulate on or near the ground surface and Achillea millifolium is common.. Soil surface cover appears higher than in any of the other rangeland types (Tables III and H i ) = Available evidence suggests that this association represents a topo­ graphic climax on the sites where it occurs. TABLE VIIo CONSTANCYj CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA ON THE PSEUDOTSUGA MENZIESIl/ JUNIPERUS SCOPULORUM ASSOCIATION, l/ Union: Taxa 2/ Cy/Cv/Fr 2/ Pseudotsuga menz'iessii: Pseudotsuga:menziessii 100/ +/ .3 Juniperus scopulo.rum: Juniperus scopulorum Rhus trilobata Antennaria dimorpha 100/19/38 100/ I/ 3 100/ 2/25 67/ +/ 3 Prunus virginiana: Prunus virginiana 100/ 2/ 5 Ribes cereum 67/ +/ 2 Rosa nutkana 100/ 5/35 Agrostis scabra 100/40/77 Fragaria virginiana 67/ 2/14 Galium aparine 33/ 3/ll Geum triflorum ,33/ 2/15 Potentilla arguta 67/ +/ + Symphoricarpos occidentalis: Symphoricarpos occidentalis 100/10/53 Agropyron’traehyeaulum 100/ 2/13 Agropyron spicatum: Agropyron spicatum 100/16/50 Agropyron smithii: Carex spp, 67/ 2/ 9 Tragopogon dubius: Achillea millifolium Aster commutatus Selaginella densa Solidago missouriensis 100/ 3/26 67/ +/ 3 100/38/60 100/ 1/10 ) TABLE VII» Continuedo ,Uninn; Cy/Cv/Fr 2/ Rock Bare Ground 6 7 / 1 / 8 0/ 0/ 0 I/ Data are mean percentages for 120 two x five dm plots distributed as -< 40 in each of three stands» 2/ Includes only those which occurred in at least two stands, i.e,, constancy of 67 per cent or more, or had mean eanooy'-coverage at least one per cent in the three stands. Other tarn are considered in Appendix Table XLIV'. , 2/ Constancy/canopy-coverage/frequency,• -t/+ indicates canopy-coverage or frequency values less than one per cent. I' - 39 - TABLE VTII,' CONSTANCY, COVERAGE, AND FREQUENCY OF TREES AND SHRUBS GREATER THAN 12 INCHES TALL IN THE PSEUDOTSUGA MENZIESII/ JUNIPERUS SCOPULORUM ASSOCIATION. l7~~ Plant Species Cy/Cv/Fr 2j Pseudotsuea menziesii ' 100/ 56/100 Pinus ponderosa 67/16/50 Juniperus scopulorum 100/30/100 Rhus trilobata 100/ +/67 Prunus virginiana 67/ 2/50 Ribes aureum ■ ' 33/ +/17 Ribes cereum 33/ +/17 Total Intercept (per cent) 106.0 l/ Data are mean percentages for six 100 ft line-intercept transects distributed as two in each of three stands„ 2/ Constancy/coverage/frequency. Frequency is the percentage occurrence among the six 100 ft lines. 4- indicates canopy-coverage values less than one per cent, Sarcobatus/Artemisia Vegetation Type Wherever dispersed clay soils containing considerable amounts of sodium occur, the vegetation characteristically is sparse and dominated by Sarcobatus vermiculatus. Footslopes and low benches along larger coulees and Missouri River bottomlands are the usual site (Figure 11), however, the type extends upward along adjacent slopes on clay slides (formed by downs lope movement of clay overlying unstable shale). Occasionally it may be found both on deep granular silty clay soils of higher outwashes or fans and in shallow upland basins. Despite some variation in community organization among the various sites, the usual prevalence of — 40 — Figure 11. Sarcobatus/Artemisia Vegetation Type. Coulee-bottom Bench Occupied by the Sarcobatus vermlculatus/Artemisia tridentata Association. Figure 12. Agropyron/Symphoricarpos Vegetation Type. Coulee-bottom Meadow Occupied by the Agropyron smithii/ Symphoricarpos occidentalis Association. — 41 — So vermiculatus and Artemisia tridentata unions seems to justify collective recognition of stands of this relatively minor type (approximately five per cent of the area) as the Sarcobatus vermiculatus/Artemisia tridentata association (Table IX). S. vermiculatus alone characterizes the shrub layer only on footslopes and benches along the Missouri River and on clay slides. The herbaceous layer typically is sparses often extremely so; however, where the association occurs in small stands on higher outwashes; it may be well-developed. Although the Agropyroh smith!! ‘union character­ istically dominates, the Poa seeunda union is conspicuous and may predominate in local stands. During spring, the latter is also well represented by Allium textile. Lomatium foeniculatum, Musineon divaricatum, Plantago spinulosa, and Thlaspi arvense. Summer forbs are typically of the Tragopogon dubius union. Those associated with A. tridentata or Xanthium strumarium unions are sparse or only locally common. Agropyron/Symphoricarpos Vegetation Type This type (Figure 12) is minor in extent, occurring only on silty clay soils of flats adjacent to water courses. Recognition is afforded by a wet meadow aspect dominated by Agropyron smith!!; however, the Symphoricarpos occidentalis union is sufficiently represented to recognize the community as the Agropyron smithli/Symphoricarpos occidentalis association (Table IX). Symphoricarpos occidentalis is the most abundant shrub and frequently occurs in dense patches. Artemisia cana and Chrysothamnus viscidiflorus are characteristically present but both are most prominent on better drained, marginal sites. Agropyron smithii TABLE IXo CONSTANCY, .CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF .TAXA ON . .. . THE SARCOBATDS VEHMICULATUS/ARTEMISIA TRIDENTATA ASSOCIATION. THE AGROPYRON SMITHIl/SYMPHORICARPOS-OCCIDENTALIS. ASSOCIATION, AND THE. XANTHlUM STRUMARIUML ASSOCIATION., l/,, ' . . . .... . . ... Association Sarcobatus Vermiculatus/' ' Agropyron smithii/ Artemisia.tridentata Symphoriearpos oceidentalis Xanthium strumarium Union:" Taxa 2/ ............ Cy/Cv/Fr 3/ Cy/Cv/Fr Cy/Cv/Fr Artemisia tridentataY Artemisia tridentata Sarcobatus vermiculatus: Sarcobatus vemieulatds Agropyron smithii; Agropyron smithii Koleria cristata Stioa viridula Poa secunda: •• Poasecunda ' Plantago,'. spinulo sa ThlasplIarVense Tragopogon duBiiis: Aster eommutatus ' Aster oecidentalls Chenopodium album Melilotus officinalis Vicia amerieana Symphoricarpos oceidentalis; Artemisia eana Chrysothamnus viseidiflorus Symphoriearpos oceidentalis Agropyrbn trachycaulum Poa epmpressa Glycyrrhiza lepidota 100/19/42 100/11/31 100/42/91 67/ 2/13 100/ 5/26 100/24/66 67/ 4/13 67/ +/ 2 67/ 4/ 3 100/ 5/23 100/ 1/10 100/ +/ 3 100/ 4-/ 4 100/88/100 100/ 3/15 67/ 4/19 67/ f/ 3 67/ 4-/ 4- 67/ 4/ 5 67/ 4/ 3 100/ 5/23 67/ 4/ + 100/ 4/12 67/ +/ 3 :■ 100/ 3/21 100/25/53 67/ 3/ 5 M t 67/ 4/ 3 67/ 2/10 67/ 4/ 7 67/ 2/ 8 TABLE IX. Continued. Association Sarcobatus vermiculatus/ Agropyron smithii/ Xahthium Artemisia tridentata Symphoricarpos oecidentalis strumarium Union: Taxa 2/ Cy/Cv/Fr 3/ Cy/Cv/Fr Cy/Cv/Fr • Distichlis stricta: Distichlis stricta 100/ 4/23 100/ l/ll Xanthium strumarium: Elvmus canadensis < 67/ 2/ 7 Hordeum .iubatum - 100/ 2/ 8 Snartiha pectinata 67/ + 100/ 2/ 4 Iva axillaris 67/ 2/11 Lepidium dehsiflorum . 100/ + / H Polygonum spp. 67/ t/ 9 100/ +/ 9 EUmex mexicanus" 100/ +/ 5 Xanthium strumarium • - 100/ 5/20 Bare Ground 100/52/98 . 100/37/95 100/ 90/100 Rock 33/ V +' 0/ 0/ 0 100/ 1/11 I/ Data, are mean percentages for 120 two x five din plots (40 in each of three stands) in ■ each association.■ - 2/ Includes only those species which occurred in at least two Stands5 i.e.5 constancy of 67 per cent or more5 or had mean canopy-coverage at least one per cent in any association. Other taxa are considered in Appendix Table ZLV. 3/ Constancy/canopy-coverage/frequency. -f-/+ indicates canopy-coverage or frequency values less than one per cent. — 44 = appears to be more abundant in this association than in any other community; however, other members of its union are poorly represented„ Poa compressa and DistieKlis strieta typically occur. The latter attains abundance only in lower spots or along the edge of watercourses. Glycyrrhiza lepidota is the most characteristic forb, but members of the Tragopogon dnbrus union are usually common. Allium textile, Lomatium foeniculatum. and Thlaspi arvense are common spring forbs. Although not analyzed in detail, Agropyron meadows on Missouri River bottomlands appear very similar in organization to this association. ' Xanthium strumarium Vegetation Type . This type (Figure 13) refers to the vegetation characteristically associated with cuts or beds of intermittent watercourses where the shale, silty clay, or gravelly substrate is typically moist and alkaline. The Xanthium strumarium union, comprised of species which tolerate or thrive under these conditions, usually.dominates the sparse vegetation growth (Tables III and IX). Xanthium strumarium is the most prominent and widely distributed species, Elymus canadensis, Spartina pectinata, Hordeum jubatum, and Rumex mexieanus are characteristic but usually restricted to marginal silty elay;deposits. Where they occur, members of the Agropyron smithii and Symphoricarpos oecidentalis unions and Distichlis strieta are also largely restricted to marginal deposits and clay banks. Melllotus officinalis-, which consistently occurs with the Xe strumarium union on these sites, often attains greater abundance than — 45 ~ Figure 13. Xanthium strumarium Vegetation Type. Coulee-bottom Cut Occupied by the Xanthium strumarium Association. Figure 14. Artemisia longifolia Vegetation Type. Steep, Loose-Shale Slope Occupied by the Artemisia longifolia Associes. “ l\.£> —- !j any other species but again largely on marginal deposits and clay banks= Where standing water persists for extended periods5, Eleocharis macrostachya and/or Scirpus paludosus may join the assqciation= / Artemisia longifolia Vegetation Type This type (Figure 14) is very limited and occurs only on the loose shale of steep cuts and slopes = Since these sites are generally similar to, but usually steeper than, those occupied by the Finns ponderosa/ Artemisia longifolia association, the latter might be expected to supplant the Artemisia longifolia associes in the absence of disturbance and/or as more stable conditions develop= This is also suggested by the vegeta­ tion composition of this community (Table X) which is similar in many respects to the understory vegetation of the Pinus ponderosa/ Artemisia longifolia association= The Artemisia longifolia union characteristically dominates the typically sparse vegetation cover (Tables III and X) = A= longifolia is usually the most conspicuous shrub; although Sarcobatus vermiculatus and members of the Juniperus scopulorum union frequently occur= S= vermiculatus is especially prominent and may predominate in local stands on south-facing slopes. The depauperate herbaceous layer is comprised largely of grasses and forbs of the Artemisia longifolia, Agropyron smithii, and Tragopogon dubius unions = Occasionally, some members of the Xanthium strumarium union attain abundance in local stands. . Agropyron spicatum may be conspicuous, especially on more stable sites. — 47 — TABLE X. CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA ON THE ARTEMISIA LONGI FOLIA ASSQ'GIES. I/ Union: Taxa 2/ Oy/Cv/Fr 2 / Artemisia longifolia: 1 0 0 / 8 / 3 5Artemisia longifolia Calambvilfa longifolia 6 7 / 3 / 7 Chaenactis douglasii 6 7 / f / 5 Mentzelia laevicaulis l p o / 4 / 8 Lactuca seriola 6 7 / + Salsola kali 6 7 / 4 . Juniperus seopulorum: 6 7 / 3 / 1 3.. Rosa arkansana Sarcobatus vermiculatus: 6 7 / 4 / 1 0Sarcobatus vermiculatus Agropyron smithii: Agropyron smithii '■ 1 0 0 / 4 / 2 6 Agropyron spicatum: 3 3 / 2 / 8Agfopyron spicatum Poa secunda: Descuriana pinnata 6 7 / 4 / + Tragopogon dubius: . . 1 0 0 / 4 / 8Chenopodium album Melilotus officinalis 6 7 / 6 / 1 3 Tragopogon dubius 1 0 0 / 4 / 2 Bare Ground 1 0 0 / 9 4 / 9 9 Rock ' . 1 0 0 / 4/ 5 l/ Data are mean percentages for 120 two x five dm plots distributed as 40 in each stand, , 2/ Includes only those which occurred in at least two stands, I ,e ,, con­ stancy of .6? per cent or more, or had mean canopy-coverage at least one per cent in the three stands. Other, taxa are considered in Appendix ■ Table X L V I , 2/ Constancy/canopy-coverage/frequency, +/+ indicates canopy-coverage or frequency values less than one per cent; a single + in place of these two values denotes occurrence in the stands but not,in the plots. — 48 — Other Vegetation Types The following vegetation types, characteristic of Missouri River bottomlands, occur on the study area but were not analyzed: The Populus/Symphoricarpos type occurs on approximately 1,000 acres where Populus sargentii and Symphoricarpos occidentails appear to be typical dominants of the tree and understory layers respectively| The Salix type comprises about 500 acres of dense thicket in which Salix interior. So bebbiana, and S„ mackenziana are common; The Agropyron/Symphoriearpos type covers about 250 acres and appears very similar to the Agropyron smithii/Symphoricarpos occidentalis association; The Sarcobatus type is restricted to footslopes and benches, and except for the paucity of Artemisia tridentata, appears similar to the Sareobatus vermieulatus/ Artemisia tridentata association. Approximately 1,400 acres of cultivated bottomland are variously dominated by alfalfa, barley, stubble, or weeds „ , POPULATIONS The distribution^ seasonal movementss and behavioral habits of mole deers elk, arid cattle as well as population trends of mule deer and elk were determined primarily by observations during 521 morning and evening periods (233 for cattle) along 11 vehicle routes which encompassed all portions of the study area (Figure I), Data were obtained from approxi­ mately 11,600 mule deer, 3»500 elk, and 25,000 cattle'observations = Mule deer and elk were classified with respect to sex,age,■young/female, and specific behavior when practicable. Locations were recorded to the nearest 0.05 mile from gridded aerial photographs and subsequently grouped into one and one-half square mile "blocksM to facilitate analyses. Supple­ mentary data were obtained through aerial surveys, hunter harvest surveys, and collections of mule deer and elk. Relocations of individually recog­ nizable mule deer and elk provided data on movements and home ranges. Mule Deer Distribution The mule deer were non-migratory and widely dispersed over the entire study area, though local population densities varied. Maximal numbers occurred on the south-central portion where close interspersion of well= developed stands of Pimus/Juniperus and Pseudotsuga/Juniperus with other vegetation types provided a relatively high security-level. Lowest densi­ ties prevailed west and north of Sand Greek. Although differences related to topography and vegetation influenced observed numbers, averages of 15.0 - 50 - The only distinct seasonal shift in distribution consisted of move­ ment onto Missouri Biver bottomlands during summer and fall as forage supplies on uplands became dessicated. It appeared -to involve only animals from adjacent slopes and possibly uplands within about two miles of the river. Mule deer were rarely observed on bottomlands after early December. The increase in numbers of mule deer observed along extensive open ridge- tops during winter and early spring (Figure 15) was probably a reflection of local movement and increased observability concomitant.with seasonal changes in food and range-use habits. Numbers of mule deer recorded in predominantly coulee areas increased simultaneously. Numbers of mule deer observed along the principal open ridgetops (Figure 15) and on Missouri Eiver bottomlands declined significantly during 1962. These areas generally were associated with main access roads and sustained a substantial portion of the annual hunter harvest. Although this appeared to directly influence the number of animals observed^ suggesting a general population decline^ there was evidence that these "declines" were at least partially only "apparent," Mule deer food, range-use, and activity habits under extremely poor range conditions dur­ ing I96O-62 rendered them exceptionally vulnerable to observation during the fall, winter, and spring. Availability of preferred forage on timbered- sites into the fall and winter of 1962-63 had an opposite effect. Increasing alertness and wariness were also evident in mule deer and and UoG mule deer respectively were recorded per observation trip along routes in these areas. OPEN RIDGETOP 3 5 - 40' ENTIRE STUDY AREA^ COULEE S 25IB 20-| 1 5' 10' 5i ^ x . * / (4) / I960 1961 1962 1963 1964 l/ Number of observation trips. 2/ Includes only those mule deer observed within "route" boundaries where the potential for observing most of the animals present existed. Figure 15. A Comparison of Mean Numbers of Mule Deer Recorded Per Observation Trip on Extensive Open Ridgetops (Roadside), Predominantly Coulee Areas (Away from Main Roads), and the Entire Study Area, 1960-64. I Vn H I “ 52 = ’■undoubtedly influenced observability» Average ®maber@ of, male deer observed, per trip in predominantly coulee areas remained about the same during 1962 as compared'to earlier years (Figure 15)« In view of decreased observabilityg this suggests that somewhat greater numbers actually used these areas during all seasons of .1962=63 than in the earlier period« Males appeared less uniformly, distributed than females s especially during summer and early fall, when single males and/or groups of two to five (occasionally more)' two-year&4)f-age and older' animals appeared to congre- gate on the same non-descript local areas. ' At least 25 such '“back.-, habitatsM were -known, for the study area. During summer, an average of 1.66 males was observed per trip through "blocks" which partially or entirely-encompassed-male-concentration areas. Only 0.24, males were observed per'trip through blocks ia other areas. Movements and Home Range- Sixteen mule deer fawns were individually marked using various combin­ ations of colored plastic tape and 2 x 3 inch flags of nylon impregnated ■ . ' , . fabric (Saflag Fabric, The Safety Flag Company of America, Pawtucket, B.I.) attached to an ear. Four were never relocated; three were last observed •- about one month after ..birth; and nine were individually relocated from Z- •to 21 times over periods of 5 to 29 months after tagging. Five Individ- . ually recognizable mule deer, one fawn and four adult males, were observed two to five times over periods of one day to four months. • A summary of movement data for recognizable animals other than fawns • observed only when less than six months old is presented in Table XI. TABLE XI. SUMMARY OF MOVEMENT DATA FOR MARKED AHD RECOGNIZABLE MULE DEER. Animal Age at First Obser­ vation Dates of Observation Number of Obser­ vations Distance Maximum First Between to Last Observ. Observ. (miles) .from Center . of Range l/ Mean Maximum Estimated Home Range (Acres) Female (D-I853) I day 6/6l - 11/63 21 1 .5 9 2/ 0 . 2 8 0.39 1.00 '3/ 300 Female 1—I LTNIA 11 mos. 5/61 - 10/63 22 1 .2 5 0.65 0.32 0.65 320 Male tJ ,D864) 4 -5 day 6 / 6 2 - 3/64 10 1 .0 9 0 . 9 3 0.38 0.62 4oo Male (D-1859) 2-3 day 6/62 - 2 /6 4 4 2.75 y 2.75 -- - --— — — — Male (D-1 8 6 9) 8 mos. 2/63 - 1 0 / 6 3 5 0.9 0 0 . 5 9 0 .4 3 0 . 5 3 300 Fawn (Recog.) 7 mos. H I 5/ 6 1 5 0 .3 4 0.18 0.11 0.18 — — — Male (Recog.) 19 mos. 1/62 - :3/62 3 . 1.00 1.00 0 .4 5 0 . 5 9 — — — Male (Recog.) Adult 5/61 - 9/ 6 1 2 0.87 OvS? -- — -- Male (Recog.) Adult 9 / 6 1 2 0 .8 4 0.84 a™ = a -- — — — = Male (Recog.) Adult 3 / 6 2 - 7/ 6 2 2 2.00 2.00 •=- = = — “ - l/ Approximte central point of the area "within which the animal was' oh served.. 2/ Maximum distance may have been influenced by hunters. Excluding this, distance was 1 ,1 2 3/ Maximum distance may have been influenced by hunters. Excluding this, distance was O.78 4/ This animal had moved 0 .4 3 mi. at 13 mos. A movement of 1 . 5 mi. occurred at about 17 mos. of age„ — 54 — Movements of marked animals relocated over periods of one or more years are illustrated in Figures 16 and I?. Home ranges of females appeared tc- encompass approximately 300 acres. Males seemed somewhat more mobile and may have had larger home ranges but few quantitative data were obtained# especially for males older than two years. Relocations of three marked males to 16# 20# and 21 months of age indicated movements through the second summer were comparable to those of females# as reported by Dasmann and Taber (1956). Subsequently# movements became more extensive during the fall and/or winter resulting in enlargement of# or establishment of# new home ranges. .Older males appeared to have greater mobility and larger home ranges. Home ranges of marked females were established, by the second summer and were partially superposed on those, of parent females. Considerable variation was evident among males. One (D-1864) ranged over an increas­ ingly larger area immediately adjacent to its home range as a fawn# at least until 21 months of age; another tJ ,1869) remained within the same general area on which it had been observed as a fawn until at least 16 months old; and the third (D-1859) was observed during the second summer on a male concentration area 0.4 mile from the tagging site and subse­ quently moved 1.5 mile during November to an entirely new area. These differences may have been influenced by social relationships during the , second summer. The three associated with another young male# an adult female and fawns# and an older adult male respectively. - 55 - ----------- — Male D-I864, June 1962-March 1964, Birth to I yr 9 mos. ---------- = Male D-1869, Feb. 1963-Oct. 1963, 8 mos to I yr 5 mos. ---- --- - - = Male D-1859, June 1962-Feb. 1964, Birth to I yr 8 mos. © = winter-spring • = summer X = fall I mile Figure 16. Movements of Three Marked "Yearling" Males on the Missouri River Breaks Study Area. — 56 — Female, D-1853, June 196l-Nov. 1963, Birth to 2 yr 5 moa. 0 = winter-spring X = fa l l Fem ale , D-1851, May 196: -Nov. Figure I? Movements of Two Marked Female Mule Deer on the Missouri River Breaks Study Area. - 57 - Differentially greater seasonal use of certain portions of home ranges was evident„ Predominant use typically occurred on brushy or timbered portions on one side of the home range during the summer and.on open ridgetop sites on the other side in winter. The.extent of use of the different portions seemed to vary between years. One of the marked females (D-1853) ranged extensively on the open ridgetop portion of her home range during the fall and winter of 1961-62 but apparently remained in the vicinity of the more timbered portion during most of the winter of 1962-63, Both marked females appeared to return to the same portion of the home range where they were born to give birth to fawns. One was observed during the fawning periods of 1962 and 1963 in the same locality where tagged ^ and one of the twin fawns of the other was tagged when one day old within ■ a few yards of where the female had been tagged as a day-old fawn. Mule deer food preferences and the quantity and quality of available ' . forage appeared to be primary factors influencing local and seasonal move­ ments within home ranges, Observations suggested that movement was most restricted during the summer ^ then increasing to winter or early spring as food habits changed to include greater representation of species from the open Artemisia/Agropyron vegetation type. That is ^ greater effort became necessary to satisfy nutritional requirements as the quantity and quality of available forage declined. The importance of the latter was indicated by yearly contrasts in movement with variations in production, and seasonal utilization of preferred species, In I960 forage production was "sub-normal^" summer-fall utilization of deciduous browse species was heavy5 and local movements increased appreciably to winter and early spring. During 1961, forage production was extremely low as a result of drought conditions, summer utilization of deciduous browse, was extremely heavy, and seasonal changes in food habits occurred much earlier than in i960. Also, fall rainfall resulted in a "green-up" of Sandberg bluegrass (Poa secunda) and junegrass (Koleria cristata) which concomitantly became important forage sources on open vegetation types. The combined effects were reflected in more extensive movement during all seasons'. During 1962-63, favorable growth conditions effected good forage production. Supplies of preferred forbs persisted into fall and winter, summer utilization of deciduous browse was light to moderate, and local movements appeared extremely restricted with only slight increase to winter and early spring. The presence of fawns, whose maximum summer movements were limited to areas about 0.5 mile in diameter, may have influenced mobility of parent females (Table H i ) . However, general observations suggested that females usually foraged independently until mid-August, and females without fawns as well as males were equally restricted in movement during the summer period. TABLE H I . MOVEMENTS OF FAWN MULE DEER DURING FIRST SUMMER AND FALL. Total Number Mean, Mean, Maximum Period Number of From First to Last Between _____________ _____ of Fawns Observations Tag Site Observation Observation June, July, Aug. 6 13 0.20 0.22 0.53 Sept.,Oct.,Nov. 4 =* 58 ™ 8 0.57 0.72 0.72 Population Trends ' ' No quantitative data are available for male deer populations, in. the Missouri River Breaks prior to 1947° Information furnished by long-time'.,- area residents suggests that, numbers declined during'the early 1900?s to' extreme scarcity during the 1920 °s and early 19300 s . The four=month - ■ hunting season with a bag limit of eight' deer j established lira 1895 was successively reduced ° By 192.3 limits 'were one month and one male-deer with i' . * ' • • , 1 1 " antlers not less than four inuhes ° From 1921 through 1930g an area eneom-. passing' .the. eastern one-half of. the study area was • closed, to deer hunting° The entire area was' closed, during. 1931-32° Hunting resumed in 1933 under regulations effected in 1923 ° .Petroleum County, lying immediately adja­ cent to the east' of the study area, remained closed to the -fomtirng of' mule" ■ deer until 1945° Hunting "was prohibited within the Fort Peek Game Range (,CoMo Russell Wildlife- Range) from 1937 through 1947. and again during 1949° Human depopulation, ..'increasingly effective law enforcement, and preda­ tor - control together with favorable deer range conditions which developed " during the period, of scarcity probably "proved conducive to the subsequent development of -the existing pqpuXatioBo Increasing numbers of deer 'were , noted during the late 1930 0Ss throughout the 1940'0S9 and into the early .19505S 0' . Hollibaugh (1944)' indicated that the "Breaks" herd was increasing at a tremendous rate undar excellent range conditions. ■ I n '1947y from " aerial surveys. Brow, estimated the male deer population as approximately five deer per square mileo That increases continued until at least 1951 was indicated by calculated post-hunting-season mule deer populations of ™ 60 ™ 3,330 in 1949, 4,850 in 1950, and 5,230 for 1951 for the Missouri Breaks of Fergus, Petroleum, and Phillips Counties (Johnson 1952)° The calculations of Brown and Brown (1950) suggest a minimal mule deer population of approx­ imately seven per square mile during the winter of 1950-51° No data indicative of population size are available for the 1952-59 period° Although mule deer numbers could not be determined precisely, the estimates presented in Table XIII are believed indicative of population trends on the study area during the 1960-64 period° Winter population estimates, believed to be conservative, were derived by ,-plotting numbers of deer observed by location and/or from aerial surveys = Trends in local densities, population structure, mortality (including hunter harvests), and changes in observability were subjectively considered° Sex and age structures were determined from ratios observed in the field from late November through January. Summer adults were computed from winter data by adding yearlings (last year’s fawns) in proportion to the sex ratios of fawns killed during the previous hunting season. Annual increments and late summer populations were calculated subsequently on the basis of September-October female;fawn classifications° . Year-to-year trends indicated decreasing mule deer numbers from I960 through 1962 and recovery during 1963 and 1964° The decline in winter populations from 1960-61 to 1961-62 appeared to be effected by low produc­ tion and/or survival of fawns, increased hunter harvest, and probable additional losses related to range conditions ° The extremely poor range conditions which prevailed during 1961 were reflected in a substantially TABLE XIII. SEASONAL AND ANNUAL TREND H NUMBERS OF MULE DEER ON THE MISSOURI RHER BREAKS STUDY -AREA, 1960-6 4 . Year Summer Populations Per Cent Increase from ..Winter Winter Populations ________________________I__________ :________ Per Cent Decrease Per Cent Change Total Male Female F a m Total Male Female FaTO Total Kill Summer Winter 196b 1,500 225 66b 615 1,100 H O 550 44o 27 . 20 “ - 1961 1,500 290 810 4oo 35 H 900 135 550 215 39 25 0 -18 1962 1,200 265 650 285 31 900 180 48o 24o 24 17 -20 0 1963 i , 4oo 34o 560 500 56 1,000 160 450 . 