Distribution, range use and population characteristics of mule deer associated with the Schafer Creek winter range, Bridger Mountains, Montana by William Floyd Steerey A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Fish and Wildlife Management Montana State University © Copyright by William Floyd Steerey (1979) Abstract: A study was conducted in the Bridger Mountains of southwestern Montana from June 1976 through April 1978. Objectives were (1) to obtain data on yearlong distribution, range use and population characteristics of mule deer associated with the Schafer Creek winter range and (2) to compare findings with those of earlier studies of deer associated with the nearby Armstrong winter range. Vegetation of the Schafer Creek study area was described as consisting of 5 habitat series, comprising 18 distinct habitat types. Series were: Idaho fescue with 2 habitat types covering 1.7% of the area; big sagebrush with 2 habitat types cov-2.6% of the area; Douglas fir with 7 habitat types covering 61.9% of the area; subalpine fir with 5 habitat types covering 30.7% of the area and; limber pine with 1 habitat type covering 2.9% of the area. Nine adult females and 3 adult males were radio-collared and monitored from the ground and 53 fixed-wing flights. An additional 17 mule deer were neck-banded. Movement from the winter range by marked animals ranged from 0 to 29 km and averaged 4.8 km. Three holding areas were delineated on two major travel corridors which were used by deer arriving on and departing from the winter range. Associations among adult females appeared to affect summer, fall and winter distributions. Home range size of radio-collared deer was: 111 ha for females and 387 ha for males during the mild winter of 1976-77; 178 ha for females and 152 ha for males summer-early fall and; 58 ha for females during the more severe winter of 1977-78. Three radioed deer summering in heavily forested habitats had significantly smaller home ranges than 8 others of broken timber habitats. Deer appeared to habitually use the same winter and summering areas. Douglas fir habitats were the most important in winter, spring and fall, and ranked second to subalpine fir in summer. Douglas fir/ Idaho fescue was the most used habitat type in winter while subalpine fir/virgin's bower appeared to be the most important summer type. Closed canopy habitats were important during spring and fall migrations. Females with fawns occurred with greater than expected frequencies in certain habitat types (PIFL/JUCO and PSME/FEID) during summer-early fall but not during the 1977-78 winter. Forbs and browse were the most important summer forage classes used while browse, grass and forbs, respectively, were the most important winter forage classes. There was an apparent 10-20% increase in population and an increase in fawn production from the 1976-77 winter to 1977-78. Winter mortality was approximately 17-19% in 1976-77, consisting primarily of fawns, and was estimated at a minimum of 15%, including 34% of all fawns in 1977-78. Noted differences between the Schafer and Armstrong areas were: more Douglas fir and subalpine fir habitats with shrub understories on the Schafer summer area; no bitterbrush and much more Douglas fir/snowberry habitats on the Schafer Creek winter range and; smaller home ranges and generally higher winter survival on the Schafer Creek winter range.  STATEMENT OF PERMISSION TO COPY In presenting this thesis in partial fulfillment of the requirements for an advanced degree at Montana State University, I agree that the Library shall make it freely available for inspection. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by my major professor, or, in his absence, by the Director of Libraries. It is understood that any copying or publication of this thesis for financial gain shall not be allowed without my written permission. ■ DISTRIBUTION, RANGE USE AND POPULATION CHARACTERISTICS OF MULE DEER ASSOCIATED WITH THE SCHAFER CREEK WINTER RANGE, BRIDGER MOUNTAINS, MONTANA by WILLIAM FLOYD STEEREY A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE ' in Fish and Wildlife Management 'MONTANA STATE UNIVERSITY Bozeman, Montana April, 1979 iii ACKNOWLEDGMENT To the following the author wishes to express his sincere appreciation for their contributions to this study: Dr. Richard J. Mackie, Montana State University, who directed the study and aided in preparation of the manuscript; Dr. Robert L. Eng and Dr. Theodore W. Weaver, Montana State University, for reviewing the manuscript; Mr. Terry Lonner, Montana Department of .Fish and Game, for his time, patience and help with computer analysis of movement data; Mr. Arnold Foss, Regional Game Manager, Montana Department of Fish and Game, for support and cooperation in supplying equipment; Mr. James Stradley, Gallatin Flying Service, for his expertise in aerial deer surveys and relocation of radio-collared deer, and friendship; and to the local landowners, especially Mr. R. Lowis, Mrs. Helen Walker, Dr. K. Goering, Mr. T. LeProwse and Mr. J. Taylor, for their cooperation. I also thank my wife Lorrie and my family for their encouragement and persistence throughout my academic career, and the many friends who have given advice and help throughout the study. The author was supported by the Montana Department of Fish and Game under Federal Aid Projects W-12O-R-S and 9. TABLE OF CONTENTS VITA . .......................................... ii ACKNOWLEDGMENT ............................................ iii LIST OF TABLES .......... . . . . . . . . . . . . . . . . . v LIST OF FIGURES .......................................... viii ABSTRACT........ '....................................... .. ix INTRODUCTION . . . . . . . . . . . . . .......... . . . . . I DESCRIPTION OF STUDY AREA ............................... 2 METHODS ........ . . . . . . . . . . . . . . . . ........ • 10 Habitat Determination.and Vegetation Analysis . . . . . . 10 Mule Deer Habitat Use, Movements and Population Characteristics . ............. . .............. H Food Habits.................................. 13 RESULTS AND DISCUSSION............................... 15 Vegetation............................. 35 Distribution, Movements and Home R a n g e .................. 53 Habitat Use ................................ ^2 Seasonal Habitat U s e .................................... 78 Winter.................................. ' • • • • • 78 Spring................................................ 83 Snmmer .......................................... 82 Early Fall ............................................ 83 Late Fall .■ ....................................... 86 Food Habits ...................... . . . . . . . . . . 91 Population Characteristics .................... . . . 98 Population Size and T r e n d ..........■ . ........... 98 Age and Sex Composition and Trends................ 102 Mortality . . ..................................... 307 APPENDIX.................................................. 309 LITERATURE C I T E D ................................. 115 Page V1. Climatological data for the U. S. Department of Commerce Weather Stations, Belgrade FAA (airport) and Montana State University .............. . . . . . . . . . . . . 7 2. Key to Figures 2 and 3 and list of major habitat series and types, of the Schafer Creek study area. . . . . . . . . 18 3. Spacial composition of the Schafer Creek study area by habitat t y p e s ................ .. ...................... 19 4. Mean percent canopy coverage/frequency of occurrence/ constancy of important plant taxa in grass and shrubland habitat types on the Schafer Creek study area .......... 20 5. Mean percent canopy coverage/frequency of occurrence/ constancy of important plant taxa in Douglas fir-grass habitat types on the Schafer Creek study area .......... 23 6. Mean percent canopy coverage/frequency of occurrence/ constancy of important plant taxa in Douglas fir-shrub and limber pine habitat types on the Schafer Creek study a r e a ................................ .............. 27 7. Mean percent canopy coverage/frequency of occurrence/ constancy of important plant taxa in subalpine fir habitat types on the Schafer Creek study area ........ . 31 8. Vegetative comparisons of Schafer Creek study area and the Armstrong area : .................................... 52 9. Seasonal home range size, in hectares, for radio-collared mule deer associated with the Schafer Creek winter range . 61 10. Seasonal average activity radius and standard diameter, in meters, for radio-collared mule deer associated with the Schafer Creek winter range . . . . . ................ 62 11. Home range size, in hectares, average activity radius and standard diameter, in meters, for neckbanded mule deer on the Schafer Creek winter range 1977-78 .................. 63 LIST OF TABLES ■Table Page Table Page 12. Monthly frequencies of use of habitat types and series by mule deer associated with the Schafer Creek winter range.................. .............................. 74 13. Seasonal frequencies of use of habitat types and series and preference indices (see text) for mule deer associated with the Schafer Creek winter range . . . . . 75 14. Seasonal frequencies of use of habitat types for mule deer associated with the Schafer Creek winter range. Data are percentages in each type as determined by aerial/ground observations and by relocation of radio- marked deer/all other deer observed ........ . . . . . . 77 15. Frequency of occurrence of mule deer fecal pellets among 2x5 decimeter plots and constancy among sites sampled as representative habitat types on the Schafer Creek study a r e a ............ 78 16. Number of fawns per 100 females recorded by habitat type on the Schafer Creek study a r e a ........................ 89 17. Percent use (U)/frequency of occurrence (0)/canopy coverage (C) of plant taxa used at summer feeding sites by habitat type............................ .. . . . 92 18. Monthly and total winter use in percent, by habitat type, of plant taxa which achieved greater than I percent use in one or more months or habitat types ........ 94 19. Winter 1977-78 food habits determined from five rumen samples collected from predator involved mortalities . . 95 20. Mean percent of all instances of use which were green grass and/or forbs at feeding sites by habitat type . . . 97 21. Monthly population estimates for mule deer on the Schafer winter range 1976-77 . . . ......................... 100 22. Monthly population estimates for mule deer on the Schafer winter range 1977-78 ............................ .. 100 vi Table Page 23. Sex and age classification of mule deer associated with the Schafer winter range determined by ground and aerial observations .................... . 104 24. Coefficients of association for 16 marked mule deer on the Schafer Creek winter range 1977-78 .......... H O 25. Mean distances between relocations, in meters, for radio-collared mule deer associated with the Schafer Creek winter r a n g e ................................ Ill 26. Average activity radii, in meters, for marked mule deer in the Bridger Mountains.................... . H 2 27. Mule deer captured and marked on the Schafer winter range 1977 and 1978 ............................... 113 • 28. Mule deer population trends and sex and age classifi­ cations determined by winter helicopter surveys involving the Schafer Creek winter range 1972-78 . . 114 vii viii 1. Map. of the Bridger Mountain Range showing major features and locations of primary study areas ........ 4 2. Distribution of habitat types occurring on the north half of the Schafer Creek study a r e a ................... 16 3. Distribution of habitat types occurring on the south half of the Schafer Creek study area . ............... 