390 28 7 ' +17 +11 1964 1,500 420 580 500 50 1,100 250 525 325 27 10 +7 +10 I — 62 — reduced increment for 1962. The effects of improved range conditions after the spring of 1962, reduced mule deer numbers, and a female segment comprised largely of older animals were manifest during 1963 in a rela- tively large fawn crop and a substantially increased late summer popula­ tion. Further increase apparently occurred during 1964 when, with continued good fawn production, the increment should have resulted in a summer population approximately equal to that of. I960.. Although harvests decreased after 1961, total annual population losses seemingly were pro­ portionately similar and the trend in winter numbers paralleled that of summer. Numbers of mule deer recorded per observation -trip (Figure 15), . probably, were more indicative of the relative observability of animals than of population trends. For example, during 1961, "trend"counts indi­ cated populations 10 to 20 per cent greater than I960 whereas all other information indicated that 1961 summer numbers were subequal to those of the previous year and that 1961 winter-spring populations were 10 to 20 per cent lower. Average numbers observed per trip during the summer and fall of 1962 suggested declines of 25 per cent and 50 per cent respectively from 1961 populations compared with calculated decreases of approximately - 20 per cent. . Indicated numbers for winter and spring were ,25 per cent and 50 per cent lower than 1961 despite evidence indicating quite- similar population levels.. Observations did not fully reflect the increase which occurred in 1963. The influence of - observability was also evidenced b y . , • ! variation in the magnitude and direction of seasonal .changes in observed numbers (Figure 15) which indicated that similar proportions of mule deer numbers were not seen in any given season of the different years „ It i should also be noted that average numbers observed per trip during 1960=61 ■ ■ i ’ and 1961-62 when the animals were relatively observable corresponded V ' ' ■ i closely to numbers observed per trip on open ridgetop sites whereas data 1 ■ for the 1962-63 and 1963-64 periods were similar to numbers observed per trip into predominantly coulee areas (Figure 15)= Elk — 63 — Distribution The study area comprised the principal range of the elk population south of the Missouri River. Adjacent "breakse s p e c i a l l y west of the area, received variable but increasing use during the four years» Some unexplained movement across the Missouri River occurred. The elk were non-migratory, but differentially utilized various por- t tions of the area during the year. Vegetational complexes and availability , . x of preferred forage, as influenced by general range conditions, annual vegetal growth, and/or prior utilization, appeared to be the primary deter­ minants of seasonal distributions and variations thereof. Physiographic and weather factors apparently were secondarily' related. Variation in use of local areas within and between seasons and years corresponded closely to changes in distribution arid intensity of grazing by domestic livestock. The Iate-spring and summer distribution (Figure 18) coincided closely Additional Winter Range 1963-64 Northrrunary Auuxnrunax nxj Winter Range Y'/S/.'" * Range 1962-63 Figure l8 . Summer and Winter Distributions of Elk on the Missouri River Breaks Study — 65 — with the distribution of more extensives well-developed. stands of the Pseudotsuga/juniperus vegetation type. Principal differences between years were those associated with use of local areas within the circumscribed area. Elk generally' dispersed over all but the extreme eastern portion of the study area during early fall. Use of Missouri River bottomlandss which appeared to increase during the years of the studys occurred largely during this period. Concentration began in late October or early November. During 1960* 1961* and 1962* a majority of the animals congregated during this period in the vicinity of cattle winter .pastures on the south-central portion of the study area. Significantly fewer elk utilized this area during 1963 when the population remained more dispersed and a segment con­ gregated in the vicinity of previously ungrazed pastures along the western boundary of the study area. Use of the south-central area generally ceased during early December when cattle were turned onto and/or snow covered the grass-dominated pastures. Winter elk distributions (Figure 18) encompassed portions of the area characterized.by extensive stands of the more open vegetation types where grasses were prominently represented and little or no livestock use. The southeastern portion* much of which was only lightly grazed by cattle* was used almost exclusively during the winters of 1960-61 and 1961-62. During the latter* elk were observed largely on the central part where* with above-average range conditions* the extreme drought had minimal influence on the quantity of forage available. Elk were more dispersed during the winter of 1962-63* evidently in response to the increased availability of forage on much of the area. Although most elk utilized the southeastern area in a distribution similar to the winter of 1960-61=, some use occurred on the western portion of the study area which had little or no prior livestock use ^ Forage conditions were generally excellent throughout the area during the winter of 1963-64» The elk' remained relatively widely dispersed with about one-third of the population wintering largely off the study area in the area lying west of Highway 191» The rather abrupt dispersal from the wint er '-.range during late March coincided approximately with the onset of the spring "green-up»** Most of the,animals returned directly to the summer range. Use of local areas during early spring appeared inversely related to range conditions and directly related to the degree of prior livestock use. ' This undoubtedly reflected the seasonal elk preference for Poa Secundaa Koleria cristata => and early spring forbs which were commonly more abundant and/or more , readily available on these sites. Movements Ten recognizable adults and three marked calves were individually relocated 2 to 19 times over periods of one day to four years. Movements of representative individuals, relocated at least 10 times during two or more seasons, are illustrated in Figures 19, 20, 21, and 22» Seasonal movements of these elk corresponded closely to and support distributional changes described above. — 67 — X - fall• - summer 0 = winter # = spring MILES Figure 19. Movements of an Adult Female Elk from July 1962 to December 1964. The Animal Was Approximately Four and One-half Years Old When Killed at Last Relocation. Figure 20 Movements of a Male Elk Calf from June to December 1962. •= summer O = winter fall 0 = spring — 68 — r-r=! Figure 21. Movements of a "Yearling" Male Elk from July I960 to April 1961. Figure 22. Movements of an Adult Male Elk from January 1961 to December 1964. The Animal Was Approximately Nine and One-half Years Ola When Killed at Last Relocation. The elk were highly mobile (Table H V ) » During springe, summers and fall ^movement typically was erratic. There were variable period of restricted movement on local areas broken by abrupt shifts involving more extensive movement. Constant "wandering" was characteristic of winter movement. In general^ movement was most limited during summer and most extensive during late winter and early spring. Rapids extensive movement occurred during all seasons. One cow-calf group moved more than five miles from morning to evening during summer5 another traveled more than seven miles overnight during winter. Maximum distances between any two reloca­ tions of a given individual were; adult females 14«5 mile; calfs 7 = 0 mile; adult male,, 13 = 2 mile; and yearling male, 11.6 mile. The extreme mobility apparently enabled elk to adjust rapidly to environmental changes and constantly utilize the most favorable portions of the range. Population Trends The elk population developed following the release of 31 animals (22 adult females, 6 calves, and 3 adult males), transplanted from Yellowstone National Park, on the south-central portion of the present study area during February 1951= No data on growth of the population were obtained prior to 1959 when Janson (Pers. Comm.) estimated a minimum of 80 elk on the area prior to a special hunting season to alleviate damage on private lands. ™ 69 ™* - 70 - ( TABLE XIV. SEASONAL .AND ANNUAL MOVEMENT OF RECOGNIZABLE ELK ON THE MISSOURI RIVER BREAKS STUDY AREA. Group Sample Mean Distance Between Relocations (Air Miles) Class Size Spring Summer Fall I/ "Winter Year Adult 4 4 ; 3 (3 )2 / 2.0 (10) 1.8 (2) 6.1 (8) 3 . 7 (2 3 ) Calf 4 3 . 3 (I) 2.2 ( 9 ) . 1 . 8 (7) 6.4 (I) 2.4 (1 8 ) Adult 2 6 . 7 (4 ) 1.6 (I) 7 . 3 (I) 3.0 ( 7 ) 3 . 3 ( 1 3 ) Yearling - 3 3.8 (4 ) 2.2 (2) 3 . 4 (8) 4 , 2 (1 4 ) —Calf combined 8 4.0 (4) 2.1 ( 1 9 ) 1.8 (9) 6 . 1 ( 9 ) 3.1 (41) Total 13 4 . 9 ( 12) 2.1 (22) 2 . 4 (1 0 ) 4 . 3 ( 2 4 ) 3.4 (68) l/ Fall data largely from Oetober-November period during which elk congregated on south-central portion of the area. 2/ Number of relocations in parenthesis. The elk on the study area during the 1960-64 period (Table XV) did not comprise a discrete population. Undetermined numbers occurred on adjacent areas during all years. Year-to-year.relationships of numbers in the various sex and age classes on the study area indicated interchange of individuals■and/or groups between the study and adjacent areas. On the basis of data from Table XV and k n o w losses, minimum contributing summer populations of 90 elk in 1959, H O in i960, and 135 during 1961 were necessary to produce the sex-age structures observed on the study area during the I96O-62 period. For example, the total of 13 yearling males for the winter of 1961-62 is not compatible with the 20 calves- for 1960-61 when sex ratios are■considered» Farther calculations suggest total summer populations in excess of 160 and 200 respectively during .1962 and I963. TABLE XV. WINTER ELK POPULATIONS ON THE MISSOURI RIVER BREAKS STUDY AREA, 1960-64, Calves:' 100 Males: 100 Year Total Males Females Calves ' - Adult Yearling 1960-61 70 8 10 32 20 . 62 56 1961-62 80 9 13 41 17 42 54 1962-63 108 12 13 54 29 . 54 46 1963-64 108 13 9 51 . 35 69 43 * The mean net recruitment to the population, based on observed popula­ tion structures, calf production (Table XV), and losses, was.approximately 35 per cent during the four years. Assuming a similar rate of increase in years prior to the study, a much larger population could exist. It seemed unlikely, however, that actual numbers were greatly in excess of those calculated.above, 'Either mortality was of greater than observed propor­ tions or emigration from the area has provided effective control over population growth. Trends in numbers of elk on the study area during other seasons appeared similar to that of winter. Spring populations generally appeared to be somewhat lower than those of previous-winters. Summer numbers apparently were subequal to those of subsequent fall and winter periods when some movement onto the study area seemed to occur, Itiiown losses - 72 - during the fall varied from two in 1962 to eleven during 1961. No legal hunting of elk occurred during the study. Cattle Distribution ■. The entire area, except portions on some Missouri River bottomlands, was subject to use by cattle. During the April !-November 30 grazing season, greatest cattle densities typically occurred on the southern one- half of the.study area; light or occasional use prevailed on the northern and extreme eastern portions (Figure 23). The only important exception occurred during 1962 when the area of greatest cattle density extended I further north;along the Wilder Trail and light use.prevailed over most of the southwestern portion. Use of local areas varied considerably during all years, reflecting changes in forage availability, water supplies, and weather conditions. The areas of greatest cattle densities were major coulee bottoms and uplands characterized by dominance of the Artemisia/Agropyron and Agropyron/Poa vegetation types associated with ridgetops, slight to moderate slopes, and well-distributed water sources. Areas of light or occasional use were those with inadequate water supplies and/or broken topography with steep slopes and limited ridgetop areas. . Access to the extreme eastern portion was also limited by fences. It appeared that grazing pressures were substantially greater where cattle could disperse from water over extensive areas of primary range types. Use of distal Winter Pastures Greatest Apr-Nov Cattle Densities North Light Apr-Nov Cattle Densities 1 0 1 miles Figure 23. Distribution of Cattle Densities, "Generalized," for the April-November Grazing Season and Livestock Wintering Areas on the Missouri River Breaks Study Area. ' - 7 4 - '. - -.. ' ' . , - portions of ridges where the dispersal area was limited was generally lew . despite sufficient water. ' ■ ■ Cattle were observed on flight w.e® areas most frequently during fall, ^following, heavy .utilisation of forage' on preferred s i t e s when rains pro- . vided-additional.water'sources and/or cooler '.temperatures "enabled mo-re',. ■ extensive, movement. . Some -use. also occurred during spring's especially In drought years $ when standing runoff water was available and/or forage was ■ relatively more .abundant, these-areas were most heavily.used during' 1961' . when reduced production and heavy use" of, forage on more intensively .grazed areas, forced cattle to' range; more widely? ' • Host cattle ,were turned- onto winter pastures and/or' removed.'.from 'the area by the end of the grazing season.Winter pastures on the'.south”., . central portion of the area were not grazed-by cattle- during ,the 'winter of 1961-62. The upland part of ! the north=4:ehtral winter pasture was usually ■ ' ' ' . r- , ■ •/ •* ,r\. < • ’; i, I , ■ • % . . grazed only during the' early >d.'nter ‘ period. Population1Trends ' Fourteen operators held permits to graze I5646 cattle (plus - calves) on or adjacent to the study area (Pars. Comm.5 BlZ5 Lewistowh5 Montana). Of these5 11 permittees with I5 205 permits ran their stock largely', on the area 1' 2 with 166 permits partially -used the area 5 and I with1175 permits usually did not use the area. Actual numbers of cattle on the study area- varied considerably within and between years, some' proportion of the total always ranging onto immediately adjacent areas. This number appeared to increase from spring to Summer5 then decreased slightly to fall.. Greatest - 75 - numbers of cattle occurred on the study area during the spring and summer periods of 1961 when most operators turned their cattle into the "breaks" where forage and water conditions were somewhat better than on adjacent areas. Lowest numbers occurred during the fall of 196l5. when most cattle were removed long before the end of the grazing'Seabon5 and during the spring of 1962, when most operators delayed turning their cattle onto the open range until early May. ' RANGE USE Mule deer, elk, and cattle observed along established routes (see previous discussion, page 49) were recorded with respect to time of day, type of activity occurrence on the various vegetation types, slopes, and exposures, and distance from known water sources„ Environmental condi­ tions (temperature, precipitation, wind direction and,velocity, cloud cover, ground,condition, and moon phase), times of surprise or sunset, as i : - well as duration and distance of travel were recorded for each observation period. Observations of mule deer and elk were recorded during all periods. Observations of cattle were recorded at weekly intervals after August I960, except during the December-March period when observations were ■ irregularly recorded whenever cattle were encountered on upland winter pastures„ All portions of the study area were usually-visited at intervals of one month or less from June I960 through September 1963, during November and December 1963, and during January and March 1964» All observational data were transferred to standard 80-column punch cards and subsequently tabulated and statistically analyzed using IBM unit records equipment and computers» Mule Deer A total df 11,381 mule deer observations^ used in evaluating range- use habits, was recorded as follows: 1960-61,■. 5,577; 1961-62, 3,431; 1962-63, 1,79%; and 1963-64, 776. \- 77 - Activity Habits The time of day of each observation was recorded to. the nearest quarter-hourj then converted to the nearest hour relative to sunrise or sunset to provide a basis for comparisons of data within and between time periods. A U animals were classified as feeding, bedding, alert,or traveling on the basis of their behavior when first observed. Mule deer typically were active and observable mostly within three-to- four-hour periods in early morning and late afternoon. They appeared more active during,mornings when an average of 26' animals was recorded per observation trip, compared to 19 per trip during evening, and "activity periods" were slightly longer. Greatest numbers were observed during the first and last hours of daylight, with 75 per-cent or more of all observa­ tions usually' occurring within two and one-half hour's after sunrise and before sunset.and more than 90 per cent within three and one-half hours. Percentage distributions for morning, evening, seasonal, and yearly periods (Figures 24 and 25) were similar despite differences in numbers of animals observed, ranger-forage conditions, weather conditions, as well as mule deer densities, food and range-use habits, and behavior. These factors generally effected variation only within the normal "activity periods." The "prolonged" activity periods of late winter and early spring reflected more nearly equal distribution of observations through normal activity periods and some extended feeding later in the day. These - 78 - Summer Fall Winter Spring 4 + 3 2 1 0 Hours before sunsetHours after sunrise Figure 24. Percentages of Total Mule Deer Observed by Hour Relative to Sunrise and Sunset During Seasons. 1960-61 1961- 62 1962- 63 1963- 64 4 + 3 2 1 0 Hours before sunsetHours after sunrise Figure 25. Percentages of Total Mule Deer Observed by Hour Relative to Sunrise and Sunset During Years. Iphenomena were most pronounced under the extremely. poor; range-forage ' conditions which prevailed during 1.961=62<>. The etShortra periods of;:fa-U : were, characterized by concentration of activity in■the first'and last hours , of daylight and sharply decreased activity later'in., the" day-5 except during- the fall of 1961 when activity was "prolonged'61 in a distribution similar to that of winter, A trend toward a fall-like distribution during all seasons occurred after the spring of -1962 as, rafige^'forage coaditions ' improved .'and mule-deer . becairid-increasingly wary=1. . The' distribution -of- activity during normal activity periods, varied with,temperature. The proportions of the.total male deer observed during - the first .and last hours of daylight,increased as temperature increased (Figure 26), The most • significant differences were' associated with sub- . z e r o temperatures, when onlyv 25 per cent, of the -animals were observed ■ during the,first and last hours of daylight and- approximately 30 per cent. of all observations were recorded more than, two' and one-half hours after • • • ‘ 1 • ,' • ' " . , . sunrise and before'sWiSet and-'with temperatures ,above'80 degreess when .. . nearly 60 per cent and less than 10 per Cent respectively of all observa­ tions occurred during.those periods„ Other weather factors undoubtedly influenced variation b u t 'significant effects were -obscured by interactions, ■ Feeding was'the. predominant-activity recorded-for mule deer observed - ■ during all hoprs of activity, periods- (Figure 27%- s e a s o n s a n d years ' . . (Figure 28,), ■ The hour' of most intensive feeding tea- between one-half and one and one-half hour'/ after sunrise and'.before .sunset (Figure 27), Decreased movement -'associated'jd.tk.this' may -have influenced the •' . ■ z' - I = 4-year means ---- before sunset FEEDING ALERT BEDDING ■BEDDING 1960-61 1961-62 1962-63 1963-64 Hours after sunrise/before sunset Hours after sunrise/before sunset Annual periods Figure 26. Percentages of Total Mule Deer Observed by Hour Relative to Sun- rise/Sunset in Relation to Temp­ erature. Figure 27. Percentages of Total Mule Deer Figure 28. Percentages of Total Mule Observed by Activity Class by Deer Observed by Activity Hour Relative to Sunrise/ Class by Years. Sunset. I g I ( ™ 81 “ characteristic decrease in numbers of animals observed during that time (Figures 24 and 25)• That type of activity influenced observability was further illustrated by the small differences in proportions observed bedding through activity periods, Numbers observed differed greatly, . v K H U B The decrease in proportions of mule deer observed feeding and the complementary increase in traveling during the four years (Figure 28) can be interpreted as an increase in wariness, This was particularly apparent along roads on major ridges. These changes appeared correlated with hunting pressures during the 1960-62 period,, suggesting selective removal of less wary animals from the previously lightly hunted population and/or that mule deer became increasingly responsive to disturbance, A summary of observed mule deer activities by seasons, and years is given in Appendix Table X L H l . A complete summary of weather factors as related to mule, deer activity is presented in Appendix Table XLVIII; Group Size Mule deer were recorded collectively when observed, within about 100 yards of each other and/or their behavior was indicative of a "group" relationship,. Groups ranged in size.from I to 25 animals, but the most common numbers were near seasonal means (Table X H ),' Groups typically were smallest in summer when mule deer were rela­ tively dispersed and using a variety of range types. Those of more than four or five animals were almost invariably comprised of males. Aggrega­ tion occurred slowly through fall and winter, concomitant with increasing - 82 - use of wOpen*' vegetation types and/or concentration on certain sites. The largest groups generally were observed in late winter and. early spring. This increase in seasonal group size apparently reflected.more strongly a greater incidence of association on mutually used areas.than changes in social relationships. Support for this statement was;.provided by observa­ tions that the composition of "groups" associated with recognizable animals changed frequently. The number of mule deer in a group reflected the loca­ tions within home ranges of' individuals on overlapping home ranges, Group sizes were somewhat larger in 1961-62 than in other years. Possibly, this was related to more concentrated use of certain vegetation types. Group sizes in general decreased in subsequent years as the animals remained more widely dispersed. TABLE W L , AVERAGE SIZE OF GROUPS FOR MULE DEER OBSERVED DURING SEASONS AND YEARS, I96O-64, Year Summer Falix Winter Spring Year 1960-61 1,5 (731)^ 2.5 (734) 3,7 (1,441) 4.4.(2,671) 3,1 1961-62 1.5 (669) 2,8 (587) 5,2 (845) 5.0 (1,330) 3,2 1962-63 1.5 (483) . 4.0 (526) 3.8 (626) 2.6 1963-64 1.6 (376) .. 2,5 (160) 3.5 (120) ■5.7 (120) 2.2 1960-64 > 1.5 2.6 I. ' 4.1 4.5 V 3.0 ■ l/ Number of mule deer in sample. == 83 “■ Use of Vegetation Types ■ ' Mule deer relations to vegetation types were evaluated by recording the type on which individuals were•first■observed» To consider bias 1 ’ resulting Trpin greater observability o n -"open" typess ^observability ' indices'" were determined by comparing average numbers recorded per observa­ tion trip with calculated "expected values"j i.e., average numbers which could have b^en recorded, based ootrdatsa; for the spring of 1961 when, maxi­ mum numbers of mule deer were observable and yearly population trends„ For example, about 78 mule deer were "expected” per observation trip during the summer of i960 in contrast to ■12 b4 ' (16 per cent - "observability index81) actually recorded0 The 84 per cent not observed evidently-were .using "timbered” types ,and/or were inactive6 . Frequencies of male deer occurrence on vegetation types receiving more than 10 per. cent of the total use- during monthly periods are plotted in Figure-290 The complete results for seasons, years, and the'four-year period are presented in Appendix Table XLI-X0 .' S m m e r (June, July, August).— ,The Fimis/juniperus type was used most intensively., in summer when it was the most important vegetation type for all mule deer activities. The Pinus ponderosa/Aaropyron spicatum associa­ tion appeared to be the most important community, -especially for feeding. Greatest use on the P. ponderosa/Juniperus scopulorum and P.'ponderosa/ ■ Artemisia longifolia associations occurred in- early- summer, though the former remained,important- for.-bedding and escape throughout the period. Artemisia/Agropyron Pinus/Juniperus Pseudotsuga/Juniperus Sarcobatus/Artemisia \ / \ ; • V J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M I960 1961 1962 1963 1964 Figure 29. Monthly Frequencies of Mule Deer Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the Total Use During at Least One Month. Use of the Art end sia/Agropyron type ranked second during three of - 85 - the four summers. , Use was predominantly on the Artemisia tridentata/ Agropyron smithii associes and the A. tridentata/A. spicatum association on smaller ridgetops and benches and/or along the edge of "timbered" types. Many forbs preferred for food were members of the Tragopogon dubius union which was best developed in these communities. Observed use of the Pseudotsuga/Juniperus type was moderate over-alls but varied from minor during the summer of I960- to relatively heavy in 1963. Most intensive use typically occurred in June during the fawning period. Over-all use of the Sarcobatus/Artemisia type was minor but reached significant proportions during some months. Other vegetation types received relatively little use because of restricted distribution and/or small total acreages. Occasionally# as in i960, abundance of preferred forbs on the Agropyron/Symphoricarpos and Xanthium types attracted intensive use. The low observability index (average 16.6 per cent) for the period suggested that use of the Pinus/juniperus and Pseudotsuga/Juniperus types was much more intensive than observations indicated, and that the relative use of the Artemisia/Agropyron type was minor. Fall (September, October, November).--Relative intensities of use on the two most heavily used vegetation types during this period varied. The Artemisia/Agropyron type, which received the highest percentage of over-all . ■ t * use for the four years, was the most heavily used type during i960 when it - 8 6 - received its greatest fall usage. Utilization of the Pinus/jtmiperus type approached that of Artemisia/Agropyron in 1961 and exceeded it during 1962 and 1963. ■< Use of individual communities in fall differed from summer, especially ' ' . / ' ' on the Artemisia/Agropyron type where activity became 'more .'intensified. on the Artemisia tridentata/Agropyron smithii associes and oh the ' , "" ■>;:- A. tridentata/A. smithii/Bouteloua gracilis association which received the greatest usage. This appeared related to utilization of: Ghrysothamnus nauseosus and occasionally, Poa seeunda during fall "green-up." The Pinus ponderosa/Agropyron spieatum association remained most important on the Pinus/Juniperus type. The P. ponderosa/ Artemisia longifolia association was used rather extensively during mid- September when A. Iongifolia was briefly important as a food source, but later received only very minor use as did the P. Ponderosa/ Juniperus scopulorum association. Use of the remaining vegetation types was minor, although for some, over-all use ip fall was greater than in summer. Differences in intensity i • of use for the,different years was largely related to precipitation and its influence ;on certain food plants« The observability index of 19«3 for all suggested continued use of the Pinus/Juniperus and Pseudotsuga/juniperus types substantially in excess of observed intensities. - 8? - Winter (December, January, February).— The Artemisia/Agropyron type received substantially greater use than any other type for the entire period except for 1962-63 when use of the Pinus/Juniperus type was closely comparableo The Art emisia/Agropyron type was especially prominent for ' feeding, receiving about two-thirds of the total. Lightest and most vari- able intensities of use were indicated for early winter, especially in ‘ ■ . .. •;/ 1962-63 when the type was of secondary importance through January. This apparently was. related to the availability of preferred. and/or "variety" forage on other vegetation types. Percentages observed on the Artemisia/ Agropyron type increased monthly, reflecting increasing mule deer depend­ ence on Artemisia tridentata and associated taxa. The A. tridentata/ Agropyron smithii associes and the A. tridentata/A.■ smithii/ Bouteloua gracilis association"remained the most important■communities. The increased importance of the latter through the winter may have been influenced by greater abundance of A. tridentata» but • v; ■ Chrysothamnus nauseosus and Phlox hoodii, which received appreciable use when available, were also more abundant here. The heavily grazed, exten­ sive open ridgetops characteristic of this association held less snow. The Pinus/Juniperus type ranked second in importance. The intensity of use for feeding varied in relation to amounts of forage available, generally decreasing throughout the winter but generally increasing during the years of study. A large percentage of the feeding, especially during the first two winters and in late winter of all years, was associated with ' 8 8 . - movement to. and from bedding sites as'well as with utilisation of ' . Juniperas scopuloram -Which, mule deer used deliberately when feeding on '■ Artemisia tfidentata. The most intensive use on the type.oceurred during and for a few days after winter, storms. T h e .Pinus:ponderosa/ . . > '■ association was, the most important community. The heavy. .'. use during and immediately after. snowstorms. ' Sarcobatus/Artemisia fl the only other type of any importance during winterj, received minor use. This occurred mostly during December and January and was predominantly on steeper slopes locally .dominated, by the type in contrast to earlier- use largely o n . coulee-bottom sites „ / ' The Pseudotsuga/Juniperus type, restricted to windswept, snow- retaining1, northerly exposures, .'received only very minor use. Most was along upper margins of' the type, transitory, or during mild, open periods„ Little or no use wa s .observed on other vegetation types during. winter.' - Observability was-mmCh increased over summer and fall, Although greater than indicated percentages probably1 used the Pinas/Jmiperus type, ■' it seems extremely unlikely that the difference was significant enough to Alter conclusions AC ,to t h e ■relative importance of the various types„ Spring "(March, 'Appil, May) Artemisia/Agro^ron type received the greatest use throughout- the period <>■" Ii was used most' interns I w l y during late'EjArbh and/or early April concdWltamt with the appearance of green plant@.o Early, spring activity ImA Centered upon the Artemisia tridentata/ Agropyron amithii/Boutelom grapills association. This appeared’related to the preeminent Poa seeunda union which was --an .important source of mule: deer- forage during the period. The relative use and importance of. the ■ .Oi m 6 n declined sharply;after. mid-April as;, forage became . ■: • ' ■ ■ ■ ■ ■ ' ' increasingly available bn other types. ■ : ' ■ / v ■' • ■ ■ Use of the Pinus/juninerus type was: relatively minor until about mid- .• ; . .;"v ' ' ■ ■' . V- ’ ; ■ April, then increased rapidly until:late May when observed' use was compar- ’ ■ ■ ■■ ■ ’able to summer o /" The Pinus ■ ponderbsa, received the ,greatest use ^ though numbers of mule ,deer observed on the Po ponderosa/Juniperus 'seopulornm association, increased considerably ,after new growth -appeared on ’shrubs A • f ■: ' Sarcobatns/ArteMsia was the only other vegetation type which 7? ■ ■ V T ■ • ■ • . ' ■ -'b'!' ■ received-’appreciable spring, use. Most, of this appeared related to utili™ -■ ' ■ ■ ■ ■; - ’ ' , ' zation of Poa secunda and associated forbs which were well"represented bn coulee-bottom sites. . “ . ' , Mule deer typically were highly observable during early spring. , Observations^ particularly for 1961 and 1962# probably reflect relative intensities of u s e .On the various' vegetation, types more.closely than for 7 ptjher periods of the year. .. . > ■ Yearlong and Year-to-Year Trends.— The results' indicated significant : • trends in mule deer relations'to -vegetation types. major type# received.maximum'use.And was.the single# ,most important type during suiraner. The moderately extensive ■ Pseudotsuga/Juhiperus type appeared to rank second.. . Relative use of both-declined slowly through fall, the latter becoming relatively unimportant except as escape cover by early winter» Though generally declining in use, the Pinus/juniperus type remained moderately important for feeding"and most important for bedding and escape until early spring. Use of both types increased'rapidly to summer levels during late spring. Relative- use and- importance of Artemisia/Agropyron, the other major type, was lowest during summer. . Use generally increased through fall, becoming of major importance during early winter. Most extensive use occurred in early spring when it was by far the most important habitat type for feeding. The moderately extensive Sarcobatus/Artemisia type received minor use during all seasona. The minor Agropyron/Symphoricarpos type was of some importance during summer and fall and little used during other periods, Other vegetation types were very minor in both area size and mule deer use. Differences between years were important. The consistent decline in \ relative seasonal use and importance of the Artemisia/Agropyron type from I960 to early fall of 1963 was complemented by commensurate increases in ■ use of other types, particularly Pinus/Juniperus and Pseudotsuga/Juniperus. The prominence: of the Artemisia/Agropyron type during i960 and 1961 . . . appeared related to over-utilization of preferred deciduous browse on "timbered" types under drought conditions and/or high mule deer populations resulting in some shifting to the former. The most significant-differ­ ences, occurring after the spring of 1962, were related to increased growth of forbs., especially Melilotus officinalis. Use of these plants by mule deer decreased utilization of preferred browse during summer resulting - 90 - in persistent availability of forage on all vegetation.types through fall and winter o .. : Weather conditions also influenced use on the- various typesc Greater percentages ,of total animals • observed were in “timbered1* types at temp­ eratures below 0® and above 80® than with more moderate temperatures for ■■■;., . ■ . ■■■- ■ . ;■■■ ■ ' summer and winter respectively <> Use of the Artemisia/Agropyron and Agropyron/Poa types.decreased progressively: with■ ‘find velocities above ' 10 mph. During winters ^ the Pinus/Juniperus ■type was used most, intensively during periods of complete snow cover. A summary, of use of vegetation . types in relation to weather conditions is presented in Appendix Table L. Use of Slopes Steepness of the slope on which each mule deer occurred was estimated and recorded by class as followss. Glass I = O0-IO61 Class 2 = H°-25®| Class 3 = 26®-35'°J and Class 4 = 36®+. The results for seasonal and yearly periods are presented in Table X H l =■ TABLE XVII. PERCENTAGES OF TOTAL MULE. DEER OBSERVED ON SLOPE CLASSES, BY SEASON AND YEARS, 1960-64. . Season. Summer Fall Winter Spring 1960-61 1961- 62 1962- 63 1963- 64 1960-64 Sample ' , . Degrees of Slope Size •. 0-10 11-25; 26-35 36#- 2,256 . 41.4 .':aW4 22.5 . 11.7 1,647 49.2 23.8 '16.9 10.0 . 2,932 49.6 21.8 . 17.9 . 10.7 4,746 64.4 . 19.9 11.8 - . 3.8 5,577 59.1 2066 " ■■ -13.3 - '■ 7.1 3,431 . 50.8 . 23.3 16.2 9.6 1,797 45.7 22.3 23.2 8.8 .776 . 50.5 23.0 20.8 $.6 lli$8i. ' : / 54.0 ' 21.8 . ' 16.2 . 7,9 - 92 - Approximately 50 per cent of the male deer were observed on slopes of 11° of more but few animals were observed on slopes steeper than 45®» Seasonally, steeper slopes were used during summer» This reflected the predominant use of timbered vegetation types. The animals were concen­ trated on level to gently sloping sites during late winter and early spring when primarily, using the Artemisia/Agropyron type. The variation between years was related to differences in use of vegetational types. Use of Exposures The direction of exposure was recorded for all mule deer observed on slopes. The results, summarized by season and year in Table ZVIIl, appeared more indicative of relative mule deer use and observability on the Pinus/Juniperus and Pseudotsuga/juniperus vegetation types and/or the relative amount of area with each exposure than distinct seasonal prefer­ ences. South, southwest, north, and northeast-facing slopes were generally most extensive on the area. Southerly-facing slopes seemed to receive more use during all seasons, especially winter. Increase use of northerly exposures during fall may have been related to use of thicker cover. The subsequent decrease in winter and spring was influenced by snow and wind. .,v. ■ ■ - , Relations to Water Distances' to the nearest stock water reservoir, well, spring-seep,. or to the Missouri River were, determined from gridded aerial photographs and recorded by quarter-mile- distance classes for all observed mule deer. Temporary water, including snow, was not considered quantitatively. - 93 - TABLE XVIIIo PERCENTAGES OF TOTAL MULE DEER OBSERVED ON VARIOUS EXPOSURES s BY SEASON AND YEARS, 1960-64» Exposure l/ oeason Size., O 0-IO0 Slope N NE E SE S SW W NW Summer 2,256 41.4 16.4 13.5 9.3 ich3: 20.3 15.3 9.0 5.7 Fall 1,642 49.2 20.6 14.5 9.8 20.4 13.7 4.7 7.8 Winter 2,932 49.6 ■18.7 8.0 7.3 .12.9 29.1 14.3 6.2 3.0 Spring 4,746 64.4 11.4 13.6 12.9 15.6 17.3 14.2 9:1 5.9 1960-61 5,577 59.1 19.3 12.9 9.7 14.4 18.7 13.3 6.4 5.1 1961-62 3,431 50.8 14.8 10.8 8.9 9.8 27.2 16.2 6.9 5.6 1962-63 1,797 45.7 9,3 11.7 12.2 11.4 26.3 14.0 10.1 4.8 - 1963-64 776 50,5 19.8 15.8 11.5 5.1- 13.9 ]4.7 12.0 7.2 1960-64 11,581 54.0 16.1 12.2 10.0 11.7 22.4 14.4 7.6 5.4 l/ Percentages of total observed on slopes steeper than 10 degrees. When all reservoirs held water, most parts of the study area were within about one mile of a water source. The extent of areas greater distances from water increased through the summer as less “permanent81 reservoirs dried. The relative percentages of observations at various distances from water (Table XfX)■indicated that use of range by mule deer decreased sharply at distances of one mile or more. Most observations, especially during summer and fall when "permanentM water sources would be most important, were within three-fourths mile of a water source. The number of observations at greater distances decreased as distance increased with negligible use at two miles or more. These relationships reflected the distribution of water sources in relation to areas used by mule deer - 94 - TABLE H X . PERCENTAGES OF TOTAL MULE DEER OBSERVED AT ONE-FOURTH H L E DISTANCE INTERVALS FROM K N O W WATER SOURCES/ B I SEASON AND YEARS, 1960-64o Season Tear Sample Size Distance Class 3/ (I) (2) (3) (4) (5) (6) (7) (8) (9) Summer 1-960 731 37.3 38.2 17.6 4.8 1.1. . T 2/ T • 1961 669 22.1 22.9 20.2 10.9 . 8.5 9.0 4.3 2.1 1962 483 8.7 21.3 28.8 18.0 8.1. 9.5 5.4 T 1963 373 17.0 19.1 24.5 17.8 8 2 ^ 9.0 4.3 Total 1960-63 2,256 23.3 86.9 21.9 11.6 . 6.0 6.4 3.2 T Fall I960 734 33.Q 29.6 j.6.5 9.8 7.4 3.0 T I 1961 587 31.3 33.6 14.5 id.2 4.1 2.7 3.6 I 1962 : 163 7.4 18.4 36.8 16.0 8.6 5.5 3.1 4.3 1963 ^ 163 4.3 20.2 33.1 18.4 5.5 10.4 2.5 4.3 1.2 Total 1960-63 1,647 27.0 29.0 19.4 11.4 6.1 3.9 2.2 T T Winter 1961 1,441 11.0 22.1 30.1 20.6 8.5 3.5 2.0 2.2 1962 - 845 37.5 21.3 17.5 16.0 3.2 1.9 2.1 T 1963 526 9.3 28.5 23.4 15.8 10.5 7.6 T 1964 : 120 22.5 ; • ■. t- 23.3 35.0 T 8.3 9.2 T Total 1961-64 2,932 18,8 22.1 25.0 19.0 7.0 4.0 2.8 1.3 Spring 1961 2,671 15.4 25.9 31.3 15.8 5.8 3.6 1.8 T 1962 1,330 14.7 21.7 29.6 18.9 7.8 4.4 2.0 T 1963 625 10.4 23.6 32.3 21.2 6.5 4.3 1.6, 1964 Tdtal 1961-64 120 2%.5 15.8 30.0 10.0 8.3 13.3 4,746 14.7 . '/ # . 2 30.9 17.3 6.6 4.2 1.7 T 1960-61 5,577 19.% 2fiG 27.3 14.8 6.1 3.1 1.5 T-'1 1961-62 3,431 24.6 23.8 22.2 15.2 6.2 4.4 2.7 T 1962-63 1,797 9.3 24.0 29.1 18.3 8.3 6.8 3.6 T 1963-64 776 16.0 15.9 27.0 V 19.4 6.5 9.9 4.0 1.0 T 1960-64 :LI,581 19.2 24.9. -V ■ 26.0 15.8 6.5 4.5 2.4 T T l/ Distance Class; (I) =..0-1/4:mile, (2) = l/4-1/2 mile ^ (3) =1/2-3/4 mile, (4) = 3/4-1 mile-,' (5) .?= Lrl 1/4 DtLle5l (6) = I 1/4-1 l/2 mile, (7) = 1 1/2-1 3/4 miles (8) = I 3/4-2 mile, (9) = 2+ mile, t I v -i ^ ^ ^ 2/, T = less than one per bent. ' Y - 95 - during different seasons and years. In 196ls when water sources were at a minimum, the percentage of mule deer observations within ■ one-half mile of water decreased and the number at greater distances increased as com­ pared to I960, • Most reservoirs were along the periphery of larger ridge- tops, Relative intensities of use of the • - . • ' • ' .. X : seasons and years largely determined the relationship between locations of mule deer observations and water supplies, The distribution of water on : the study area even during the most arid years was hot a significant factor in determining mule deer distribution. Elk Elk activity habits, use of vegetation types, slopes, and exposures as well as relations to water were evaluated using methods described for mule deer. The small number, irregular distribution, and gregarious behavior of elk was - reflected in observations for only 280 observation trips (54 per cent of the total) at the low frequency of 1.8 groups (12,4 individuals) per trip. A total of 3,489 elk observations, used in evaluating range-use habits, was recorded as follows: I96O-6I, 1,307, 1961-62, 1,111; 1962-63, 815; and 1963-64, 256. Activity Habits Like mule deer, elk were active mostly within three-to-four-hour periods in early morning and late afternoon. Slightly greater numbers (58 per cent of the total) were observed during mornings when an average of 15 elk was recorded per observation trip, compared to 10 per trip — 96 — during evening, and “activity periods" were slightly longer. During the four-year period, greatest numbers were observed during the first and last ■; hours of daylight. Approximately 78 per cent of all observations occurred within two and one-half hours after sunrise and before sunset, and 94 per cent within three and one-half hours. Variation in the distribution of observations during normal activity periods for feeding elk is shown in Figures 30 and 31« The slopes of curves are exaggerated in some instances because of the concentrated nature of observations due to the gregarious habits and the distribution of elk. The consistency of "peaks" for the two-hour category suggested a true relationship and that elk feeding was at a high level through/at that time during some seasons. Activity periods were "shortest" in fall when feeding was concen­ trated in the first and last hours of daylight, and "longest” in late winter when most elk were observed between one and one-half and two and one-half hours, after sunrise and before sunset. Feeding was the predominant activity for elk observed within three i x ’ • • and one-half hours after sunrise and before sunset (Figure 32). Higher ■ '• . ■ ' ' percentages were observed feeding in spring than during other seasons. The indicated decrease in proportions feeding and increased percentages alert and traveling during 1962-63 and'1963-64, as compared to earlier years, were related to changes in range use and further, in 1963-64, to undue influence of. concentration of observations in summer. Pe rc en ta ge • Pe rc en ta ge - 97 - summer fall winter spring 4 + 3 2 1 0 Hours before sunsetHours after sunrise Percentages of Total Feeding Elk Observed by Hour Relative to Sunrise and Sunset During Seasons. 1960-61 1961- 62 1962- 63 1963- 64 4 + 3 2 I 0 Hours before sunsetHours after sunrise Figure 31 Percentages of Total Feeding Elk Observed by Hour Relative to Sunrise and Sunseft During Seasons. Alert 80 •= Bedding Feeding Traveling § UO ■ 1960-61 1961-62 1962-63 1963-64 SeasonsHours after sunrise and before sunset Years Figure 32. Percentages of Total Elk Observed by Activity Class, During Normal Activity Periods, Seasons, and Years. - 99 - Elk food and range-use habitss range-forage conditions^ and weather^ doubtless influenced elk activitiess but these relationships were largely obscured by interactions0 A complete summary of elk activities as recorded by seasons and years is presented in Appendix Table LI. ' Elk activity- relations to various environmental conditions are summarized in Appendix Table L U . Group Size Observed groups ranged in size from I to 66 animals„ Seasonal and yearly averages (Table XX) varied in relation to social relationships and distributions of elk use on the area. The high average during the winter of 1961-62 was associated with extreme concentration on the southeastern portion of the study area. That of the fall of 1962 was related to aggre­ gation on winter pastures on the south-central portion during November. Single individuals and small groups were most common in late May and early June. "Average" groups were common only in mid-summer and spring prior to the calving-period. The largest and most variable group numbers occurred during fall and winter. Groups including more than 30 elk were rare aggregations associated with feeding and/or flight. The composi­ tion of groups associated with recognizable elk changed frequently during all.seasons. - 100 - TABLE XX. AVERAGE SIZE O F GROUPS YEARS, 1960-64. FOR EL K OBSERVED FCNS:0« v I L v R D v AND Tear . Summer Fall Winter Spring Year 1960-61 4.0 (194)^ 7»5 (378) 7 . 9 (3 9 7 ) 6 . 1 (3 3 8 ) 6.4 1961-62 3.7 (145) . 6 . 7 ( 2 6 3 ) 14.6 (484) . ' 7 » 5 ( 2 1 9 ) . 7 . 9 1962-63 4 . 2 (1 8 9 ) 12.6 ( 2 78) 10.8 (1 7 3 ) 4 . 7 (1 7 5 ) 6.8 1963-64 5 . 9 ( 1 3 6 ) 6 . 4 (3 2 ) 8.2 ( 7 4 ): 4.6 (14) 6.4 1960-64 4<;3. •. 8.1 10.4 6.0 6 . 9 l/ Number of elk in s a m p l e „ U s e of Vegetation Types Frequencies of elk occurrence on vegetation types receiving more than ' ... ■ ' 10 per cent of the total use during monthly periods are plotted in Figures 33 and 34» The variation in these data reflected both small monthly samples in which single observations often influenced results and actual trends in use of the various t y p e s „ A complete summary of results for seasons5 years, and the four-year period is presented in Appendix Table I I I I . f E l k population and distributional phenomena on the study area pre­ cluded the use of observability indices as a measure of undetected use of timbered vegetation types. iV i I \ A . T J A S O N D J F M A M J J A S O W D J F H A M J J A S O N D J F M A M J J A S O N D I960 1961 1962 1963 _ Finns/ Juniperus _ Pseudotsuga/ Juniperus _ Artemisia/ Agropyron Figure 33- Monthly Frequencies of Elk Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the Total Use During at Least One Month, (see Fig. 3^ also) 1 1 101 Sarcobatus/Artemisia Agropyron/Symphoricarpos 30' \ ; A v J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M A M JJ A S O N D J I960 1961 1962 1963 1964 Figure 34. Monthly Frequencies of Elk Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the Total Use During at Least One Month. (see Fig. 33 also) 102 ™ 103 “ Summer (June, July, A u g u s t ) - U s e and importance of vegetation types varied widely during summers, Use of the Sarcobatus/lrtemisia type for all activities varied from minor to heavy during the four summers = As a feeding area, the type ranked first in over-all summer usage for the four years and in 1962 and 1963 but ranked fifth in 'I960 and 1961» It was used in association with the Agropyron/Symphoricarpos type, which received moderate to heavy use in all years, and the. XantMum,type, which occasion­ ally received heavy use when abundant growth of forbs occurred. These "coulee-bottom" types collectively comprised the major.feeding areas in i summer, particularly in July and August. The Artemisia/Aeropyron type ranked second in average summer usage for total elk activities and for feeding, but use varied significantly between years in relation to the abundance of preferred food plants on other types. It was the most important type for feeding in i960 and 1961 but its impor- 1 * tance in 1962 and 1963 was minor or moderate. The Artemisia tridentata/ Agropyron smithii associes and the A. tridentata/A. spieatum association were the most important communities, reflecting distributions on smaller ridges, benches,--and-a-long—the -edge-of timbered types and development of the Tragopogon dubius union. Greatest summer usage .occurred on timbered types. The Pjnus/ Juniperus and Pseudotsuga/Juniperus types were moderately important for feeding and more important than other types for bedding. The former typically was used most intensively during June. The Pinus 'ponderosa/ Agropyron spieatum association appeared to be the most- important community. “ 104 = The Pseadot saga/J m i p e m s type "was used in. June for. ealvingo late July and ■ e^rly August’ use.seemed related to increased utilization of browse- and ■ persistence of forbs 0 • ' ' . Other vegetation types were of little importance during the summer= • • • - . . . " Fall (September, October, November) o— The Artemisia/Agropyron type was the most heavily used vegetation type during this period., when grasses became increasingly important .in the diet ,of elk. Use generally increased as the season progressed, but varied in. relation to!the availability of various grass species and the abundance o f .fofbs on other types. . The Artemisia tridentata/Agropyron smithii associes was■the. most / important community' on the Artemisia/Agropyron type during i960, whereas ' the A. tridentata/A. smithii/Bbuteloua gracilis association appeared to be used more intensively during years of “green-up.8I' The Agropyron/Poa type was heavily used- in October and November of 1961 and 1962, following “green-ups81' of Poa secmnda and Koleria cristata on.winter pastures in the south-central'.portion-'of-, the area. • ' The more'intensive use- in'1962 reflected increased elk numbers on the area and possibly, greater availability of Poa-secunda' and Koleria'cristate ,after the' pastures -were cut for hay. Over-all use of the Pinus/jmiperus- type was moderate. For feeding it was very.:minor .in 1961 but- prominent in 1962 and 1963 when fdrbs, . especially Melilotus officinalis., were abundant. The Pinus pbnderosa/ . Agropyron spieatum association was' most important, but the P-. ponderosa/ Juniberus scopulorum association received intensive use in 1962.- =- 10 5 “ The Po ponderosa/Artemisia longifolia association received only minor use, largely in September. Other types generally received only minor use during fall. Moderate feeding on the"Agropyron/Symphoriearpos type in 1961 reflected utilization of Chrysothamnus viscidiflorus and grass, primarily during September, Data for the fall of 1963 were obtained only during September, Winter (December, January, February),— The Artemisia/Agropyron type was most important three of the four winters and for all winters combined, The Artemisia tridentata/Agropyron smithii associes was generally used most intensively. The A, tridentata/A, spicatum association ranked second. It was especially important during 1961 when elk use was concentrated on areas with better range conditions. Use on the A, tridentata/A, smithii/ Bouteloua gracilis association varied in relation to availability of grasses. Use was influenced by prior grazing and snow cover, The Pinus/juniperus type ranked second,. The intensive use in the- winter of 1961-62 reflected attraction to the Finns ponderosa/ Agropyron spicatum association on steeper, semi-open slopes where grasses persisted in' relative abundance. Other communities received only occasional use", Sarcobatus/Artemisia was the only other type of importance, receiving moderate use during all winters except 1963. Use was largely on steeper slopes, in contrast to predominant use of this type on coulee-bottom sites during other seasons, where grasses were relatively abundant , - 106 - Spring (March, April, M a y ) „--The received the greatest use throughout the period. The most intensive use, o c curring' during late March and early April, coincided approximately w i t h the onset of the spring "green-up.11 The Artemisia tridentata/Agropyron smithii/ Bouteloua gracilis association was the most important community, reflecting ' ~ " _ ' V.*. . ''v-: seasonal elk preferences for grasses and early developing forbs of the Poa secunda union. The A. tridentata/A. s m i t h i i .as'soeies received increas­ ing use throughout spring. Relative use on the Artemisia/Agropyron type declined sharply during the last week of M a y as most adult females dis­ persed prior to calving,. Over-all use of all other vegetation types was minor or negligible, though use of some types was significant in some m o n t h s ; e.g., the Sarcobatus/Artemisia type in late March and/or early April and the P s eudotsuga/Juniperus type in late May b y adult females. Yearlong and Year-to-Year Trends.— Yearlong elk relations to vegeta­ tion types were characterized by significant tr e n d s . The Sarcobatus/ Art e m i s i a , Agropyron/Symphoricarpos» and Xanthium types, which individ­ ually received variable use b y elk, collectively comprised the most important area for feeding during summer. U s e of these ‘'coulee-bottom8* types was minor after early fall. Relative use of the Artemisia/Agropyron types, which received the most intensive use over-all, generally increased from very minor in mid-summer, through fall and winter-. It was by far the most important habitat type for feeding throughout winter and spring, with - 107 - the most intensive use occurring in early springe The Pinna/Juniperus type was of moderate importance through summer and fall and of major importance during winter when lightly or ungrazed stands of grass on semi- open, steeper slopes were important forage sources. It received only minor use during spring. Stands of Pseudotsuga/Juniperus received impor­ tant use by female and calf elk during late May and early June, and were relatively important for feeding in late summer. This type was especially important for bedding during summer and early fall. The Agropyros/Poa type was important only during October and November in years of fall "green-up.*' Use of the Artemisia longifolia type -was minor at all times. The important differences in use of vegetation types between years were largely related to pfSci^itation and its influence on forage supplies and feeding habits of elk. The prominent use on the Artemisia/ Agropyron type from I960 through tlie spring of 1962 was associated with drought conditions. The decreased use and importance of this type and increased or markedly different use of other vegetation types after the spring of 1962 reflected increased growth of forbs, especially Melilotus officinalis. Weather conditions generally did not appear to influence relative use of the various. vegetation types, except during winter when complete snow cover seemed to effect significantly greater use on the Firms/Juniperus type and/or decreased use of the Artemisia/Agropyron type. A complete summary of use I of vegetation types in relation to weather conditions is presented in Appendix Table..LIV . “ 108 — Use of Slopes The distribution of elk observations on slopes of varying steepness is presented in Table XXI. Approximately 42 per cent of all elk occurred on slopes steeper than 10 degrees» Less than one per cent were observed on slopes of 45 degrees or more. , ' ■ TABLE XXI. PERCENTAGES OF TOTAL ELK OBSERVED ON SLOPE CLASSES, BY SEASON AND YEARS, 1960-64. Degrees of Slope ■ Size 0-10 11-25 . 26-35 35+ Summer 664 49.5 18.4 17.6 . 14.5 Fall 951 70.7 11.5 . 8.7 10.0 TVinter 1,128 45.0 25.0 20.9 9.0 Spring ' 746 67.8 15.5 16.0 8.6 1960-61 1,307 57.8 15.8 . 17.0 9.5 1961-62 1,111 56.2 21.8 12.2 9.9 1962-63 815 62.2 18.4 . 14.5 4.9 1963-64 256 50.8 12.1 28.1 9.0 1960-64 3,489 57.8 18.0 15.7 8.6 Use of slopes varied during seasons and years in relation to relative frequencies of use on the various vegetation types. Seasonally greater use of steeper slopes largely reflected use on the Pinus/Juniperus and Pseudotsuga/Juniperus types in summer and the Pinus/Juniperus and t; - Sarcobatus/Arteanisia types in winter. The high percentages observed on level to gently sloping sites in 'fall and spring correspond to intensive - 109 - Use of Exposures The distribution of elk observed on slopes of various exposures dur­ ing seasons and years is presented in Table XXII„ Although the results partly reflect relative amounts of area with each exposures northerly exposures appeared to receive especially intensive- use in summer when most elk ranged in proximity to the Pseudotsuga/Juniperus type. Greater use of west and northerly exposures was evident during winter when upper portions of slopes dominated by the Pinus ponderosa/Agropyron spicatum were impor­ tant feeding areas. Cold winds and/or snow depths which prevailed in winter did not appear to influence use of northerly exposures by elk. Exceptionally intense use appeared on south, southeast, and east-facing slopes in spring. TABLE XXII. PERCENTAGES OF TOTAL ELK OBSERVED ON VARIOUS EXPOSURES, BI SEASON AND YEARS, I96O-64. Season Sample Size, % Op-IO0 Slope Exposure l/" ’ SW~ N NE E SE S W NW Summer 664 49,5 26.0 19.1- 3.3 4.4 14'. 6 12.4 5.0 11.3 Fall 951 70,7 16,9 20.9 0.7 3.2 24.5 21.2 4.7 6,7 Winter 1,128 45,0 22,1 21,1 7.9 4.0 7.9 13.1 12.9 10:7 Spring 746 67.8 19.2 12.4 14.5 26.9 16.2 4.3 4.7 1.7 1960-61 1,307 57.8 13.8 19.7' 5.6 10.2 24.6 14.2 4.8 7.1 1961-62 1,111 56.2 32.2 16.1 7.2 8.9 2.5 7.4 .15.1 10.5 1962-63 815 62.2 19.6 25.2 3.7 4.7 11.0 17.6 ■ 5.3 11.3 1963-64 256 50.8, 19.5 14.8 12.8 0.0 19.5 18.1 2.7 2.0 1960-64 3,489 57.8 ' 21.6 19.2 6.5 7.6 14.0' 13.1 8.1 8.6 l/ Percentages of total observed on slopes steeper than 10 degrees. - H O - Relations to Water Relative percentages of elk observed at various distances from water are presented in Table XXIII. Most observations during-summer and fall were within three-fourths mile of a water sources though greatest numbers typically occurred between one-fourth and thre'e^fdurths mile. Use at distances of one mile or more was only minor at'all times and consider- 5, ably lower than was recorded for mule deer. ' Year-to-year relationships reflected the distribution of water in relation to areas used by elk in the same manner as described for mule deer. The percentages of elk observations within one-half mile of water decreased and the number at greater distances increased in the summer of 1961, as compared to 1960# when water sources were at a minimum. Use at greater distances was also higher in 1962 and 1963 when.elk used "timbered" and "coulee-bottom" vegetation types more intensively than in prior years when intensive use occurred on the Artemisia/Agropyron type in proximity of reservoirs. Use on areas more distant from "permanent" water sources was greatest during winter and early spring when elk were distributed on the south­ eastern portion of the study area where few reservoirs existed. Yearly differences were related to the degree and/or location of elk concentra­ tion. - Ill - TABLE XXIII. PERCENTAGES OF TOTAL ELK OBSERVED AT ONE-FOURTH MILE DISTANCE INTERVALS. FROM KNOW WATER SOURCES, BI SEASON AND YEARS, 1960—64o Season Year Sample Size Distance Class l/ (9)(I) (2) (3) (4) (5) (6) (7) (8) Summer i960 194 25.3 51.5 21.1 2.1 1961 145 20.0 46.2 15.2 6.9 3.4 3.4 4.8 1962 189 7.4 34.9 20.6 21.7 4.8 8.5 2.1 1963 . 136 13.2 50.7 32.4 2.9 T 2/ Total 1960-63 664 16.6 45.5 22.0 8.9 2.1 3.3 1.7 Fall I960 37& 15.6 15,1 38.4 16.9 9.8 4=2 1961 263. 35.0 36.1 12.9 9.9 6.1 1962 278 15.5 40.3 38.1 2.9 3.2 1963 32 3,1 53.1 18.6 6.3 18.8 Total 1960-63 : 951 20.4 27.9 31.8 10.9 6.5 1.7 T T Winter 1961 397 . '1.5 ■ 9.1 41.6 25,4 4.3 12.6 5.5 1962 484 19.0 19.2 38.6 10.7 2.3 4.3 A-o 3- 1.7 1963 173 8.7 40.5 23.1 27.7 1964 74 58.1 14.9 24.3 2.7 Total 1961-64 1,128 13.3 15.0 24.3 25.1 9.9 3.5 6.2 2.7 Spring 1961 338 1.5 4.7 11.2 32.0 39.6 6.5 A® A 1962 219 18.3 22.4 39.7 11.0 8.7 V- 1963 175 21:7 29.7 28.6 10.9 6.3 2.9 1964 14 50.0 35.7 14.3 Total 1961-64 746 11.1 15.7 24.4 20.2 19.4 6.2 2.7 T 1960-61 1,307 8.6 13.7 19.9 26.1 20.8 4.2 5.0 1.7 1961-62 1,111 22.8 27.4 29.7 10.1 2.9 4.1 2.4 T 1962-63 815 13.5 36.8 28.8 14.2 3.6 2.6 T 1963-64 256 23.8 27.3 30.9 10.9 1.2 2.7 T 2.3 ■ 1960-64 3,489 $5.4 24.4 25.9 17.1 9.5 3.6 3.0 T 1 T I l/ Distance Class: (I) --0-1/4 mile, (2) - 1/4-1/2 mile, (3) = 1/2-3/4 mile, (4) ='3/4-1 mile, (5) = 1-1 1/4 mile, (6) - I 1/4-1 l/2-mile, (7) = I 1/2-1 3/4 mile, (8) = l;:3/4-2 mile, (9) = 2+ mile. ' . 2/ T = less than one per cent. - 112 ~ Cattle R a n g e - u s e ,characteristics of cattle were evaluated in the same manner as those of mule deer and elk. The total of 233 observation trips typi­ cally included one morning and one afternoon each week during the grazing season, except during July and August, I960 and after September '1963= A total of 25,125 cattle observations, used in evaluating range-use habits, was recorded as follows: 1960-61, 6,5625 1961=62, 7,4665 1962=63, 6,2915 and 1963-64, 4j788o Activity Habits Cattle activity habits resembled those of mule deer and elk in that feeding was most intensive during three-to-four-hour periods in early morning and late afternoon, but most cattle also fed sporadically between these two periods= For the four-year period, greatest numbers were observed during the first and last hours of daylight= Approximately 67 per cent of all cattle and 76 per cent of feeding cattle observed during observation periods were recorded within, two and one-half hours after sun­ rise and before sunset, wi t h 84 per cent and 89 per cent respectively observed within three and one-half hours = These percentages were "high" for total daily activities because observations usually were not contin­ uous through mid-day during late spring, summer, and early fall.= Percentages of the total that were feeding among cattle observed during morning, evening, seasonal, and yearly periods are shown in - 113 - Figures 35# 36, and 37» Periods of intensive activity were shortest during summer and longest in winter» . . The duration of intensive activity as well as the extent of feeding later in the day varied in relation to range-forage conditions= Intensive activity was most prolonged during drought yearsa 1960-61 and 1961-62# when cattle were relatively active throughout the day as compared with 1962-63 and 1963-64 whdn decreased proportions observed feeding more than two and one-half hours after sunrise and before sunset reflected improved forage conditions. Feeding predominated the recorded activity for cattle within three and one-half hours after sunrise and before sunset= ' Feeding was more intensive in early morning than late evening. Proportions observed bedded increased sharply between two and one-half hours after sunrise and one and one-half hours before sunset. Traveling was a minor activity at all times. Propor­ tions traveling were highest between two and one-half and three and one- half hours after sunrise and between one-half hour before and after sunset when greatest numbers of cattle moved toward reservoirs and evening feed­ ing areas, respectively. Greater numbers were observed traveling during 1961-62 when extensive movement occurred between feeding and bedding (reservoir) areas. Complete summaries of cattle activities as recorded by seasons and years and in relation to recorded environmental conditions are presented in Appendix Tables LV and LVI, 100 90 80 70 60 g 50 I 30 20 10 Feeding (A.M.) Feeding (P.M.) N \) Bedding (A.M.) Bedding (P.M.) Si spring / / Traveling Hours, after sunrise/before sunsetHours after sunrise/before sunset 100 90 80 70 S I 30 20 10 Figure 35. Percentages of Cattle Feeding, Bedding, and Traveling by Hour Rela­ tive to Sunrise/Sunset. Figure 36. Percentages of Cattle Feeding by Hour Relative to Sunrise/Sunset During Seasons. X \ ------------ = 1960-61 1--------- -- 1961-62 — — - — - = 1962—63 .......... = 1963-64 0 1 2 3 4+ Hours after sunrise/before sunset Figure 37. Percentages of Cattle Feeding by Hour Rela­ tive to Sunrise/Sunset During Years. - 115 - Group Size Average numbers of cattle observed per group (Table XXIV) varied in relation to the degree of dispersal and/or use of certain vegetation types as well as the type and duration of activity. Group sizes, generally were smallest during drought years, especially 1961-62,.'when- cattle were rela­ tively dispersed, seeking forage on normally less-used areas and timbered vegetation types, and fed more intensively during much of the day. Larger groups prevailed during the last two years when cattle were relatively concentrated during most periods with abundant forage in proximity to water sources, heavier seasonal use on ridgetops, and less intensive feeding. TABLE XXIV. AVERAGE SIZE OF GROUPS FOR CATTLE OBSERVED DURING SEASONS AND YEARS, 1960-64. Year Summer Fall Winter Spring Year 1960-61 7.5 (548)^ 5.7 (2,879) '■ 8.3 (411) 7.4 (2,724) 6.6 1961-62 4.9 (3,519) 4.5 (2,340) 6.1 (154) 7.4 (1,435) 5.1 1962-63 7.1 (3,379) 5.2 (1,298) 4.7 (71) 11.2 (1,543) 7.2 1963-64 13.6 (3,254) 8.5 (1,388) 11.7 (82) 64.O (64) 11.6 1960-64 , 7.1.. - 5.5: . 7.4 . .8.2 6.7 I/ Number of cattle in sample. Use of Vegetation Types Relative intensities of cattle use on vegetation types which received more than 10 per cent of the total use during months are illustrated in Figure 3 8 « A complete summary of cattle use on vegetation types for feed­ ing and all activities during seasons and years is in Appendix Table LVII. Artemisia/ Sarcobatus/ Juniperus «^1 l\ V \ . A/ I ' \//\\ • \ A A\ ; I f-J V A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J I960 1961 1962 1963 1964 Figure 38. Monthly Frequencies of Cattle Occurrence on Vegetation Types Receiving More Than Ten Per Cent of the Total Use During at Least One Month. Summer (June, July5 August) „ — The Artemisia/Agropyron type was most — 117 “ important in summer during all years, Relative intensities of use gener= ■ ally decreased through the period, being lowest in August„ Use was pre­ dominantly on the Artemisia tridentata/Agropyron smithii/Bouteloua gracilis association. Use of the A. tridentata/A. smithii assoeies'varied in rela­ tion to the distribution of water on the area, it was used more intensively during 1961 when cattle ranged further to obtain forage. The A. trldentata/ A. spicatum association received only very light use. The Pinus/Juniperus type ranked second, receiving moderate use for feeding and some additional use for bedding and loafing. It typically was used most intensively during mid and/or late summer. The PinUs ponderosa/Agropyron spicatum association on. slopes of less than about 25 degrees was the most important community. The P= ponderosa/ Juniperus seopulorum association received some use for both feeding and bedding. The P. ponderosa/Artemisia longifolia type was only rarely used by cattle. The Sarcobatus/Artemisia type ranked third for feeding during three of the four summers and for all years combined. Over all, it was of moderate importance, but use reached significant proportions during August of some years. Use on the Agropyron/Symphoricarpos and Agropyron/Poa types was minor during all years. The former received its greatest use during August when green forage persisted. Other vegetation types received very minor use at all times. - 118 - Fall (Septembers October, November) .