17 4. Aerial photo showing boundaries of the total Schafer Creek winter, range and principle winter area together with travel corridors, holding areas and general locations of. marked mule deer after departure . from the winter range........... 54 5. Activity centers and polygon home ranges of radio- collared mule deer on the Schafer Creek winter range 1977-78 ............. 57 6. Summer-early fall activity centers and polygon home ranges of radio^collared mule deer associated with the Schafer Creek winter range . . ....................... 58 7. Mean distance between relocations for radio-collared mule deer associated with the Schafer Creek winter range by season.............. .............. i o/ 8. Shifts of activity centers by three female mule deer on the Schafer Creek winter range 1977-78 . ........... 70 9. Observed fawn and male ratios on the Schafer Creek study area January 19.77' — April 1978 ........... • • • ^2 10. Percent visual relocations for radio-collared mule deer on the Schafer. Creek study area by season . . . . . . . 85 11. Averaged periodic estimates of fawns per 100 adults on the Schafer Creek winter range ...................... 105 LIST OF FIGURES Figure Page ix ABSTRACT A study was conducted in the Bridger Mountains of southwestern Mon­ tana from June 1976 through April 1978. Objectives were (I) to obtain data on yearlong distribution, range use and population characteristics of mule deer associated with the Schafer Creek winter range and (2) to compare findings with those of earlier studies of deer associated with the nearby Armstrong winter range. Vegetation of the Schafer Creek study area was described as consisting of 5 habitat series, comprising 18 distinct habitat types. Series were: Idaho fescue with 2 habitat types covering 1.7% of the area; big sagebrush with 2 habitat types cov- 2.6% of the area; Douglas fir with 7 habitat types covering 61.9% of the area; subalpine fir with 5 habitat types covering 30.7% of the area and; limber pine with I habitat type covering 2.9% of the area. Nine adult females and 3 adult males were radio-collared and monitored from the ground and 53 fixed-wing flights. An additional 17 mule deer were neck- banded. Movement from the winter range by marked animals ranged from 0 to 29 km and averaged 4.8 km. Three holding areas were delineated on two major travel corridors which were used by deer arriving on and de­ parting from the winter range. Associations among adult females appear­ ed to affect summer, fall and winter distributions. Home range size of radio-collared deer was: 111 ha for females and 387 ha for males during the mild winter of 1976-77; 178 ha for females and 152 ha for males summer-early fall and; 58 ha for females during the more severe winter of 1977-78. Three radioed deer summering in heavily forested habitats had significantly smaller home ranges than 8 others of broken timber habitats. Deer appeared to habitually use the same winter and summering areas. Douglas fir habitats were the most important in winter, spring and fall, and ranked second to subalpine fir in summer. Douglas fir/ Idaho fescue was the most used habitat type in winter while subalpine fir/virgin's bower appeared to be the most important summer type. Closed canopy habitats were important during spring and fall migrations. Fe­ males with fawns occurred with greater than expected frequencies in cer­ tain habitat types (PIFL/JUCO and PSME/FEID) during summer-early fall but not during the 1977-78 winter. Forbs and browse were the most important summer forage classes used while browse, grass and forbs, res­ pectively, were the most important winter forage classes. There was an apparent 10-20% increase in population and an increase in fawn produc­ tion from the 1976-77 winter to 1977-78. Winter mortality was approxi­ mately 17-19% in 1976-77, consisting primarily of fawns, and was esti­ mated at a minimum of 15%, including 34% of all fawns in 1977-78. Noted differences between the Schafer and Armstrong areas were: more Douglas fir and subalpine fir habitats with shrub understories on the Schafer summer area; no bitterbrush and much more Douglas fir/snowberry habitats on the Schafer Creek winter range and; smaller home ranges and generally higher winter survival bn the Schafer Creek winter range. INTRODUCTION Ecology of mule deer (Odocoiteus hem-vonus Rafinesque) in the Bridget Mountain Range was first studied by Wilkins (1957) in the vicinity of the Armstrong winter range in 1955-56. Since 1971, studies have continued on the Armstrong area (Schwarzkoph 1973, Bucsis 1974, Hamlin 1974, Mackie et al. 1976, Pac 1976, Morton 1976, Mackie and Stewart 1976, Hamlin 1977, Mackie and Knowles 1977, Youmans 1977, Mackie et al. 1978, Youmans in prep.), In 1975, they were expanded to compare ecological characteristics of mule deer populations and deer ranges throughout the Bridget Mountain complex. Observations by Mackie, Hamlin and Mundinger (1976) and Mackie, Pac and Jorgensen (1978) have indicated possible range-related differences in population characteristics of mule deer associated with different winter ranges in the Bridget Mountains. One of these differ­ ences was apparently higher fawn production and/or survival among deer wintering along the southern portion of the west slope of the Bridget Range as compared with the Armstrong area on the northern portion. This study was established during - the summer of 1976 to determine mule deer range, range use and population characteristics on the south­ ern. area and to compare findings with similar information from the Armstrong area. . Data were obtained through full-time field work during the summers of 1976 and 1977 and the winter of 1977-78 and part-time, intermittent studies at other times from June 1976 to April 1978. DESCRIPTION OF THE STUDY AREA The Bridger Mountain Range (Fig. I) lies immediately northeast of Bozeman, Montana, extending northwesterly in a gently curving arc for approximately 37 km. McMannis (1955) describes the Bridger Range as being formed primarily of steeply dipping Paleozoic sediments (limestone) with a narrow strip of Precambrian metamorphics along the west front. This strip becomes wider south of a fault passing through Ross Pass and includes much of the area on which this study was conducted. The study area was located on the west slope of the Bridgers approximately 13 km north of Bozeman and 12 km due east of Belgrade, Montana. The area is bounded on the east by the Bridger Divide, on the north by the Jones-Ross Creek Divide, on the west by the Gallatin Valley floor and on the south by the Middle Cottonwood- 2 Walton-Sypes Creek Divide. It encompassed approximately 51 km . 2 The Schafer Creek winter range comprised approximately 7.2 km as a strip about 1.6 km wide between Middle Cottonwood Creek and Miser Canyon (Figs. I and 4). This strip occurs on a mountainous convex­ ity, or toe, that extends westward into the valley floor, as does the intensively studied Armstrong area, 14.5 km north. Elevations of the study area range from about 1525 m, where the mountains meet the valley floor, to 2793 m on Saddle Peak along the Bridget Divide. Elevational limits of the winter range are 1525 m to 2190 m; however, the majority of winter deer use occurs below 31980 m. Five major permanent streams occur on the area; Middle Cottonwood, Schafer, Bostwick, Truman and Jones Creeks, All flow in a general east to west direction; and all, except Schafer Creek, originate along the Bridger Divide. This creates a series of distinct east-west ridges and extensive steep slopes of northerly and southerly aspect. Typical of the interior are many limestone outcrops, ledges and talus slopes. At low elevations on the winter range (1525 m to 2190 m) forest cover consists of stands of Douglas fir (Pseudotsuga menziesH (Mirbel) Franco.) on north-facing slopes. Douglas fir extends upward to domi- nate all aspects of the summering area to an elevation of about 2195 m. At higher elevations subalpine fir (Abies lasioearpa (Hook) Nutt.) and limber pine (Pinus fZexiZis James.) predominate as either relatively open forest or dense clumps of trees with mountain meadows between them. The vegetation on the southerly and westerly slopes of the winter range tends to be open shrub-grassland or open Douglas fir-bunchgrass communities. Dominant shrubs were.big sagebrush (Artemisia tridentata Nutt.) and Rocky Mountain juniper (Juniperus soopulorian Sarg.). Domi­ nant bunchgrasses were bluebunch wheatgrass (Agropyron spicatum (Pursh) Scribn. and Smith) and Idaho fescue (Festuea idahdensis Elmer). The 4MEAGHER CO LEGENDGALLATIN CO. CREEK DIRT ROAD ■“ PAVED HIGHWAY '■■■ COUNTY LINE * £ MOUNTAIN PEAK CITY MULE DEER WINTER RANGE MILES IW IlSALL SCHAFER CREEK WINTER RANGE INGSTONBOZEMAN ,Figure I. Map of the Bridget Mountain Range showing major features and locations of primary study areas. 5open Douglas fir-bunchgrass communities occurred in all drainages at elevations less than 2073 m; above this bunchgrasses are replaced by elk sedge (Carex geyevi . Approximately 86 percent of the study area is federally owned and administered by the U.S.D.A., Forest Service, Gallatin National Forest. Sixty-five percent of the Schafer Creek winter range is privately owned. At present access into the area is through private lands; however, a U.S. Forest Service access road following Middle Cottonwood Creek is under consideration. Livestock are grazed at least seasonally over the entire area. The National Forest lands include four cattle allotments permitting a total of 103 head of cattle from July I through October 15 in the area from Jones Creek south to Middle Cottonwood Creek (R. Duncan, U.S. For. Ser., Bozeman Ranger District (Pars. Comm.)). There has been no extensive logging on the area, which is currently^classified as non-commerical forest-land. The area is relatively inaccessible and human activity is very minor throughout the year, except during hunting season when there is moder­ ate hunting pressure in the Middle Cottonwood drainage. In addition to mule deer, the area is also used by small numbers of elk (Cewus canadensis neisoni), mountain goats (Oreamnosamericanus),- moose (Aloes aloes shirasi) and black bear (Ursus amerioanus). Climatological data (U.S. Department of Commerce) from two U.S. Weather Bureau Stations, Belgrade FAA (airport) and Montana State 6University at Bozeman, for the period of January 1977 through April 1978 are presented in Table I. Although both of these stations are not on the study area (Fig. I), they may be indicative of climatic conditions. The 30 year mean annual temperature was 5.4°C and 6.2° C, and precipitation was 34 cm and 47.5 cm for Belgrade and Bozeman, respectively. The annual temperature for 1977 was 5.7°C and 7.1°C, and precipitation was '39.7 cm and 52.7 cm for Belgrade and Bozeman, respectively; 1977 was therefore somewhat warmer and wetter than average. Approximately the same amount of precipitation occurred during the period January-April 1978 but temperatures were colder than the same period for 1977. This resulted in greater snow depth and more days of snow cover in 1978. Both January-April periods were warmer and considerably drier than normal as indicated by 30 year means. Table I. Climatological data of the U.S- Department of Commerce'; weather stations, Belgrade FAA (airport) and Montana State University weather stations. Month Weather Station Temperature °C 30 Yr, Mean Mean Precipitation cm . 30 Yr Mean Mean Snowfall cm Maximum Snow Depth cm Days of Snow on Ground,5 cm, Jan 77 Bel 1 -10.3 - 8.7 1.70 1.68 33.3 20.3 31 MSU 2- , - 6.5 - 6.2 ' 1.52 2.34 34.3 22.9 31 Feb Bel - 2.6 - 4.9 .23 1.04 2.5 2.5 10 MSU 1.0 - 3.2 .15 1.65 2.5 7.6 12 Mar Bel — 1.8 - 2.2 3.94 2.21 39.4 10.2 20 MSU — 0.6 - 1.2 6.07 3.66 68.6 20.3 29 • April . Bel 8.1 5.1 .89 2.95 3.6 2.5 I MSU 8.9 5.5 1.45 4.52 14.5 20.3 6 May Bel 9.9 10.4 6.50 5.33 5.1 MSU 10.2 10.4 9.75 6.78 8.6 June Bel 17.3 14.3 5.89 6.78 MSU 17.3 14.2 5.61 8.18 July Bel 19.1 19.2 2.16 2.79 MSU 18.6 19.1 5.69 3.30 Aug Bel 17.6 18.3 3.51 2.92 MSU 17.7 .18.3 5.41 3.48 Sept. Bel 12.4 12.4 7.62 3.53 MSU 13.3 12.9 7.32 4.47 Table I. (continued) Month Weather Station Temperature °C 30 Yr Mean Mean Precipitation cm 30 Yr Mean Mean Snowfall cm Maximum Snow Depth cm Days of Snow on GroundJi2.5 cm Oct Bel 7.1 6.9 2.59 2.54 ■ MSU 8.4 7.7 3.66 3.71 Nov Bel - 1.7 0.7 2.06 2.06 13.5 MSU ■ - 0.2 0.3 3.02 3.20 30.5 20.3 11 Dec Bel - 7.2 - 5.8 .2.59 1.50 29.0 20.3 22 MSU - 3.8 - 3.8 3.02 2.11 32.3 20.3 22 Jan 78 Bel - 8.8 - 8.7 2.06 1.68 19.1 20.3 31 MSU - 5.4 - 6.2 2.62 2.34 34.8 30.5 31 OO Feb Bel - 6.7 - 4.9 2.03 1.04. 19.8 22.9 28 MSU . - 3.2 - 3.2 2.21 1.65 38.4 27.9 28 Mar Bel 0.7 - 2.2 .99 2.21 9.9 20.3 19 MSU 2.9 - 1.2 1.88 3.66 11.7 25.4 19 April Bel 6.7 5.1 1.88 2.95 2.5 2.5 I MSU 7.2 5.5 2.36 4.52 3.6 30 yr Bel ■ 5.4 34.00 Mean MSU 6.2 47.50 1977 Bel 5.7 39.70 Mean MSU 7.1 42.70 Jan-Apr Bel - 2.7 7.87 30 yr~X MSU - 1.3 12.17 .Table I. (continued) Temperature °C Precipitation cm Weather 30 Yr 30 Yr Snowfall Month Station Mean Mean Mean Mean cm: Maximum ■ Snow Depth cm Days of Snow on Ground 2.5 cm Jan-Apr Bel - 1.6 6.76 1977 X MSU 0.7 9.19 Jan-Apr Bel - 2.0 6.96 1978 X MSU 0.4 9.07 1Belgrade FAA, airport 2Montana State University at Bozeman METHODS Habitat Determination and Vegetation Analysis Plant communities of the study area were tentatively delineated through field reconnaissance on both the winter range and the summer range. Final classification of habitat types followed Pfister, et al. (1974, 1977) for forest types, and Mueggler and Handl (1974) for shrub and grassland types. Some additions were necessary where vegetative char­ acteristics encountered were not described by these authors. Tentative plant communities and habitat types were outlined on copies of aerial photographs in the field, and later refined to construct a habitat type map of the study area. A planimeter was used to measure the land areas of the various habitat types. Vegetative characteristics of habitat types on the winter range were measured using 33 permanent transects established on representa­ tive sites within one to four stands of each type. Transects consist­ ed of ten points 15 m apart, in a straight line perpendicular to the fall line of the slope. Any deviations from this brought about by local conditions were outlined on the back of field data sheets to facilitate future relocation of the transect points. The points were used as the center of a 4 x 10 dm frame to estimate shrub canopy cover. A 2 x 5 dm frame, . located in the lower right corner of the larger plot frame, was used to estimate canopy coverage of grasses and forbs. Canopy coverage estimates followed Daubenmire (1959). Coverage classes were: class I = 0-1%; class 2 = 1-5%, class 3 = 5-25%; 11 class 4 = 25-50%; class 5 = 50-75%; class 6 = 75-95%; and class 7 = 95-100%. Tree and shrub densities were estimated using the point-centered quarter technique (Gottorn and Curtis 1956). Forest canopy cover was estimated at two points along each permanent transect using a spherical densiometer (Lemon 1956). Four readings were made at each point, then averaged for a stand estimate. Exposure and percent slope were esti­ mated using a Keuffel and Esser pocket transit. Elevations were deter­ mined in the field with a Thommen pocket altimeter and later cross checked with a U.S.G.S. topographic map. Common and scientific names of plants encountered and collected followed Booth and Wright (1966), Booth (1972), and Hahn (1973). Mule Deer Habitat Use, Movements, and Population Characteristics Both ground and aerial observations of mule deer were recorded as to time, habitat type where deer were first seen, physiognomy of the immediate area if nontypical for that habitat type, number of animals, age, sex when possible, activity, marking, exposure, relative slope, and elevation. All observations were recorded on copies of aerial photographs and later assigned Universal Transverse Mercator (UTM) grid system coordinates to facilitate movement and habitat usage analysis by computer. 12 To provide data on year-long movements and habitat use, 12 adult mule deer (3 males, 9 females) were fitted with radio transmitter collars in March 1977, while they were on the winter range. These deer were relocated weekly from the air from April 1977 through April, 1978, and periodically from the ground throughout the year. Some attempt was made on nearly all relocations to achieve visual contact, to con­ firm the exact location, determine the habitat type being used, and obtain supplemental information about associations with other deer and/or fawns. In addition there was a maximum of 26 neckbahded.mule . deer on the study area during spring and summer 1977. Nine of these were marked in the spring of 1975 (Mackie Pers. Comm.) and 17 in March and April, 1977. Neckbands were made of ''Armorbite" material painted with identifying symbols (Mackie, et al. 1976). When considering movements, home range calculations, and habitat use, seasons were determined for radioed animals as follows: spring began when an animal was first relocated off of the winter range and ended when that animal was established on its summer range; summer- early fall covered the entire period of time an animal was on its summer range. Summer lasted until the first snows and/or killing, frosts, and early fall ended with, the animal's departure from the summer home range. Late fall covered the time of migration to the winter range. Winter extended from the time an animal became established on the winter range to the time of its departure. 13 To relate seasons to other mule deer, average seasonal dates were determined from movements of radio-collared animals with the follow­ ing results: Winter 1976-77 Ended April 30 Spring May I - June 4 Summer June 5 - August 26 Early Fall August 27 - November 14 Late Fall November 15 - December 20 Winter 1977-78 December 21 - April 30 In using this method, seasons would probably vary slightly from year to year due to weather conditions which influence deer movements. Home range and movement data were analyzed by the minimum area (Mohr. 1949) method. Also, relative use of areas by mule deer was documented as the areas were habitat typed. Population estimates were derived as Lincoln (Overton and Davis 1969) and Schnabel (Overton 1965) indices from observations of marked and unmarked animals during winter and early spring. Aerial and ground observation provided estimates of fawn:female, fawn:adult, and male: female ratios for the population. Food Habits Food habits of mule deer were determined by examination of 24 feeding sites and the contents of five rumens of mule deer found dead 14 on the winter range. Feeding sites were examined as soon as possible, or practical, after being used by mule deer. Instances of plant use were recorded as described by Cole and Wilkins (1958). Rumen samples were analyzed according to the technique of Wilkins (1957). When practical, availability of plant species present at feeding sites on the summer range was determined by recording canopy cover and fre­ quency of species in 10 to 20, 2 x 5 dm frames at the site. On the winter range, availability was estimated from data collected at the nearest permanent transect of the habitat type in which the feeding site occurred. RESULTS AND DISCUSSION Vegetation On the winter range, three series of habitat types (h.t.s here­ after) were recognized. These included an Idaho fescue grassland series comprised of two h.t.s, a big sagebrush shrub grassland series,. also with two h.t.s, and a Douglas fir tree series comprising four types, one of which has two phases. In addition, a mock-orange (PhLZadelphiis Iewissi Pursh.) h.t., was identified to occur locally within larger stands of other h.t.s, on the winter range. Three series were found to occur on the summer range: Douglas fir, subalpine fir, and limber pine which together comprised a total of 13 h.t.s, one of which occurred as two distinct phases. Eighteen h.t.s (Table 2) were then identified and described for the entire study area. Of these, four, PSME/SYAL, FSME/VAGL, ABLA/CLPS, and ABLA/VAGL, comprised two-thirds of the area; and -PSME/SYAL alone covered 43.8 percent of the winter range. Table 3 lists the h.t.s and the areas they comprise on the winter and summer ranges as well as the entire study area as a whole. Figures 2 and 3 depict the distribu­ tion and relationship of h.t.s on the area; while Tables 4, 5, 6 and 7 present data showing understory species composition as the mean canopy coverage and frequency of occurrence of plant taxa among all frames read and their constancy of occurrence among all sites sampled for each h.t. 16 17 Figure 3. Distribution of habitat types occurring on the south half of the Schafer Creek study area. 18 Table 2. Key to figures 2 and 3 and list of major habitat series and types, of the Schafer Creek study area. KEY: Arabic numerals denote habitat series. Letters denote habitat types within series. 1. Festuca idahoensis Series IA: Festuca idahoensis/Agropyron spicatum IB: Festuca idahoensis/Agropyron smithii 2. Artemisia tridentata Series 2A: Artemisia tridentata/Festuca idahoensis 2B: Artemisia tridentata/Agropyron spicatum 3. Pseudotsuga menziesii Series 3A: Pseudotsuga menziesii/Festuca idahoensis 3B: Pseudotsuga menziesii/Agropyron spicatum 3C: Pseudotsuga menziesii/Calamagrostis rubescens- Calamagrostis rubescens phase 3D: Pseudotsuga menziesii/Carex geyeri 3E: Pseudotsuga menziesii/Symphoricarpos albus- Calamagrostis rubescens phase 3F: Pseudotsuga menziesii/Symphoricarpos albus- Agropyron spicatum phase 3G: Pseudotsuga menziesii/Physocarpus malvaceus- Calamagrostis rubescens phase 3H: Pseudotsuga menziesii/Vaccinium globulare- Vaccinium globulare phase 4. Abies lasiocarpa Series 4A: Abies lasiocarpa/Arnica cordifolia 4B: Abies lasiocarpa/Calamagrostis rubescens 4C: Abies lasiocarpa/Carex geyeri- Pseudotsuga menziesii phase 4D: Abies lasiocarpa/Clematis pseudoalpina 4E: Abies lasiocarpa/Vaccinium globulare 5. Pinus flexiT is Series ,5A: Pinus flexilis/Juniperus communis FEID/AGSP FEID/AGSM ARTR/FEID ARTR/AGSP PSHE/FEID PSME/AGSP PSME/CARU PSME/CAGE PSME/SYAL-C PSME/SYAL-A STAEMNTOEF PSME/VAGL ABLA/ARCO ABLA/CARU ' ABLA/CAGE ABLA/CLPS ABLA/VAGL PIFL/JUCO 19 Table 3. Special composition of the Schafer Creek Study Area by habitat type. , 2 km 5.89e Hectares Acres % Entire1 Area % WR2 COCO Entire Area Principal 50.8 19.6 5083.5 12556.4 — — — Winter Range Total 2.3 1.4 230.0 915.0 4.5 32 ■ Winter Range 7.2 2.8 714.8 1765.6 14.1 — — Summer Range 42.8 16.5 4282.9 10578.8 84.4 — — SERIES FEID Series 0.9 0.3 86.8 214.4 1.7 12.1 — — ARTR Series 1.3 0.5 131.1 323.8 2.6 18.4 — PSME Series 31.4 12.1 3143.0 7761.6 61.9 67.9 61.0 ABLA Series 15.6 6.0 1561.0 3854.5 30.7 — 36.4 PIFL Series 1.5 0.6 149.7 369.3 2.9 — 3.5 HABITAT TYPES FEID/AGSM 0.2 0.1 15.2 37.5 0.3 2.1 —— FEID/AGSP 0.7 0.3 71.6 176.9 1.4 10.0 — ARTR/FEID. 1.0 0.4 99.9 246.9 2.0 14.0 — ARTR/AGSP 0.3 0.1 31.2 77.1 0.6 4.4 — PHLE/SYAL IOCCURS ON SMALL LOCAL SITES: OFTEN LESS THAN .5 HA IN SIZE PSME/FEID 3.6 1.4 358.5 885.6 7.1 12.4 6.3 PSME/AGSP 1.5 0.6 146.2 361.1 2.9 8.2 2.0 PSME/CAGE 1.7 0.7 171.6 423.9 3.4 — — 4.0 PSME/CARU 1.8 0.7 178.6 441.2 3.5 — 4.2 PSME/SYAL-C 7.8 3.0 776.5 1918.0 15.3 36.3 12.0 PSME/SYAL-A 2.4 1.0 238.4 588.9 4.7 7.2 3.4 PSME/VAGL 9.6 3.7 955.5 2360.1 18.8 3.5 21.7 PSEM/PHMA 0.8 0.3 ■ 81.5 201.3 1.6 — — 1,9 ABLA/ARCO 0.3 0.1 27.2 67.1 0.5 — 0.6 ABLA/CLPS 7.3 2.8 726.1 1793.5 14.3 — 17.0 ABLA/CARU 0.6 0.2 56.0 138,3 1.1 1.3 ABLA/CAGE 0.5 0.2 48.2 . 