— The Artemisia/Agropyrom type was by far the most important vegetation type during fall, for feeding as well as all activities. The Artemisia tridentata/Agropyron smithii/ Bouteloua gracilis association ,remained the most important community, but use on the A. tridentata/A. smithii associes increased as precipitation provided additional water sources and/or cattle movement became more extensive with cooler temperatures„ The A= tridentata/A = spicatum assoc­ iation received only very minor use= The Pinus/juniperus type was moderately important, ranking second in three of the four years = Use of individual communities was similar to summer, though slightly increased use on the Finns ponderosa/ Juniperus scopulorum association occurred late in the period = Over-all use on the Sareobatus/Artemisia type was about equal to that on the Pinus/juniperus type, largely because of relatively intense use during I960= The type ranked third and was of minor importance during- other years. The Agropyron/Symphoricarpos and Agropyron/Poa.types received minor and very minor use respectively in all years = Use of other types was negligible = Winter (December, January, February)=— Cattle use of upland vegetation types during this period was largely -confined to winter pastures on the north and south-central portions of the area= Winter use on the former area was predominantly on the Artemisia/Agropyron type= Use varied through the winter on the south-central area ^ but both the Agropyron type and the Agropyron/Poa type were important« The Pinus/Juniperus type was of minor importancey but use occasionally reached significant, proportions oh the north-central pasture in early winter . This use was almost entirely on the- Pinus poaderosa/ Agropyron spicatum association. The Sarcobatus/Artemisia, type also received minor Use5 primarily in December and/or January, Use on,other vegetation types was negligible during winter. Spring (March, April, May) ,— Cattle use during March was entirely on winter pastures oh the south-central portion of the area. Less than one per cent of the total recorded use in spring occurred during the month. The Artemisia/Agropyron type received, the most intensive use during April and May, The Artemisia tridentata/Agropyron smithii/ Bouteloua gracilis association was the most important community, especially during April, The A, tridentata/A,' smithii association received important . use during some years, particularly during May, The Sarcobatus/Artemisia type was of only-minor importance for the entire period,.but received significant use during .April in 1962 and 1963, Over- all, the Pinus/JuhiPerus type was also of relatively minor importance, though moderate intensities of - use were observed during April and May of 1962, The Agropyron/Svmphoricarpos and Agropy&fea/Poa types received only vay minor, use during spring. Other vegetation types were used only occasionally. - 1 2 0 - Year^to=Year Treads.=Gattle relations to vegetation types were significant in that a single type ^ ArtemisiaZAgropyron0 was used most intensively throughout the springy Sramners falls graairig season and. relatively few types received important use. 'Seasonallys cattle use on the. Artemisia/Agropyron type varied, only slightly. Monthly usage was more varied evidently in relation,to the distribution of water'and the relative availability of forage on other types. .Use on this type was lowest.in Augusts then generally increased through November. The SarcobatUsZ Artemisia and Agropyron/Symphoricarpos types which received moderate and minor use respectivelys collectively comprised the Seconds most important • 1 " ‘ ’ W f ir > .A-' 'I'- area for feeding. Use. on these types was most intensive during early spring and late summer, , The Pinus/Juniperus type was of moderate import­ ance over-all. Relative intensities of use generally increased from spring through early falls then decreased to the end of the gracing, season. The • Agropyron/Poa type received very minor use throughout the period. Only occasional or transient use occurred oh other vegetation types.. "• Relative intensities of cattle use on the various vegetation.types on winter pastures were similar t©'that-during.the gracing season, except on the south-central portion of the area where the'Agropyron/Poa type' , received important use during, some winters.. Differences in use of Artemisia/Agrbpyron between years' were minor. Some differencess largely related to precipitation and its influence on plant growth and water supplies occurred on. certain other types.. The most important of these occurred in the spring of X962s when relatively high - 121 - intensities of use occurred on the Pinus/Juniperus and Sarcobatus/ Artemisia types during April, when cattle numbers were low and forage conditions somewhat better on these types, and during early fall of I960 when the Sarcobatus/Artemisia type received greater use than in other years. Weather conditions generally did not appear to influence relative use on the various vegetation types by cattle. A summary of use of vegetation types in relation to weather conditions is presented in Appendix Table L H I I . Use of Slopes Percentages of total cattle observed during seasons and years on slopes of varying steepness are presented in Table XXV. Use was predomi-, nantly on level to gently sloping sites and decreased sharply on steeper slopes during all periods. Slopes steeper than 25 degrees received only very minor use by cattle. This plus the increased use on steeper slopes in winter, when cattle were confined to pastures, and during 1961-62, when forage supplies on normally grazed sites were minimal, suggested' that steepness of slope exerted important influence on the distribution of cattle use on the area. Use of Exposures The distribution of observed cattle use on slopes by direction of exposure (Tabid X X H ) appeared largely related to relative amounts of area with each exposure. Over-all use was only slightly greater on northerly than on southerly exposures. Increased use on upper portions of - 122 - north-facing slopes, where grasses typically, were abundant, occurred in fall and during 1961-62. TABLE XXV. PERCENTAGES OF TOTAL CATTLE OBSERVED ON SLOPE GLASSES, BY S O K S b Y K Y J F O K I S y 1 9 6 0-6 4 . Season Sample Size Degree of Slope 0-10 11-25 . 26r35 35+ Summer ■.1 0 ,7 0 0. 8 2 . 2 13.2 4.2 AU. Fall 7,923 82.0 12.7, ' . 4.3 1.0 Winter 718 80.4 ' 9.4 1 0 . 1 T Spring ' 5,766 8i;4 14. 6 ■ 3.6 0.3 1 9 6 0 - 6 1 6 , 5 6 2 83.1 11.7 4.6 T 1961-62 7,466 1 7 . 8 5.8 1.0 1962-63 6,291 83.3 12.9 3.5 ' T 1963-64 . 4,788 88.5 8.9 2.3 T 1960-64 25,125 81.9 13.3 4.2 T l/ T = lessi than one per cent. TABLE XXVI. PERCENTAGES OF TOTAL CATTLE OBSERVED ON VARIOUS EXPOSURES, BY SEASON AND YEARS, 1960-64. Season % SctfUpl© . r\ 0 i r\c1 Exposure l/ _. U —XU Slze Slope N NE E SE S SW W NW Summer 1 0 , 7 0 0 82. 2 19.9 17.0 5.9 6 . 8 22.3 14.4 7.1 5.7 Fall 7,923 8 2 . 0 29.7 ' 11.4 5.9 5.0 19.9 1 4 . 0 4.5 9.5 Winter 718' 80.4 13.1 19.0' 8.0 12.4 2 1 . 2 1 6 . 1 0.0 10 . 2 Spring 5,766 81.4 -"13.8-23.9 9.1 10.3 15.0 15.0 '2.7 '10.3 1 9 6 0 - 6 1 6 , 5 6 2 83.1'""'117^ 2 o-19.2. 7 . 6 6.6 18.0 15.7 3.5 12.1 1961-62 7,466 75.5 2 6 . 0 13.8 5.5 10.3 19.8 11.3 5.8 6.5 1962-63 6,291 83.3 19.1 19,9 8 . 4 4.9 16.1 19.1 5.1 7.4 1963-64 4,788 88.5 19.2 16.3 5.7 1.8 30.3 14.0 5.5 6.5 1960-64 25,125 81.9 21.4 16.8 .6.7 7.2 19.8 14.4 5.0 8.1 I/ Percentages of total observed on slopes steeper,than 10 degrees. Relations to Water Relative percentages of cattle observed during seasons and years at various distances from known water sources are presented in Table X X H I . Most observations occurred within three-fourths mile of a water source. Greatest numbers typically were observed within one-fourth mile of water, and only minor use was recorded at distances greater than one mile. During summer, when permanent water sources were most important, use was predom­ inantly on sites less than one-half mile from water. More than two-thirds of all observations were within this distance during 1961 when the number of water sources was minimal. This was in contrast to observations for mule deer and elk suggesting that the distribution of cattle use on the area was more closely related to the distribution of water. Cattle use at greater distances from water increased during fall as water supplies and/ or cattle movement increased. The concentration of cattle use near water sources in spring reflected increased availability of water as well as the predominant use on more intensively grazed sites early in the period. In 1962 when most cattle were turned onto the area during May, distributions with respect to water were similar to summer. — 124 — TABLE ZXVIIo PERCENTAGES O F TOTAL CATTLE OBSERVED A T ONE-FOURTH MILE DISTANCE INTERVALS FROM KNOWN WA T E R SOURCES, B I SEASON AND YEARS, 1960-64. Season Year Size C D (2) (3) (4) (5) (6) (7) (8) (9) Summer 19602/ 548 26.6 17.2 18.4 25.7 9.1 . 1.6 1.3 1961 3,519 46.2 24^3 13.2 8.1 3:6 1.4 2.3 T 3/ 1962 3,379 31.8 22.1 22.8 15.1 4.9. .2.3 T T 1963 3,254 37.8 25.8 26.6 8.8 1.0 Total 1960-63 10,700 38.1 23.7 20.6 11.4 3.5 1.3 1.0 T T Fall I960 2,879 29.8 9.0 19.6 12.7 '15.4 6.5 6.3 T T 1961 ' 2,340 35=4 26.9 15.1 10.0 4.4 4°1 3.6 T 1962 1,298 19.3 14.6 33.4 14.3 10.0 2.0 2.9 3.5 1963 1,388 39.8 7.5 30.5 8.1 9.1 3.2 T 1.2 Total 1960-63 7,923 31.5 15.0 22.4 11.4 10.1 4.5 3.8 1,1 T Winter 1961 411 59.9 20.7 13.1 3.6 1.7 1.0 1962 154 56.5 38.3 3.9 1.3 1963 71 42.3 42.3 , 2.6 9.0 3.8 1964 82 86.6 1.2 8.5 2.4 1.2 Total I96I-64 718 6O.3 16,3 7.9 12.1 2.2 T T T Spring 1961 2,724 38,8 37,8 11.1 6.4 3.3 T 1.7 1962 1,435 34.6 17.4 24,7 19,8 2.0 1,6 1963 1,543 47.2 29,4 13.5 8.1 1.2 T 1964 64 100,0 Total 1961-64 5,766 40,7 30,1 15.0 10,1 2,4 T T T 1960-61 6,562 35,2 22.4 15.6 10,6 9.0 3.3 3.6 T T 1961-62 7,466 40.8 23,3 15,7 11.6 3,6 2.3 2.2 T 1962-63 6,291 47.2 29,4 13.5 8.1 1,2 T 1963-64 4,788 40,1 19,7 26,9=; 8,5 3.3 1.0 T T 1960-64 25,125 37:3 22,2 19.5 11.1 5.3 2.2 1,8 T T I/ Distance Class: (I) = 0-1/4 mile, (2) = 1/4-1/2 mile, (3) = !/2-3/4 mile, (4) = 3/4-1 mile, ($) = 1-1 l/4 mile, (6) = I 1/4-1 l/2 mile, (7) = I 1/2-1 3/4 mile, (8) = I. 3/4-2 mile, (9) = 2* mile, 2/ August only. 2/ T = less than one per cent. ,FOOD HABITS ■ Food habits, of mule deer, elk, and cattle were determined primarily by recording frequencies of plant use. by taxon at -feeding sites as des­ cribed by Gole (1956) and others. The^ requisite.minima during my study for recording.data were 50 instances of recent plant use on one vegetation type by one of the animals studied. Use of a rooted stem for grass and grass-like plants, an individual leaf or twig for' trees and- shrubs, and individual leaves or stems for forbs' each constituted.one'instance of use = Data generally were obtained for each of the three animals on ctBajorto •vegetation types at intervals of one month or less and on other types -as opportunity afforded = Use of a taxon was computed as a percentage of the total instances: of. use bn each site= Percentages for various sites were aggregated and averaged within vegetation types and/pi* time periods= Data, from feeding site examinations were supplemented by analyses of one quart rumen samples from each, of 37 mule deer, 12 elk, and 18 cattle =: Rumen samples for mule deer were from animals killed by hunters or collected specifically for this study. A U .cattle samples were from ani­ mals found dead= The. deaths of most of those from which rumen samples were collected' during spring and early summer of 1963 were attributed to sweetclover (Melilotus officinalis) bloat. Elk samples were represented by animals found dead as. well as intentionally .collected= Separations and measurements of items in the rumen samples were performed by personnel of the Montana Fish and Game Department Wildlife Laboratory using standard ■ techniques = I assisted in identification, of plant materials = Relative volumes of plant taxa in each,rumen sample were expressed .as percentages = 126 - Percentages were aggregated and. averaged for season's by year. Mule Deer A total of 40,982 instances of plant use was recorded at 263 mule deer feeding sites between June i960 and March 1964° The number of instances of use at each site varied from 50 to 500 but typically was between 100 and 200 (average 156)= Smaller samples were characteristic of "minor'" vegeta­ tion types and early summer when mule deer group sizes' were small. Data for use of taxa which individually received at least ten per cent of the total instances of plant use for at least one vegetation type during at least one season are presented in Table XXVIII. Those taxa which did not meet thepe requirements are considered in Appendix Table LIX. Summer (June, July, August).--A total of 8? mule deer feeding sites was examined during the four summers ^ 27, 27, 23, and 10 for I960, 1961, 1962, and 1963 respectively. Use was recorded on 62 taxa, including 44 forbs, 12 shrubs, and 6 grasses or sedges. Nine taxa, five forbs and four shrubs, received an average of more than one per cent of the total instances of use during summers. Forbs predominated the use on all but timbered vegetation types. Melilotus officinalis was by far the single most important taxon, accounting for 44.5 per cent of the total combined summer use. It occurred on 87 per cent and was used on 70 per cent of all mule deer feeding sites. Other forbs individually were of only minor importance. Those taxa which received the greatest use were Comandra umbellatum, Glycyrrhiza Iepidota9 and Tragopogon dubius. - 127 - TABLE XXVIII. MJLE DEER USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1960-64 COMBINED„ Vegetation Type; Plant Taxa Artemisia/Agropyron: Poa secunda • Grass Total Artemisia tridentata Chrysothamnus nauseosus Browse Total Comandra UmbeLIatum Lomatium foeniculatum Melilotus officinalis Forb Total Agropyron/Poa; Poa secunda Grass Total Chrysothamnus nauseosus Browse Total Fritillaria pudica Melilotus officinalis Tragopogon dubius Forb Total Pinus/juniperus; ■ Grass Total Artemisia cana Artemisia longifolia Artemisia tridentata Chrysothamnus viseidiflorus Juniperus seopulorum Rhus trilobata Symphoricarpos spp. Browse Total Melilotus officinalis Forb Total Summer Fall Winter Spring (18-14)1/ (11-15) (27-27) (27-38) 6/T 2/ 27/17 15/1 59/12 .hE 36/18 37/2 85/16 27/10 85/56 70/32 n/T 54/26 41/22 4/T 17/6 73/37 96/88 79/35 33/6 27/T 30/10 52/10 94/85 54/36 15/T 22/3 94/94 73/45 44/10 89/49 (1-1) (1-4) (1-7) 100/79 100/1 100/81 100/83 100/81 100/12 100/100 100/17 100/100 100/18 100/17 (38-42) (17-37) (20-2$) (22-45) 13/T 28/2 20/T 50/T 6/5 15/10 23/3 3/T 22/11 25/6 14/T 11/7 40/8 50/13 22/20 35/5 9/1 33/7 80/18 50/20 84/44 83/33 85/14 50/10 60/10 56/11 70/18 41/6. 92/64 100/83 100/86 100/66 58/24 22/2 20/7 9/2 79/36 67/15 55/14 68/26 128' - TABLE XXVIII. Continued. Vegetation Type: Plant Taxa Summer Fall Winter Spring Pseudotsuga/juniperus: Grass Total Artemisia tridentata Chrysothamnus viscidiflorus •' Juniperus scopulorum Prunus virginiana Pseudotsuga menziesii Rhus trilobata Rosa spp. Symphoricarpos spp. Browse Total Melilotus officinalis • Forb Total Sarcobatus/Artemisia: Poa secunda Grass Total Artemisia tridentata Chrysothamnus viscidiflorus Sarcobatus vermiculatus Browse Total Melilotus officinalis Musineon divaricatum Sphaeralcea coccinea Tragopogon dubius Forb Total Agropyron/Symphoricarpos: Grass Total • Artemisia cana Chrysothamnus viscidiflorus Symphoricarpos spp. Browse Total Glycyrrhiza lepidota Melilotus officinalis Forb Total Xanthium: Symphoricarpos spp. Browse Total Glycyrrhiza lepidota Melilotus officinalis Forb Total (10-24) (6-14) (6-17) (4-23) . 20/T . 20/T 17/T 25/T 68/15 25/T 10/T . . 20/18 68/29 25/T ' 10/T : Ao/i 100/30 50/23 80/14 40/5 68/4 17/7 50/24 • 90/44 ' • . 80/21 100/4 50/7 80/6 40/4 50/T 75/10 80/11 100/32 68/5 50/11 100/82 100/86 100/98 100/81 50/11. 40/8 . 17/1 25/10 79/36 67/15 55/14 68/26 ■ ■ (6—11) (4-17) (8-18) (9-21) 25/19 12/10 56/16 ' • 25/19 12/10 56/16 50/30 89/47 25/25 12/12 11/T .25/T ■ 75/25 44/T 50/2 . 50/25 100/80 89/53 83/71 50/26 25/5 56/11 25/19 50/17 . 50/T 12/T 22/T 100/98 . .100/56 50/8 89/28 (5-9) (607) (1-2) (2-9) 40/1 50/1 .33/4 100/26 33/30 100/74 100/48 60/36 50/46 50/41 60/37 83/81 100/100 100/95 80/3 33/16 50/1 8O/56 33/2 100/62 50/19 50/4 (6-9) (1-3) 50/24 . 50/29 68/3. . 100/19. 50/24 100/74 100/71 • 100/93 - 129 - TABLE XXVIII. Continued. Vegetation Type: Plant Taxa Summer Fall ■- Winter Spring Artemisia loneifolia: (3-10) (3-8) Artemisia loneifolia 100/89 Rosa spp. . 33/10 , 33/2 Browse Total 67/13 . 100/93 Glvcyrrhiza lepidota 68/24 Melilotus officinalis 68/50 33/T ' Forb Total 100/87 100/7 ■ l/ In parenthesis: The number of feeding sites on this vegetation-The total number of taxa used on the type. 2/ Per cent frequency of use among all feeding sites on this vegetation type/aggregate mean percentage of the total instances of use for this vegetation type. T = less than one per cent. - 130 - Browse ranked second in total use on feeding sites during summer. It was the. most important forage class used on the Pinus/juniperus and Pseudotsuga/juniperus vegetation types. Rhus trilobata was the most intensively used taxon on the latter and ranked second over-all, Symphoricarpos spp. (S. alba and S. occidentalis) which received about 10 per cent of the total summer use, Rosa spp. (R. -ark'ansana and R, nutkana), and Prunus virginiana ranked third, fourth,, and fifth respectively for all _ -v , vegetation types combined. The latter was important only on the f' , 'S ‘ ' - I , M ' , Pseudotsuga/juniperus type where it was the second, most intensively used taxon. Symphoricarpos spp, and. Rosa spp, received prominent use.on both timbered types. Symphoricarpos occidentalis, which.was used intensively on Agropyron/Symphoricarpos, was the only shrub receiving significant use by mule deer on a non-timbered vegetation type during summer. Use on grass and grass-like plants was negligible at all times during summer. Analyses of six mule deer rumen samples from the summer of 1963 (Table XXIX) showed similarly used taxa but greater use of browse than feeding site data. The predominance of browse,, including the slightly greater volume, of Rhus trilobata as compared to Melilotus officinalis, in the rumen samples may represent a truer relationship between forbs and browse in the mule deer diet than feeding site examinations. Numbers of feeding site examinations corresponded generally to relative intensities of observed mule deer use of timbered types. This, together with the observed predominant use of browse on timbered types, was circumstantial ITABLE XXIX0 FREQUENCY OF OCCURRENCE AND VOLUME EERCENTAGES OF PLANT TAXA AND FORAGE CLASSES AMONG 37 MULE DEER RUMEN "SAMPLES. Taxa l/ Forage Class Summer 1963 Fall (Oct-Nov) Winter Spring 'i960 11961 1962 Total 1961-62 1962-63 Total 1963 (6) 2/ ' (I ) (6 ) (10) (17) (4) (4) (8) (6) Gramineae Family 33/T 3/ 100/43 70/23 77/29 100 /3 100 /19 100 /11 67/25 Grass & Grass-like 33/T “ IOO/43 70/23 77/29 100 /3 100 /19 100 /11 83/25 Artemisia cana . 30/5 1/3 50/T 25/T Artemisia tridentata 100/15 D b h E 4l/5 100 /63 100 /21 100 /42 83/5 -Chiysothamnus spp. 100/95 83/25 to /7 ; 77/19 75/2 75/16 75/9' 33/8 Juniperus scopulorum 100/9 60/3 71/5 100 /1 6 75/9 : 87/13 .33/5 Prunus -virginiana 83/14 D b h E 5/T ' 17/T Pseudotsuga menziesii : 17/T D b h E ll/T 50/2 . 25/T 17/T Rhus triloibata 100 /26 33/T 20/T . 23/T 25/T 25/T 25/T' 17/T Rosa spp. IOO/1 3 100/T 17/T 40/1 . 47/T 50/T . 25/T 67/15. Symphoricarpos spp. 83/2 33/T 50/3 41/1 25/T . 13/T 33/2 Unidentified browse ■ 17/T lOO/l 67/4 6 0 /1 65/2 50/T. 25/7 .. 37/3 ' 67/1 . - Browse Total IOO/59 IOO/.99 100/57 100/21 100/39 ]D0/95 .100/53 100/73 100/5.4 Allium textile 'Aster spp. ■ Cirsium spp. Coiiiandra 'umbellatum ■" -'family • Medlcago spp... Melilotus' officinalis' ‘ Tragonogon duUius ' Unidentified fdrbs -ForU Total 33/2 100/25" t i 100/36 : ■ -;. . 26/1 : u / T " " 17/T V HeU M shE ' 10/9' - 5/5. .2 0 /5 1 1 /3 30/16. iT/9 :PPH.A 67/T 100 /2 1 ' 88 /1 3 Y R R.A ' 6 7 /1 100/54 88 /3 2 :Vr;'a5/5; :'13/3 . 33/5/ 75/1 .. 25/T 25/5 5 0 /2 75/15 50 /2 100 /28 :'37/ T . 17/T iuhE H 67/3 13/3 33/4 63/9 83/2 75/27 100/30 :6 TABLE XXIX, Continued. l/ Taxa having volume percentages of one per cent or more of the total during any seasonal period. Others occurring during seasons included: Carex spp. (S), Artemisia frigida (W) Artemisia ludoviciana (F), Antennaria spp. (F), Conringia orientalis ( s ) , Crepis occidentalis (S), Descuriana pinnata (s), Gaura, coccinea (S), Geum triflorum ( s ) , Glycyrrhiza lepidota (FMr), Lactuca pulchella (Su), Linium rigidum (SuJT^ Lomatium foeniculatum ( s ) , Microseris nutans ( 8 ) , Potentilla arguta (Su), Senecio canus (Ehi), Thlaspi arvensis (8), Vicia americana (S Su), Yucca glauca (F), Petalostemum (F), and Ericaceae family (F). 2/ Humber of rumen samples in parenthesis. 3/ Per cent frequency/per cent of total volume. T = less than one per cent. - 133 - evidence for greater than observed importance of shrubs. This suggested that Rhus trilobate was the most intensively used individual taxon and that use of other shrubs was also relatively greater in su m m e r . Fall (September, October, November),— Data were obtained at 48 mule deer feeding sites; 22, 18, 4; and 5 for i960, 1961, I962, and 1963 respectively. Mule deer use was recorded on 53 taaa; 33, forbs, ,12 shrubs, and 8 grasses. Fifteen of these, five forbs, nine shrubs, and one grass, received more than one per cent of the total combined use. Use of browse increased from summer and predominated use for all types combined as well as on Agropyron/Symphoricarnos, Agropyron/Foa, Artemisia longifolia, and both timbered types, Over-all, use on Rhus trilobata, Chrysothamnus viscidi,florus. Symphoricarpos spp,, and Melilotus officinalis, a forb, was about equal. Rhus trilobata was used at intensities similar to summer during September and early October, Considering the observed and/or,inferred importance of the Pinus/Juniperus and Pseudotsuga/Juniperus vegetation types for use by mule deer, it pro­ bably was the single, most important taxon, at least in September. Although Chrysothamnus v iscidiflorus,received rather intensive use on both of the timbered vegetation types as well as on Sa r cobatus/Artemisia and, Agropyron/Symphoricarpos, this taxon was abundant and/or widely distri­ buted only on the latter. Chrysothamnus n a u seosus, which was abundant, widely distributed, and intensively used on the Artemisia/Agropyron type, appeared much more important and may have been the single, most important taxon during October and November. Symphoricarpos spp. received intensive — 134 — use on the Agropyron/Symphoricarpos type during September and early October as well as on both timbered types throughout the p e r i o d . Other shrubs individually received only minor use, occurred largely on minor vegetation types, and/or were used, intensively during.only part of the period. Those which appeared of some importance included: R o s a s p p . in September; Artemisia longifolia which was used intensively :d.uring September, largely on the minor Artemisia lon g i f o l i a •type and Pihus ponderosa/ Artemisia longifolia association; Artemisia cana in October and November; and Artemisia tridentata as well as Juniperus scdpulorum in November, Melilotus officinalis, the most intensively used forb, was one of the more important forage sources during fall,- especially. in-September, Use of other forbs generally was very minor b u t .varied in' relation to avail­ ability of M e l i l o t u s . Aster commutatus and Tragopogon dubius were most consistently and widely used. Glycyrrhiza. Iepidota received intensive use in early fall, largely on the Agropyron/Symphoricarpos. vegetation type. Other taxa receiving somewhat greater use included Artemisia ludoviciana and Eriogonum multiceps during October and Phlox ho o d i i -in November. Use of grass by mule deer during the fall varied considerably. Intensive use was restricted to Poa secuhda and was related to fall "green-up." It occurred" only during October and November. The greater discrepancy between rumen samples and feeding site data during fall was related to collection of most samples during late October and November in years of intensive fall grass use. - 135 - Winter (December, January, February).--Sixty-two mule deer feeding sites were examined during the four winters as follows: 196l, 32; 1962, 16; 1963, 9; and 1964, 5» Use was recorded on 39 taxa; 14 forbs, 16 shrubs, and 9 grasses. Of these, 15— including 4 forbs, 10 shrubs, and I grass— received more than one per cent of the total instances of use for all years. Browse was by far the most important forage class. Artemisia tridentata, the single, most important taxon> received about 33 per cent of the four-year total winter use. It was used most inten­ sively during mid-winter and periods of snow cover, Use of Chrysothamnus nauseosus, which ranked second, was most intensive in early winter and declined to minor levels late in the period. Juniperus scopulorum ranked third. It appeared to receive some use in association with feeding on Artemisia tridentata, but was also used inten­ sively when mule deer used the Pihus/Juniperus vegetation type during, periods of heavy snow and cold temperatures. Symphoricarpos sppj and Rhus trilobata' received frequent, though minor, use on the Pinus/ Juniperus type throughout the winter. Use on other shrubs usually was very minor and/or restricted to minor or moderately used vegetation types. Chrysothamnus viscidiflorus was used mostly on the Pseudotsuga/Juniperus and Sarcobatus/Artemisia types which received only minor winter use by mule deer. Forbs collectively were of only minor importance during most winters. Taxa used most consistently included Melilotus officinalis, Tragopogon d u b i u s , Artemisia frigida, and Ph l o x hoodii. The latter was - 136 - used relatively intensively during snow-free periods and during February= None of these received m o r e than three per cent of the total winter use= Wi n t e r use of grasses was very minor over-all but varied in relation to “green-up*1 and snow cover. Intensive use occurred only on “green" Poa s ecunda, mostly during December, Other, ,species received only incidental and/or use of seedheads. Volumetric percentages for winter rumen samples'indicated forage class and individual taxon usage similar to,that recorded at feeding sites. The slightly greater use of grasses reflected collections only during “green-up“ years. Spring (March, April, Ma y ) , --A total of . 6$ feeding site examinations was made during spring; including 34 in 1961, 20 in 1962, 10 in 1963, and I during March 1964« The use of 65 taxa, comprised of 44 forbs, 16 shrubs, and 7 grass and grass-like plants, reflected the flux in availability as well as mule deer forage preferences throughout the period. Twenty-one taxa, 9 forbs, 10 shrubs, and 2 grasses, received more than one per cent of the total instances of use recorded during spring. Browse was the most important forage class for all years combined. Artemisia tridentata was the most important taxon over-all, largely because of intensive use during March. Its use and importance generally declined sharply during April to very minor by early May, Juniperus seopulorum ranked second in importance among shrubs and third among all plants. Trends in use of this as well as other winter browse plants which received only minor use in early spring, paralleled that of Artemisia tridentata. - 137 - This was reflected in a 56 per cent decrease in recorded use on shrubs from March through May, Rhus trilobata„ Ribes Cereums Rosa spp„s and Symphoricarpos spp, were of only very minor importance for the entire period3 but received increasing use monthly, Ribes cereum received some intensive use during April, Forbs, which ranked second in combined data, typically received only minor use during March but were about equally used as compared to browse during April and predominanted recorded use in May, Phlox Sioodii9 which received minor Use5, was the only forb with significant use during March, Although no taxon received more than six per cent of the total instances of use for the entire periodseveral members of the Poa secunda union received intensive use during April and early May, The more important, and most consistently used taxa. included Allium textile, Lomatium foeniculatum, Microseris nutans, and Fritillaria pudica, Use of this group declined by mid-May and was supplanted by increasing use of taxa from the Tragopogon dubius Unionjl especially Comandra umbellatnm,, Tragopogon dubius „ and Melilotus officinalis s but including Arnica Sororiaj, Crepis occidentails., Gaura coccinea, and Vicia americana, Grasses were used more intensively by mule deer during spring than other seasons but typically received only moderate Use51 largely during April,, Poa secunda was the only taxon receiving intensive use. The very minor use of Agropyron smlthii and occasional use of Koleria cristata appeared largely incidental, to use of Poa, Rumen samples for the spring period generally reflected trends in use of forage classes and individual taxa similar to those indicated by feeding site data. The undue prominence of Rosa spp. in the rumen data reflected influence of a sample comprised largely of that item. Yearlong Trends.--The results indicated significant - yearlong changes in mule deer use of various forage classes and individual taxa for food = Shrubs (browse) comprised the most important forage source averaging approximately 62 per cent of mean monthly instances of. use during the four-year period. The mean seasonal percentage of browse in rumen samples was 56 per cent. Use of browse was lowest in Mayj, increased gradually through summer and sharply through fall to a maximum in Januaryjl then decreased through March and April to the minimum by late April and/or early May. Trends in use bf forbs5 collectively ^ complemented changes in use of browse. This forage class comprised approximately 32 per cent of the total instances of plant use averaged over months and 31 per cent of rumen samples averaged over seasons. Most intensive use occurred in early May, the lowest in January. Over-all, grasses and grass-like plants were of minor importance. Yearlong averages were about six per cent of mean monthly total instances of plant use and fourteen per cent of mean seasonal rumen sample volume percentages. The discrepancy between rumens and feeding site data was related to concentration of rumen collections in periods of more intensive grass use. Use of grass was negligible during summer and early fall, increased in late fall in years of "green- up," declined to a minimum during mid-winter, 'increased sharply during - 139 - late Ma r c h and/or early April to maximum yearly, intensities of use in early ■ : ' April, then decreased to m i n i m a l .summer levels by. mid-May. Melilotus officinalis was the single, most important taxon in relative use on feeding sites on all vegetation types from mid-June through - - ■■■ V-:.';'3- September. Subsequent yearly use was generally minor but varied in rela- tion to late summer-early fall growth and persistence.of- first-year stems and/or the abundance of dry seed stalks. It was used most intensively after flowering commenced. Rhus trilobata ranked second in use on all feeding sites from early June through late September or early October, but was b y far the single, most important taxon on the intensively used Pinus/ Juninerus and Pseudotsuga/Juniperus vegetation types,and m a y have been most important over-all. Subsequent fall, winter, and spring use was generally minor, but varied in relation to prior summer-early fall use and/or relative intensities of mule deer use.on the Pinus/juninerus type during these periods. Symnhoricarnos a l b a . occidentalis. R o s a ^arkansana„ ,R. n u t k a n a . and Prunus vireiniana were other shrubs .receiving relatively important use by mule deer throughout the summer and early fall. Trends in use of these deciduous shrubs generally paralleled use on Rhus trilobata.. -■ ' except that Svmphoricarpos spp. remained important throughout fall and into early winter and Prunns vireiniana. was only rarely used in winter and early spring. Early summer use on each consisted largely of leaves. Use of twigs increased progressively and predominated late summer and subse­ quent p e r i o d s . / — 140 — One of four major seasonal changes in the yearlong food habits of mule deer occurred during early fall when use on Ghrysothamnus nauseosus and C. viscidiflorus increased sharply concomitant to decreased availability and use on deciduous shrubs and forbs. The former appeared most important. The abundance of Co viscidiflorus only on the minor Agropyron/ Symphoricarpos type precluded important u s e . . There was.some evidence to suggest that this taxon had been more abundant and probably of greater importance in previous years. Both Chrysothamnus nauseosus and C. viscidiflorus were used intensively only after flowering. "Leaders" which had previously been "clipped" appeared more preferred than those which held seedheads. Artemisia cana and Artemisia longifolia received some use though never significant because of limited ,abundance and/or distribution. Poa secutida was moderately important in late fall of some y e a r s . Use of "fall" browse taxa, especially C. nauseosuss remained important into early winter, then declined to only minor levels in late winter and early spring.. The second major seasonal change in mule deer food habits occurred concomitant to decreased availability of Chrysothamnus nauseosus apd the .onset of winter conditions in late fall. It was characterized by sharply,increased use of Artemisia tridentata and' Juniperus scopulorum. The former prevailed as the single, most important taxon on feeding sites on the intensively used Artemisia/Agropyron vege­ tation type as well as all types combined through March. U s e on Juniperus scopulorum was consistently of minor or moderate importance. - 141 - Although relatively unimportant over-all. Atriplex n u ttallii, Sarcobatus vermiculatus, Pseudotsuga menziesii, and Pinus ponderosa received intensive use on some feeding sites during some winters. The third and most marked seasonal change in mule deer food habits occurred in late March and/or early April. This coincided with the onset of spring ■ . ■ "green-up" when the sudden, significant decline in mule deer use of Artemisia tridentata was complemented by a marked increase in use of grass, largely Poa secunda. on feeding' sites. Use of this taxon declined after mid-April in most years with increasingly intensive mule deer use of early forbs of the Poa secunda union. These, collectively, were most important during late April and early M a y but decreased as increasingly .important use of forbs of the Tragopogon dubius union and deciduous shrubs resulted in the last major seasonal change to food habits characteristic of summer. Trueblood (i960) reported generally similar trends in food habits of mule deer in the Missouri River Breaks of Valley County, Montana. Janson and Picton (1959) found Pseudotsuga menziesii and Chrysothamnus spp. to be the principal items in 14 rumen samples from mule deer collected during February 1958 from the Missouri River Breaks of Fergus County, west of the present study area. > ■ Year-to-Year T r e n d s .— Relative intensities of mule deer use of forage classes (Table XXX) and individual taxa (Table XXXI) varied significantly between and within years. This was largely related to extensive differ- ences in precipitation and its influence on vegetal growth. Conditions ranged from severe drought and extremely poor plant growth during 1961 to TABLE XXX. A COMPARISON OF PERCENTAGES OF MOLE DEER USE OF GRASS, BROWSE, AND FORBS AT FEEDING SITES ON VARIOUS VEGETATION TYPES DURING SEASONS AND YEARS, 1960-64. L B A I h . y Y D s v . y v I s o . v L B E R . L g g A r y I v Forage AGROP l/ JUNIP l/ JUNIP l/ ARTEM l/ COMBINED uiass Su Fa, Wr mT D2/Su Fa Wr Sp Su Fa Wr Sp Su Fa Wr Sp Su Fa Wr Sp Grass and Grass-like 1960-61 O O T ^ 9 T O ' 0 2 O O O I T T . TL 0 T 6 1961-62 T 65 5 16 ' T 5 3 2 O -=•■=• O O 0 26 T 18 T 17 3 .14 1962-63 O 2 3 39 T .T T 30 O 2 — — O 0 0 45 87 T I 12 36 1963-64 O O I — ™ T T O — — T O T = — 0 — — ■= ™ ™ = T T T — = 1960-64 T 18 2 16 T 2 T 7 T T T T 0 19 12 19 T 5 2 13 Browse 1960-61 O 57 92 4o 77 92 100 68 72 100 100 90 •a =• = 1» 100 52 46 81 96 56 1961-62 14 22 94 33 84 79 80 75 99 100 100 I 34 96 78 54 90 91 54 1962-63 2 O 64 21 4l 36 .53 43 49 64 — 43 2 0 28 9 25 25 52 31 1963-64 O 2 95 — =■ 46 86 79 = =• 76 ■ 63 93 5 = — =• = = =• 45 65 88 1960-64 6 37 88 LTNCO 64 83 86 66 82 86 98 81 x 2 25 80 53 ■43 72 88 52 Forbs 1960-61 100 ^3 8 50 23 8 T 29 4 0 0 9 0 4o 51 :19 ■ 4 38 1961-62 85 14 2 52 16 16 17 23 1 = = 0 0 99 4i 4 4 45 ■ 13 6 32 1962-63 98 98 32 4o 59 63 47 27 51 34 = = 57 98 ICO 27 4 75 74 36 33 1963-64 100 98 4 •= = '54 i4 21 ™ = 23 37 6 = ™ 95 = = * “ ■=■ = = 54 35 12 = =• 1960-64 94 45 10 49 36 15 14 26 18 14 2 19 98 56 8 28 56 23 10 35 l/ Vegetation types receiving 10 per cent or more of the total observed mule deer use during " at least one season; Artemisia/Agropyronj Pinus/Juniperusj Pseudotsuga/juniperusj and Sarcobatus/Artemisia. 2/ Summer, fall, winter, and spring respectively. 3/ T = less than one per cent. St fT ” 143 " well above "normal81 rainfall and exceptionally good plant growth during 1962» They were "below normal" in I960 ^ and "above normal" in 1963« All classes of plants were affected but changes in composition and abundance of forbsj especially Melilotus officinalis (Table XXXII) were most signifi­ cant. TABLE XXXI. A COMPARISON OF RELATIVE INTENSITIES OF MULE DEER USE OF SOME IMPORTANT FOOD PLANTS DURING SEASONS AND YEARS, 1960-64. I/ Taxon 1960-61 Su/Fl/Wr/Sp 1961-62 1962-63 Su/Fl/Wr/Sp 1963-64 Su/Fl/Wr/Sp Melilotus officinalis 45/13/ Q/ T 36/ I/ 0/ T 58/62/19/14 37/24/ 7/-- Rhus trilobata 29/20/ 6/ 6 36/ 6/ 7/ 2 10/ 9/ T/ 4 22/23/ 3/-- Svmnhoricarnos son. 10/15/10/ 6 10/13/ 2/ I 8/ 9/ I/ 3 14/8/11/— Chrysothamnus son. 0/27/19/1 0/30/ 9/ 5 0/ 0/28/ 4 .0/ 5/34/“” Artemisia tridentata 0/ 6/33/27 0/ 3/49/24 0/ 0/13/16 0/ 0/18/— Juninerus scopulorum 0/ T/ 9/ 6 T/ 7/16/18 0/ 0/ 3/ 4 0/ T/ 2/-- Number of taxa used 21/20/20/45 30/37/24/50 34/15/31/41 26/2I/I9/IO l/ Percentages of total instances of use on all vegetation types combined. TABLE XXXII. PERCENTAGES OF MULE DEER FEEDING SITES EXAMINED DURING SUMMER AND FALL SEASONS ON WHICH MELILOTUS OFFICINALIS WAS ABUNDANT,' SCATTERED, AND ABSENT, 1960-63." Relative Abundance I960 1961 1962 1963 Summer Fall Summer Fall Summer Fall Summer Fall Abundant 59 23 ' 30 0 61 100 90 60 Scattered 19 4 16 40 30 0 10 40 Absent 22 73 54 60 9 0 0 0 ~ 144 — Decreased availability of forbs during I960 and 1961 was reflected in increased use of browse as well as 'unseasonable use of certain taxa as compared with other y e a r s „ These phenomena were most pronounced during 1961 when summer-fall use of Mo officinalis and other forbs was lowest and browse, especially Rhus t r I lobata0 was most intensively used in summer= As a result of sharply decreased availability,, fall use of "summer" browse taxa was relatively minor and was supplanted by "early,cs intensive use on "fall" browse, especially Ghrysothamnus spp== Subsequently, with decreas­ ing availability and use of these taxa, Artemisia tridentata and and Juniperus scopulorum received unseasonable, intensive use in October= Although feeding sites showed minor use of A= tridentata during both i960 and 1961, this taxon comprised an average of 12 per cent for three rumen samples from October 1961 while absent from the one sample for 1960» Winter use of both the latter and J= scopulorum was relatively high, as compared to i960, and use of most other "preferred" browse was low= These taxa were also-used exceptionally late into spring, early May, in 1961 = The most significant differences appear in comparisons of data for 1960-61 and 1961-62 with that for subsequent y e a r s » Forbs were moderately abundant during early summer of 1962, but greatly increased in late summer and fall following growth of first-year stems of Melilotus officinalis. This taxon, in particular, was intensively used during summer and fall and received moderate use in winter on the timbered vegetation types where it persisted in relative abundance = Relative intensities of use on deciduous browse,, especially Rhus t r i l o b a t a „ F& decreased significantly during summer as compared to previous years. Use of the latter may have been influenced by severe defoliation by a larval Lepidoptera during that summer and/or undue sampling, bias favoring liOpenli vegetation types. Moderate to high levels of use occurred on feeding sites on both of the timbered vegetation types. Fall use.,was at intensities ' ■ ■ -Vj"- similar to summer. Significantly ^ fall use was not detected on Chrysothamnus spp„, Artemisia tridentata, and Jnmiperas seopulomnm at feeding sites. Minor percentages of Chrysothamnus spp, and J, scopulorum and only a trace percentage of A, tridentata occurred in 10 rumen samples collected during the period (Table XXIX) ^ though Chrysothamnus spp, was prominent in several samples collected prior to .intensive use of grass in late October, The substantial decrease in use of A, tridentata and J, scopulorum and the greatly increased use on Chrysothamnus spp, during winter5 as compared with previous years, was more significant than changes in use during fall. The repeated occurrence of the same phenomena under somewhat similar conditions during,the winter of I963--64 suggested that both A, tridentata and J, scopulorum were only secondarily preferred winter forage species. Their importance during 1960-61 and 1961-62 m a y .have been unduly exaggerated due to. excessive and/or unseasonal utilization of Chrys01hamnus spp, and other more preferred shrubs; e.g., Symphoricarpos spp,, Forbs, especially Melilotus officinalis, were exceptionally abundant again during 1963 and received intensive use during summer and fall. The relatively intensive use of Rhus trilobata at feeding' sites, despite the ' 7 Z abundance of M= officinalis, was further evidence that this taxon was — 146 - equally or more important as a summer-fall forage'source« This further suggested that the decreased use of IL trilobata on feeding sites during 1962 was exaggerated by some factor other than preference for Mo officinalis 0 Some yearly variation in mule deer food -habits was also influenced during fall, winter, and spring by the relative availability of green" grass, especially Poa secunda. The increased occurrence of grass in both the feeding site data and rumen samples for the fall and winter periods of 1961-62 and 1962-63 reflected "green-ups" in early October and late October respectively. This certainly influenced intensities of use on Chrysothamnus spp. and may also have influenced use of other forage taxa. The relatively high use of grass for the spring of 1963 reflected early growth and greater abundance of Poa secunda and its intensive use during March as well as April. The relatively light use of grass during the spring of 1961 was related to the generally reduced herbaceous growth of that year. In addition to changes in intensities'of use of important taxa, variation in forage conditions influenced differences in the number of taxa used during seasons and years (Table XXXI). This was most significant during spring and fall of driest years when supplies of seasonally pre­ ferred taxa were minimal. The significance of seasonal and yearly differences on observability of mule deer as well as population trends was discussed earlier. - 147 - Elk A total of 285,030 instances of plant use by elk was recorded at 178 feeding sites between June I960 and January 1964= The number of instances of use at each site varied from 50 to 750 but usually was between 100 and 200. The average number was 157= The smallest samples occurred in late spring and early summer when single individuals and small groups of elk were common. Data for elk use of taxa which received at least 10 per cent of the total instances of plant use for at least one vegetation type during at least one season are presented in Table XXXIII= Taxa which received lesser intensities of use are considered in Appendix Table LX= Summer (June3 July3 August),--A total of 64 elk feeding sites was examined during the four summers 5 23, 14, 19, and 8 respectively for I960, 1961, 1962, and 1963» Use was recorded on 71 taxa, including 49 forbs, 10 shrubs, and 12 grass and grass-like plants. Eleven taxa, six forbs, three shrubs, and two grasses received an average of more than one per cent of the total instances of use during summer. Forbs were the most important forage source on all but the Pseudotsuga/Juniperus vegetation type and comprised 75 per cent of the total combined summer use. Melilotus officinalis, the predominant taxon, accounted for more than two-thirds of the use on forbs and received approx­ imately 53 per cent of the total instances of use for the period. It was used on 88 per cent of all elk feeding sites. Glycyrrhiza Iepidota9 which — 148 — TABLE XXXIII. ELK USE OF TAXA RECEIVING AT LEAST TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE AT FEEDING SITES ON AT LEAST ONE VEG­ ETATION TYPE DURING AT LEAST ONE SEASON, 1960-64 COMBINED. Vegetation Types Plant Taxa Artemisia/Agropyron: Agropyron smithii Koleria cristata Poa secunda Grass Total Melilotns officinalis Forb Total Artemisia tridentata Browse Total Pinus/jnniperus: Agropyron smithii Muhlenbergia cuspidata Stipa viridula Grass Total Arnica sororia • Melilotus officinalis Forb Total Juniperus scopulorum Symphoricarpos spp. Artemisia longifolia Browse Total Pseudotsuga/juniperus Agropyron smithii Grass Total Artemisia ludoviciana Melilotus officinalis Forb Total Chrysothamnus viscidiflorus Juniperus scopulorum Prunus virginiana Pseudotsuga menziesii Symphoricarpos spp. Artemisia tridentata Browse Total Summer Fall Winter Spring (29-26)1/ (17-27) (17-19) (19-29) 42/13 2/ 88/43 100/70 100/54 33/6 82/17 71/8 53/4 17/T 76/10 76/5 90/20 67/23 100/81 100/86 100/81 92/59 18/7 12/T 21/2 100/77 53/13 35/2 90/13 18/2 41/10 58/8 8/T 29/6 47/11 58/8 (15-35) (9-19) (11-22) (5-16) 33/13 44/24 91/64 100/49 13/T 56/11 18/1 20/T 22/T 36/2 60/14 47/18 89/45 91/78 100/77 20/11 87/62 22/20 9/T 100/74 44/22 4.6/ 6 40/13 11/11 36/3 20/5 33/3 44/10 27/5 7/T 11/11 9/1 20/T 47/8 56/32 73/17 40/10 (10-24) (3-12) (1-16) 10/T 67/22 100/3 40/T 67/31 33/16 100/5 60/26 100/2 70/32 67/18 . ' 100/6 100/13 100/12 80/37 33/2 100/19 100/21 33/14 100/T 33/33 100/42 100/67 68/51 100/90 TABLE XXXIII„ Continued„ Vegetation Type; Plant Taxa Summer Fall Winter Spring Sarcobatus/Artemisia: (4-13) (3-7) (8—l6) (3-16) Agropyron smithii 25/20 33/30 75/57 100/61 Poa secunda 100/40 62/16 68/25 Grass Total 50/25 100/73 88/77 100/86 Melilotus officinalis 100/69 33/2 Forb Total 100/73 ' 33/2 25/1 100/11 Ghrysothamnus viscidiflorus 33/17 Artemisia cana 12/12 Browse Total 5Q/2 67/25 62/22 67/2 Agropyron/Symphoricarpos: (13-25) (5-14) (4-18) Agropyron smithii 36/2 60/20 100/62 Poa compressa 20/7 75/15 Grass Total 43/3 100/36 100/84 Aster commutatus 14/T 40/12 Glycyrrhiza lepidota 100/34 60/12 Helianthus maxamilliana 7/T 20/10 Melilotus officinalis 93/53 40/15 50/T Forb Total 100/94 80/59 '75/16 Xant hium; (7-18) (1-2) (1-4) (2—10) Agropyron smithii 33/4 100/£9 Distichlis stricta 100/12 Grass Total 33.6 100/12 100/93 GIycyrrhiza lepidota 83/37 100/60 100/12 50/1 Melilotus officinalis 100/46 100/33 50/4 Xanthium strumarium 33/T 100/60 Forb Total 100/92 100/100 100/71 50/7 Artemisia longifolia 17/T 100/17 Browse Total 67/3 100/17 Artemisia longifolia; Lactuca serriola Melilotus officinalis Forb Total (3-10) 67/24 100/57 100/100 - 150 - TABLE XXHIIo Continued Vegetation Type; Plant Taxa ■ Summer Fall Winter Spring Agropyron/Poa: ( 5 - 1 3 ) 100/13 100/57 100/74 60/22 100/24 Agropyron smithii Poa secunda Grass Total Melilotus officinalis Forb Total l/ In parenthesis s The number of feeding sites on this vegetation-The total number of taxa used on the type. 2/ Per cent frequency of use among all feeding sites on this vegetation . type/aggregate mean percentage of the total instances of use for this vegetation type. T = less than one per cent. received moderate use, was the only other forb of importance. It was especially important on the Agropyron/Symphbricarpos type which was used intensively by elk during July and August. Aster eommutatus 9 Lactuca serriola, Microseris nutans, and Tragopogon dubius received minor Usejl largely during June. Comandra umbellatum, Helianthus maximiiliana, and Petalostemum purpureum also received minor use later in the period. Browse5 which received about 15 per cent of the total summer Use5 was used most intensively on the Pseudotsuga/juniperus vegetation type during July and especially5 August. The most important taxon was Prunus virginiana. Symphoricarpos spp. ranked second. Rhus trilobata received only minor use throughout the periods but was used on both of the timbered vegetation types. Rosa spp. and Ribes spp. also received minor Use5 largely on the Pseudotsuga/Juniperus type. Grass and grass-like plants generally constituted only a minor per­ centage of the total use for elk during' the summer» Agropyron smithii was the most intensively used taxon» It ranked third in importance over alia mostly because of exceptionally intensive use during one summer= Koleria cristata received minor Use9 primarily on the Artemisia/Agropyron and Pinus/juniperus vegetation types= Other grasses received only very minor use at all times= The slightly decreased use on forbs through the summer was comple­ mented by increased use of browse= Grass use was consistently low through- . out = The contents of one elk rumen9 collected during July 196.3 3 was comprised almost entirely of Melilotus officinalis (Table XXKIV). Prunus virginiana= Rosa spp=9 and Symphoricaroos spp= were present but only in minor or trace amounts= Fall (September9 October9 November)=--Forty-three elk feeding sites were examined as follows; ■ I960=, 23; 1961, 12; 1962, 7; and 1963, 1= Use was recorded on 35 taxa, including 18 forbs, 11 shrubs, and 16 grass and grass-like plants= Sixteen of these, 5 forbs, 5 shrubs, and 6 grasses, received more than one per cent of the total combined fall use= Use of grass increased significantly from summer, constituting about 62 per cent of the total fall use= It was intensively used on all but the minor Xanthium vegetation type= Agropyron smithii„ the single, 'most important taxon, received approximately 30 per cent of the total fall use= - 152 - TABLE XXXIVo FREQUENCY OF OCCURRENCE AND VOLUME PERCENTAGES OF PLANT TAXA AND FORAGE CLASSES AMONG 12 ELK RUMEN SAMPLES, Taxa Srammer Fall Winter Spring Forage class l/ 1963 1961 1962 1963 Total 1963 1963 (I) 2/ (5) (I) (3) (9) (I) (I) Gramineae (Total) 100/872/% 100/24 100/68 97 82 Artemisia frigida 33/6 11/2 T . Melilbtus officinalis 96 100/15 33/5 10 Solidago sp. 33/3 ll/T Yucca glauca 67/2 22/T Unidentified forbs T 60/T T 44/T 2 I Forb Total 97 60/1 T 100/26 66/8 2 12 Artemisia cana .20/T 3 3 / H 22/6 Artemisia longifolia : 33/4 11/1 Artemisia tridentata "■’-80/6 . T . 55/3 Chrysothamnus spp. 40/1 67/17 44/6 Prunus virginiana I 33/13 11/4 T Rosa sup. I 40/T 33/T 33/T T Symphoricarpos spp, T 100/2 33/T I Unidentified browse T 60/4 T 67/T 66/2 T 4 Browse Total ■ 3 100/12 I 100/48 1 ■ 100/24 I 6 l/ Only those taxa which comprised at least one per cent of the total volume percentage for at least one season for one year. Others occurring^ with season of use in parenthesiswere: Aster sp. (SuF)9 Cirsium sp. (F), Glycyrrhiza lepidota (SuF), lafeiat§e^family (Su)5 Lomatium foeniculatum (S)5 Menzelia laevieaulls td os'Microseris nutans (S)5 Tragopogon,..dubius (SuF) 9 Cpmpositae family (W)5 ■ Juniperus scopulorum (FW)»- Pimfs ponderosa (F)5 Rhus trilo'bata "(F); and Ribes sp, (S). 2/ Number of rumen samples in parenthesis, 2/ Per cent frequency of occurrence among rumen samples/average percentage of total volume each sample. .1 - 153 - This occurred primarily during October and November,■ A= smithii received only minor use during September, Poa secunda ranked second among grasses for the entire period but was used intensively only when "green," Its Koleria cristata, the only other grass ^o;receive ■important use throughout intensively used grass during September, Other grasses received only very minor use. Forbs ranked second, Melilotus officinalis was the most intensively were equally important during September5 when forbs were the most important forage sources but received only minor use later. Other forbs individually received only minor use and/or were used mostly on minor vegetation types. Use on shrubs comprised about 15 per cent of the combined fall total. The most intensive use of browse occurred during September. Symphoricarpos spp. and Chrysothamnus viscidiflorus received consistent ^ though generally minor3 use throughout fall. Use of other taxa was usually very minor, Artemisia longifolia was of some importance in September and both Artemisia tridentata and Juniperus seopulorum received minor use in late fall of some years. Rumen samples indicated similar high levels of grass use during fall,, greatest use occurred on the fall, was also used most intensively when "green51 and in late fall. Muhlenbergia cuspidata received only minor use over-all, but was the most used taxon for the entire period. and Aster commutates but suggested that browse" may be used to a greater extent than indicated by feeding site data, at least during some years. - 154 - Winter ( D e c e m b e r January5, February) „--Data were obtained at 38 elk feeding sites; 22 in 1961* 9 in 1962, 3 in 1963, and 4 in I964. Elk use was recorded on 39 taxa, including 14 Dorbs9 13 shrubs, and 12 grass and. grass-like plants „ Ten51 consisting of one forb, five shrubs, and four grasses, received more than one per cent of the total combined winter use. Grass was by far the most important forage class. Agropyron smithii0 the single, most important taxon, received about 62 per cent of the total use. Poa seeunda and Koleria cristata, which ranked second and third respectively, were used most intensively during early and late winter in years of fall 11 green-up.1,1 Agropyron spicatum was used relatively frequently but in only minor amounts. All other grasses received only occasional use during the period. Shrubs constituted about 17 per cent of the total combined instances of use for winter. Nearly one-half of this was on Artemisia tridentata. Artemisia cana, Juniperus scopulorum, and Symphoricarpos spp. received minor use, mostly during January. This reflected sharply increased use of browse by elk during periods of snow cover. Rouse (1957) reported important use of browse by.elk in winter only during periods of severe cold and snow cover. Other shrubs were used only occasionally, primarily in ,December and/or January. The contents of the single rumen sample collected in late winter was comprised a.lmost entirely of grass. - 155 - ' Spring (March, April, May).— A total of 33 elk feeding sites was examined during spring; 19 in I960, 9 in 1962, and 5 in 1963 „ Elk use was . ' .■ ' recorded on 50 taxa, including 34 forbs, 5 shrubs, and TL grass and grass­ like plants. Eleven, five forbs, one shrub, and'five grasses, received one per cent or more of the total spring vuse. „ . . . Grass continued from winter as the predominant forage for elk, consti­ tuting approximately 82 per cent of the total, combined spring use. Agropyron smithii. which remained the most important taxon, received about 50 per cent of the total use. Poa seeunda received moderate use and was the second, most important source of forage throughout the period. Other grasses received only minor use. Those used more intensively included Agropyron spicatum in March, Stipa viridula in. March and April, Poa compressa in April but only on the Agropyron/Symphoricarpos vegetation type, and Koleria cristata throughout the period. Use of grasses generally decreased monthly throughout spring. Forbs, which ranked second in the combined data, received only minor use during March, increased in relative use and importance during April, and were used rather intensively during May. Although no taxon individ­ ually received more than two per cent of the total .combined spring use, several members of the Poa seeunda union. Allium textile. Lomatium foeniculatum, and Microseris nutans„ received relatively intensive use during April and May. Decreased use of this group after mid-May was complemented by relatively intensive elk use of several members of the - 1$6 - Tragopogon dubius union# especially Arnica Sororia9 Gomandra nmbellatnm, Lactnca puleheIlaQ Melilotus officinalis. Taraxacum officinale, Tragopogon dubius, and Vicia americana. Spring use of shrubs was only minor# about five per cent of the total# and consisted largely of use of Artemisia tridentata during March, and early April, Juniperus scopulorum also received some use during March, Use of browse was negligible during May, The single rumen sample collected during early May showed similar taxa used and about the same percentages of grasses# forbs# and browse as feeding site data. Yearlong Trends = — Yearlong elk preferences for forage classes and individual taxa for food were characterized by significant trends,. Grass comprised the most important forage# averaging 58 per cent of mean monthly instances of use during the four-year period, ■The mean seasonal percentage of grass in elk rumen samples was 62 per cent. Use of grass was lowest during summer# increased sharply through September to October when grass became the predominant forage# then gradually•through late fall and early winter. Grass use typically decreased during January in relation to increased use of other forage during periods of complete snow cover# but subsequently increased to the yearly maximum in February, Use was inten­ sive through mid-April# then decreased sharply through late April and May to relatively low levels by early June, Trends in use of forbs# which ranked second over-all# generally complemented changes in use of grass. - 157 - Yearlong averages for use of forbs were 29 per cent of mean monthly instances of plant use and 30 per cent of mean seasonal rumen sample volume percentages. Use of forbs was most intensive during s u m m e r w h e n this forage class was by far the most important source of forage for elk. Use declined sharply throughout fall and early winter# increased, slightly in January# then decreased to minimal in February. Use of forbs increased slightly through March and sharply through April and May to maximal summer levels during June. Over-all use of browse was only moderate. This forage class comprised approximately 13 per cent of the total instances of plant use averaged over months and 8 per cent of rumen samples averaged over seasons. Seasonal differences in intensities of use on shrubs# collec­ tively# were not extensive. Use of browse was lowest during late April and May# when only occasional use occurred# then increased to near maximum during August and September. Use decreased slightly through November but increased during December to the yearly maximum during January. Subse­ quently# elk use:' 0f browse again declined slightly through March# then sharply to the minimum in late spring. Melilotus officinalis was the single# most important taxon in relative use on feeding sites on all but the Pseudotsuga/Ju n i p e r u s ,vegetation type from early June through early September. It persisted as the most import...- tant forb# receiving moderate use# throughout the fall when dry stems and/or abundant growth of first-year stems were available. Subsequent winter and spring use was always very minor or negligible. Glvcyrrhiaa lepidota was the second-ranking forb and forage source ■ during September on intensively used "coulee-bottom61 Sitesjl but was used only 6 - . rarely during other periods. Prunus virginiana ranked third and was the most important browse plant for summer USes especially during July and ' . August. Its use and importance declined after, .September when only oceas™ I ■ . ■: V'S - : - • ■ . ional use occurred. Svmnhoricarnos so. (mostly S . occidentalis) was the f : I I, — . . . . . . . . . . . . . . . . . . ;r’ V ; . Seconds most important browse plant for seasonal use. It received.con- summer and early fall. It received important use during Julys Augusts and Sistents (though generally.,minors use throughout summer as well as during fall and;early winter. Rhus trilobata consistently received very minor use through summer and only occasionally during the remainder of the year. Summer elk use on shrubs consisted largely of leaves. Use of twigs increased and predominated after late summer. A major change in the yearlong food habits of elk occurred during late September when increased use on Muhlenbergia cuspidata Q Koleria Cristata9 Poa secunda. and other grasses complemented decreased use of forbs and deciduous browse, M. cuspidata was used intensively and was the most important forage source only during late September. Use of Agropyron smithii„ which received only minor use during summer and early . fall, increased very sharply during October when this taxon became the. Singles most Important food source, for elk. Extremely intensive use of A. smithii prevailed! throughout late fall, Winters and early spring. Its relative use and importance declined gradually through April and early May then very sharply during mid and late May to near minimum by early June. Poa seeuhda ranked as the seconds most important source of elk forage - 159 - during the entire period of intensive use on A= smithii. Moderate intensi­ ties of use occurred during late fall and early winters primarily following •'green-ups5c' as well as throughout spring= Mid and late winter use varied in relation to the availability of-"green" blades and/or snow cover = Koleria cristata = which received minor use throughout most of the year was the only other grass receiving prominent use during the fall-winter™ spring period = Its greatest importance occurred after "'green-up*1 in late fall= Poa compressa received intensive use only on coulee-bottom sites during September and April= Stipa viridula received minor use throughout fall as well as in March and Apriljl and Agropyron spicatum received very- minor use throughout winter and spring= Among Shrubsjl Chrysothamnus viscidiflorus received considerable use during September and relatively minor use later, in ,the period= This taxon was used only occasionally after early winter = Artemisia longifolia received some . intensive use during September but was used only rarely in other periods= Use of Ahtemisia tridentata increased during October and/or November and persisted at moderately low levels throughout winter and early spring= The most intensive use on this taxon occurred during periods of complete snow cover = Use declined sharply and was negligible by early April= The second major change in the yearlong diet of elk occurred during late April and May= It coincided with increased availability of forbs and resulted in this forage class supplanting grass as the most important elk forage = The change was marked by increasingly intensive use of early forbs of the Poa secunda union through April and early May and subsequently^ by increasingly important use of forbs of the Tragopogon dubius union in late May and early June as food habits became characteristic of summer. Yearlong trends in elk food habits reported for other studies.vary between areas in relation to vegetational characteristics and/or seasonal availability of the various forage classes. Generallyas reported here, the results have indicated preference and intensive use of forbs and leaves of deciduous shrubs during summer. Extreme differences have been reported for winter; e.g. =, differences in relative use of grass and browse which range from more than 99 per cent grass (Morris and Schwartz, 1957) * to 90 per cent browse (Young, 1938). Lovaas (1963) presented data showing relative use of forage classes by elk ranging from 69 per cent grass and 23 per cent browse during an "open" winter to 9 per cent grass and 91 per cent browse during a "severe" wiziter as evidence of elk preference for grass when available. The intensive use of grass by elk in early fall, when "summer" browse remained available, as well as daring winter indi­ cated that the importance of grass in the fall-winter-spring diet of elk in the Missouri River Breaks resulted from seasonal preference for this class of forage. Year-to-Year Trends.