119.1 1.0 — 1.1 ABLA/VAGL 7.0 2.7 703.1 1736.6 13.8 — 16.4 PIFL/JUCO PSME/FEID & 1.5 0.6 149.7 369.8 2.9 3.5 PSME/SYAL-A 2.4 0.9 235.5 581.7 4.6 — 5.5 ITotals may not equal 100% due to rounding and planimeter errors.' 2WR=Winter Range 3SR=Summer Range Table 4. Mean percent canopy coverage/frequency of occurrence/constancy of important1 plant taxa in grass and shrubIand habitat types on the Schafer Creek study area. Habitat Typesz FEID/AGSP FEID/AGSM ARTR/FEID ARTR/AGSP PHLE/SYAL # Sites 3 2 3 5 I TAXA # Frames 30 20 30 50 10 GRASSES AND SEDGES: Agropyron smithii 4/.50/1.0 Agropyron spicatum 7/.37/1.O 2/.15/1.0 13/.47/1.0 20/.58/1.0 10/.10/1.0 Bromus carinatus I/.03/.33 2/.07/.33 Bromus tectorum 9/.73/1.O 4/.40/1.0 I/.13/1.0 12/.72/1.0 5/.20/1.0 Bromus spp Tr3/.10/.5 Danthonia uriispicata 3/.15/.50 Festuca idahoensis 8/.23/.67 6/.25/1.0 11/.40/1.0 3/.08/.60 Koeleria cristata 4/.30/1.O 2/.25/1.0 4/.50/1.0 6/.26/.80 Poa pratensis 13/.30/.33 43/.80/1.0 4/.13/.33 NDO Poa sandbergii I/.03/.33 I/.23/1.0 I/.12/.80 Poa spp 5/.27/.67 Stipa Comata 2/.07/.67 Unidentified grasses I/.05/.50 TOTAL GRASSES4 51/1.0/1.0 70/1.0/1.0 34/.87/1.0 38/.92/1.0 15/.30/1.0 FORBS: Achillea millefolium 7/.50/1.O 5/.35/1.0 6/.33/1.0 3/.24/1.0 2/.10/1.0 Alyssum alyssoides 4/.90/1.O I/.10/.50 2/.43/1.0 3/.82/1.0 Apocynum androsaemifoli"um 2/.10/.33 Artemisia Iudovieiana 2/.10/1.O 7/.30/.50 I/.02/.20 Aster pansus 4/.10/.50 Aster spp I/.03/.33 Balsamorhiza sagittata 13/.30/1.0 7/.10/.50 11/.33/1.0 17/.32/.80 2/.10/1.0 Brodiaea grandiflora Tr/.03/.33 Cerastium arvense 11/.67/1.0 9/.63/1.0 7/.58/1.0 Table 4. (continued) Habitat Types # Sites TAXA . . # Frames FEID/AGSP 3 30 FEID/AGSM 2 20 ARTR/FEID 3 30 ARTR/AGSP 5 50 PHLE/SYAL I 10 FOKBS: (continued) Chrysopsis villosa 4/.27/1.O ■ 7/.10/.50 I/.03/.33 Collomia linearis Tr/.13/I.O I/.14/.80 Comandra umbellata I/.03/.33 5/.30/1.0 3/.17/1.0 2/.12/.60 Epilobium paniculatum I/.20/.67 I/. 20/.50 I/.03/.33 I/.18/.60 Galium triflorum I/.02/.20 7/.30/1.0 Gaura coccinea I/.03/.33 3/.15/.50 Helianthella uniflora 3/.05/.50 Hieracium cynoglossoides I/.03/.33 Lactuca serriola I/.05/.50 2/.20/1.0 Linum perrene 2/.03/.33 Tr/.05/.50 Lithospermum ruderale I/.10/1.0 N> Lupinus spp 2/.10/.67 Tr/.10/.50 8/.23/1.0 ' ' 4/.16/.60 H Monarda fistulosa Tr/.05/.50 , Phacelia linearis I/.33/1.0 I/.23/.67 I/.40/1.0 Polygonum douglasii Tr/.13/1.0 I/.04/.20 Solidago missouriensis 4/.10/1.0 Tragopogon dubius 4/.23/.67 6/.45/1.0 2/.17/.67 I/.14/1.0 Unidentified forbs Tr/.07/.33 Tr/.06/.40 TOTAL FORBS 42/1.0/1.0 43/.95/1.0 43/.97/1.0 36/.98/1.0 26/.60/1.0 SHRUBS AND TREES: Amelanchier alnifolia 2/.10/1.0 Artemisia tridentata 24/.60/1.0 11/.36/1.0 Clematis ligusticifolia 8/.20/1.0 Philadelphus lewisii 35/.50/1.0 Prunus virginiana 6/.20/1.0 Rhus radicans I/.10/1.0 Rhus trilobate 9/.10/1.0 Table 4. (continued) Habitat Types FEID/AGSP FEID/AGSM ARTR/FEID ARTR/AGSP PHLE/SYAL # Sites 3 2 3 5 I TAXA # Frames 30 20 30 50 10 SHRUBS AMD TREES Rosa nutkana (continued) 6/.15/.50 Tr/.04/.40 2/.10/1.0 Rubus idaeus 2/.10/1.0 Symphoricarpos albus 7/.10/1.0 TOTAL BROWSE 00/ 00/ 00 6/.15/.50 24/.60/1.0 . 11/.40/1.0 54/.70/1.0 Selaginella densa Moss I/.03/.33 I/.04/.40 14/.40/1.0 Lichens Tr/.06/.40 TOTAL VEGETATION 66/1.0/1.0 84/1.0/1.0 76/1.0/1.0 70/1.0/1.0 63/.70/1.0 Bareground 31/.80/1.0 8/.50/1.0 18/.70/1.0 22/.72/1.0 00/ 00/ 00 Rock 2/.23/1.0 4/.25/1.0 3/.13/.67 ' 6/.38/1.0 50/.70/1.0 Litter 69/1.0/1.0 84/1.0/1.0 80/1.0/1.0 69/.98/1.0 60/7.0/1.0 1 Taxon which obtained a minimum coverage of .5% or frequency of .05. 2See Table 2 for habitat type abbreviations 3Less than .5% canopy cover or frequency of .05. ttMean of the total plant class among sites. Table 5. Mean percent canopy coverage/frequency of occurrence/constancy of important1 ■ plant taxa in Douglas fir-grass habitat types on the Schafer Creek study area. Habitat Type2 PSME/FEID PSME/AGSP PSME/CARU PSME/CAGE # Sites 5 6 I 2 TAXA # Frames 60 60 20 ' 40 GRASSES AND SEDGES: Agropyron spicatum 11/.42/1.O 13/.40/1.0 5/.20/1.0 Bromus carinatus I/.07/.40 7/.35/1.0 7/.25/1.0 Bromus tectorum 2/.23/.60 6/.35/1.0 Calamagrostis rubescens 14/.35/1.0 2/.08/.50 Carex geyeri I/.05/.20 56/.95/1.0 54/.80/1.0 Carex spp I/.07/.30 Elymus cinereus Tr?/.05/1.0 I/.03/.50 Festuca idahoensis 11/.52/1.0 Koeleria cristata 3/.18/.60 I/.12/.50 Poa pratensis I/.02/.20 Poa sandbergii 2/.12/.67 Poa spp I/.13/.20 I/.03/.50 Stipa comata 3/.07/.20 Stipa occidentalis I/.02/.20 TOTAL GRASSES 4 33/.92/1.0 22/.68/1.0 73/1.0/1.0 63/.90/1.0 FORBS: Achillea millefolium 5/.33/1.0 3/.25/1.0 8/.33/1.0 Agoseris galauca I/.02/.20 I/.10/1.0 Alyssum alyssoides I/.35/1.0 I/.32/1.0 Alyssum desertorum I/.02/.20 Antennaria racemosa 14/.30/1.0 I/.03/.50 Antennaria spp I/.03/.40 Aquilegia flavescens I/.03/.50 Arenaria congesta I/.08/.40 Arnica cordifolia 11/.45/1.0 I/.13/1.0 Table 5. (continued) # Sites TAXA # Frames Habitat Types PSME/FEID 5 60 PSME/AGSP 6 60 P SME/CARU I 20 PSME/CAGE' 2 40 FORBS: (continued) Artemisia campestris I/.03/.17 Artemisia frigida I/.05/.17 Artemisia ludoviciana I/.02/.17 • Aster conspicuus 4/.10/1.0 4/.15/1.0 Aster novae-angliae 18/.50/1.0 Balsamorhiza sagittata 14/.28/1.0 11/.22/.50 8/.15/1.0 8/.20/1.0 Cerastium arvense 9/.52/1.O 3/.18/.83 2/.10/1.0 Chrysopsis villosa 2/.07/.40 3/.20/.83 Cirsium Undulatum I/.03/.40 3/.20/.83 Comandra umbellata I/.03/.40 I/.02/.17 Disporum trachycarpum I/.02/.20 ' Epilobium paniculatum Tr/.05/.4Q I/.05/.50 Eriogonum umbellatum I/.02/.20 Fragaria vesca 13/.45/1.0 I/.05/.50 Fragaria virginiana 4/.13/.50 Helianthella uniflora I/.05/1.0 2/.05/1.0 Hieracium cynoglossoides Tr/.05/.40 I/.07/.33 3/.20/1.0 Tr/.05/.50 Lupinus spp 3/.07/.20 5/.15/1.0 5/.13/1.0 Osmorhiza chilensis 2/.10/1.0 Phacelia hastata I/.08/.40 Tr/.07/.33 I/.03/.50 Phacelia linearis I/.12/.40 Tr/.10/.67 Tr/.08/.50 Polygonum douglasii I/.12/.60 Tr/.05/.33 Potentialla spp I/.02/.20 Solidago mis souriensis I/.05/.20 Thalictrum venulosum I/.08/.50 Tragopogon dubius . I/.13/.60 Table 5. (continued) Habitat Types PSME/FEID PSME/AGSP PSME/CARU PSME/CAGE # Sites 5 6 I 2 TAXA # Frames 60 60 20 40 FORBS: (continued) Valariana spp 3/.15/1.0 Viola nuttallii Unidentified forbs I/.03/.40 Tr/.07/.50 Tr/.05/1.0 I/.08/.50 TOTAL FORBS 33/.95/1.0 20/.75/1.0 59/1.0/1.0 37/.85/1.0 SHRUBS AND TREES: Amelanchier a.lnifolia Artemisia tridentata 6/.15/.60 I/.02/.17 Tr/.05/.50 Berberis repens Juniperus communis Juniperus scopulormri 5/.08/.40 4/.03/.33 I/.05/1.0 2/.05/.50 Prunus virginiana Pseudotsuga menziesii I/.03/.40 Tr/.05/1.0 Tr/.10/.50 Rhus trilobate Spiraea betulifolia 2/.03/.33 8/.40/1.0 Symphoricarpos albus I/.05/.40 2/.12/.50 2/.05/1.0 I/.05/.50 TOTAL BROWSE 12/.33/1.0 7/.22/.83 2/.10/1.0 11/.45/1.0 Selaginella densa 6/.20/.60 2/.07/.33 Moss Lichens I/.10/.40 2/.05/.50 Woodsia scopulina I/.05/.40 PRIFC YMnMIFIpRr 64/1.0/1.0 44/.92/1.0 82/1.0/1.0 74/1.0/1.0 Table 5. (continued) TAXA Habitat Types PSME/FEID # Sites 5 # Frames 60 PSME/AGSP 6 60 P SHE / CARU I 20 PSME/CAGE 2 40 Bareground 24/.60/1.0 15/.53/1.0 8/.35/1.0 ' 4/.28/1.0 Rock 7/.30/1.0 22/.65/1.0 3/.25/1.0 8/.33/1.0 Litter 67/1.0/1.0 69/1.0/1.0 87/1.0/1.0 81/1.0/1.0 1Taxon which obtained a minimum coverage of .5% or frequency of .05. 2See Table 2 for habitat type abbreviations. 3Less than .5% canopy cover. 4Mean of the total plant class among sites. Table 6. Mean percent canopy coverage/frequency of occurrence/constancy of important1 plant taxa in Douglas fir-shrub and limber pine habitat types on the Schafer area. Creek study # Sites TAXA # Frames Habitat Types2' PSME /SYAL CARD Phase 7 70 PSME/SYAL AGSP Phase 2 30 PSME/VAGL 5 80 PSME /PHMA 2 60 PIFL/JUC0 2 40 GRASSES AND SEDGES: Agropyron caninum 2/.08/.50 Agropyron spicatum 11/.33/1.0 Tr3/.03/.50 Bromus carinatus 2/.27/1.0 I/.05/.50 Bromus tectorum I/.01/.14 2/.30/1.0 Calamagrostis rubescens 30/.53/.86 2/.03/.50 28/.63/1.0 34/.60/1.0 2/.05/.50 Carex geyeri 3/.09/.29 14/.33/.50 2/.08/.40 8/.20/1.0 Carex spp I/.04/.29 Tr/.07/.50 Festuca idaoensis 6/.17/1.0 Poa sandbergii Tr/.07/.50 Poa spp I/.10/.57 2/.13/.50 I/.15/1.0 TOTAL GRASSES4 39/.79/1.0 42/.97/1.0 30/.70/1.0 41/.72/1.0 4/.35/1.0 FORBS: Achillea millefolium 2/.27/1.0 I/.02/.50 5/.20/.50 Agoseris gIauca 5/.33/.50 Allium spp I/.08/.50 Alyssum alyssoides I/.10/1.0 Anemone multifida 3/.20/.50 Antennaria racemosa 6/.40/.71 12/.49/1.0 5/.30/1.0 2/.13/.50 Antennaria spp I/.07/.50 Aquilegia flavescens ■ 2/.15/.50 Arabis nuttallii I/.40/1.0 Arenaria congests Tr/.07/.50 Arnica cordifolia I/.13/,57 5/.23/.10 11/.36/1.0 10/.47/1.0 Table 6. (continued) # Sites' TAXA # Frames Habitat Types PSME/SYAL CARU Phase 7 70 PSME/SYAL AGSP Phase 2 30 PSME/VAGL 5 80 P SME/PHMA 2 60 PIFL/JUC0 2 40 FORBS: (continued) Arnica latifolia I/.05/1.0 Artemisia michauxina I/.08/.50 Aster conspicuus 2/.10/.43 3/.12/1.0 6/.38/.50 Aster spp I/.03/.50 Astragalus, miser 3/.15/1.0 Astragalus spp. 2/.01/.50 Balsamorhiza sagittata 5/.13/.50 Bupleurum americanum 2/.18/.50 Campanula rotundifolia 4/.35/1.0 Cerastium arvense 3/.33/1.O Tr/.08/.50 Chimaphila umbellata 2/.06/.40 Chrysopsis villosa 2/.03/.50 Cirsium foliosum 3/.13/.50 Claytonia perfoliata I/.27/.50 Collinsia parvifIora 3/.37/.50 Collomia linearis Tr/.07/.50 Crepis atrabarba 2/.17/.50 Disporum trachycarpum 2/.14/.86 ■ I/.03/.50 7/.30/1.0 Erythronium grandiflorum ■ I/.07/.50 Frageria vesca 6/.40/.86 . 3/.10/.50 I/.04/.60 5/.33/1.0 Frageria virginiana 2/.03/.50 Frasera speciosa 2/.10/.50 Galium boreale 9/.65/1.0 Hedysarum sulphurescens I/.03/.20 Helianthella uniflora I/.03/.50 Heuchera parvifolia Tr/.06/.57 Hieracium albiflorum 3/.38/.80 Table 6. (continued) Habitat Types PSME/SYAL PSME/SYAL PSME/VAGL PSME/PHMA PIFL/JUCO CARU Phase AGSP Phase # Sites 7 2 5 2 2 TAXA # Frames 70 30 80 60 40 FORBS: (continued) Hydrophyllum capitatum I/.10/.50 Linum perenne Tr/.08/.50 Lomatum cous 2/.25/1 .0 Lupinus sulphureus 2/.06/.43 Lupinus spp 2/.07/.50 I/.01/.20 4/.07/1.0 Myosotis sylvatica 2/.03/.50 Osmorhiza chilensis I/.05/.40 Penstemon spp I/.05/.50 Phacelia linearis Tr/.10/.50 Pyrola secunda 3/.25/.80 I/.03/.50 Pyrola virens I/.01/.20 2/.23/.50 Sedum lanceolate Tr/.07/1.0 I/.08/.50 Senecio canus I/.13/.50 Senecio spp 3/.25/1.0 Smilacina racemose I/.03/.29 Taraxacum officinale I/.05/.50 Thalictrum venulosum 2/.05/.50 Townsendia parfyi I/.10/.50 Tragopogon dubius I/.07/.50 Trifolium haydenii 4/.18/.50 Valariana spp I/.03/.25 2/.13/.50 Viola nuttallii I/.03/1.0 Zigadenus elegans 4/.18/.50 Unidentified forbs I/.07/.29 TOTAL FORBS 22/.80/1.O 28/.90/1.0 37/.81/1.0 36/.87/1.0 43/.98/1.0 Table 6. (continued) # Sites TAXA # Frames Habitat Types PSME/SYAL CARU Phase 7 70 PSME/SYAL AGSP Phase 2 30 PSME/VAGL 5 80 PSME/PHMA 2 60 PIFL/JUCO 2 40 SHRUBS AMD TREES: Acer glabrum 3/.04/.43 I/.04/.40 Amelanchier alnifolia . 2/.05/.60 I/.05/1.0 Arctostaphylos uva-ursi 2/.03/.20 Berberis repens I/.04/.43 I/.10/.80 I/.07/1.0 Tr/.05/.50 Clematis Columbians I/.06/.43 2/.07/1.0 Juniperus communis 9/.15/1.0 Lonicera utahensis 2/.04/.60 Physocarpus malvaceus 22/.38/1.0 Prunus virginiana I/.01/.14 I/.03/.50 Pseudotsuga menziesii I/.10/.60 Rosa spp . I/.03/.20 Shepherdia canadensis I/.01/.20 4/.08/.50 Spiraea betulifolia 12/.53/1.O 14/.60/1.0 23/.62/1.0 I/.08/.50 Symphoricarpos albus 15/.59/1.O 10/.30/1.O I/.03/.40 Vaccinium membranaceum 31/.78/1.0 TOTAL BROWSE 30/.84/1.O 10/.40/1.O 48/.91/1.0 49/.93/1.0 16/.13/.50 Selaginella densa 3/.10/1.O Moss 14/.83/1.0 11/.30/1.O 26/.79/1.0 59/.93/1.0 Lichens 4/.43/1.O Woodsia scopulina 3/.26/.43 3/.20/1.0 TOTAL VEGETATION 75/1.0/1.0 66/1.0/1.0 74/.99/1.0 88/1.0/1.0 52/1.0/1.0 Bareground I/.06/.57 15/.43/1.0 I/.03/.40 00/ 00/ 00 12/.30/1.0 Rock 4/.19/.86 2/.19/.80 I/.03/.50 21/.75/1.0 Litter 74/1.0/1.0 74/1.0/1.0 85/1.0/1.0 78/1.0/1.0 78/1.0/1.0 (one site) 1Taxon which obtained a minimum of .5% or frequency of .05. giU.Cdi.1 Vi- L lie U V UCtO. y J-O-It V U.-L 0 . 0 0 O L U V l l g O O - U C O • PLess than .5% canopy cover. 