— Intensities of elk use of forage classes (Table IXM) and individual taxa (Table XXXVI) varied significantly between years. These differences were largely related to precipitation and its influence on plant growth (see discussion, page .141, and Table XXXII). TABLE XXXV, A COMPARISON OF PERCENTAGES OF ELK USE OF GRASS, BROWSE, AND FORBS AT FEEDING SITES ON VARIOUS VEGETATION TYPES DURING SEASONS AND YEARS, 1960-64. Forage ARTEM/AGROP l/PINQS/jUNIP l/PSEUD/JUNIP l/ SARCO/ARTEM l/ All Ifrpes Combined Class Su Fa Wr Sp 2/Su Fa Wr Sp Su Fa, Wr Sp Su Fa Wr Sp Su ‘ Fa Wr Sp Grass and .... Grass-like 196b-6l 9 92 9T 85 0 61 76 80 I 31 T 63 75 85 2 664 .82 82 1961-62 Tl 81 69 Tl 65-31 82 IbTs -- T 4 O = = = ■= = = 88 — T5 88 •• 44 67 75 79 1962-63 4 4i 64 83 = = T == = 5 = = 0 93 -- -- T 44 44 86 1963-64 5 69 82 -- 13 -- 79 = »=J 2 ap a = = = = = =J = =• = = 4 69 83 = = 1960-64 23 81 86 81 ir as Er TT T 31 5 = =■ 25 73 77 86 11 62 78 82 Browse 1960-61 0 3 T 4 9 19 16 O 92 51 2 37 25 4 20 16 H 3 1961-62 0 11 31 12 T 66 18 25 100 =• = = = = = 4 — 22 T 16 22 24 ll 1962-63 0 O 34 4 15 0 — = =* 31 =- = 8.9 = =* 0 T == = = 9 T 53 3 1963-64 T 22 = == 0 — 16 = = 12 = = = = = = = = == 10 = = ■ 3 22 14 C= = 1960-64 T 6 ll 6 4^ COCO CO 10 6T 51 90 ““ 2 25 22 2 14 T 15: 17 '- 5 Forts I96O-6I 9T 5 3.11 91 20 ,8 20 T 18 91 0 0 ii' 78 : 20 7 '15 1961-62 29 8 c DE 35 3 T I T = = = = 8 — '3-11 40 12 I . 10 1962-63 96 59 2 13 84 92 -- = = 69 = = 6 CS) = 100 ' 90 56 3 10 1963-64 95 9 2 — 87 — 5 = = 86 = = = = = = =J = == 0 = = 93 9 - 3 = = 1960^64 TT 13 2 13 74 22 6 13 32 .18 6 ™ = T3 .2 1 11 ■75 • 23 5 13 & ■ H l/ Vegetation Types I . Artemi8la/Agropyronj Pirius/Juniperusj Pseudotsuga/Juniperusj and Sareobatus/ArtertNsTa. 2/ Summer, fall, winter, and spring respectively. 3/ T = less than one per cent. TABLE XXXVI. A COMPARISON OF RELATIVE INTENSITIES OF ELK USE OF SOME IMPORTANT FOOD PLANTS DURING SEASONS AND YEARS, 1960-64. 1/ ' Taxa 1960-61 1961-62 1 9 6 2 - 6 3 . 1963-64 j? I 1S 1 U y Su/Fa/¥r/Sp Su/Fa/Wr/Sp Su/Fa/Wr/Sp Melilotus officinalis 6 5/ 6/ 0 / T 9/ T/ 0/ T 5 6/56/ T/ 5 8 5/ 8/ l/__ Prunus virginiana ilh Eh nh l 7/ 0/ 0 / O 3/ 0 / 0 / O 0 / 0 / 0 /-- Symphoricarpos spp. 5/ 6/ 2/ 0 6/ T/ 0/ O .3/ 0/ 0/ O Eh lh lhp Agropyron smithii T/4 0/6 5/68 32/24/58/41 Eh 1/26/32 1/67/77/- Poa secunda 4/ 6/ 9/12 Eh24/ 2/11 T/33/ 4/33 T/ I/ I/- Artemisia tridentata 0/ 4h Eh n 0/ 2/1 7/ 8 0/ 0/34/ T - . 0/22/ 8/__ Number of taxa used 31/32/2 1 / 3 9 36/2 5/1 7 / 1 9 . . 39/14/1 5/ 33/ 7/20/-- l/ Percentages, of total .instances, of use on;all' vegetation types' combined. ';-'y 2/ Summer, fallj winter, and spring respectively.. ‘ . . . 7b' “ 163 — The decreased availability of forbs during I960 and 1961 was reflected in increased use of grasses and/or browse, by elk. The most important difference occurred during the summer of 1961 when use of Melilotus officinalis and other forbs was lowest„ Grass5 especially Agropyron Smithii5 received intensive use and was the principal source of elk forage„ Shrubs appeared to be used more intensively during summer in i960 and 1961 as compared to 1962 and 1963= Greater use of browse occurred during the summer of I960 when decreased use of forbs was comple­ mented by increased use on shrubs, rather than grass. During the summer of 1961, elk browsed intensively in June but only rarely in late summer when the most intensive use of shrubs occurred in other years. The extreme abundance of forbs during the summer and fall of 1962 as well as the spring and summer of 1963 was reflected in increased use of this forage class5 especially Melilotus officinalis, and decreased use of browse as compared to earlier years. Use of grass5 as well as browse., declined during the fall of 1962 coincident to intensive use of M, officinalis, Some yearly variation in fall food habits of elk was also influenced by the availability of "green" grass5 especially Foa secunda, Use of this taxon increased sharply and use on Agropyron smithii declined following "green-ups” during October in 1961 and 1962, The decreased use of grasS5 especially A= smithii, and intensive use of browse5 particularly Artemisia tridentata, during the winter of 1962-63 was influenced by persistent snow cover and sold temperatures during January and early February. Cattle A total of 184 cattle feeding sites was examined between April 1961 and January 1964# mostly during the April-Noyember grazing season. During winterp feeding sites were examined only when cattle were ranging freely on upland pastures and using only range forage. The number of instances of plant use recorded at each site varied from 60 to 780 but typically was between 200, and 300 (mean 237)» The total number for all sites was 43#655 = Data for use of taxa which individually received at least ten per cent of the total instances of plant use for at least one vegetation type dur­ ing at least one season are presented in Table XXXJII. Taxa which received less intensive use are considered in Appendix Ta,ble LXI. Spring (Aprils May) .-Thirty-seven cattle feeding sites were examined during spring after cattle were turned onto open range. This total included 10 in 1961# 12 in 1962# and 15 in 1963= Use ,was recorded on 48 taxa; 28 forbs# 6 shrubs# and 14 grass and grass-like plants. Ten of ■ these# five forbs and five grasses# received an average of more than one per cent of the total instances of use during spring. Grass was the most important source of forage on all vegetation types# comprising 82 per cent of the total combined spring use. Agropyron smithh.# the predominant taxon# accounted for approximately two-thirds,of the use on grasses. It was used on all sites examined. Poa secunda# which ranked - 16$ - TABLE X X X H L . CATTLE U S E O F TAXA WHICH I N D l H D U A L L Y RECEIVED AT LEAST TEN ' PE R CENT O F THE TOTAL INSTANCES O F PLANT US E FOR AT LEAST ONE VEGETATION TYPE DURING AT !LEAST ONE SEASON, 1961-63 C O M B I N E D „ Vegetation Type: Plant Ta%a Artemisia/Agropyron: Agropyron smithii Mnhlenbergia cnspidata Poa secimda Grass Total Browse Total Melilotus officinalis Forb Total m u s / Jumiperus s Agropyron smithii Poa seeunda Stipa Yiridula Grass Total Browse Total Melilotiis officinalis eras: Stipa'' firidula ' Grass Total Browse Total Melilotus officinalis Forb Total Spring ■ Summer ; Fall Winter (17-32)^ (35r60) -- (22-31) (6-20) 100/53 2/ 100/35 100/47 100/55 5/2 65/15 61/5 ' 17/T 100/15 57/1 65/8 100/10 100/79 IOO/65 100/78 100/83 24/T 46/3 56/6 , 100/14 47/10 65/23 65/14 33/1 94/20 94/31 95/15 67/3 (9-24) (22-45) (8-24) (4-19) 100/45 100/37 100/57 100/32 44/1 45/3 75/3 75/33 77/17 23/T 38/T , 50/15 89/9 95/14 87/13 75/5 100/79 100/63 100/83 100/94 11/T 54/2 75/4 100/5 33/9 73/31 (4-19) 100/19 75/13 100/49 50/4 75/43 100/46 62/9 . 25/T Agropyron smithii Poa seeunda Grass Total Sarcobatus Termiculatus Browse total Melilotus officinalis Forb Total (7-20) 100/49 43/3 100/55 100/11 100/11 71/31 85/33 (7-25) 85/38 71/13 85/55 85/19 85/22 71/19 100/23 (4—12) 100/47 100/34 100/83 25/T 25/t 75/8 100/17 - 166 - TABLE XXXHI. Continued. Vegetation Type; Plant Taxa Spring Smuner Fall Winter Agropyron smithii Grass Total Browse Total Melilotus officinalis Forb Total Xanthium; Agropyron smithii Spartina peetinata Grass Total • Browse Total Chenopodimm album Melilotus officinalis Forb Total. ($-22) (11-35) X4-20) 1 00/# ; 10/72; : 100/81 100/88 ' 100/75 ■, 100/89 40/1 73/2 ' 75/2 60/7 72/15 100/5 80/11 100/22 100/9 (6-26) . (2-16) 100/33 100/35 50/10 50/2 100/51. ■ 100/46 5P/T ’■ • 50/T 33/3 - 100/15 100/38 100/31 100/49 '. 100/53 Artemisia longifo-lia; Agropyron smithii Calomovilfa longifplia> Grass Browse Total Agropyron/Poa; Agropyron smithii Hordemm .jubatum Poa secmda Stipa viridula Bouteloua gracilis Grass Total :Browse Total Melilotus officinalis Forb Total (2-7) (1-3) 100/10 .. ■ 100/82 ■ 100/91 100/8 ■(5-21)■ . (1-5) 100/57 100/51 100/33 30/11 100/13 80/T. •' 100/63 100/13 . 80/l0 ' 100/T 100/95 ' . 40/3 100/67 ' 100/99 100/2 100/4 40/1 100/24 100/31• ■ 100/T (1-10) 100/63 . 100/6 • 100/11 100/86 100/10 100/13 I/ In parenthesis; The number of feeding sites on this Vegetation=-The total number of taxa used on the type« 2/ Per cent frequency of use among all feeding sites on this vegetation type/aggregate mean percentage of the total instances.of use for this vegetation type. T = less than one per cent.. - 167 - secondj, was used at moderate intensities and on about 85 per cent of the siteso Other grasses received only minor use though some ^ especially Koleria cristata and Stipa viridula, were used relatively frequently. Forbs received about 18 per cent of the total spring use but increased in importance throughout the period. Melilotus-officinalis was the most intensively used forb, largely because of exceptional use during one year. All other taxa. received only very minor use. Allium textile9 Lomatium foeniculatum, and Microseris nutans were the most important individual taxa during April and early May. Tragopogon dubius and Vicia americana, in addition to Melilotus officinalis, were more intensively used in mid and late May. Shrubs were only rarely used by cattle during spring. Analyses of six cattle rumen samples (Table XXXVIII) ^ collected during the spring of 1963, showed similar taxa but slightly greater use of grass than feeding site data. Summer (June, July, August).--A total of 91 feeding sites was examined during the three summers; 23, 30, and 38 respectively for 1961, 1962, and 1963. Cattle use was recorded on 95 taxa, including.62 forbs, 10 shrubs, and 23 grass and grass-like plants. Eleven taxa, three forbs, one shrub, and seven grasses, received an average of one per cent or more of the total combined summer use. y- 168 - TABlE X H H I l o FREQUENCY OF OCCURRENCE AND VOEJME PERCENTAGES OF PLANT TAXA AND FOIiAGE GLASSES AMONG CATTLE RUMEN SAMPLES» Taxa Spring Summer Forage Class 1963 1962 1963 Total l/ (4) (8) (12) Cyperaceae family 17/T 2/ 38/T 25/T Gramineae family 100/90 100/92 100/71 100/78 Grass and Grass-like Total 100/90 100/92 100/71 100/78 Gamelim misrocarpa 25/T 8/T Gomndra umbellatum. 17/T 62/T 4 V T Compositae family 25/T 12/T 8/T Legummosae family 83/10 100/26 8/T MelilotUB officinalis 25/T 75/18 Thlaspi arvense 33/T 12/T 8/T Vicia amerieana ■ 12/T 8/T Unidentified forbs 33/T 75/6 38/T 50/2 Forb Total 100/10 75/6 100/27 92/20 Artemisia tridentata $0/T . 62/T 42/T Ghrvsothamnus viseidiflorus 12/T 8/T Prunus vireiniana1111 111. in *A* KTw A1M M tM Pinus ponderosa 33/T 12/T 50/T 8/T 33/T Rhus trilobata 33/T 25/T 62/T 50/T Ribes eereum IJ^/T 8/T Rosa sppo 33/T 25/T 38/T 25/T Symphoriearpos sppo Unidentified browse 17/T 25/T 38/T 25/T 100/T 75/1 100/T 92/T Browse Total 100/T 75/1 100/2 92/2 / l/ Number of rumen samples in parenthesis„ 2/ Per cent frequency of occurrence among rumen samples/average percentage of total volume for each sample„ — 169 “ Use of grass decreased, from spring., constituting about 64 per cent of the total summer use. Agropyron smithii remained the most important ■■■■-,taxonj, predominating in use for vegetation types that were used most intensively by cattle. It received about 4P per cent of the total use for all types combined and wa,s used on 99 per cent of ,all: feeding sites. . .Use of Muhlenbergia cuspidata, .which ranked second among.grasses and third over-alls was moderate on the Artemisia/Agropyron type.and only minor on the Pinus/Juniperus type. It was used on less than one-half of all cattle feeding sites. Stipa viridula received only minor use particularly on the intensively used Artemisia/Agropyron vegetation-type, but was used on. about two-thirds of the sites examined. Use was most intensive in late summer and on the Pinus/Juniperus tjpe. Other grassesindividuallys received only very minor use. Of these,, Bouteloua gracilis, which received minor use during early summer and moderate use in August on the Artemisia/Agropyron vegetation type, was of greatest. importance. Although grass was the most important forage throughout the period, forbs received moderate to relatively intensive use at feeding sites, especially during June and July. Melilotus officinalis was the most important forb and ranked second over-all for the three summers. It accounted for approximately 26 per cent of the total combined summer use. Use on other forbs was generally minor, but varied in relation to their abundance as well as the relative abundance and use of M. officinalis. Comandra umbellatum and Vicia americana were used most consistently and of some importance, especially in early summer. Tragopogon dubius was - 170 - Shrubs5 collectively as well as individuallys were of only minor importance. Sarcobatus vermiculatus, the most consistently used browse plant 5 received only one per cent of the total summer use. It was used largely on the Sarcobatus/Artemisia vegetation type. Rhus trilobate and Symphoricarpos sp. received relatively frequent3 though very minor5 use on the Pinus/Juniperus vegetation type. The contents of 12 cattle rumen samples^ collected during the summers of 1962 and 1963 (Table XXXVIII) were comprised of similar taxa but greater percentages of grass as compared'to feeding site data. i‘ t Fall (September, October, November) .--Data were obtained at 45 cattle feeding sites; 14 in 1961, 14 in 1962, and I? in 1963. Use was recorded on 60 taxa, including 31 forbs, 10 shrubs, and 19 grass and grass-like plants. Ten of these, one forb ,two shrubs, and seven grasses, individually received one per cent or more of the total instances of plant use during fall. Grass was by far the most important forage class. Use increased from summer, constituting approximately ?6 per cent of the combined, fall total. Agropyron smithii, the single, most important taxon, comprised about two- thirds of the total use on grasses. It was used on 98 per cent of all I feeding sites. Use on Poa secunda was minor over-all but was relatively intensive and important, especially in October, during years of fall 11 green-up = Muhlenbergia cuspidate, Bouteloua gracilis, Koleria -cristate. used frequently at feeding sites but constituted only a small percentage of the total use. -V - 171 - and Stlpa viridula received relatively minor# though frequents use at feeding sites, Mi cuspidata and S, viridula were used most intensively during September; B, gracilis was more important in November, Other grasses were of only minor importance throughout the period, Forbs ranked second throughout fall, largely-'because ■ of intensive use of Melilotus officinalis, the only important taxon, during 1962 and September 1963, Other taxa were used only occasionally or on minor vege­ tation types, Cattle use of shrubs increased from summer but remained of only minor importance. Browse constituted approximately 8 per cent of the total combined fall use and was used on 64 per cent of the feeding sites, Sarcobatus vermiculatus and Chrysothamnus nauseosus were the only taxa receiving intensive use. The latter was used most intensively during September, S, vermiculatus received'minor use throughout the period, Symphoricarpos spp, received minor use on the Pinus/juniperus and Agropyron/Symphoricarpos vegetation types, • Winter (December, January)■,— Eleven cattle feeding sites were examined during early winter; six in 1961 - 62 , three in 1962-63, and two in 1963 -64 , Use was recorded on 27 taxa, including 9 forbs, 9 shrubs, and 9 grass and ' I . grass-like plants. Twelve of these, I forb, 3 shrubs, and 8 grasses, received one per cent or more of the total instances of use. Grass,the predominant forage, constituted about 87 per cent of the total winter use, Agropyron smithii, the most intensively used taxon. - 172 - received nearly 50 per cent of the combined total use= Agropyron spicatum ranked second ^ but was used intensively only on the Pinus/Juniperus vegetation type= Its.greatest use occurred during periods of complete snow cover in January= Poa secunda received-moderate- use9 mostly during. early winter in years marked by fall 11 green-up.51 -Bouteloua gracilis „ Koleria Cristata5 and Stipa viridula received,minor, but frequent, use throughout the period= . Shrubs ranked second, receiving about-10 per cent of the total winter use. Artemisia tridentata, which was used bn 46 per cent.of all sites, Chrysothamnus nauseosus, and Sarcobatus vermieulatus were the most important browse plants, but individually received only very minor use. Use of forbs was very minor during winter. Most of that recorded was on Melilotus officinalis. Yearlong Trends.— Grass comprised the most important forage for cattle throughout the year, constituting approximately 76 per cent of the total instances of plant use averaged over months for the three-year period. Use of grass was most intensive during April, decreased gradually to the yearly minimum during June, then increased consistently to near maximum levels by November. Intensities of grass use on winter pastures were similar to those of November and/or April. Trends.in use of forbs.by cattle complemented changes in use of grass. . This forage class comprised about 18 per cent of the total instances of use on feeding sites averaged for months. Maximum use of forbs occurred during June; the minimum in ■ • . j- . - 1?3 - D ecember<, Browse was of only minor importance at all times. Use on shrubs constituted approximately five per cent of the mean monthly instances of plant use. Relative intensities of use of this forage class were lowest in April and increased gradually through summer and early fall to a maxi­ m u m of ten per cent of the total instances of use for October, Browse use on winter pastures was similar to that in October, Agropyron smithii was the single, most important taxon in relative use on cattle feeding sites throughout the year. Trends in use of A. s mithii, which comprised between one-half and two-thirds of the use on grass and grass-like plants during all periods, paralleled those described for the forage class, Poa secunda ranked second in use on feeding sites. - I : '' ' during spring. Subsequently, it received only occasional use during . summer and early fall. Fall use varied from very minor to intensive in relation to fall precipitation and "green-up." Other grasses were of only minor importance during most periods. Those which were frequently used on feeding sites inc l u d e d : Stipa viridula which received more intensive use in late spring as well as late summer and early fall; Muhlenbergia cuspidata which was of some importance only during summer and i early fall; Bouteloua gracilis, which received minor use throughout but was of greater importance in late' summer; and Koleria cristata which received minor use during most periods. Agropyron spicatum received important use only on winter pastures. Us e of all other grasses was always very minor and/or occurred largely on minor vegetation types. - 1 7 4 - The decline in relative use of grasses in late April was complemented by increased use of early forbs, especially Allium t e x t i l e , ' Lomatium foeniculatum, and Microseris nutans of the Poa secunda union in late April and early M a y and Comandra umbellatum» Vlcia americana« and Melilotus officinalis of the Tragopogon dubius union in late M a y and early' June. M. officinalis was the single, most important forb used by cattle i from June through September as well as later in the fall and winter when first-year growth and/or dry stems persisted. Use of other forbs in summer, fall, and winter was only very minor and/or occurred only on minor vegetation t y p e s . Sarcobatus vermiculatus« the most important shrub, received only minor use throughout most of the year. Rhus trilobata, occasionally received minor use on the Pinus/Juniperus vegetation type during dummer. Svmphoricarpos spp*, .was frequently used at feeding sites on t h e .Pimis/ Juniperus and Agropyron/Symphoricarpos types during summer and.,, fall,but constituted only a minor, percentage of the total use. U s e on Chrysothamnus nauseosus occurred after flowering in August, increased to some importance during early fall, then generally declined. Minor use of this taxon also occurred on winter pa s t u r e s . Artemisia tridentata received minor use on winter pastures, mostly during periods of complete snow cover. Year-to-Year Trends .— Cattle food habits were less affected by. yearly differences in precipitation and vegetal growth than those of mule deer U ■ — 175 — or elk but somfe important variation in use of forage classes (Table XXXIX) and individual taxa (Table XL) was evident. Grass use was consistently high at cattle feeding sites throughout 1961 and the spring of 1962 when plant growth was minimal. Subsequent increased growth and availability of forbs ^ especially Melilotus officinalis, during the summer of 1962 was' directly reflected in greater representation of this forage class as well as $£■, officinalis in recorded use for the summer and fall of that year. Further increases in abundance during 1963 influenced more intensive use of forbs during spring* and especially* summer when forbs comprised the most important forage class and JJ. officinalis was the single* most important taxon at cattle feeding sites. Comparisons of mean percentages of total forbs and M1, officinalis in rumen samples for 1962 and 1963 (Table XXXVIII) show increased use of these items during 1963, The greater percentages for grass; than for forbs in the.samples for the summer of 1963 was in contrast to feeding site data. The maximum percentage of forbs in a single sample was 82 per cent but the percentage in most was between 10 and 15 per cent. This suggested that the relative use and importance of forbs may have been unduly exaggerated at feeding sites. Although differences were not extreme* higher percentages of use of browse at feeding sites during 1961 as compared with 1962 and 1963 suggested slightly greater use of browse by cattle during periods of poor range-forage conditions.. TABLE XXXIX. A COMPARISON OF PERCENTAGES OF CATTLE USE OF GRASS, BROWSE, AND FORBS AT FEEDING SITES ON VARIOUS VEGETATION TYPES DURING SEASONS AND YEARS, I96O-63. Forage KIEOfhKqIbT ih TDYLShALYDT i/ SKIgbhKIEOf ih All Types Combined Class Sp Su Fa 2/ Sp Su Fa Sp Su Fa Sp Su Fa Wi Grass 1961 90 86 97 88 92 93 70 35 89 -87 81 90 1962 85 70 65 86 65 70 77 61 80 86 69 70 78 1963 ' 64 45 85 62 42 89 88 38 62 73 46 80 90 1961-63 79 66 78-. 79 62 84 82 55 56 82 64 76 87 Browse 1961 Enh s T O 5 5 30 39 - 6 6 11 9 1962 .1 3 4 O I 2 O 10 T T 4 2 16 1963 . -O T 4 • T 2 7 I .5 18 T I 6 2 1961-63 T 3 6 T 3 4 T 12 22 T ' 3 7 10 Forbs 1961 • 10 .9 2 12 3 2. O 26 ■ 10 8 . T 1962 14' 26. .31 ' 14 34 28. 23. 29 20- . ' -13 .27:0 2 8 ' 6 1963 36 54. 11 - 38 56 -4 '- 11 57' 20 . - 26 '...5 2 0 )14' 3 3 / 1 7 : 8 1961-63 - 20 31. 16 21 35 12 17 33 22 18 : 3 . ■ . '■ -IU ■ - l/ Vegetation Tyrpes: Artemisia/Agropyronj Pirns/JuniperusJ and Sarcobatus/'Artemisia. 2/ Spring, summer, arid fall'respectively. ■ . ' ■ 3/ T = less than one per cent. - 1 7 7 - TABLE XL. A COMPARISON'OF RELATIVE INTENSITIES OF CATTLE USE OF SOME IMPORTANT FOOD PLANTS DURING SEASONS AND.TEARS, 1961-63. I/ Taxa 1961 1962 ■ 1 9 6 3 Sp/Su/Fa Sp/Su/Fa Sp/Su/Fa Melilotus officinalis T/ T/ 5 ,: 2 / 1 ? / # : 19/48/11 Agropyron smithii 5 8 / 4 7 / 4 5 = 6 2 / 4 8 / 4 1 - . 4 8 / 3 0 / 6 1 Poa secunda 16/ T/10 . " 14/ 2 / 1 4 18/ T/ I Number of taxa used 2 3 / 4 4 / 3 0 3 2 / 5 9 / 3 3 3 7 / 6 9 / 4 5 l/ Percentages of total instances of use on all vegetation types combined. The influence of fall precipitation and 11 green-tip11 on cattle use of ' • - 7. ' individual grasses was indicated by relatively intensive use of Poasecunda when "green" during fall periods in 1961 and 1962 as compared to only very minor use of this taxon plus increased use of Agropyron smithil during the fall of 1963 when "green-up" did not occur. RANGE RELATIONS Interspecific relations between grazing animals _may connote "competition" in which the welfare of one or more of the animals is adver­ sely affected. The extent to which two species graze the same area and prefer the same forage was considered by Inlander (1958) as the main factor controlling interspecific competition. Cole (1958a). listed-similar criteria, but included requirements that forage plants be in limited supply or deteriorating as a result of combined use. Smith and Inlander (1953) differentiated the latter as "forage" competition as opposed to "land use" competition which they associated with common use of a forage species which was adequately abundant to meet the requirements of both animals, but where removal of one would permit increased'numbers of the other. Cole (op.eit.) recognized circumstances under which certain big game species may benefit from heavy livestock grazing. Data from the present study were evaluated with respect to these considerations which require comparisons of the distributions of animal use on the area in time and space in relation to vegetation types, topo­ graphy, and water sources, as well as forage preferences and the composi­ tion, abundance, and intensity of use of forage plants. Elk and Mule Deer Relations Mule deer and elk were most similar in habits between April and September. The distribution of elk during this period overlapped the dis­ tribution of greatest mule deer densities. Both animals used the Artemisia/Agropyron vegetation type, especially the Artemisia tridentata/ *' ■ *''V Agropyrdn smithii/Bouteloua gracilis association., intensively during early spring. After Inid-May5, mule deer use became concentrated on timbered vegetation types as did some significant summer elk use. Some forbs and Poa seeunda were used by both animals during April and May5, but grass, primarily Agropyron smithii, was the most important elk forage, Forbs. became important for elk as well as mule deer in late spring and summer when similar taxa were used by both, Melilotus officinalis was one. of the most important mule deer forage sources and the single, most important elk forage from June through early .September, Browse, a very important summer forage for mule deer, ,was of only moderate importance in the . >( summer diet of elk, Rhus trIlobata8- the most important mule deer summer browse plant, received only minor use by elk, PrUnus virginiana9 the most important summer brows© for elk, received relatively minor use by mule deer. Browse became more important for mule deer during September when grass was used increasingly by elk, Chrysothamnus nauseosus„ used inten­ sively by mule deer, was of little importance to elk. Dissimilarities in range use and food habits of mule deer and elk during late fall and winter afforded less opportunity for association and conflict. Grass, the most important elk forage after September, received little use by mule deer. During winter, the Artemisia/Agropyron vegetation type was used intensively and the Pinus/Junlperus and Sarcobatus/Artemisia types received moderate use by both animals, but food habits differed. Grass was the major item in the elk diet and browse was the most important food for mule deer, Artemisia tridentata, the most important winter deer - 179 - — 180 = browse under prevailing conditions, received only minor use by elk as did Juniperus seopulorum and Symphoriearpos spp» which received moderate or minor use by mule deer. Although these results indicate probable conflicts between mule deer and elk, complete evaluation of the interaction requires consideration of forage conditions as well as other range use factors. Numbers, of elk on the study area apparently were not excessive with respect to forage, as indicated by the average ratio of 57 calves per 100 females for the four-year period. Mule deer probably contended with forage deficiencies and intra­ specific competition during most years as a result of excessive utiliza­ tion of forage. Measurements of nine utilization and condition trend sample units using methods described by Cole (1958b) showed that an average of 58 per cent (40 per cent to 81 per cent yearly) of the available "leaders" on Rhus trilobata was used annually during the four-year period. Relatively low levels of "leader use" during 1962-63 and 1963-64, 40 per cent and 53 per cent respectively, reflected "buffering" by the unusual abundance and use of fprbs during those years. Despite this,. more than 80 per cent of all plants of R= trilobata were "severely hedged." More than 95 per cent of the plants were "decadent" and/or reduced to "resprouts." The annual average percentage of leader use on Chrysothamnus nauseosus for six sample units was 85 per cent (77 per cent to 89 per cent yearly). More than 95 per cent of the plants were "decadent" by the spring of 1964° Leader use on Chrysothamnus Viscidlfkms - 181 - averaged more than 80 per cent for the four years on one upland site. The average percentage of leaders used on Artemisia, tridentata on five sample - units was 24 per cent (8 per cent to 32 per cent yearly) for the four-year periods bat utilization of this extremely abundant plant varied in relation to the availability of other forage. Prunus virginiana was reduced to clumps of resprouts and plants taller than six feet on which'browse was largely unavailable to mule deer. The extreme utilization and deteriorated condition of important summer-fall forage plants indicated that summer and fall range condition was equally as important as winter range to the wel- : fare of mule deer. The condition of the winter range was directly related to the intensity of summer-fall use of certain "key" species. Additional utilization of any of the more important summer and fall browse plants by . other animals would have adverse effect on the mule deer forage supply and population. \ The home range behavior and restricted movement of mule deer may also have affected interspecific relations. In contrast to mule deer=, elk moved extensively in response to changes in availability of forage during all season's and thus were capable of substantial impact on forage supplies of individual deer. It seems apparent that direct competition of elk with mule deer must have occurred during the April-September periods especially early spring,, of most years. In comparison with the April-September period=, conflict between elk and mule deer during late fall and winter would be of minor “ 182 — The evidence indicated that elk were more "efficient11 competitors than mule deer. Elk intensively used most vegetation types ^ topographic sites, as well as all classes of forage during the year and/or as necessary under the various conditions which prevailed during the four-year period. Mule deer consistently used a limited number of vegetation types and were strongly restricted in their diet of browse and forbs, except for use of a single grass species when "green" and when other forage was limited. importance related to occasional elk use of mule deer browse plantss other than Artemisia trIdentata0 on local areas» . Elk and Cattle Relations The distributions of intensive elk and cattle use overlapped through­ out the April-November grazing season. The two animals were extremely similar in habits during spring and fall but generally dissimilar through summer. Elk habits during winter were alike those of cattle but the distribution of elk was restricted to the southeastern and/or other portions of the study area where only occasional cattle' use had occurred previously. Both animals used grass intensively and in about the same proportion during spring. Use on Agropyron smithii, the most important forage source for both, was also very similar. The Artemisia/Agropyron vegetation type was used most intensively by elk as well as’ cattle. Habits diverged after late May when forbs generally became the most impor­ tant source of elk forage and the timbered vegetation types received significant elk use. Although some forbs were commonly used by both animals ^ elk typically utilized•only minor amounts;of grass which continued as the most important cattle forage,.. Habits of the two species became increasingly similar during September and were closely comparable during October and November, These data indicated opportunity for direct ..and intensive interaction X .' . ■■'. between elk and cattle during early spring and fall, especially on the Artemisia tridentata/Agropyron smithii/Bouteloua gracilis association, in early spring. Conflict of a less direct nature could.result because of intensive use of Agropyroh smithii, the favored winter food of :both_ animals, by cattle during .the grazing season, and by elk in fall