4Mean of the total plant class among sites. Table 7. Mean percent canopy coverage/frequency of occurrence/constancy of important1 plant taxa in subalpine fir habitat types on the Schafer Creek study area. Habitat- Types2 ABLA/ARCO ABLA/CLPS ABLA/CARU ABLA/CAGE PSME Phase ABLA/VAGL # Sites I 2 2 I 4 TAXA # Frames 25 40 ■ 60 20 • 80 GRASSES AMD SEDGES: Agropyron spicatum Tr3/.08/1.0 I/.02/.50 Bromus carinatus 2/.10/.50 1/1.5/1.0 Calamagrostis rubescens 2/.10/.50 21/.60/1.0 I/.05/1.0 2/.09/.25 Carex geyeri 4/.10/.50 37/.62/1.0 38/.75/1.0 12/.30/.75 Elymus cinereus Festuca idahoensis .5/.13/1.0 I/.05/1.0 - 6/.21/.50 Poa spp 2/.12/1.O ' I/.15/.50 I/.10/1.0 2/.06/.25 Unidentified grasses 3/. 19/ .'50 TOTAL GRASSES4 2/.20/1.O 6/.35/1.0 60/.97/1.0 39/.85/1.0 24/.64/1.0 FORBS: Achillea millefolium 4/.20/1.O 2/.08/1.0 4/.23/.50 7/.40/1.0 Agoseris glauca 3/.15/1.0 Allium brevistylum 3/.28/1.O I/.10/.50 Anemone multifile I/.08/1.0 Antennaria racemose Tr/.08/.50 4/.28/1 .0 I/.05/1.0 I/.09/.50 Aquilegia falvescens 4/.20/1.O 4/.10/.50 3/.10/.50 Arabis nuttallii 4/.60/1.O ' 2/ .08/.50 Arnica cordifolia 25/.76/1.0 20/.60/1.0 12/.52/1.0 22/.65/1.0 30/.84/1.0 Arnica latifolia 4/.15/1.0 I/.15/1.0 Aster conspicuus 3/.10/.50 9/.27/1.0 7/.19/.75 Aster engelmannii 7/.23/.50 I/.02/.50 Aster novae-angliae Aster spp 3/.12/1.0 8/.40/.50 4/.17/.50 Astragalus miser I/.08/1.0 Table 7. (continued) Habitat Types ABLA/ARCO ABLA/CLPS ABLA/CARU ABLA/CAGE ABLA/VAGL PSME Phase # Sites I 2 2 I 4 TAXA # Frames 25 40 60 20 80 FORBS: (continued) Balsamorhiza sagittata I/.02/.50 6/.25/1.0 Besseya wyomingensis I/.03/.50 Campanula rotundifolia Tr/.08/1.0 Castilleja miniata 2/.08/1.0 Catillleja pulchella 2/.10/1.0 Cerastium arvense I/.02/.50 Tr/.05/1.0 Chimaphila umbellata 2/.10/.75 Cirsium foliosum 3/.08/.50 Claytonia lanceolata 6/.80/1.0 Collinsia parviflora I/.05/1.0 Delphinium bicolor I/.25/1.0 Erigeron spp I/.05/1.0 Erythronium grandiflorum I/.05/1.0 I/.03/.25 Fragaria vesca 6/.25/1.0 I/.05/1.0 I/.06/.25 Frasera speciosa 4/.10/1.0 I/.03/.25 Galium boreale 10/.32/1.0 9/.45/1.0 5/.20/1.0 I/.04/.25 Galium triflorum I/.04/.50 Geranium viscosissimum I/.03/.50 Goodyera oblongifolia 2/.07/.50 3/.19/.75 Habenaria spp Tr/.05/1.0 Hieracium albiflorum 2/.13/.50 2/.23/.75 Hieracium cynoglossoides I/.02/.50 2/.10/1.0 Hydrophyllum capitatum I/.15/1.0 Lomatum cons Tr/.05/1.0 Lupinus caudatus 2/.45/1.0 Lupine spp Mertensia oblongifolia I/.04/1.0 I/.03/.50 3/.13/.50 Table 7. (continued) Habitat Types ABLA/ARCO ASIA/CLPS ABLA/CARU . ABLA/CAGE PSME Phase ABLA/VAGL # Sites I 2 2 I 4 TAXA # Frames 25 40 60 20 80 FORBS: (continued) Osmorhiza chilensis 4/.28/1.0 I/.07/1.0 4/.26/1.0 Pedicularis paysoniana Pedicularis racemosa I/.04/1,O 4/.13/.50 Potentialla spp I/.12/1.O Tr/.05/.50 Pyrola secunda Ranunculus spp 5/.24/1.O '2/.18/1.0 4/.21/1.0 Senecio crassulus 7/.24/1.O 2/.20/.50 Silene parryi I/.04/1.O Taraxadum officinale I/.08/1.O ' I/.05/1.0 l/.03/.50w Thalictrum occidentale Thalictrum venulosum 7/.20/1.O 27/.48/1.0 15/.37/1.0 9/.29/1.O^ Trifolium haydenii 9/.48/1.O 9/.23/1.0 Valariana dioica Valariana spp Viola nuttallii 4/.20/1.O I/.08/1.0 I/.02/.50 2/.15/1.0 Zigadenus elegans Zfgadenus venosus 2/.08/1.O I/.10/1.0 TOTAL FORBS 62/.88/1.0 70/.93/1.0 50/.97/1.0 46/.95/1.0 48/.95/1.0 SHRUBS AND TREES: Abies lasiocarpa 16/.40/1.0 9/.20/.50 2/.07/.50 5/.05/1.0 5/.24/1.0 Acer glabrum Berberis repens I/.03/.50 I/;10/1.0 I/.04/.25 Clematis columbiana 2/.10/1.0 2/.10/.25' Clematis spp Lonicera utahensis I/.03/.50 2/.03/.25 Table 7. (continued) Habitat Types ABLA/ARCO . ABLA/CLPS ABLA/CARU ABLA/CAGE ABLA/VAGL PSME Phase # Sites . I 2 2 I 4 TAXA # Frames 25 40 60 20 80 SHRUBS AND TREES: (continued) Picea spp I/.05/.50 Pseudotsuga menziesii I/.11/.50 Ribes setosum 2 / .04 /1 .O Rosa spp I/.04/.25 Shepherdia canadensis I/.01/.25 Spiraea betulifolia I/.03/.50 11/.48/1.0 9/.30/1.0 Symphoricarpos albus I/.05/.50 2/.04/.25 Vaccinium membranaceum 39/.76/1.0 Vaccinium scoparium 2/.05/.25 TOTAL BROWSE 17/.44/1.O 11/.25/.50 16/.55/1.0 5/.05/1.0 50/.86/1.0 Selaginella densa 3/.40/1.0 Moss 13/.48/1.O 3/.28/1 .0 .5/.25/.50 I/.10/1.0 13/.51/1.0 Lichens I/.35/1.0 Polystichum lonchitis Tr/..01/. 25 TOTAL VEGETATION 76/1.0/1,0 75/.98/1.0 . 80/1.0/1.0 70/1.0/1.0 78/.99/1.0 Bareground 00/ 00/ 00 00/ 00/ 00 3/.15/1.0 7/.30/1.0 I/.06/.50 Rock 12/.48/1.0 I/.18/1.0 4/.20/1.0 9/.50/1.0 Litter 85/1.0/1.0 91/1.0/1.0 88/1.0/1.0 80/1.0/1.0 95/1.0/1.0 1Taxon which obtained a minimum coverage of .5% or frequency of .05. 2See Table 2 for habitat type abbreviations. 3Less than .5% canopy cover. tfMean of the total plant class among sites. 35 Festuoa idahoensis/Agropyron smithii, (FEID/AGSM) (Idaho fescue/western wheatgrass h.t. (Mueggler 1974)) This h.t. occurred at the base of the mountains on gentle roll­ ing or flat footslopes of the winter range at elevations below 1585 m. In most areas the original Idaho fescue/western wheatgrass (Agropyron snrlthii Rydb.) community has been replaced by agricultural crops or seeded pasture or is grazed heavily by domestic livestock. Bucsis (1974) also described this h.t. in the Bridget Mountains 14.5 km north on the Armstrong winter range. It was a relatively minor h.t. , currently making up only 2.1 percent of the winter range (Table 3). Low-growing vegetation was well developed, with grasses having a mean canopy coverage of over 69 percent (Table 4). Major grasses were Idaho fescue, western wheatgrass, cheatgrass brome (Bromus teotovum L.), and Kentucky bluegrass (Poa pvatensi-s L.). The latter, possibly due to grazing influences (Bucsis 1974), was by far the predominant grass, averaging 43 percent canopy cover for the two. sites sampled. Principle forbs wane western yarrow (Achillea mille­ folium L.), cudweed sagewort (Artemisia ludovioiana Nutt.), arrowleaf balsamroot (Balsamorhiza sagittdta :(pursh) Nutt.), golden-aster (Chryso- psis villosa (pursh) Nutt.), and common salsify (Tragopogon duhius Scop.). Shrubs occurred as small thickets of nootka rose (Rosa nufkana Presl.) and chokecherry (Prunus virginiana L.) or as stringers of willows (Salix spp) along ditches and small streams. 36 Festuoa idahoensis/Agropyron sp-Loatum (FEID/AGSP) Idaho fescue/bluebunch wheatgrass h.t. (Mueggler 1974) This was the typical grassland type of the area. It was found throughout the winter range, but was especially prevalent north of the mouth of Bostwick Canyon where it appeared to replace big sage­ brush dominated h.t.s. It occurred on southerly and westerly ex­ posures at elevations between 1890 m and 1645 m. It comprised 10 percent of the winter range. Small stands may also be found on the summer range as openings on steep dry southerly and westerly exposures within larger stands of other h.t.s. Grass cover was primarily bunch grasses such as Idaho fescue, bluebunch wheatgrass and Junegrass (Roetccria oristata (L.) Pers.). Important forbs were western yarrow, arrowleaf balsamroot, field chickweed (Cerastium arvense L.), pale alyssum {Alysswn atyssoides L.), and common salsify. Shrubs were of minor importance, occurring as very widely scattered big sagebrush- and/or Rocky Mountain juniper plants. Artemisda tridentata/Festuoa idahoensds (ARTR/FEID) Big sagebrush/Idaho fescue h.t. (Mueggler 1974) The aspect of this type was a sagebrush grassland with conspicuous amounts of Idaho fescue and bluebunch wheatgrass and widely scattered Douglas fir and Rocky Mountain juniper. The type was generally found on the winter range at elevations from 1768 m to 2360 m. It comprised 14 percent of the winter range. The big sagebrush/Idaho fescue h.t. 37 appeared similar to the big sagebrush/bluebunch wheatgrass h.t.. ; however, Idaho fescue had a canopy coverage of 5 percent or more (Mueggler 1974) and the total vegetative cover was better developed as a result of greater shrub and forb cover (Table 4) than in the AR.TR/ AGSP h.t. Also, ARTR/FEID generally occurred at higher elevations and/ or in more mesic situations, such as the heads of the small east-west drainages along the mountain front and along ridges that collect considerable amounts of snow. Important grasses were bluebunch wheatgrass, Idaho fescue and Junegrass. Conspicuous forbs were arrowleaf balsamroot, field chick- weed and lupine (Lupinus spp ) . Big sagebrush was the only conspicuous shrub. Avtem-Lsia tridentata/Agvoipyvon spioatum (ARTR/AGSP) Big sagebrush/Bluebunch wheatgrass h.t. (Mueggler 1974) The aspect of this type was very similar to the ARTR/FEID h.t. Important differences between the two h.t.s included a less conspicuous occurrence of arrowleaf balsamroot, more open vegetation with bare ground more prevalent, and generally drier site conditions for the ARTR/AGSP h.t. (Table 4). The ARTR/AGSP h.t., was found on the winter range south of Bostwick Creek at elevations between 1615 m and 1890 m on steep, well drained, south to west slopes. Rock outcrops and loose, unstable soils were common site characteristics. The type comprised 4.4 percent of the winter range. Principal grasses 38 recorded were bluebunch wheatgrass, Junegrass, and cheatgrass brome. Major forbs included: arrowleaf balsamroot, field chickweed, lupine and pale alyssum. Big sagebrush was the dominant shrub with nootka rose occurring in lesser amounts. PhtladeVphus Iew-LsiyL/Symphov-Laarpos albus (PHLE/SYAL) Mockorange/Common snowberry h.t. This was a distinct, though very minor type on the study area. It occurred on steep, stable talus slopes of westerly and southerly exposures at the bases of major slopes and small draws below 1753 m. It resembled the A.oer glabrim/Philadelphus lewisii h.t. on the Arm­ strong winter range (Bucsis 1974). The total area occupied was not measured because stands generally were found on small (less than .5 ha) local sites within other h.t.s. Scattered Douglas fir trees may provide a tall open overstory but the patchy dense shrub understory 1-1% m in height, is character­ istic; The presence of soil at the surface was limited and exposed rock made up over 49 percent of the substrate at the one site sampled (Table 4). This explains the limited forb and grass cover. Wherever the type occurred, mockorange was the most abundant shrub. Other shrub species were numerous and included common snowberry (Symphori- aarpos albus (L) Blake), skunkbush sumac {Rhus trilobata Nutt.), chokecherry, and Rocky Mountain maple;{Acer glabrum). Important grasses were bluebunch wheatgrass and cheatgrass brome. Forb diversity 39 was great with many species occurring among different sites but with only three to four being important at any one site. Conspicuous forbs included arrowleaf balsamroot, bedstraw (GaZtiumtrtfidum L.), golden- rod (Sotidago spp ) , and western yarrow. Pseudotsuga mensiesii/Festuoa idahoensis (PSME/FEID) Douglas fir/Idaho fescue h.t. (Pfister et al. 1977) This type was similar in appearance and vegetational composition to the PSME/AGSP and PSME/SYAE-AGSP h.t.s. The presence or abundance of one or more species often was the criteria used in selecting one type over another. The PSME/FEID and PSME/SYAL-AGSP types occurred on less droughtIy sites with PSME/AGSP occurring on steep warm dry sites along the mountain front and on some of the major south slopes along the lower portions of the larger canyons. This type comprised 12.4 percent of the winter range and 7.1 percent of the study area. The understory was dominated by Idaho fescue and bluebunch wheat- grass while needle-and-thread (Stipa oomata Trin & Rupr.) and Junegrass occurred in lesser amounts (Table 5). Cheatgrass brome was common. The major shrubs were big sagebrush and Rocky Mountain juniper. The greater abundance of big sagebrush in this type also separated it from . the two similar open-canopy Douglas fir types. Principal forbs were arrowleaf balsamroot, field chickweed, western yarrow, lupine, golden- aster, pale alyssum, along with many species occurring in minor amounts (Table 5). 