on.live­ stock wintering areas. Intensive use of Muhlenbergia euspidata,. • Poa secunda, and Koleria cristata by elk under various, conditions., somewhat lessened the conflict on cattle pastures. Actual competition was,also . limited by the low numbers, distribution, and movements of elk, .Elk.pre- c^ferred areas which had received little, if any, prior use by cat tie...during the same year and moved extensively in response to. changes in forage availability and/or the distribution of domestic livestock grazing. They, readily used steeper and"the timbered vegetation types in deference to d ridgetops and bottomlands which had received recent use by livestock. The opportunity for elk-cattle interaction increased significantly ■■ . : ; ■ ■ ■ ■ ;• , v . :v <■ during periods of severely reduced forb growth when elk ..intensively utilized grass, especially Agropyron smithii. during summer as well as - 133 - . other seasons. Male Deer and Cattle Relations The overlap between areas of intensive cattle, grazing and greater r male deer densities indicated opportunity for interactions bat food and range-use habits differed substantially, . -Grass comprised the major portion of the cattle diets whereas browse and/or forbs were the most important foods of male deer. More extensivea level or gently rolling ridgetop and coulee-bottom areas were used most intensively by cattle. Mule deer range use was largely on the timbered vegetation types on steeper slopes. The greatest overlap was in April and early May on the Artemisia/Agropyron vegetation types especially on the Artemisia tridentata/Agropyron smithii/ Bouteloua gracilis association. Poa seconda and several forbsa the occurrence of which was probably related to heavy grazing by cattle in previous years a were commonly used by both animals 9 but Agropyron smithii. the most important source of cattle forage was used only occasionally by mule deer, Forbs received moderate use by cattle. Little overlap in food habits occurred in summer. The degree of overlap increased after late summer when cattle use on the Pinus/juniperus type increased slightIya possibly because of intensive grass utilization on primary range sites during spring and summer. Mule deer use on the Artemisia/Agropyron, type became increasingly intensive during fall. Common use occurred on Chrysothamnus nauseosus9 an important browse plant for mule deer. Moderate use of this plant by cattle may have increased its palatability for mule deer because of the latterts preference for clipped stems. The opportunity for association and conflict between male deer and cattle during winter was limited by restriction of cattle to pastures where only minor use of browse occurred. CONCLUSIONS AND RECOMMENDATIONS FOR MANAGEMENT Rhus trilobata and Chrysothamnus nausebsus were "key" mule deer, forage species« Agropyron smithii was a "key" species for both elk and cattle„ The abundance ^ intensity of Uses and condition of these plants on "key" range sites should serve as bases for evaluating range conditions„ Melilotus officinalis was extremely important to mule deer and elk« Its heavy use during years of abundance relieved the drain on key species» This "exotic" apparently has become established as a part of the vegeta­ tion of the Missouri River Breaks but because of extreme fluctuations in abundance should not be considered as a basic forage source for management„ "Key" range areas for mule deer were the Pinus/Juniperus vegetation type on moderate to steep slopes and the Artemisia/Agropyron type on smaller ridgetops and along margins of more extensive ridges, . Small to moderately extensive ridgetops dominated by Artemisia/Agropyron in areas of "light" cattle use appeared to be "key" areas for elk, "Primary" range areas for cattle were extensives unbroken ridgetops as well as coulee- bottoms within about one mile of a dependable water source. Mule deer should be- more'rigidly controlled by adequate hunter har­ vests and maintained a t ■levels which will permit improvement and replace­ ment of deteriorated browse plants as well as sustain deer populations, Greater than existing numbers of elk will not be. compatible with management to sustain mule deer range and populations. Whether existing numbers are compatible is not clear. The relative importance of elk with respect to mule deer as well as other game species and land uses must be - 187 - .evaluated and established as a basis for management„ If mule, deer are . deemed the most important species, elk should not.be permitted to increase beyond existing numbers or be. reduced in number from existing population levels o The potential impact of elk range use on the biota of. the Missouri River Breaks should be recognized arid -given foremost consideration in the management of existing game populations, in introduction "of new animal species to the area, and in relation to introduction of elk into other areas along the Missouri River or similar habitat in eastern Montana where "multiple-use" is the object of management, In multiple-use management programs, livestock numbers and management should be considered on the basis of the forage available, with "proper" utilization of Agropyron smithii, on "primary” range areas rather than..by "type" or the entire area. Winter grazing should be restricted from areas where smaller ridges and timbered coulees are closely interspersed. Big game-live stock interactions were most intensive during April and early May. Consideration should be given to changing the date of onset of livestock grazing to mid-May when plant growth is more advanced. This would eliminate conflict during early spring and reduce possible competi­ tion during subsequent periods. The development and control of water sources in association with "primary" cattle range areas could effect better cattle distribution and lessen opportunities for overlapping range use and conflict with game. Water development should not be considered for terminal portions of larger ■ X ■ --r - 188 - ridges or on smaller ridges where the area available for cattle dispersal on “primary*1 range types is limited. Allowable utilization with respect to forage production and repro­ duction of''“key" browse and grass species should be evaluated experimentally Experiments should also be designed and conducted to ascertain the time and v conditions required to restore “range condition,“ especially browse in areas where existing plants are in deteriorated condition, reproduction is lacking, and/or previously existing plants have been removed by.excessive use. Until such time as more quantitative criteria are established, usually accepted criteria for “proper" use should be rigidly enforced. SUMMARY A study of the food habits s range use and relations of mule deer ^ Blkjl and cattle was conducted during 1960=64 on a YSsOOO acres area in the Missouri River Breaks in northcentral Montana to provide criteria for management on ponderosa pine/juniper and related vegetation types. Physiographic ^ climatic,, and land-use characteristics of the area were described. Canopy-coverage analyses of three stands of each of 12 rangeland communities provided data on vegetal characteristics of the Artemisia/ Agropyron vegetation type ^ including the A. tridentata/A. spieatum assoc­ iation, the A. tridentata/A o smithii associes, and the A. tridentata/ A. smithii/Bouteloua gracilis association; the Agropyron/Poa type (A. smithii/P . secunda associes); the Pinus/Juniperus type, including the ' Po ponderosa/A. spieatum association, the P . ponderosa/j . scopulorum association, and the P. ponderosa/Artemisia, longifolia association; the Pseudotsuga/Juniperus type (P. menaiesii/j. scopulorum association); the Sarcobatus/Artemisia type (S. vermieulatus/A-. tridentata association); the Agropyron/Symphoricarpos type (A. smithii/S. occidentalis association); the Xanthium strumarium type (association); and the Artemisia longifolia type (associes). Observations totaling approximately 11,600 for mule deer, 3$500 for elk, and 25,000 for cattle provided information- on distributions, movements, population trends, and range use. - 190 - Mule deer were distributed over the entire area„ Variation in sea­ sonal distribution reflected differences in food and range-use habits in relation to availability of preferred forage. Seventy-six relocations of 10 recognizable mule deer over periods of I day to 29 months indicated that mule deer movement was largely restricted to,relatively small home ranges. Relocations of two marked females over periods of 29 months indi­ cated movement within approximately 300 acres. Relocations of three males to 16s 20=, and 21 months of age suggested that males moved more extensively after their second summer. Mule deer population estimates indicated a decline in numbers during 1960-62 and recovery.during 1963-64$ and suggested a mean winter density of about 10 mule deer per square mile for the 1960-64 period. Comparisons of trends in estimated mule deer numbers with trends in numbers of mule deer observed per trip along vehicle routes indicated that the latter reflected changes in "observability" rather than numbers of animals on the area. Elk were generally distributed on the south and west-central portion of the area during summer and the southeast portion during winter. Local distributions varied 'in relation to forage conditions. Sixty-eight reloca­ tions of 13 recognisable elk within periods of I day to 4 years showed average movement of 3°4 miles between relocations| the maximum was 14°5 miles. The extreme mobility enabled elk to adjust rapidly to environ­ mental conditions and utilise the most favorable portions of the range. Minimum counts of elk on the study area during winters were 70 in 1960-61=, 80 in 1961-62$ 108 in 1962-63$ and 108 in 1963-64. - 191 - Greatest cattle densities occurred on the southern one-half of the area during the April-November grazing season. Use of local areas varied in relation to forage availability, water supplies, and weather conditions. Winter grazing was restricted to pastures on the northern and southcentral portions of the area. Numbers of cattle on the area varied considerably within and between years. Approximately 90 per cent of the total numbers of mule deer, elk, and cattle observed were recorded within three and one-half hours after sunrise and before sunset. Intensive activity was concentrated in the first and last hours of daylight. The “prolonged11 activity periods of late winter and early spring were characterized by persistence of activity through two and one-half or three and one-half hours after sunrise and before sunset and some activity in mid-day. Approximately 54, 70, and 87 per cent respectively of all observed mule deer, elk, and cattle were feeding. Feeding was the predominant activity among mule' deer observed during all hours as well as. among elk and cattle observed within three and one-half hours after sunrise and before sunset. Decreased proportions of mule deer observed feeding through the four years were indicative of increased wariness. ■Mean numbers of animals observed per group were 3.0 mule.deer, 6.9 elk, and 6=7 cattle. Average group sizes varied in relation to intensities of use of the Artemisia/Agropyron vegetation type and/or certain range sites. Variation in numbers of mule deer in groups also reflected the . - 192 - locations within home ranges of individuals on overlapping home ranges„ More than 47 per cent of the mule deer observed during summer were on the Pinus/Juniperus vegetation type. Use on this as well as the Pseudotsuga/Juniperus type ^which was indicated as second in importance^ declined after early fall concomitant with increased; use on Artemisia/ Agropyrono The latter was shown to be the most important type for mule deer use from early December through mid-Mayg receiving about 6? and ?6 per cent respectively of total use for winter and spring. The low "observability*1 of mule deer during summer and fall indicated greater than observed use of timbered vegetation types during those periods. Intensi­ ties of mule deer use on the various vegetation types were shown to vary between years in relation to influences of precipitation and/or mule deer populations on abundance of certain forage plants. During summers elk use was predominantly on coulee-bottom and timbered vegetation types with more than 50 per cent of the total feeding elk observed on the Sareobatus/Artemisla« Agropyron/Symphoricarpos, and Xanthium types collectively. The Pseudotsuga/Juniperus type was especially important for calving during June and for bedding, throughout summer. After September^ use on these types declined concomitant to increased use on the Artemisia/Agropyron type. The Iatter5 which predominated elk use from October through May5 received nearly 50 per cent of elk use for feeding during fall and winter and Si per cent of that for spring. The Pinus/ Juniperus type was indicated as moderately important in fall and received more than one-third of the total use for winter when open and semi-open slopes were used intensively. The Sarcobatus/Artemisia type received some important use on slopes in winter and on coulee bottoms in early spring. The Agropyron/Poa type received important elk. usage during October and November in years of fall "green-up." The Artemisia/Agropyron type received intensive elk use during all season's under drought conditions which prevailed from I960 through the spring of 1962. Approximately 60 to 70 per cent of the total cattle observed during all seasons were on the Artemisla./Agropyron type. The Sarcobatus/Artemisia and Agropyron/Symphoricarpos types in coulee bottoms collectively received moderate cattle usage and comprised the second, most important area for cattle use during the grazing season. The Pinus/Juniperus type was shown to receive minor use by cattle during winter and spring and moderate usage in summer and fall. Some yearly differences in seasonal use of vegetation types occurred in relation to differences in precipitation and its influence on plant growth and water supplies. Use of slopes of various degrees of steepness- closely reflected animal use of vegetation types. Approximately 50, 42, and 18 per cent respectively of the total mule deer, elk, arid cattle were observed on slopes steeper than 11 degrees. Percentages of total mule deer, elk, and cattle observed on slopes of various exposures appeared to reflect relative amounts of area with each exposure more strongly than animal preferences. Distances at which mule deer and elk were observed from water were largely related to the distribution of water sources with respect to areas - 194 - used by the animals during different seasons and years„ The distribution of water sources <, even during the most arid years, was not a significant factor in determining either mule deer distributions-or the distribution of elk within seasonal ranges» Consistent observations of most cattle within one-half mile of water indicated that the distribution of cattle use on the area was closely related to the distribution of water. Data from 263 mule deer feeding site.examinations showed that forbs were the most important forage during summer. Use of forbs constituted 56 per cent of the total combined instances of plant use in summer, 23 per cent during fall, 10 per cent during winter, and 35 per cent during spring. Browse constituted 43 per cent of.the total use for summer, and 72, 88, and 52 per cent respectively of the total use for fall, winter, and spring when shrubs, were the most important forage source. Use of grass was minor, amounting to a trace, 5, 2, and 13 per cent of respective seasonal use. Melilotus officinalis was the principal summer forb. ■ Rhus trilobata, Chrysothamnus nauseosus. and Artemisia tridentata respec­ tively were indicated as the most important browse plants for summer-early fall, late fall-early winter, and late winter-early spring. Poa Secunda9 when green, was the only grass to receive significant use by mule deer during any season. Significant changes in the diet of mule deer occurred during September, late November or early December, late March or early April, and late May. Use of forage classes and individual taza varied significantly between years in relation to precipitation and its influence on plant growth. Intensive and unseasonally early use of browse during I960 and 1961 was associated with decreased availability of forbs„ Inten­ sive use of Helilotus officinalis in years of abundance' “buffered" use on important browse species= Examination of I?S elk feeding sites showed /that forbs were the most important forage during summer. This forage class constituted 75 per cent of the total combined instances of plant use in summer, 23 per cent in fall, 5 per cent in winter, and 13 per cent in spring.' Grass use was minor during summer, comprising 11 per cent of the total, but was most important during fall, winter and spring when it comprised 62, 78, and 82 per cent respectively of the total seasonal use. Browse amounted to 14, 15, 17, and 5 per cent of the summer, fall, winter, and spring use respectively. Melilotus officinalis was the most important summer"forags. Agropyron smithii was the single, most important taxon in use from October through May. Significant yearlong changes in the diet of elk occurred during September and late May or early June. Agropyron smithii was used intensively throughout the summer of 1961 and browse received increased summer use during I960 as well as 1961 when forb growth was minimal. Examination of 184 cattle feeding sites showed.that grass was the ■ predominant forage throughout the year, constituting 82,. 64, 76., and 87 per cent of the total instances of plant use for spring, summer, fall, and winter respectively. Forbs comprised 18, 33, 17, and 3 per cent respec­ tively of total use during these seasons. Browse was a minor item in the - 195 - diet of cattle at all times, amounting to a,trace, 3, T3. and 10 per cent of the total use for respective seasons. Agropyron smithii was the predominantly used taxon throughout the year. . Comparisons of food and range-use habits and behavior of mule deer and elk indicated conflict between April and September, especially during April and early May. Heavy utilization and deteriorated condition of important mule deer browse plants aggravated conflict/occurring after midi- May. The evidence indicated that elk were more efficient competitors than mule deer. Considerable overlap in habits of elk and cattle afforded opportunity ■ for direct conflict during early spring and fall as well as less direct conflict resulting from use of elk wintering areas in summer■by cattle and use of cattle wintering areas by elk in fall. Competition was less­ ened by low numbers, distribution, and movements of elk. The potential for competition between mule deer arid battle was limited by broad differences in food and range-use habits. The greatest opportun­ ities for conflict occurred on the Artemisia/Agropyron vegetation type in April and early May and during late summer and fall. Overlap in range use increased during the latter period and common use on Chrysothamnus naissosus occurred on the Artemisia/Agropyron type. Abundance, intensity of use, and condition of "key ^ species Rhus trilobata, Chrysothamnus nauseosus, and Agropyron smithii on "key" range sites should serve as criteria for evaluating.range conditions as 197 well as regulating animal numbers in mule deer, elk, arid '.'multiple-use*' ■ ■ , ; . . . . ■ management programs. ...y . ;:v; V - 198 - APPENDIX rPA-RTl-F. XLI. CONSTANCY ^ CANOPY-COVERAGE ^ AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 67 PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN ONE PER CENT IN ANY COMMUNITY ON THE ARTEMISIA/AGROPYRON TYPE, l/ JL tridentata/ JL tridentata/ A. trldentata/ Combined Community Au, snicatum A. smithii A. smithii/ Type Association Associes .Bf gracilis Totals Association Union: Taxa Cy/Cv/Fr g/ Cy/Cv/Fr Cy/Cv/Fr Cy/Cv/Fr Agropyron spicatum: Carex filifolia Muhlenbergia cuspidata■ Artemisia tridentata: Aster canescens Aster occidentalis Astragalus bisulcatus Linum rigidum Orobanche ludoviciana Psoralea argophylla Tragopogon dubius: Bahia, oppositifolia ■ Collomia linearis Gaura coccinea Grindelia squarrosa Lactuca pulchella Phacelia linearis. Poa secunda: Astragalus agrestis Astragalus missouriensis Lepidium ■ dens if lorum Lomatium foeniculatum 33/ V + 33/ 33/ V 11/ +/ + V 3 5 22/ +/ 3 33/ +/ + 11/ +/ + 33/ + / + 11/ + / + 33/ +/ h 11/ +/ I 33/ V 6 11/ V 2 33/ + / 2 11/ +/ .+ 33/ V 2 : 11/ + / + 33/ V 9 33/ V . u / + / 3 33/ +/ + ' n / + / + 6 33/ + / 11/ V 2 33/ + / + 11/ .+/ + +■ 33/ +/ 11/ + / + 33/ + / 6 33/ V 8 2 33/ +/ 5 33/ + / 2 11/ V + 33/ +/ 2 11/ V + 33/ + / + 11/ V + 33/ + / 2 11/ + / + 1 9 9 TABLE XLI0 Continued. Community A. tridentata/ A. spicatum Association A. tridentata/ A. smithii Associes A. tridentata/ A. smithii/ B. gracilis Association Combined Type Totals Union: Taxa Cy/Cv/Fr 2/ Cy/Cv/Fr Cy/Cv/Fr Cy/Cv/Fr Sarcobatus vermiculatus: Sarcobatus vermiculatus 33/ +/ k 11/ +/ I Xanthium strumarium: Lactuca serriola Polygonum erectum ■ Iva axillaris CU + CO CO 33/ +/ 2 33/ +/ 2 33/ +/ + XL/ +/ + 33/ +/ + 11/ +/ + l/ Data are mean percentages for 120 two x five dm plots (40 in each of three stands) in each community„ i ?/ Constancy/canopy-coverage/frequency. +/+-.indicates canopy-coverage or frequency values less than one per cent. 200 — 201 — f TABLE XLIIo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY. OF LESS THAN 6? PER CENT OR MEAN CANOPY-COVERAGE OF LESS THAN•ONE PER CENT ON THE AGROPYRON SMITHIl/POA SECUNDA ASSOCIATION« l/ • ' Union: Taxa Cy/Cy/Fr 2/ Agropyron smithii: Bouteloua gracilis Koleria cristata Schedonnardus paniculatus 33/+/ + 33/ +/ 2 33/+/ + Artemisia t r i dentata: Artemisia tridentata Gutierrezia sarothrae Aster occidentalis Plantago spinulosa ■ ' ■ 33/ +/ + 33/ +/ 5 33/ +/ +■ . 33/ +/ 3 . Poa secunda: Mertensia lanceolata 33/ + / . 2 Tragopogon d u b i u s : Cirsium undulatum Tragopogon dubius 33/ t/ + 33/ +/ + Xanthium strumarium: Conringia orientalis 33/ t/ 3 l/ Data are me a n percentages for 120 two x five dm plots distributed as 40 in each of three stands. 2/ Constancy/canopy-coverage/frequency, 4-/+ indicates canopy-coverage or frequency values less than one per cento TABIxE XLIII.. CONSTANCY, CANOPY-COVERAGE AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS. THAN 6j PER CENT OR MEAN CANOFY-COVERAGE OF LESS THAN ONE PER. CENT ON THE PINUS/jUNIPERUS TYFEc-, l/ . .L ' . . Pinus ponderosa/ Finns ponderosa/ Pinus ponderosa/ Combined Association Agropyron spicatum J. Scopulorum Artemisia longifolia Type Total Association Association Association Union: Taxa Cy/Cv/Fr 2/ Cy/Cv/Fr Cy/Cv/Fr Cy/Cv/Fr Artemisia tridentata: Psoralea argophylla OJ'coCO + 'coCO Poa secunda: Astragalus agrestis 33/ +/ 3 33/ V 1 Poa secunda 33/ +/ 3 33/ +/ + 22/ +/ 1 Juniperus scopulorum: Danthonia unispicatum 33/ +/ 1 11/ +/ + Poa compressa 33/ +/ 2 11/ +/ + Tragopogon dubius: Collomia linearis 33/ +/ + 33/ +/ + 22/ +/ + Helianthus petiolaris 3/ +/ + 11/ v + Senecio cams 33/ +/ + 11/ +/ + Spha,era,Icea coccinea 33/ +/ 3 33/ +/ + 22/ +/ 1 Prunus virginiana: Rihes setosum 33/ +/ + 11/ +/ + Galium aparine 33/ +/ 5 11/ +/ 2 - Artemisia longifolia: Convulvulus sepium 33/ +/ +- • n / +/'+ ■ ‘ Yucca glauca ■ . 33/ +/ + 11/ +/ + Xanthium strumarium: Cirsium arvense 33/ +/ 2 11/ +/ + Polygonum sp, 33/ +/ 2 u / +/ + Salsola kali 33/ +/'2-- 11/ +/ + l/ Data are mean percentages for 120 two x five dm plots (40 in each of three stands) in each association. 2/ Constancy/canopy-coverage/frequency, + = canopy-coverage or frequency less than one per cent. 202 - 203 - TABLE XLIV' CONSTANCY, CANOPY-COVERAGE, AND- FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6 ? PER. CENT OR M E A N CANOPY-COVERAGE O F LESS THAN ONE P E R CENT ON THE PSEUDOTSUGA/JUNIPERUS TYPE, l/ Union: Taxa Cy/Cv/Fr 2/ Symphoricarpos occidentalis: Poa compressa 33/ +/ + Agropyron sm i t h i i : Stipa viridula 33/ +/ 2 Poa secunda: Descuriana pinnata 3 3 / V .5 Tragopogon d u b i u s : Artemisia ludoviciana 3 3 / t/ + Cirsium undulatum 33/ +/ + Lactuca pulchella 33/ + / t Melilotus officinalis 33/ t/ 3 Senecio canus 33/+/ + . Taraxacum officinale 33/ V + Tragopogon dubius 33/ f/ + Vicia americana . 33/ + / 4 l/ Data are mean percentages for 120 two x five dm plots' distributed as 40 in each of three stands„ ' ' 2/ Constancy/canopy-coverage/frequency. +/+ indicates canopy-coverage or frequency values less than one per cent„ , TABLE XLVo CONSTANCY, CANOPY-COVERAGE AND FREQUENCY AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY OF LESS THAN 6? PER CENT 0% MEAN CANOPY-COVERAGE O F LESS THAN ONE PER CENT ON THE SARCOBATUS VERMICULATUS/ARTEMISIA TRIDENTATA ASSOCIATION, THE AGROPYRON SMITHII/ SYMPHORICARPOS OCCIDENTALIS ASSOCIATIONT=OfhTHE . XANTRIUM STRUMARIUM ASSOCIATION, vI/'_ ; Association Sarcobatus vermiculatus/ Artemisia tridentata Agropyron smithii/ Symphoricarpos occidentalis Xanthium strumarium Union: Taxa ............. ■■ ■ Cy/Cv/Fr 2/ Cy/Cv/Fr Cy/Cv/Fr Artemisia tridentata: Gutierrezia sarothrae 33/ t/ 6 Sarcobatus vermiculatus: Atriplex nuttallii 33/ +/ I Agropyron smithii: Bromus japonicus 33/ 4/ 2 Tragopogon d u b i u s : Artemisia ludoviciana Grindelia squarrosa Symphoricarpos occidentalis: Rosa arkansana Xanthium strumarium: Atriplex dioica ■ ‘ 33/ 4/ .9 % : 33/4/ + 33/ 4/15 33/ 4/ 4 I/ Data are mean percentages for 120 two x five d m plots (40 in each of three stands) in each association. 2/ Constancy/canopy-coverage/frequency. -f/f indicates canopy-coverage or frequency values less than one per cent. 2 0 4 - 205 - TABLE XLVIo CONSTANCY, CANOPY-COVERAGE, AND FREQUENCY-AND DISTRIBUTION BY UNIONS OF TAXA HAVING CONSTANCY . OF LESS THAN 6? PER CENT OR CANOPY-COVERAGE OF LESS THAN ONE PER CENT ON THE ARTEMISIA LONGIFOIIA ASSOCIES. l/ Union: Taxa Cy/Cv/Fr 2/ ' Artemisia longifolia: ■ Mirabilis linearis Yucca glauca ■■ ' • . . 33/ + / t ■ 33/ +/ + Agropyron smithii: Stipa viridula . 3 3 / + / + . Poa secunda: Poa secunda 33/ +/ 3 Tragopogon d u b i u s : Artemisia ludoviciana Campanula rotundifolia Solidago missouriensis Thermopsis rombifolia Gaura coccinea 33/ + / + 33/ +/ + 33/+/ 4 . 33/ +/ 3 - 33/ + / 7 ; Xanthium strumarium: Conringia orientalis Polygonum spp. Iva axillaris 33/ + / 2 ' 33/ + / 2 33/ +/ 7 ' l/ Data are mean percentages for 120 two x five dm plots distributed as 40 in each of three stands» 2j Constancy/canopy-coverage/frequency. +/+ indicates canopy-coverage or frequency values less than one per cent. - 206 - TABLE XLVII. PERCENTAGES OF TOTAL MDLE DEER OBSERVED BI ACTIVITY CLASS DURING SEASONS AND YEARS, 1960-64. Activity Year Summer Fall ■ Winter Spring Yearly Averages Feeding 1960-61 55 (400)2/48 (353) 54 (739) 66 (1,727) 562/ 1961-62 48 (312) 57 (318) 46 (343) 61 (758) 53 1962-63 42 (195) 44 (70) 50 (244) 51 (311) 47 1963-64 39 (139) 49 (77) 42 (44) 31 (36) 40 1960-64 Combo 47 51 50 62 54 Alert 1960-61 30 (220) 38 (273) 33 (452) 25 (644) 31 1961-62 34 (223) 29 (161) 48 (386) 27 (334) 34 1962-63 31 (143) 29 (45) 33 (163) 28 (170) 30. 1963-64 29 (104) 32 (51) 28 (30) 27 (32) 30 1960-64 Comb= 31 33 37 26 31 ■ Traveling 1960-61 11(79) 12 (90) 6 (90) 8 (198) 9 1961-62 11 (71) 10 (56) 2 (17) 10 (123) 8 1962-63 19 (87) 24 (38) 13 (66) 18 (108) 18 1963-64 23 (83) 17 (27) 10 (10) 37 (43) 22 1960-64 Comb, 14 13 7 10 11 Bedding 1960-61 4 (29) 2 (12) .6 (87) 2 (48) 3 1961-62 7 (49) 4 (24) 4 (31). 2 (30) 4 1962-63 9 (42) 2 (4) 3 (16) 3 (20) 5 ' 1963-64 9(32) 2 (3) . 20 (21) 5 (6) 8 1960-64 Combo 7 3 6 2 4 I/ Number of mule deer in parenthesis„ 2/ Mean of seasonal percentages 0 . Jc. TABLE XLVII. AVERAGE NUMBERS OF. MULE DEER OBSERVED BY ACTIVITY CLASS FER TRIP UNDER VARIOUS ENVIRONMENTAL CONDITIONS. ' - 207 - Condition Number of Trips Feeding Bedding ..Alert and Traveling Total Temperature -Oe 20 18.4 (57)^ 2.1, (6). ; . 12.1 (37) 32.6 r-320 76 17.4 (51) 1.9(6) . - 14.5 (43) 33.9 53-45" 91 17.9 (57) 1.1 (4) 12.0 (39) 31.0 46-60® 175 8.9 (56) 1.1 (5) • 7.6 (39). 19.5 61-80® 96 8.6 (51) T (4) 2/ ■ 7.6 (45) 16.9 81°+ 63 3.4 (45) -T (6) 3.6 (49) 7.4 Precipitation None 455 U . 7 (54) 1.1(5) • 8.9 (41) 21.8 Light Rain 30 12.7 (60) 1.2 (6) 7.2 (34) 21.2 Rain 24 2.5 (52) ■ ■ T (4) 2.7 (44) 5.7 Light Snow 6 19.2 (44) 1.6 (9) 15.0 (47) 36.8 Snow 6 10.0 (60) 1.0 (6) ■ 5.7 (34) 16.7 Tfind Velocity Calm 185 14.4 (54) i.i (4) 11.4 (42) 26.9 -10 mph 208 11.8 (57) , 1 . 0 ( 5 ) . 8.1 (38) 21.0 11-20 mph 67 9.2 (49) 1.6.(8) 8.1 (43) 18.8 21-30 mph 31. 10.1 (56) . -'I- (4) 7.2 (40) 18.0 31+ mph 30 6.5 (51) T (6) 5-5 (43) 12.8 Cloud Cover Clear 186 11.1 (51) ■ 1.1 (5) 9.6 (44) §1.8 Partly 201 12.2 (55) 1.2 (5) 8.8 (40) 21.0 Mostly 61 8.7. (52). I M (7) 6.8 (41) 16.6 Overcast 73 16.7 (61) T (3) 10.0 (36) 27.6 Ground Condition Dry 345 11.1 (55) T (4) 8.3 (41) 20.2 Damp 64 15.0 (58) 1.5 (6) 9.3 (36) 25.9 Muddy 24 12.2 (62) . T- (4). 6.6 (34) 19.6 Scat. Snow 41 13.8 (45) 2.7(9) 13.2 (46) 30.9 Snow Cover 46 13.2 (51) 1.2 (5) - 11.4 (44) 26.1 Moon Phase New 135 11.8 (53) 122 (5) 9.2 (41) 22.2 First £ 114 12.0 (56) 1.0 (5) . 8.5 (39) 21.4 Full 139 12.3 (54) 1.0 (4). , 9.4 (42) 22.7 Last £ 131 12.1 (54) 1.2 (5) . 9.1 (41) 22.5 l/ Percentage of total mule deer observed under each- set.of. conditions in parenthesis. 2/ T = less than one per cent. TABLE XLIX. PERCENTAGES OF TOTAL MBLE DEER OBSERVED ON VEGETATION TYPES BY ACTIVITY, SEASONS, AND YEARS, 1960-64. Season Year Average Sample Number Size Observ. Per Trip Observ­ ability Index I/ Percent­ age Feeding a/ L B A I h . AStoP AGROP/ PINUS/ POA JDNIP PSEBD/ JBNIP SARCO/ ARTEM AGROP/ SYMPH XANTH ARLO 2/ Feed Totl Feed Totl Feed Totl Feed Totl Feed TotA Feed Totl Eeeu Totl Iteed Totl 6/ 3/ Summer i960 731 12.4 16 .0 54 .9 36.3 37.2 T 0.5 39.7 47.2 5.3 3.8 5.0 3.7 6.3 3.8 4.8 3.2 2,4 1.5 1961 669 15.2 19.5 4 7 .6 33.6 30.0 T 0.6 34.2 44.2 13-5 13.0 9.3 7.0 6.0 4.3 1.5 T T T 1962 483 10.3 16.3 4 1 .7 22.9 22.2 1.0 T 52.9 53.8 9.5 12.0 11 .0 9 .5 2.4 1.7 T T 0 0 1963 373 11.1 14.6 38.8 20.9 20.4 0 T 44.6 45.6 ■24.5 22.8 5.0 5.6 3.6 2.7 T T T T 1960-63 2,256 12.3 16.6 47.3 31.0 29.1 T T 41.2 47.5 11.0 11.4 7.5 6.3 5.1 3.3 2.4 1.0 1.3 T Fall I960 734 14.7 21.6 48-4 47-5 47.7 5.5 3-0 22.7 29.2 8.2 8.7 2.5 4.0 9.8 5.6 1.4 T 2.5 1-4 1961 587 18.3 28.0 56 .8 34.7 35-1 5.5 3.6 31.1 33-9 3.8 7.2 11 .5 10 .1 5.5 3-9 1 .4 1 .0 3.0 3.1 1962 163 7.4 13.2 44 .5 22.4 28.8 0 1.2 39.5 40.5 11.8 11.7 26.3 16.0 0 1.8 0 0 0 0 1963 163 9.4 14.5 48.7 11.9 21.5 0 0 35.7 4 1 .1 34-5 25.2 4.8 5.5 6.0 3.1 0 0 7.1 3.7 1960-63 1,647 13.6 19.3 51.0 37.3 38.7 4.6 2.7 29.2 33.2 9.3 10.1 8.5 7-5 6.7 4.4 1.1 T 3.0 2.1 Winter 1961 1,441 27.7 39.1 54.0 63.5 57.7 1.5 T 21.5 28.3 2.1 1.9 9.5 10.0 0 T T T T T 1962 845 33.8 70.4 45.6 73.8 59.8 T T 22.2 32.4 0 T 4.0 7.6 0 0 0 0 0 0 1 1963 526 22.9 47.7 49.8 59.9 43-0 0 0 33-3 42.4 5.2 44-6 1 .5 10 .1 0 0 0 0 O O M 1964 120 17.1 31.7 41.9 79.2 63.3 0 0 11 .3 30 .0 3.8 4 .2 5.7 2.5 P 0 0 0 O O r o 1961-64 2,932 27.6 47.2 50.3 66.8 55.9 T T 23.6 32.1 2.1 2 .0 8.1 9-0 0 T T T T T I Spring 1961 2,671 58.1 100.0 65.9 ’ 82.0 77.5 0 0 11.6 16.7 T T 4.8 4.0 1.4 1.3 0 0 0 . 0 1962 1,330 47.5 100.0 60.8 72.0 62.3 T 1.5 18.6 26.5 1.3 1.1 5.4 6.7' T • 1.0 T T T T 1963 625 20.2 42.1 51.0 51.5 51:8 ■ T T 21.1 27.0 2.1 4.2' 10 .0 12 .5 1.8 T T T 3.6 3.0 1964 120 30.0 55.5 30.7 100 .0 66 .7 0 0 . 0 26 .7 0 4.2 0 2 .5 0 0 0 0 . 0 0 1961-64 • 4,746 43.6- 74.4 61.7 7 6 .2 69;6 T T- . 14.4 21.0 T 1.3 ' 6.5 5.8 1.3 1.1 T T T T 1960-61' 5,577 27.0 44.2 59.1 68 .3 63 .2 T T 18.5 25.3 2.2 2.4 5.6 -5.5 2.6 1.9 T ' T T . T. 1961-62 3,431 26 .6 54.2 ■ 53.7 58.9 50.7 1.4 1-3 24.4 32.7 3.6 4.3 6.9 7-5 2.4 1.9 T T T T 1962-63 1,797 14.7 29.8 47.6 44.8 39.2 0 T 33.9 40.0 5-6 7.1 12.5 11.3 1.2 T T T 1.3 1.1 1963-64 776 12.5 29.1 ■ 40.1 38.1 34.4 0 T 31.1 39-3 20.6 17.5' 4.4 4-6 3.2 1.9 T T 2.2 1.7 1960-64 ' 11,581 22.3 39.4 54.5 61.1 53.8 T T 23.1 30.7 4.0 4.7 6.8 6 .9 2.4 1.8 T T T T \J Percentage of total mule deer expected to be observed per trip. 7j Percentage of total mule deer observed feeding. 2/ Vegetation types: Artemisia/Agropyron; Agropyron/Poa; Pinus/Juniperus: Pseudotsuea/Juniperus: Sarcobatus/Artemisia: Agropyron/Symphoricarpos; Xanthium: and Artemisia longifolia. respectively. 4/ Feeding = the percentage of feeding mule deer observed on this type. 2/ Total — the percentage of total mule deer observed on this type, T = less than one per cent. ~ 209 — TABLE Lo PERCENTAGES O F TOTAL MU L E DEER OBSERVED ON VARIOUS VEGETATION TYPES IN RELATION TO W E A T H E R CONDITIONS. • Condition Sample Size L B A I h . L n B R y d AGROP/POA y PINUS/ JUNIP y v I s o . JUNIP' v L B E R . L B A I h d AGR0P/SYMPH, XANTH. Temperature -0° 654 50.2 39.8 ' 1.5 8.6 1-32' 2,575 62.4 27.3 2.2 8.1 33-45* 2,825 64.2 27.1 3.2 5.5 46-60° 3,425 46.2 34.9 7.4 11.5 61-80° 1,623 50.5 32.2 5.4 • 11.8 81°-t 468 35.9 4.2 . 9.0 13.0 Precipitation None 9,914 53.4 32.3 4.9 9.3 Light Rain 636 53.3 30.7 5.2 10.8 Rain 34 41.2 38.2 5.9 14.7 Light Snow 886 70/3 22.2 1.6 5.9 Snow 100 45.0 29.0 8.0 18.0 Wind Velocity Calm 4,984 56*5 . 31.2 4.8 . 8.6 -10 mph 4,382 56.9 29.8 4.4 8.9 11-20 mph 1,262 49.7 29.3 6.2 ■ 14.8 21-30 mph 559 42.4 . 45.2 8.9 11.4 31+ mph 383 37.6 42.3 6:5 13.6 Cloud Cover Clear 4,048 47.0 36.1 6.0 10.8 Partly 4,494 59.8 28.6 3.7 7.9 Mostly 1,012 44.8 35.8 7.0 12.4 Overcast 2,016 63.2 26.5 3.0 7.3 Ground Condition Dry L 6,970 52.0 32.2 5.6 10.2 Damp 1,659 65.1 26.9 2.8 5.2 Muddy 471 47.1 28.9 10.6 13.4 Scat. Snow 1,268 59.3 32.2 1.0 7.5 Snow Cover 1,202 53.3 33.8 3.5 9.4 l/ Vegetation types: Artemisia/Agropyron and Agropyron/Poa combined| Finns/Jun i p e r n s ; Psendotsuga/jnniperus; and Sarcobatns/Artemisia, Agropyron/Symphoricarpos„ and Xanthiton combined„ ™ 210 == TABLE LI o' PERCENTAGES OF TOTAL ELK OBSERVED BY ACTIVITY CLASS DURING SEASONS AND YEARS, 1960=64» Year Summer Fall Winter Spring Yearly- Averages Feeding 1960-61 78 (151)2/74 (267) 67 (253) 79 (253) 74^/ 1961-62 54 (72) 85 (203) 76 (342) 75 (144) 72 1962-63 70 (114) 58 (140) 56 (79) 70 (HO) 63 1963-64 60 (81) 4 (I) 78 (48) 14 (2) 39 1960-64 Combo 66 70 70 74 ' 70 Alert 1960-61 6 (13) 20 (76) 20 (75) .12 (41) 15 1961-62 16 (21) 2 (5) 22 (99) 4 (9) . H I962-63 12 (21) 30 (71) 26 (37) 26 (42) 24 1963-64 12 (16) 96 (23) 0 (0) 86 (12) 48 1960-64 Combo 11 20 20 15 16 Traveling 1960-61 10 (21) 4 (15) 8 (33) 6(22) 8 1961-62 20 (26) 10 (24) 0 (0) 2 (5) 8 1962-63 10 (18) 8 (19) 16 (22) 0 (0) 8 1963-64 18 (24) 0 (0) 3 (2) 9 (0) 5 1960-64 Combo 14 6 6 4 8 Bedding 1960-61 4 (9) I (5) 4 (14) I (4) 2 1961-62 10 (14) . 2 (6) 2.(8) 17 (33) 8 1962-63 6 (11) 4 (IQ) 2(3) 3 (5) 4 1963-64 9 (12) 0 (0) 18 (11) 0 (Q) 6 I96O-64 Combo 7 2 4 6 4 l/ Number of elk in parenthesiso 2/ Mean of seasonal percentages 0 TABLE LII. AVERAGE NUMBERS OF ELK OBSERVED BY ACTIVITY CLASS PER TRIP UNDER VARIOUS ENVIRONMENTAL CONDITIONS. Condition Number of Trips Feeding Bedding Alert and Traveling Total Temperature -0° 20 . 8.2 (76)^ - 2 . 0 ( 1 8 ) T. (.5)2/ 10.7 1-32" 76 6.7 (66) - T.. (8) 2.6 (26) 10.0 33-45° 91 . 7.3 (68) . 1.1 (10). 2.4 (22) 10.9 46-60° 175 3.1 (65) T (5) 1.4 (29) 4,8 61-80° . 96 3.1 (72) T (8) T (20) 4.3 81°+ 63 3.0 (72) " T (6). , T (21) 4.2 Precipitation None 455 4.6 (6?) T (7) 1.7 (25) 6.8 Light Rain 30 2.2 (52) 1,0 (26) 1.0 (23) 4.3 Rain 24 7.3 (84) T (5) 1.0 (11) 8.8 Light Snow 6 3.5 (88) •, T (12) . 0.0 (0) 4.0 Snow 6 3.0 (90) 0.0 (0) T (10) 3.3 Wind Velocity Calm 185 5.1 (63) T . (7). 2.3 (29) 8.0 -10 mph 208 4.0 (71) ' T (10) . 1.0 (18) 5.5 11-20 mph 67 4.4 (66) T . (12) 1.4 (21) • 6.6 21-30 mph 31 6.3 (79) ■ T (8) . 1.0 (12) 8.0 31+ mph 30 3.4 (62) T (2) 2.0 (36) 5.6 Cloud Cover Clear 186 4.4 (65) T . (4) . 2.1 (31) 6.7 Partly 201 . '4.5 (75) T (11) T (15) 6.0 Mostly 61 2.2 (46) T (17) 1.8 (37) 4.9 Overcast 73 7.1 (72) T (9) 1.9 (19) 9.9 Ground Condition Dry 345 3.7 (70) . . T (8) 1.2 (23) 5.3 Damp • 64 4.9 (64) 1.1(15) 1.6 (21) 7.6 Muddy 24 3.9 (64) T (9) 1.7 (28) 6.3 Scat. Snow 41 6.5 (64) T (6) 3.1 (30) 10.3 . Snow Cover 46 8.9 (71) 1.0 (8) 2.6 (21) 12.6 Moon Phase New 135 5.9 (71) ■ •. T (7) . 1.8 (22) 8.4 First £ 114 4.1 (66) T (10) 1.5 (24) 6.2 Full 139 3 . 9 ( 5 9 ) T , (10) 2.0 (31) 6.5 Last £ 131 4.3 (76) T (7) T (16) 5.6 l/ Percentage of total elk observed under each set of conditions in parenthesis. 2/ T = less than one per cent. TABLE LIII. PERCENTAGES OF TOTAL ELK OBSERVED OH VEGETATION TYPES BI ACTIVITY, SEASONS, AND YEAflS, 1960-64. Season Year Average Sample Number Size Observ. Per Trip Percent­ age Feeding I/ L B A I h. AGROP AGROP/ POA - Y D o v. JDNIP y v I R o. JDNIP SAHCO/ ARTEM AGROP/ SYMPH XANTH AflLO 2/ Feed Totl Feed Totl Feed Totl Feed Totl Feed Totl Feed Totl Feed Totl Feed Totl a/ a/ Summer i960 194 3.3 77.8 3 0 .4 3 0 .7 0.0 0.0 18.3 22.2 0.0 7.9 12.0 5.8 13.6 13.2 24.6 19.0 1.0 1.1 1961 145 3.3 54.1 3 2 .9 3 3 .1 0.0 0.0 17.1 28.3 13.3 10.3 5.1 7.6 31.6 20.7 0.0 0.0 0.0 0.0 1962 189 4.0 69.5 3 .3 4 .9 0.0 0.0 22.3 29-3 22.3 44.0 39.1 5-4 12.6 14.7 T T 0.0 1.1 1963 136 4.0 60.9 13.6 14.7 0.0 0.0 0.0 27.2 9-9 11.8 55.6 33.1 17 .3 n . o 0.0 0.0 3-7 2.2 1960-63 664 3.6 66.9 19.8 20.6 0.0 0.0 17.0 26 .6 H . 9 19.4 24-8 11.8 18.1 14.8 7.4 5.7 T 1.1 Fall I960 378 9.6 73.5 64.7 57.9 0.0 0.0 17.9 25.9 4-5 4.8 4.5 2.4 5.9 6.3 2.3 2.6 T 0.0 1961 263 8.2 85.2 41.8 56.3 34.2 24.0 1.6 1.9 2.5 6.5 5.6 4.9 13.5 5.3 T 1.1 0.0 0.0 1962 278 12.6 58.3 10.1 39.6 51.3 29.1 31.6 24.8 0 .0 1 .4 5.1 4.0 1.9 1.1 0.0 0.0 0.0 0.0 1963 32 1.9 4.1 0 .0 3 .1 0.0 0.0 25-0 6.3 50.0 81.3 0.0 0.0 25.0 9.4 0.0 0.0 0.0 0.0 1960-63 951 7.8 70.6 46.2 50.3 22.0 15.1 13.2 18.3 3-5 6.8 5.0 3.5 8.7 4.6 1.3 1.4 T 0.0 Winter 1961 397 7.6 67.4 50.2 49.4 0.0 T 34.3 36.0 T T 13-3 12.6 T 1.0 1.3 0 .0 0.0 0.0 1962 ' 484 19.4 76.1 34.5 38.6 0.0 0.0 48.2 47.1 0.0 0.0 17.3 14.3 0.0 0.0 0.0 0 .0 0.0 0.0 1963 173 7.5 56.0 75.0 52.6 0.0 0.0 12.0 27.2 0.0 12.7 13.0 7.5 0.0 0.0 0.0 0 .0 0.0 0.0 , 1964 74 10.6 78.6 69.6 67.5 0.0 0.0 23.2 24.3 3.6 2.7 3.6 5.4 0.0 0.0 0.0 0 .0 0.0 0.0ro 1961-64 1,128 10.5 70.3 47.6 46.5 0.0 T 36.8 38.7 T 2.3 14-2 12.1 T- , T T 0 .0 0 .0 0 .0 K Spring 1961 338 7-3 79.0 86.5 68.6 0.0 T ■■ 4.1 19.5 2.1 2.7 3.9 6.8 3,0 ■ 2.1 T T 0.0 0.0 1 1962 219 7.8 75.3 74.6 73.1 0.0 0.0 10.7 11.4 2.8 3.7 5.6 3.7 5.1 7.8 1.1 .T 0.0 0.0 1963 175 5.6 70.0 57.3 59.4' 0.0 0.0 8.9 10.3 7.3 8.0 25.8 21.1 0.0, T 0.0 0.0 T T 1964 14 3.5 14.2 0.0 . 0.0 0.0 0.0 100.0 100.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.-0 0.0 0.0 1961-64 746 6.8 74.6 ,81.0 66.5 0.0 0.0 5*9 . 16.5 2.8 4.2 . 6.7 9.1 3>0 3.4 T T T T 1960-61 1,307 6.3 73.8 63.7 54.1 0.0 T 19.5 26 .8 1 .6 3 .4 8.3 7.1 3.6 4.6 3.1 3-6 T T-. 1961-62 1,111 8.6 75.2 ' 42.5 48.9 12.1 5.7 22.4 26.9 3-0 3.6 10.0 9.1 9.5, 5.5 T ■ T 0.0 0.0 1962-63 815 6.6 63 .1 28.3 38.8 13 .6 10 .0 20.3 23.2 9.5 14.9 22.9 8.8 5.0 3.8 T ■ T T- T 1963-64 256 4.1. 56.8 33.1 27.7 0.0 0.0 11.9 27.7 10.6 17.2 31.1 19.1 11.3 7.0 0.0 0.0 2.0 1.2 1960-64 3,489 6.7 70.6 51.5 46.9 5.4 4 .2 20.2 26 .1 3-4 7.2 : n . 6 9 .0 5.8 4.9 1.8 1.5 T T l/ The percentage of total elk observed that were feeding. 2/ Vegetation types: Artemisia/AEropyron; Agropyron/Poa; Finns/J n n i p e m s : Pseudotsnga/J n n i p e m s ; Sarcobatps/Artamisia; Agropyron/Svmphoricarpos; Xanthinm: and Artemisia longifolia, respectively. 2/ Feeding = the percentage of feeding elk observed on this type. 4/ Total = the percentage of all elk observed on this type, T = less than one per cent. — 2.13 — TABLE LIV „ PERCENTAGES OF TOTAL ELK OBSERVED ON VARIOUS VEGETATION TYPES IN RELATION TO WEATHER CONDITIONS„ Condition Sample Size ARTEM/AGROP, AGROP/POA y PINOS/ JUNIP PSEUD/ JUNIP vLB ER.L B A I hd AGROP/SYMPH XANTE. Temperature -0° 214 37.8 29.0 33.2 1-32° 759 58.9 29.4 3.4 8.3 33-45° 989 64«4 24.7 3.7 7.2 46-60* 834 47.4 26.4 . • 11.4 14.9 61-80* 415 -40 oO 18.6 . 17.6 23.9 81°+ 261 18.8 34.1 .6.9 40.0 Precipitation None 3,093 53.2 26.5 6.7 13.7 Light Rain 125 16.0 28.8 33.6 21.6 Rain 24 33.0 4.2 62.5 Light Snow 210 48.1 19.5 32.4 Snow 20 10.0 90^0 Wind Velocity 56.1Calm 1,468 20.1 10.2 12.9 -10 mph 1,153 53.0 29.0 5.5 12.5 11-20 mph 442 45.7 26.8 6.8 19.7 21-30 mph 242 33.5 42 01 T 2/ 24.0 31+ mph 16? 34.7 29.9 3.0 32.3 Cloud Cover Clear 1,252 57.8 25.1 7.2 9.9 Partly 1,212 50.8 25.7 5.3 18.2 Mostly 293 42 0 4 31.1 19.1 7.4 Overcast 715 43.5 27.8 5.1 23.2 Ground Condition Dry 1,838 47.8 25.3 9.8 17.1 Damp 486 68.7 10.9 9.5 10.9 Muddy 147 31.3 34.0 4.8 29.9 Seato Snow 422 74.9 18.7 3.1 3.3 Snow Cover 579 34.7 46.3 T 18.6 ?oa'combined:l/ Vegetation types; Artemisia/ Finns/Juniperus; Psendotsnga/Jianipems I and Sarcobatus/Artemisia, Agropyron/Sympboricarpos» and JCanthinm combinedc g/ T = less than one per cent, TABLE LVo PERCENTAGES OF TOTAL CATTLE OBSERVED BY ACTIVITY CLASS DURING SEASONS AND YEARS, 1960-64« l W Year Summer Fall Winter Spring Yearly l/ Averages Feeding I96O-6I 88 (439)2/ 84 (2,183) 88 (301) 86 (1,731) 87 1961-62 76 (2,171) 89 (1,792). 84 (117) ■78 (708) 82 1962-63 83 (2,000) 90 (1, 022) 98 (67) . 93 .(1,064) 91 •1963-64 86 (2,037) 92 (1,026 100 (53). 100 (38) 94 1960-64 Comb. 82 88 90 86 86 Traveling 1960-61 2 (8) 6 (156) T (2)2/ 2 (51) . 2 1961-62 6 (158) 2 (55) 0 (P) 6 (53) 3 I962-63 4 ( n o ) 2 (20) 0 (0) 2 (20) 2 1963-64 2 (40) 4 (51) 0 (0) 0 (0) 2 I96O-64 Comb. 4 .4 T 2 2 Bedding 196O-6I 10 (48) 9 (245) 10 (36). 11 (226) 10 1961-62 18 (502) 7 (158) 15 (2i) 16 (149) 14 1962-63 12 ( 287) 7 (86) I ( I ) 4 (56) 6 1963-64 12 (280) 3 (37) 0 (0) 0 (0) ■ 4 I96O-64 Combo 14 8 10 10 10 3/ Mean of. seasonal percentages. - 2/ Number of cattle in parenthesis. 2/ T = less than one per cent. - 215 - TABLE LVI. AVERAGE NUMBERS OF CATTLE OBSERVED BY ACTIVITY CLASS PER TRIP UNDER VARIOUS ENVIRONMENTAL CONDITIONS. Condition Number of Trips Feeding Bedding Traveling, Total Temperature -O= I • 57.0 (97)V 2.0 (3) 0.0 (0) 59.0 1-32= 19 46.8 (77) 11.9 (19) 1.3 (2) 59.5 33-45° 35 ,85.7 (82) 16.3 (15) 2.4 (2) 104.4 46-60« 90 76.0 (70) 26.0 (24) 3.7 (5) 106.0 61-80= 53 86.1 (75) 22.6 (19) 5.2 (5) 113.3 81=+ 35 101.7 (75) 28.4 (21) 4.2 (2) 134.1 Precipitation None 204 83.2 (75) 24-2 (21) 4.1 (3) ' 111.4 Light Rain 18 75.8 (81) 16.4 (17) i-9 95.1 Rain 8 35.5 (83) 7.0 (16) 41.6 Light Snow 3 93.7 (89) 11.7 (H) 0.0 (0) 105.3 Snow 0 0.0 (0) 0.0 (0) 0.0 (0) 0.0 Wind Velocity Calm 70 84.1 (75) 24.1 (21) 4.2 (3) 112.4 ' ■ -10 mph 101 87.6 (75) 23.9 (21) 3.9 (3) 115.3 11-20 mph 35 78.8 (71) 28.4. (25) 3.5(3) 110.3 21-30 mph 14 51.4 (85) 8.1 (13) 1.2 (2) 60.8 31+ mph 13 53.3 (81) 9.6 (15) 2.8 (4) 65.8 Cloud Cover Clear 87 84.3 (75) 24.6 (21) 3.9 (3) 112.7 partly 83 85.7 (73) 26.7 (23) 4.2 (3) 116.5 Mostly 31 70.7 (76) 17.6 (19) 4.5 (5) 92.9 Overcast 32 71.1 (83) 13.4 (15) 11.9 (I) 85.3 Ground Condition Dry 176 86.6 (75) 24.5 (21) 4.2 (3) 115.3 Damp 31' 72.0 (74) 22.1 (23) 3-1 (3) 97.0 Muddy 14 70.1 (81) 15.0 (17) 1,4 (2) 86.4 Scat. Snow 7 44.3 (73) 15.4 (25) T (I) 59.1 Snow Cover 5 28.0 ( 86) 4.4 (13) . 0.0 (0) 32.4 Moon Phase New 59 81.3 (76) 20.4 (19) 5.0(5) 106.6 First 5 46 65.6 (71) 21.6 (23) 4.4 (5) 91.6 Full 60 80.6 (75) 23.2 (21) 2.4 (2) 106.1 Last 5 68 92.0 (76) 25.6 (21) 3.3 (3) 120.8 l/ Percentage of total cattle observed under each set of conditions in parenthesis 2/ T = less than one per cent. < TABLE LVII. PERCENTAGES OF TOTAL CATTLE OBSERVED ON VEGETATION TYPES BT ACTIVITY, SEASONS, AND YEARS, 1960-64. Season Tear Sample Size Average Percent- Number age ARTEM/ Observ. Feeding AGROP Per Trip l/ ' AGROP/ POA PINUS/ JUNIP PSEUD/ JUNIP SARCO/ ARTEM AGROP/ SYMffl XfiNTH " ARLO 2/ Feed Totl Peed Totl Feed Totl Feed Totl Feed Totl Feed Totl Feed Totl Feed Totl Summer 1960^/ 548 60.9 88.6 3 ^ 2 39^1 11.3 10.4 28.2 2 6 .1 E T 16.2 15.7 4.0 6 .0 2.9 2.6 0.0 0.0 1961 3,519 140.8 76.6 61.3 60.3 3.8 4*4 13-3 17.5 T T 11.8 9.2 8.3 6 .9 1.1 T 0.0 0.0 1962 3,379 130,0 83.4 57.8 58.3 9.1 9.3 17.2 15.0 T T 8.7 11.1 5.6 4 .9 T T T T 1963 3,254 141.5 86.4 66.1 63.3 3.7 3.7 14.716.5 1.7 1.5 8.6 10.8 5.0 3 .9 T T 0.0 0.0 1960-63 10,700 127.9 82.2 59.4 59.5 5.7 6.0 15.5 16.9 T 1.0 11.2 10.6 6.5 5 .3 T T T T. Fall I960 2,879 82.3 84.4 ' 55,7 55.4 1.6 1.3 10.5 U . 5 T T 25.5 25.5 5.9 5 .7 0.0 0.0 T T 1961 2,340 137.6 89.3 65.7 64.9 3.1 3.5 17.6 17.4 T T 6.9 7.3 5.5 5 .7 T T T T 1962 1,298 92.7 90.6 66.9 67.1 3.4 2.9 16.4 16.7 T T 9.5 9.6 3.4 3 .2 T T 0.0 0.0 1963 1,388 106.8 92.1 72.2 69.9 1.9 1.7 8.7 10.5 4.7 4.2 6.1 7.6 5.0 5 .0 T T 0.0 0.0 1960-63 7,923 100.3 88.1 ,64.3 62 .7 2.4 2.3 13-5 13.9 T 1.1 13.7 14.3 5.2 5 .2 T T T T Winter 1961 411 31.6 88 .7 62.3 53.3 2 6 .8 35.0 3.1 5.1 0.0 0.0 7.5 6.3 T T 0.0 0.0 0.0 0.0 1962 154 51,3 84.7 82.6 66.9 0.0 .0.0 12.8 26 .6 0.0 0.0 4.6 6 .5 0.0 0.0 0.0 0.0 0.0 0.0 1963 71 35.5 98.5 87.7 82.1 0.0 0.0 12.3 17.9 0.0 0.0 0.0 0.0 0.0 0,0 0.0 0.0 0.0 0.0 1964 82 27.3 100.0 13.4" 13.4 82.9 82.9 1.2 1.2 0.0 0.0 2 .4 2.4 0.0 0.0 0.0 0.0 0.0 0.0 1961-64 718 34.2 89.9 65 .6 54.8 22.2 29.2 6.9 10.6 0.0 . 0.0 5.2 5.2 T T 0.0 0.0 0.0 0.0 Spring 1961 2,724 133.8 86.2 80.2.77.0. 7.4 7.2 8.3 H . 7 T T 1.7 1.5 2.0 1.8 T. T 0.0 0.0 1962 ■ 1,435 130.4 77.8 54.7 55.1 3-2 6.3 15.3 12.3 0.0 0.0 20.6 19.7 5.7 5,9 T T 0.0 0.0 1963 1,543 96.4 93.3 64.0 57.4 7.9 8.0 6.4 7.6 T T . 14.8 20.9 5.5 4.9 1.0 T T T ■1964 64 64.0 100.0 100.0100.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 1961-64 5,766 115.3 86 .4 70.2 66.6 6 .5 7.1 9.3 10.6 T T 9.6 11.2 3.4 3.6 T T T T 1960-61 6,562 83 .1 85.7 64.7 62.9 6 .4 6.6 10.7 12.4 T T 13.7 13.5 3.8 3.7 T T T T 1961-62 7,466 133.3 81.3 62.5 60:9 3 .3 4"4 15.1 16.7 % T 11.3 10.6 6.6 6.2 T T T T 1962-63 6,291. 108.5 87.7 61.7 60.2 7.4 7 .6 14.3 13-6 T T 10.3 13.0 5 .0 4-5 T T T T 1963-64 4,788 129.4 88 .5 '67.4.64.8 4.7 4 .4 12.4 14.3 2.5 2.2 7.6 9.6 4 .8 4.1 T . T 0.0 0.0 . 1960-64 25,125 109.2 85.4 • 63 .6 62 .0 5.4 5 .8 13.2 14.3 T T 11.4 11.8 5.3 4.7 T T T . T I/ The percentage of the total cattle observed that were feeding. 2/ Vegetation types: Artemisia/Agropyron: Agropyron/Poa: Pinus/Jnniperus: Pseudotsnga/Juniperhs: Sarcobatus/Arteaisia: Agropyron/Symphoricarpos: Xanthium; and Artemisia IongifoIla, respectively. 2/ Feeding = the percentage of feeding cattle observed on this type. ■ kj Total = the percentage of all cattle observed on this type, T = less than one per cent. y August only. - 217 - TABLE L H J I « PERCENTAGES OF TOTAL CATTLE OBSERVED ON VARIOUS VEGETATION TYPES IN RELATION TO WEATHER CONDITIONS. Condition Sample Size ARTEM/AGROP, AGROP/POA lJ PINUS/ JUNIP PSEUD/ ■ JUNIP vL B E R.LBA Ih AGROP/SYMPH XANTE. Temperature -Oe 59 69.5 22.0 ■ 8.5 1-32' 1,132 74.4 14.1 ■ 11.5 33-45* 3,654 70.7 16.8 6.6 ' 11.8 46-60* 9,507 67.6 15.2 . T 2/ 16.3 61-80" 6,088 67.2 10.6 1.6 20.6 81*4- 4,716 64.7 15.9 T 19.1 Precipitation None 22,730 68.1 14.7 T 16.6 Light Rain 1,714 59.5 13.8 3.4 23.3 Rain 379 68.1 7.7 24.2 Light Snow 333 89.7 4.5 ' 5.7 Snow Wind Velocity Calm 7,829 69.0 15.3 T 15.3 -10 mph 11,682 69.3 13.8 T 18.2 11-20 mph 3,937 71.6 11.4 2.3 14.6 21=30 mph .852 55.4 19.9 T 24-.4 314- mph 856 57.2 22.0 T 20.3 Cloud Cover Clear 9,788 66.2 16.0 . T 17.2 Partly 9,691 69.7 13.3 T 16.2 Mostly 2,944 67.8 12.6 T 18.3 Overcast 2,733 66.0 14.3 - 2.2 17.5 Ground Condition Dry 20,294 67.5 14.1 . T 17.5 Damp 3,074 69.6 16.5 T - 13.7 Muddy 1,212 62.1 13.6 1.5 22.8 Scat 0 Snow 414 81.6 13.0 5.4 Snow Cover 162 67.9 25.9 6.2 I/ Vegetation t y p e s s Artemisia/Agropyron and Agropyron/Poa combined; Finns/Jtmiperus; P sendotsnga/Jnniperns; and Sarcobatus/Artemisia, Agropyron/Symphoricarpos, and Xanthinm combined = 2/ T = less than one per cent. /= 218 — TABLE LIXo MJLE DEER USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST ONE PER CENT BUT LESS THAN TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1960-64 COMBINED„ l/ Vegetation Types Plant Taxa Summer Fall Winter Spring Artemisia/Agropyron: Agropyron smithii Allium textile Aster sppo Fritillaria pudica Microseris nutans Phlox hoodii Sphaeralcea eoceinea Tragopogon dubius Yucca glauca Artemisia cana Chrysothamnus viscidiflorus Juniperus seopulorum Symphoriearpos spp0 Pinus/Juniperus s Agropyron smithii Poa seeunda Allium textile Arnica sororia Artemisia frigida Aster commutatus Comandra umbellatum Fritillaria pudica Microseris nutans Phaeelia linearis Phlox hoodii Tragopogon dubius Vicia americana Yucca glauca Chrysothamnus nauseosus Pinus ponderosa Prunus virginiana Ribes sppo Rosa spp 0 / 18/T2/ 15/T % 4/T 7/2 44/6 .hE 36/3 7/2 30/5 30/6 6/2 4/T 11/T 9/T 11/2 37/T 4/2 4/2 7 2 26/2 7/2 6/T 7/2 V T .hE Dbh E 27/1 6/T D bhE 23/5 36/4 9/1 6/T 20/4 11/T 39/4 5/T 9/T V T .hE 18/1 27/6 3/T .hE 41/4 5/1 9/T .hE 9/1 34/2 28/T 35/2 50/3 21/1 6/T 5/T 14/1 5/3 17/3 18/23/T 10/3 15/T 18/2 10/1 11/T 1Q/T 18/1 13/1 5/T 14/3 63/8 39/4 45/2 . 45/3 - 219 - TABLE LIX. Continuedo Vegetation Types Plant Taxa Summer Fall Winter Spring ■ Allium textile Aster commutatus Glyeyrrhiza lepidota Laetuea pulchella Laetuca serriola Tragopogon dubius Vicia amerieana Artemisia longifolia Ribes cereum Sareobatus/Artemisia s Agropyron smithii Allium textile Aster caneseens Chenopodium album Cirsium undulatum Comandra umbellatum Eriogonum multiceps Helianthus petiolaris Lomatium foenieulatum Mieroseris nutans Monolepis nutalliana Vicia amerieana Artemisia longifolia Atriplex nutallii Rosa spp, J-O/T 10/1 40/2 25/1 50/2 25/2 50/1 10/1 40/4 . 33/T 25/T 20/2 . 17/T 25/2 20/T 20/T 17/T 25/4 50/1 25/T 33/2 67/7 16/2 33/2 25/1 ■ ■ 12/T 12/1 ll/T 17/T 25/2 25/1 12/T 11/T 33/6 33/2 50/3 50/2 . 25/2 12/6 11/T 17/T 17/1 - 25/7 11/3 Aster eommutatus Chenopodium album Lactuca pulchella Rosa sppo Sareobatus vermieulatus 20/2 40/1 33/2 40/2 50/2 50/T 50/T 50/6 Xant h i u m ; Ghenopodium album Chrysothamnus v iseidiflorus Rosa spp= 17/1 17/2 50/4 100/7 - 220 - TABLE LIXo Continuedo Vegetation Types Plant Taxa Summer Fall Winter Spring 33/3 33/3 33/4 33/2 33/1 33/2 33/1 33/1 . 100/1 100/4 100/1 100/1 100/3 100/1 Artemisia longifolia; Chenopodium album Collomia linearis Eriogonum multiceps Polygonum spp. Chrysothamnus viseidiflorus ;ropyron/Poa; Agropyron smithii Conringia orientalis Lomatium foeniculatum Taraxacum officinalis l/ Other taxa used by mule deer but comprising less than one per cent of the mean seasonal use on any vegetation type with seasons of recorded use in parenthesis were; Grasses; Agropyron spicatum (FWS)* Agropyron trachycaulum (Su)jl Hordeum ,jubatum (WS)j, Koleria cristata (FWS)s Muhlenbergia euspidata (FW)s■Stipa viridula (SuFWS)s Bouteloua gracilis (FW)s Calamonilfa longifolia (SuW)s and Carex (SuFS). Forbs; Achillea millifolium (FS)s Astragalus bisulcatus (SuFW)s Astragalus gilviflorus (Su)s Astragalus missouriensis (FS)s Atriplex rosea (Su)s Atriplex argentia (Su)s Atriplex dioica (F)s Calchortus nutallii (S Su)s Camalina microcarps (S Su)s Convulvulus arvensis (Su)s Crepis occidentalia (S Su)s Descurainia pinnata (S)s Euphorbia helioscopia (Su)s Gaura coccinea (S SuF)s Grindelia squarrosa (FWS)9, Lappula redowski (S)s Liastris punctata (SF)s Lomatium orientale (S)s Mirabilis linearis (Su)s Melilotus alba (F)s Mertensia longiflora (S)s Oenothera caespitosa (S)s Opuntla polyeantha (Su)s Penstemon nitidis (S Su), Petalostemum candidum * - ' v Salsola kali(Su), (S), Polygonum erectum (Su) „ Polygonum lapathifolium (Su)9 ________ ______________ (FW)s Solidago gigantea (SuF)s Stephanomeria runcincata (S SuF)s Thermopsis rombifolia (S) Thlaspi.arvense (S Su), and Viola nuttallii (S)„■ Browse; Gutierrezia sarothrae (Su), Ribes setosum (SuWS)s (S Su), and Juniperus horizontalis (¥S) <, Ribes aureum 2/ Per cent frequency of use among all feeding sites on this vegetation type/aggregate mean percentage of total recorded instances of use for this type. T = less than one per cent. t t . - 221 - TABLE LXo ELK USE OF TAXA "WHICH INDIVIDUALLY BECEIVED AT LEAST ONE PER CENT BUT LESS THAN TEN PER CENT OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASON, 1960-64 COMBINED 0 I/ Vegetation Type; Plant Taxa Artemisia/Agropyron; Agropyron spieatum Stipa Tiridnla Boutelona gracilis Allium textile Aster eommutatus Comandra umbellatum Lomatium foeniculatum Microseris nutans Oenothera caespitosa P etalostemum purpureum Tragopogon dubius Vieia amerieana C hrysothamnus viscidiflorus Juniperus seopulorum Agropyron spieatum Koleria cristata Poa seeunda Calamoyilfa longifolia Carex spp. Artemisia ludoyieiana Aster eommutatus Microseris nutans Gaura eoeeinea Prunus virginiana Rhus trilobata Rosa spp, Artemisia tridentata Summer Fall Winter Spring 8/T g/ l2/T 47/1 10/T 4%/2 35/4 6/T 32/2 17/T 53/4 29/1 5/T 68/3 5/T8/T 18/3 ■12/T • 25/4 18/1 . 11/T 8/T 42/2 25/5 42/1 8/2 5/T 8/2 25/2 12/T ■ ' 12/T 26/T 25/1 6/T 5/1 6/1 6/T 12/2 13/T 11/T 36/1 60/8 33/2 56/4 36/7 40/T 20/T . 4 4 / 2 64/3 60/5 2 2 / 1 9/T 7/1 11/T 9/T 20/2 18/3 33/2 22/T 27/1 33/3 13/1 7/1 9/3 20/3. 27/2 20/T 20/T 11/1 27/2 40/5 Agropyron spieatum Agropyron traehycaulum Muhlenbergia euspidata Stipa viridula Artemisia frigida Aster eommutatus Glycyrrhiza lepidota 10/T 33/1 100/T 20/T 67/5 100/1 33/3 33/T 100/1 60/2 33/2 100/T 20/1 / - 222 - TABLE LX« .Continuedo Vegetation Type; Plant PseudotsugaZj-Uniperas: cent <, Taraxacmri. officinale Tragopogon dubius Rhus trilobata Ribes sppo Rosa S p p 0 Artemisia cana Distichlls strieta - Muhlenbergla euspidata Carex spp„ Allium textile Gamalxna mieroearpa Microseris nutans Vieia amerieana Sarcobatus Termieulatus Artemisia tridentata Atriplex nuttallii Summer ■ , Fall . Winter Spring ' '■ ! ' ' ■ ■ . ■■ . _________ 10/1 10/T . 100/3 60/4 • ' 33/2 . 100/T 40/2 80/3 , -100/1 25/2 12/2 / 33/2 25/2 12/T 67/3 53/2 33/1 50/T ' — 33/1 50/1 33/T 50/2 67/8 . 50/3 38/7 33/T 33/2 Distichlis strieta Poa secunda Stipa viridula Allium textile Artemisia biennis Chenopodium album Helianthus Laetuea pulei: lapathif olium Taraxacum officinale Thlaspi arvense Tragopogon dubius Tlseidiflorus spp0 Xa m t h i u m : Agropyron traehycaulum Poa eompressa Poa secunda 20/1 40/7 50/4 50/3 14/2 14/1 25/1 7/T 20/7 25/T 38/T 20/3 50/4 25/2 7/T 50/3 50/3 50/3 36/1 40/5 . 50/T 50/8 17/2 50/8 50/2 /“ 223 “ TABLE LIo Continued„ ■Vegetation Types Plant Taxa Summer Fall Winter Spring Xanthiums eont„ Stipa viridula Helianthus maxamilliana Rumex spp=, spp. 17/3 33/1 33/1 Artemisia IongiTolias Chenopodium albtan ' HeliantMs petiolaris Vicla amerleana Agropyron/Poas ' Koleria erlstata Bouteloua gracilis Artemisia 33/8 33/8 33/2 100/2 60/1 . 60/2 50/6 50/9 50/2 l/ Other taxa used by elk but comprised less than one per cent of the total use of any vegetation type were ^ with season of use in parenthesis s Grasses^ Bromus .japonieus (Su)s B, tectorum (F)s Panicum capillare (FW)s Sporobolus cryptandrus (FW)s Carex filifolia (FWS)s CarexTgeyeri (Su), ForbsI Achillea millifolium (SuFS)s Astragalus bisulcatus (Su)s Astragalus missouriensis (F)s Calochortus nuttallii fe Su)s Ohaenacti& douglasii .(-S)^ -Chenopodium gigantospermum (Su)s Cv glaucum (Su)s Collomia linearis (Su)s Conringia orientalis (S Su)s Crepis oeeidentalis (S Su)s Fritillaria pudica (S)s Geum triflorum (Su)s Grindelia squarrosa (W)s Lepidlum densiflorum (S)s Mentzelia albicaulis Mo laevicaulis (Su)s Mirabilis linearis (Su)s Monalepis nuttalliana (S)s Optmtia polyeantha (WS)s Penstemon nitidis (S)s Phlox hoodii (FWS)s Polygonum douglasii (Su)s Po erectum (Su)s Ranunculus cymbalaria (S)s Rumex crislus (F)s R= mexicanus (Su)s Salsola kali (Su)s Seneeip imtegerrimus (Su)s Solidago missouriensis (Su)s Sphaeraleea eoecinea (S SuF)s Thermopsis rombifolia (Su)j Viola nuttallii '(iSu)s Bahia oppositifolia (Su)s Hymenoxys richardsonii (S Su)s Iva axillaris (Su)s Liastris punctata (Su)s Yucca _____ Browse3 Ghrysothamnus nauseosus (FWTs Gutierrezia sarothrae (W), Pinus ponderosa (W)s Ribes aureum (Su)s Ribes cereurn- (SuF). Ribes setosum (Su). 2/ Per cent frequency of use among all feeding sites on this vegetation type/aggr^gate mean percentage of the total instances of> use for this vegetation type. T — less than one per cento — 224 “ TABLE LXIo CATTLE USE OF TAXA WHICH INDIVIDUALLY RECEIVED AT LEAST ONE PER CENT BUT LESS THAN TEN PER CENT. .OF THE TOTAL INSTANCES OF PLANT USE FOR AT LEAST ONE VEGETATION TYPE DURING AT LEAST ONE SEASONj, I96O-64 COMBINED. I/ Vegetation Types Plant Taxa Spring .Summer , Fall Winter Artemisia./Aeropyron: ' Koleria cristata Stipa viridnla Bontelona gracilis Carex spp. Chrysothamnns nanseosus Gntierrezia sarothrae Sarcobatns vermicnlatns Artemisia tridentata Allinm textile Comandra nmbellatnm Lomatinm foenicnlatum Vicia amerieana Finn's/ Jnhjperns; Bromns tectornm Koleria cristata Mnhlenbergia cnspidata Poa compressa Carex spp. Chrysothamnns nanseosus Rhus trilobata spp Allium textile Microseris nutans 5 9 / 1 ^ 49/1 . . 87/5 67/6 71/3 74/3 - 83.5 50/3 23/2 57/7 . 73/5 100/7 .hE 25/1 . 30/1 17/T inhE . 3/T 43/5 17/3 .hE 9/T 4/T 50/1 12/1 13/1 50/4 67/4 17/T.hE 29/1 9/T 65/4 6/T 18/T . 43/2 4/T 53/3 .hE P 59/1 45/3. 4/T - • : 25/1 67/3 45/1 87/4 50/2 33/T H A 22/1 59/5 4/T . 27/1 . 50/T 50/1 37/4 25/3 » ■ 12/T 50/3 36/1 T/T 25/T 31/3 63/3 50/T 55/1 67/3 9/T Agropyron smithii Koleria cristata Mnhlenbergia cnspidata 'Garex spp. Pruntis virginiana Aster commntatns 75/5 100/1 75/5- 25/4 25/3 50/1 Sarcobatns/Artemisia: Koleria cristata Carex spp. ' Atriplex nnttallii 25/1 29/T 29/1 29/1 14/T , 29/2 TABLE LXIo Continuedo - 225 - Vegetation Type: Plant Taxa Spring Summer- Fall Winter Sarcobatus/Artemisia: cont, Allium textile 100/5 Atriplex dioica Lomatium foenieulatum 75/1 Thlaspi arvense 50/1 Agropyron/Symphoricarpos: Distichlis strieta 20/T Poa compressa 60/3 Poa secunda 20/1 Stipa viridula 20/T 20/T Chenopodium album Glycyrrhiza lepidota Thlaspi arvense 80/1 Tragopogon dubius 40/T Xanthiixm: Distichlis strieta Hordeum ,jubatum Scirpus paludosus Glycyrrhiza lepidota Helianthus marimilliana Laetuea pulchella Rumex mexieanus Tragopogon dubius Artemisia longifolia: Artemisia longifolia 14/1 • -14/T 55/1 75/3 18/T 18/T 75/1 18/T .75/3 45/1 . 75/1 18/1 • 75/1 54/1 18/T 75/1 45/1 25/T 83/3 100/5 33/T 50/1 17/2 . 100/1 83/2 17/1 . 67/1 50/T 17/T 50/3 33/T 50/2 100/8 ropyron/Roa: Koleria eristata Muhlenbergia cuspidata Artemisia frigida Lomatium foenieulatum Polygonum spp» Vicia americana 50/T 40/T . 20/3 100/1 100/5 100/1 50/1 20/1 : 40/1 . 100/1 50/T 60/1 . , 1V 100/T - 226 - TABLE LXI. Continued. l/ Other taxa used by cattle but comprising less than one per cent of the mean seasonal use on any vegetation type ^ with seasons of recorded use in parenthesis9 were: Grass and grass-like; Agropyron eristatum (Su)s Agropyron traehycaulum (S SuF)s Bromus ,japonicus (Su), Puccinella nuttalliana (Su)s Schedonnardus paniculatus (SuF)s Carex filifolia (SuF). Forbs;' Amaranthus graecizans (Su)s Artemisia ludovlciana (SF)s Aster caneseens (SF)s Aster oecidentalis (Su)s Astragalus agrestls (Su)s Astragalus bisuleatus (S SuF)s Astragalus missourlensis (Su)s Caloehortus nuttallii (S)s Camallna microearpa (Su)s Chenopodium gigantospermum (SuF)s Cirsium spp. (Su)s Conringia orientals (Su)s Convulvulus sepium (Su)s Crepis oecidentalis (S Su)s Deseuriana pinnata (S)s Erysimum ineonspieuum (S Su)s Fritillaria pudica (S)s Geum triflorum (Su)s Grindelia squarrosa (S SuF)s Haplopappus nuttallii (W)s Helianthus petiolaris (SuF)s Lactuca serriola (Su)s Lappula redowski (S)s Linum rigidum (Su)s Mertensia longifolia (S Sn). Oenothera eaespitosa (Su)s Penstemon nitidis (Su)s Petalostemum eandidum (Su)s Petalostemum purpureum' (SuF)s Phacelia linearis (Su)s Phlox hoodii (S SuFW)s Plantago spinulosa (S Su)s Polygonum ereetum (Su)s Polygonum lapathifolium (Su)s Iboialea argophylla (Su)s Psoralea eseulenta (Su)s Rumex erispus (F)s Salsola kali (SuFW)s Solidago missouriensis (SuF)s Solidago gigantea (Su)s Spaeralcea ooeelnea (S SuF)s Stephanomeria runeincata (Su)«,' Taraxacum officinale (S Sn) s Thermopsis rombifolia (SuF)s Xanthium strumarium (Su)s Bahia oppositifolia (SuF)s Musineon divarieatum (S)s Gaura coeeinea (Su% Hymenoxys richardsonii (Su)s Iva axillaris'(SuF). Shrubs; Chrysothamnus viscidiflorus (SF)s Eurotia lanata (Su)s Artemisia eana (F). 2/ Per cent frequency of use among all feeding sites on this vegetation type/aggregate mean percentage of total recorded instances of use for this type. T = less than one per cent. LITERATURE CITED B r a t m jl D.L, and L.E. B r o t m« 1950» Mid-winter deer Surveyj, Fort Peck Game Range= Mont= Fish and Game P-R= Qtr= 10(1)= Cliffj, E = P= 1939» Relationships between elk and mule deer in the Blue Mountains of Oregon = Trans M= Am, Wildl= Conf=% 4:560-569 = Coles G=F= 1956o The pronghorn antelope - Its range use and food habits in central Montana with special reference to alfalfa = Mont = Fish and Game Dept= and Mont= Agr= Expt= Sta= Tech= Bull. $16= 63 pp. = 1958a= Big game-live stock competition on Montana's mountain rangelands = Mont= Wildl==, April:24-30» ' 1958b= Range survey guide. Montana Fish and Game Dept. Multilith. 18pp. , Cowans I . McT= 1947» Range competition between mule deer=, bighorn sheep=, and elk in Jasper Parks Alberta. Trans. M= Am. Wildl= Gonf = 3 12:223-227» Gouplands R = T= 1950= Ecology of mixed prairie in Canada. Ecol= Monog.=, 20:271-315. . 1961=. A reconsideration of grassland classification in the northern Great Plains of Morth America. J= Ecol= 5 49:135-167» Dasmann=, W= 1949» Deer-livestock forage studies on the interstate winter deer range in California. J= Range Mgmt=s 2(4):206-212. Dasmanns R.F. and R=D. Taber = ,1956= Behavior of Columbian black-tailed deer with reference to population ecology. J. Mamm.=, 37(2) :i143-164. Daubenmires R. 1952. Forest vegetation of northern Idaho and adjacent Washingtons and its bearing upon concepts of vegetation classification. Ecol. Monog.s 22(4)»301-330» f . 1959. A canopy-coverage method of vegetational analysis. ■ Northwest Sei.=, 33:43-64» Dealys J.E. I960. The densiometer for measurement of crown intercept above a line transect. U.S. Dept, of Agr.9 Pacific N.W. For. and Range Expt= Sta0j, Res= Note 199» 5 PP» DeNios R=M= 1938= Elk and deer food and feeding habits. Trans = N= Am= Wildl= Conf = ,s 3:421-427. — 228 = ■ Giesekers LoFo 19,38» Soils of Fergus County= M o n t « Agr» Sxpto Sta. e s : : c 355<. 82 ppo o o CoBo Manifolds AoTo Strahorns and O 0F 0 Bartholemew0 1953° Soil surveys central Montana, U,S,.Dept, of Agr, and Mont, Agr, Sxpt, Sta, Series 1940s No, 9° 133 PP° Hs=nsons HoC, and W, W h i t m a n , 1938, Characteristics of major grassland types in western North D a k o t a , E c o l „ M o n o g os' 8:57-114° Hollibaughs°Go 1944° Fergus Unit Quarterly Report for the 2nd quarter, Montana Fish and Game Dept, Typewritten, 8 pp, Jansons R=G0 and H 0D, Pieton, 1959° Big game surveys and investigations, deer surveys phase, P-R Completion Report, Project W~74-R“4° Montana Fish and Game Dept, Multilith, 30 pp, Johnson, D 0E, 1.952° Population studies of deer - Fort Peek Game Range and vicinity, P-R Completion Report, Project 1-R, Montana Fish and Game Dept, Multilith, 8 pp, Jonas, R 0J 0 1964, Ecology and management of Merriam1s turkey in the Long Pines, southeastern Montana, Unpubl0' thesis (PhD), Montana State College, 118 pp, Inlander, 0, 1937, Utilization of browse by wildlife, Trans, N, Am, Wildl, Conf,, 2:276-287, , 1955° Deer and cattle range relations in Utah, For, Sei,s 1(2):130-139° , 1958, Techniques in studying competition between big game and livestock, J, Range Mgmfe,, 11:18-21, , and D,E, Jeffery, I964, Deer, elk, and cattle range relations on summer range in Utah, Trans, N, Am, Wildl, Gonf,, 29:404-417° Kimball, T,L, and A,G, Watkins, 1951° The Kaibab North cooperative deer- livestock forage relationships study, Aris, Game and Fish Comm, 77 PP° .Lovaas, A,L° 1963° Gallatin big game studies, P-R Completion Report, Project W-98-R-4° Montana Fish and Game Dept, Multilith, ■ 37 pp, Lemon, P,E, 1956, A spherical"densiometer for estimating forest over­ story density. For, Sci,, 2(4):314-320, - 229 - Morrisj, MoS= and J=E= Schwartz= 1957» Mule deer and elk food habits on the National Bison Range = J= Wildl= Mgpit = s 21(2) ; 189-193« Rouse5 R=A= 1957» Elk food habitS5 range Use5 and movements. Gravelly Mountains, Montana= Unpubl= thesis (M=S = ), Montana State College = 29 PP » Smith5 J = G= and O= Julander= .1953» Deer and sheep competition in Utah = J= Wildl'= Mgmt„5 17(2): 101-112 = Stevens5 D=R= 1964= Elk and livestock relationships in the Crow Creek Drainage5 Elkhorn Mountains, Montana= Unpubl= thesis (M='S = )5 Montana State College = 68 pp. Trueblood5 R= I960= Big game surveys and investigations = P-R Completion Report5 Project W-76-R-5= Montana Fish and Game Dept= Multilith= 30 pp= U = S= Department of Commerce Weather Bureau = 1943-64= Climatological data Montana Monthly Summary= U=S= Government Printing Office, Washington5 D= C= Weaver5 J=E= and F=E= Clements= 1938= Plant ecology= 2d ed= McGraw-Hill Book Co= New York. Wright5 J=C= and E=A= Wright= 1948= Grassland types of south central Montana= Ecol=, 29(4):449-460= Young5 V=A= 1938» The carrying capacity.of big game range= J= Wildl= Mgmt=, 2(2):131-134» 3 MONTANA STATE UNIVERSITY IIBRaBTFC 3 1762 1001 0980 8 DATE DUE •5 v ? - A I J J U ? UJJ'; A Jm ■ ■ Vv /-T -+ I * ? =SftflR NOV 2 $ ^002 Demco1 Inc. 38-293 /9 ^ Zj/ A' i M