40 Pseudotsuga menztestt/Agropyroh sptcatum (PSME/AGSP) Douglas fir/Bluebunch wheatgrass h.t. (Pfister et al. 1977) Douglas fIr/bluebunch wheatgrass h.t. was the dry counterpart of the Douglas fir/Idaho fescue h.t. Understory vegetation was less well developed with a greater amount of bare ground,and big sagebrush was absent except for widely scattered individuals or small groups of plants. This type was restricted to the mountain front except in Jones Creek where it occurred on the north side of the creek from the mouth of the canyon upstream for approximately 2 km. Elevations ranged from 1494 m to 2042 m. The type comprised 2.9 percent of the entire study area and 8.2 percent of the winter range. The over­ story consisted of scattered Douglas fir with an occasional limber pine and had a canopy coverage averaging 69.5 percent for two sites measured. The major grass was bluebunch wheatgrass, with cheatgrass brome and Sandberg bluegrass also conspicuous. Idaho fescue was not encountered in plots on four of six of the sites sampled. The princi­ pal shrub was Rocky Mountain juniper with lesser amounts of skunkbush sumac and common snowberry. The more abundant forbs included arrow- leaf balsamroot, golden-aster, field chickweed and pale alyssum. Pseudotsuga menziesit/Calamagrostis Tubesoensi Calamagrostis rubesoens phase (PSME/CARU-CARU Phase) Douglas fir/pinegrass hit. pinegrass phase (Pfister et al. 1977) This type was found only on the summer range. This type was 41 measured as 4.2 percent of. the summering area, but this did not include many sites, too small to measure, scattered across the area. Locally it occurred in all major drainages on minor slopes of all exposures and small benches at elevations between 1950 m and 2316 m. Typically, stands closely resembled Pfister et al.'s (1977) pinegrass phase; however, elk sedge usually was the dominant understory species with a canopy coverage slightly higher than that of pinegrass (Calcomagvost-is vubesoens Buckley) (Table 5). Pac (1976) also described this type and phase for the Bridget range. The overstory usually was dominated by Douglas fir with scattered limber pine occurring on calcareous sites and occasional whitebark pine ! on sites with a granitic substrate. Understory species included elk sedge, pinegrass, aster spp, heartleaf arnica, raceme pussytoes and woodland strawberry. Shrubs were of minor importance.. Pseudotsuga menzi-es't't/Carex geyeri (PSME/CAGE) Douglas fir/Elk sedge h.t. (Pfister et al. 1977) This was another relatively open Douglas fir type found on dry, steep slopes of the summer range on all but east and northeast expo­ sures. It comprised 4 percent of the summering area at elevations between approximately 1950 m and 2360 m. The PSME/CAGE and PSME/ CARU habitat types are very similar, the former probably occurring on sites where pinegrass can not achieve a coverage of five percent or more (Pfister et al. 1977 ) . Total vegetation and forb cover 42 of PSME/CAGE was less developed compared with PSME/CARU (Table 5). Douglas fir was dominant in the overstory with scattered white- bark pine. The understory was made up primarily of elk sedge which had a mean canopy coverage of 53 percent in the two stands examined. Principle forbs were western yarrow, arrowleaf balsamroot, showy aster {Aster aonspiauus Lindl.) and lupine. White spiraea {Spiraea betulifolia Pall.) and Oregon grape {Berberis repens Lindl.)' were the major shrubs. Pseudotsuga mens-tesH/Symphorioarpos albus (PSME/SYAL) Douglas fir/Common strawberry h.t. (Pfister et al. 1977) This type is one of the more common h.t.s of Montana (Pfister , et al. 1977). It was well represented on the study area as well as in portions of the Bridger Range described by Bucsis (1974) and Pac (1976)., It covered approximately 20 percent of the entire area at elevations ranging from 1524 m to abotit 2195 m. Two phases of this type were identified based on vegetative • composition of the understory (Pfister et al. 1977). These were the pinegrass phase and the bluebunch wheat,grass phase. The pinegrass phase typically occurred on the north slopes of the winter range where it was found in all drainages. It comprised 36.6 percent of the winter range. On summer range, it occurred on all exposures and in most instances gradually lapses into forest habitat types characterised by Vaeeinium understories. The overstory 43 in all cases is a closed canopy of Douglas fir, which had a mean canopy coverage of 9.5 percent for seven sites measured. Lodgepole pine QKnus eontovta Cougl.) was present in small numbers. The striking features of the understory were a low, evenly distributed shrub layer of common snowberry and white spiraea and a well developed grass layer, primarily of pinegrass with a lesser amount of elk sedge. Characteristic, forbs of this phase were woodland strawberry (FragaTia vesaa L.), raceme pussytoes (Antennavia racemosa Hook.), showy aster, sulfur lupine (Lupinus sulphureus Dougl.), rough-fruited fairybell (Disponm trashyoarpwn (Wats.) B. & H.) and Rocky Mountain woodsia CWoodsia soopulina D.C. Eaton.). The bluebunch wheatgrass phase was characterized by a much more open overstory than the pinegrass phase and occurred on drier, south­ erly and westerly exposures. It covered 7.2 percent of the winter range and 4.7 percent of the entire study area. Typically this phase was associated with Douglas fir/bunchgrass h.t.s in an intricate mosaic. The PSME/SYAL-AGSP phase occurred where trees were more clustered with either the PSME/FEID or PSME/AGSP h.t.s being found where the tree cover was more open and/or the site more droughty. This probably reflects a transitional zone between types and/or environments (Pfister et al. 1974), and made delineation of these types difficult. The overstory.consisted of Douglas fir interspersed with widely scattered limber pine. Shrubs included common snowberry, scattered 44 thinly over the more open areas and in patches under areas with a well developed Douglas fir canopy. A number of other species occurred in small amounts. Important grasses were elk sedge, bluebunch wheatgrass, and Idaho fescue. Pinegrass was present in small amounts where a good snowberry cover occurred. A diversity of forbs occurred with both moist and dry forms present. Principal forbs were arrowleaf balsam- root, heartleaf arnica {Arm'ioa OovdifoUa Hook.),. Woodland and Virginia strawberry (Fvagevia VyLvginicma Duch,) and field chickweed. Pseudotsuga menziesii/Physooavpus mdlvaoeus3 Calamagvostis vubesoens Phase (PSME/PHMA, CARD Phase) Douglas fir/Ninebark h.t. pinegrass phase (Pfister et al. 1977) This type was represented on the study area by a single stand located in the lower third of Middle Cottonwood Canyon. This stand comprised 1.6 percent of the area and ranged in elevation from 1585 m to 1860 m . The overstory was comprised of a closed canopy of almost pure Douglas fir, with some lodgepole pine and a few limber pine trees found on rock outcrops and near ridges. The shrubby understory was made up of a patchy layer of ninebark (Fhysocavpus malvaoeus (Greene) Kuntze.), I-Ih m tall, with a lower layer of white spiraea and common snowberry evenly distributed between the taller patches. Pinegrass, together with a lesser amount of elk sedge, formed a well developed grass layer. Important forbs were heartleaf arnica, rough-fruited fairybell, woodland strawberry, raceme pussytoe, 45 rattlesnake plantain (Goodyera oblongiftora Raf.) and lupine. Pfister et al. (1977) indicated that this phase may be transit tional to the PSME/CARU or PSME/AGSP habitat type. Considering the amount of common snowberry and pinegrass present the stand observed could also represent a transition to the PSME/SYAL-CARU phase h.t. which was prevalent on forested north slopes in the vicinity of the study area. South of the study area, however, ninebark is a more common plant. Pseudotsuga menzies-id/Vaoeindum gtobutave3 Vaoodndum globulare Phase (PSME/VAGL, VAGL phase) Douglas fir/Blue huckleberry h.t, Blue huckleberry Phase (Pfister et al. 1977) Other than the combined phases of PSME/SYAL this was the most abundant h.t. on the area (19% of the study area) . Combined with Abdes Zasdoeavpa / Vaoodndum globuZare h.t., the Vaoedndum- dominated types covered 32.5 percent of the study area (Table 6). The Douglas fir/blue huckleberry type was found in all major drainages, primarily on the summer range, on all exposures and declivities, and at eleva­ tions ranging from 1707 m to 2195 m. All stands encountered were typical of the Vaeedndum gtobulave phase (Pfister et al. 1977) . Douglas fir was the indicated climax species in the overstory, but usually shared or ranked second in dominance to lodgepole pine. Understories were dominated by an evenly distributed shrub layer 46 comprised of thinleafed huckleberry (Vaociniimi membranacewn Dougl.) with lesser amounts of white spiraea. Pfister et al. (1977) accepts V. membvanaoeim as being V. globulare. A well developed pinegrass layer occurred under the shrub layer. Conspicuous forbs of the type were raceme pussytoes, heartleaf arnica, side-bells wintergreen (JPyvola secunda L.) and white hawkweed (Hieraoiim albiflorum Hook.). A ground moss was also an important part of the vegetations I composition Abies lasiooavpa/Arnica oordifolia (ABLA/ARCO) Subalpine fir/Heartleaf arnica h.t. (Pfister et al. 1977) The ABLA/ARCO type was a very minor h.t. which covered less than one percent of the area. Like other subalpine fir h.t.s, it was found only on the summer range. Only a single site at the head of Bostwick Creek was encountered. This site was a gentle north slope with a steep incline to the east and a lesser incline on the west. The elevation ranged from 2256 m to 2438 m. Vegetational composition was very diverse, though there was a marked paucity of grasses. Subalpine fir and spruce (Picea spp.) co­ dominated in the overstory with Douglas fir and limber pine occurring in lesser amounts. Together, these species formed a closed canopy. Spruce was more abundant in this type than in any other. Over 15 per­ cent of the ground cover was shrubby subalpine fir, with other shrubby species being of only minor importance. Twenty-three species of forbs were encountered in the. 25, 2 X 5 dm plot frames examined (Table 7). 47 The more conspicuous forbs. were heartleaf arnica, northern bedstraw, haydens clover {Tvi-foli-um haydenti Porter.), western meadow rue iThaltatWm ooeddenbdle Gray.) , and thickleaf groundsel {Senee-to cvassuZus Gray.). Spotted frasera (,FTaseva speaiosa Dougl.) was also quite noticeable but was not encountered in sampling. Abies lasiocavpa/Calcmagvostis vubescens (ABLA/CARU) Subalpine fir/Pinegrass h.t. (Pfister et al. 1977) Subalpine fir/pinegrass h.t. appeared to replace PSME/CARU (Pfister et al. 1977) at higher elevations and/or on cooler, more moist sites. Where noted, it was limited to west-through-south exposures at elevations of 2164 m to 2438 m . This was a fairly minor summer range type, comprising only 1.3 percent of the summer­ ing area. The aspect was one of open forest with a well developed grass understory. Pfister et al. (1977) reported a luxuriant.mat of pihegrass and elk sedge in young open stands, as was the case for the site near the head of Bostwick Creek. In all cases, overstory and reproduction were dominated by Douglas fir, with scattered mature or young subalpine fir throughout. Lodgepole pine was a minor component of the overstory. A grass layer of pinegrass and elk sedge character­ ized the understory. White spiraea was the dominant shrub with lesser amounts of Oregon grape. Western meadow rue and heartleaf arnica were conspicuous forbs with showy aster, woodland strawberry and northern bedstraw found in lesser amounts (Table 7). 48 Abies lasioeappa/Cavex Qeyevi3 Pseudotsuga menziesii Phase (ABLA/CAGE, PSME Phase) Subalpine fir/Elk sedge, Douglas fir phase (Pfister et al. 1977) This type may occur on some of the driest sites on which sub- alpine fir occurs (Pfister et al. 1977). It was a minor type, cover­ ing only 1.1 percent of the summer range, and was found only in the head of Middle Cottonwood drainage on a major, steep 'south-facing slope between 2164 m and 2500 m. Pfister et al. (1977) recognized two phases of this type, an elk sedge phase in which subalpine fir and lodgepole pine are the only major tree components* and a Douglas fir phase which occurs on warmer sites has a greater forb component, and is dominated by Douglas fir. Only the Douglas fir phase occurred on the area; however, subalpine fir was well represented in the overstory with lesser amounts of whitebark pine, lodgepole pine and limber pine. Shrubs were of minor importance. Important forbs included heartleaf arnica, arnica {Avniea latifolia Bong.), western yarrow, arrowleaf balsamroot, and houndtongue hawkweed (HievaciiM oynoglossoides Arv.). Abies lasiooavpa/Clematis pseudoalpina (ABLA/CLPS) Subalpine fir/Virgin’s bower h.t. (Pfister et al. 1977) This was one of the four major h.t.s on the study area compris­ ing 14.3 percent of the study area (Table 3). Stands occurred at elevations from 2134 m to the Bridger crest, where trees become stunted at about 2713 m. Typically the type was found on steep, dry, south to northwest slopes on calcarious substrates. Its aspect 49 was either one of clumps of trees interspersed with forb meadows, or of open forest having a well developed forb understory interspersed with suhalpine fir^spruce thickets, Subalpine fir is indicated as the climax species (pfister et al, 19.77) but in stands on the study area this species shares the overstory with Douglas fir, limber pine and spruce. The abundance of Douglas fir decreased with elevation while limber pine increased in dominance on the drier south slopes and convex west, slopes. Subalpine fir and spruce achieved greatest densities on concave west slopes and northerly aspects. Heartleaf arnica. Arnica Zatifotia, spotted frasera, northern bedstraw, western meadowrue, Hayden’s clover and mountain sweetroot (Osmorhiza chiZensis Hook & A.) were principle forbs. Shrub and grass species sampled appeared to be inconsistent (Table 7) in occurrence. The interspersed forb meadow portions of this type resemble Pac's (1976) description of of the Krummholz h.t. elsewhere in the Bridger mountains. Abies Zasioearpa/Vaocinivm gZobuZare (ABLA/VAGL) Subalpine fir/Blue huckleberry h.t. (Pfister et al. 1977) Subalpine fir/blue huckleberry was another of the major h.t.s, covering nearly 14 percent of the study area. Stands occurred in all major drainages and on all slopes and exposures over a wide eleva- tional range from 1890 m to 2530 m. This type is similar to PSME / VAGL with which it shared many of the same species, except for the occurrence of subalpine fir, which moved in at higher elevations and on 50 cooler, more moist sites. Douglas fir was the.dominant tree in most stands with subalpine fir and lodgepole pine occurring in about equal amounts as co-dominants. This type also seemed to occur on sites that were beyond the ecological tolerance limits of pinegrass. While total grass cover was about the same as in the PSHE/VAGL h.t., there was a shift from pinegrass to elk sedge (Table 7). As in the PSME/VAGL h.t. thinleaf huckleberry and white spiraea formed a distinct shrub layer. At seeps and in other wet areas, dense patches of mountain alder (Atnus sinuata (Regel) Rydg,.). were- common. Other important species found were heartleaf arnica, showy aster, rattlesnake plantain, western meadow rue, mountain sweetroot, and side-bells wintergreen. Pinus flexilis/Junipevus communis (PIFL/JUCO) Limber pine/common juniper h.t. (Pfister et al. 1977) The PIFL/JUCO h.t. occupied approximately 3 percent of the area, occurring on generally convex, dry west to south slopes at elevations above 2073 m. It appeared similar in composition to stands of the ABLA/CLPS h.t. on drier sites, but apparently was too dry and/or warm for subalpine fir. The transition between these two h.t.s seems to be very broad with some widely scattered subalpine fir occurring in a PIFL/JUCO stand. Among stands examined the overstory generally was dominated by limber pine with Douglas fir as a co-dominant. . At higher elevations almost pure stands of limber pine were found. The under-* story was the most diverse encountered of any of the habitat types 51 sampled. Conspicuous shrubs included common juniper (Juniperous oormunis L.) throughout the stands and Canadian buffaloberry (Shepherdia canadensis Nutt.) in all but the highest stands. Forbs were very diverse, many occurring in small amounts. However, northern bedstraw, roundleaf harebell (Campanula rotundifclia L.), milkvetch (Astragalus miser Dougl.), Nuttall rockcress (Arabis nuttallii Robin.), and Amiea latifolia appeared to be consistently common (Table 6). General vegetative characteristics of the study area resembled those described for the Bridger Range in the vicinity of the Armstrong winter range, about 14.5 km to the north (Fig. I), by Schwarzkoph (1972), Bucsis (1974) and Pac (1976). The principle differences ob­ served were the lack of antelope bitterbrush (Purshia tridentata (pursh.) DC) habitats on the Schafer winter range and the lack of both whitebark pine (Pinus dlbioaulis engelm.) habitats and extensive alpine areas on associated summer range. Vegetational data also indicated several, possibly important differences between the two areas. Summer range on Schafer Creek included approximately 80 percent more area covered by Douglas fir and subalpine fir habitat types (Table 8). It included more area dominated by shrub understory (62 % vs 44 %) and less with grass-forb understory (36 % vs 49 %). Because shrub-dominated habitats tended to have well developed overstories, the Schafer Creek range may also have included a higher proportion of closed-canopy forest than the Armstrong area. 52 Table 8. Vegetative comparisons of Schafer Creek study area and the Armstrong area. Habitat Schafer Creek % Armstrong % Summer Range1 ■ . Douglas fir 61 36 Subalpine fir 34 20 Shrub understory 62 44 Grass-forb understory 36 49 Winter Range2 Closed and semi-closed forest 48 48 Grassland 12 14 Shrubland 18 36 Open Douglas fir and/or Rocky 21 3 Mountain juniper Snowberry understory 44 . 7 -1 Armstrong data from Pac (1976). 2Armstrong data from Bucsis (1974). Winter range on the two areas appears to include proportionately similar amounts of both closed or semi—closed forest types and grass­ land types (Table 8). Low shrub (big sagebrush, bitterbrush) habitats appeared about twice as extensive on the Armstrong winter range, cover­ ing 36 percent of that area (Bucsis 1974) as compared with 18 percent of the Schafer Creek winter range. Open Douglas fir-juniper habitats, however, were more abundant on the Schafer area. There also was a dis­ tinct difference in the occurrence of snowberry habitat, which comprised nearly 44 percent of the Schafer Creek winter range but only 7 percent of the Armstrong winter range (Table 8). 53 Distribution, Movement and Home Range Dispersal from the winter range in 1977 began in late April and extended through mid-May. All radio-marked deer which eventually left the winter range had done so by May 22. Figure 4 shows general areas in which marked mule deer were observed after they left the winter range. The approximate average movement from the winter range to these points was 4.8 km. Dispersal distances ranged from essen­ tially zero for one female, which remained on the winter range, to 29 Ion for a yearling male. Except for this buck, which may not have returned to the winter range, the average movement was 3.7 km. rRro major routes (Fig. 4) were used by deer leaving the winter range in spring and returning during fall. The north ridge of Bost- wick Canyon was used by deer traveling to the north end of the study area and into the Bostwick drainage. The ridge forming the Schafer- Middle Cottonwood Creek and Bostwick-Middle Cottonwood Creek divides was used by deer moving to the south and into the Middle Cottonwood drainage. Travel corridors used during spring appeared to be wider and less well defined than those followed during the fall migration. Robinette (1966) pointed out the tendency of deer to "tarry" at certain points along migration routes while Bertram and Remple (1977) described definitive holding areas which were considered as "key" spring and fall habitats for migrating mule deer of fhe North Kings herd in California. Three, such concentration or "holding" areas. Figure 4. Aerial photo showing boundaries of the total Schafer Creek winter range and principal winter area together with travel corridors, holding areas and general locations of marked mule deer after departure from the winter range. 55 (Bertram and Remple 1977, Pac 1976) were identified; one on the Bostwick route and two on the Middle Cottonwood route. The Bostwick holding area was large and linear, with apparently lower deer den­ sities than the two smaller Middle Cottonwood areas. The eastern Middle Cottonwood area was used primarily in fall while the others received both spring and fall use. Endurixig snow cover and the lack of early spring growth on the eastern Middle Cottonwood holding area probably reduced its early spring use by deer. Pac (1976) iden­ tified and described similar holding areas along migration routes used by mule deer associated with the Armstrong winter range; however, these areas typically were on southerly and westerly slopes typified by relatively open habitat types. On the Schafer Creek study area, northerly slopes and Vacoin-Iwn habitat types were common components of holding areas. All deer with continuously functioning radio-collars appeared to follow the same routes in spring and fall with the exception of one adult male. This animal (ch 4-4)'followed a circular route using the Bostwick corridor for spring travel and the Middle Cottonwood corridor in the fall. Movements of radio-collared deer appeared to generally represent movements of the entire winter range herd in the timing of departure and arrival on the winter range. Though deer were able to cross the Bridget divide anywhere along the east boundary of the study area, only one marked animal (ch 3-7) 56 used the east slope during the study. This doe moved to the head of Middle Cottonwood creek in early June and subsequently was relocated June 16, 2.5 km northeast on the east side of Saddle Peak. On June 30 the doe was back at the June 16 location to remain on the west slope apparently for the duration of the study period. Immediately following the termination of field studies, two marked animals were known to have crossed the Divide. One, a female radio-collared in April 1978, moved 15.7 km east from the winter range to summer at the head of Weasel Creek. The other, a 2% year-old male, was shot by a hunter 7 km southeast in Beasley Creek (Mackie pers. comm.). In October many tracks were observed crossing Ross Pass into the head of Jones Creek. Hamlin (1974) also observed this and concen­ trations of deer north of Ross Pass. Winter and summer-early fall activity centers and. polygon home - . ranges of radio-collared deer are shown in Figures 5 and 6, respect­ ively. These data indicated some association among certain adult females throughout the year, except during the fawning and early summer periods, as documented by Miller (1974) among blackball deer. Five of nine radioed females (chs 3-3, 3-6, 3-7, 3-8 and 4-3), and a neckbanded female (number 2277) shared, the same portion of the winter range and subsequently moved to the same area in the head of Middle Cottonwood Creek to summer. Summer activity centers all fell within a circle with a radius of 240 m and winter 1977-78 activity 57 LEGEND Mountain front Stream - Geographic activ ity center Channel 3 -2 Channel 3 -9 Channel 4 -1 Channel Channel Channel O 3 -3 1 3 -8 3 -6 , 3 - 7 1 4 -3 - — — '® 3 -5 --------- — - > ($) Figure 5. Activity centers and polygon home ranges of radio-collared mule deer on the Schafer Creek winter range 1977-78. 58 •U H — 3-9 CMMd »IIMIl 3-4 CtKMMIC ICIIVItI CEIIH © 3-3 — © 3-6 — © 3-7 — © 3 - 6 — © 4- 3 — ® Figure 6. Summer-early fall activity centers and polygon home ranges of radio-collared mule deer associated with the Schafer Creek winter range. 59 centers of all six females coincided (Figs. 5 and 6). The mean coefficient of association (Knight 1970) of these females for the winter of 1977-78 was .65 with a range of .51 to .82 for the 15 possible associations. Two other radio-collared females (chs 3-2 and 4-1), with a coefficient of association of .62 for late winter 1 1976-77, spent spring through fall in close proximity to one another on and just east of the winter range. For the winter of 1977-78 these two animals had a coefficient of association of .56. One of these females was also apparently associated with a neckbanded adult female and a male neckbanded as a fawn. All three were observed to­ gether April 1977. Later, during the summer, the male was observed bedded within 500 m of the two females which were bedded within. 50- 100 m of one another. Upon return to the winter range, their mean coefficient of association was .62, when the male was one and one . half years old. Appendix Table 24 lists coefficients of association for 16 marked mule deer observed regularly during the winter of 1977- 78. It may be possible that close associations among some animals result from parent-offspring relations which in turn result in development of traditional group movement and seasonal distribution patterns. Hamlin (pers. comm.) has observed neckbanded and radio- collared mule deer, in the Missouri River breaks, to form apparent family groups during late summer-early fall. These groups last 60 through the winter and into May and form again the following late summer. Home range size, standard diameters (Harrison 1958) and average activity radii (Hayne 1949) by season for radio-collared deer are listed in Tables 9 and 10 respectively. Table 11 lists winter 1977-78 home range size, average activity.radii and standard diameter for 13 neckbanded deer for which there were more than 10 observations. Home ranges include all points of relocation unless otherwise specified. Though the number of males for which data were available is low, it appeared generally that, males had larger home ranges, activity radii and standard diameters than did females (Tables 9,/10 and 11). Exceptions to this were radioed males, which had smaller average home ranges and standard diameters in summer; however,-values for. these males fell within the range of female values (Tables 9 and 10). Neckbanded adult males also had significantly (P<.05) smaller average home ranges as well as activity radii and standard diameters than females during winter (Table 11). If data for one neckbanded adult male, relocated five times in summer, are included in the cal­ culations for radio-collared males during summer, only home range size was smaller than females during that period. The tendency for males to have larger,home ranges than female deer is well documented in the literature. Table 9. Seasonal range size in hectares for radioed collared mule deer associated with the Schafer Creek winter range. Late Winter 1976-77 Spring Migration Summer Early Fall Late Fall & Hunting Season Winter 1977-78 Females ha ha ha ha ha ha 3-2 200 6 50 40 37 68 4-1 100 20 73 126 30 77 3-9 100 381 39 29 -' 93 3-3 100 92 37 51 53 60 3-6 100 186 207 113 2 43 3-7 100 102 120 93 23 44 3-8 100 156 53 335 I 58 4-3 100 537 100 121 7 48 3-5 100 62 73 50 28 34 X 111 171 84 106 23 58 Males 4-4 60 765 66 658 292 - 3-4 700 101 37 67 28 - 4-2 400 39 - - - . - X 387 302 52 363 160 — X all 180 204 78 153 50 58 Table 10. Seasonal average activity radius and standard diameter, in meters, for radio- collared mule deer associated with.Schafer Greek winter range. Late Winter Channel 1976-77 Spring Summer Early Summer- Late Winter Fall Early fall Fall 1977-78 AAR1 SD2 AAR SD AAR SD AAR SB AAR SD AAR SD AAR SD 3-2 200 500 170 390 470 1030 300 700 350s 850 360 780 340 780 4-1 200 510 240 560 420 960 590 1300 5804 1260 300 680 310 800 3-9 200 540 1340 2780 310 730 250 550 280 640 - - 350 800 3-5 • 200 500 720 1480 360 840 550 1270 450 1100 990 2020 250 650 3-3 200 420 590 1370 270 690 610 1500 380 1080 490 1080 260 590 3-6 400 900 1110 2380 640 1750 490 1120 600 1510 . 230 520 240 540 3-7 200 520 1030 2180 530 1520 500 1100 540 1380 430 860 240 530 3-8 500 1130 1430 3030 320 710 840 2170 540 1480 440 920 270 610 4-3 400 990 1930 4110 360 950 580 1220 450 1080 490 1030 250 560 X $ 278 668 951 2031 409 1020 523 1214 463 1153 466 986 279 651 4-2 500 1080 1000 2150 - - - - - - - - - — . 4—4 300 580 2480 5340 430 930 1260 3070 750 2050 1130 2420 - - 3-4 500 1090 790 1730 430 1020 460 1030 530 1220 360 860 - - X OjT 433 971 1423 3073 430 . 975 860 2050 640 1635 745 1640 - - X 317 730 1069 2292 . 413 1012 585 1366 495 1241 . 522 1117 279 651 1 Average Activity Radius 2 Standard Diameter 3 Spring-Summer-Fall ^Summer-Fall 63 Table 11. Home range size, in hectares, average activity radius and . standard diameter, in meters, for neckbanded mule deer on the Schafer Creek winter range, 1977-78. Animal Number Home Range (ha) Average Activity Radius . (m) Standard Diameter (m) Number of Observations Adult Females 2217 30 200 480 35 2277 60 300 580 39 2077 60 400 870 27 2157 40 200 570 21 2085 50 300 670 26 ■ 2075 50 300 760 22 2287 20 300 650 14 9 44 286 654 . 26 Adult Males 2045 10 . 200 380 20 2067 30 400 740 15 2237 20 200 470 19 X 20 267 530 15 Yearling Males 2207 70 300 780 39 • 2167 50 300 670 21 2087 40 . 300 770 25 X . 53 300 740 28 X Cf 37 283 635 23 . • X All 41 285 645 25 For the entire year (March 20, 1977 to April 27, 1978) the 2 radio-collared deer collectively ranged over an area of 49 km (standard diameter of 4.8 km) around a common geographic activ­ ity center . situated near the head of Jackrabbit Creek on the 64 upper limits of the winter range. This represents the minimum area required for the Schafer winter range herd to maintain itself. This minimum area, of course, increases with the inclusion of the marked animals known to have used areas east, of the Bridger Mountain divide during the summer of 1978. A closer representation of the true "herd range" size would be obtained with more marked animals. The overall mean spring home range size of 204 ha was partially dependent on the distance any particular deer migrated away from the winter range. Also inflating spring home range size was the tendency of deer to wander enroute to their respective summer ranges. Average activity radii for females and males was 951 m and 1423 m respectively (Table 10). These averages were much larger than those reported for deer associated with the Armstrong winter range (Appendix Table 26), but fall within the range of spring average activity radii reported by Hamlin (1974) and Mackie et al. (1976). Summer home ranges averaged 84 ha for females and 52 ha for two adult males (one radio ceased to function in May). Pac (1976) reported a mean home range of 107 ha for five radio-collared females associated with the Armstrong range for approximately the same annual time period. Other studies reported 311 ha for a female on a prairie-breaks area of Montana (Knowles 1975), and 40 ha for mule deer summering in the moun­ tains of western Montana (White 1960). Average activity radii for this time period were 409 m for females and 430 m for males. 65 Three adult females (chs 3-2, 3-6 and 3-7) moved off of their "normal" home range during June and returned by June 30. One of these females moved approximately one km west, another moved one km northeast arid the third moved over the Bridger divide as previously mentioned. None of these areas was used during the remainder of the summer. In all cases, these movements occurred at the time of expected fawn drop, and it is possible that they were associated with fawning. If this was the case, the fawns would have had to make substantial moves while still quite young, particularly that of the female crossing the divide. Dood (1978) reported fawns being able to make extended moves while very young on a prairie-breaks area. Mean home range sizes, average activity radii and standard diam­ eters increased for both males and females during early fall. By this time the growing season had ended, first snows had occurred, and deer made rapid short-term moves to areas that were snow free. Also the two radioed females that summered on the winter range may have been displaced during the first three weeks of hunting season. An adult female had ranged in the vicinity of an aspen patch since July where it was joined by another female on October 12. Both deer were relocated in this area October 18, four days prior to hunting season. . By October 27, after hunting began, both moved approximately .5 km north to a heavily forested north slope along Schafer Creek. On November 16 both had returned to the pre-hunting location. 66 For the summer-early fall period the average home range size was 173 ha for all radio-marked deer (females 178 ha, males 152 ha). Three deer (females 3-2 and 3-9 and male 3-4) which summered in pre­ dominately closed-canopy habitats, had significantly smaller home ranges (P<.05) (Fig. 6) and moved significantly less (P<.05) between relocations (Fig. 7) than eight others which summered on areas of broken timber cover. The restricted usage of a heavily forested area my one male caused the mean summer-early fall home range size of males, to be less than that for females; however, the average activity radius, standard diameter and mean distance between relocations were larger for males, (Table 10, Fig. 7 and Appendix Table 25); indicating males were more mobile within their home ranges than were females. Summer-early fall activity radii were larger than Pac (1976), Hamlin (1974) and Schwarzkoph (1973) have reported for deer in the Bridget Mountains (Appendix Table 26), while home range size may be more comparable (Pac 1976). All radio-collared females returned to the same summer areas in 1978 as used in 1977 (Pac pers. comm.). The average late fall home range size was 23 ha for females and 160 ha for males. .These were the smallest home ranges recorded for any season and probably reflect the directness of movement towards the winter range, concentration on holding areas and a limited number of relocations. As noted earlier, travel corridors were apparently much narrower in fall than in spring though the same general routes were 67 meters females --------males deer of closed canopy hbts all other deer Figure 7. Mean distance between relocations for radio-collared mule deer associated with the Schafer Creek winter range by season. followed. An adult, 5 year-old, male (ch 4-4) had a large fall home range of 290 ha, possibly due to rutting movements. This buck moved an average of 1.5 km between fall relocations (Appendix Table 25) and was in a different area of the Middle Cottonwood drainage at each relocation. Another older buck, about 8h years old (ch 3-4) did not exhibit such drastic fall movements averaging only .6 km between relo­ cations, with all being on or very close to its summer home range. Average activity radii increased, during late fall, from the means for the summer, but decreased from early fall (Table 10). Standard diameters also decreased after early fall. 68 Average home range size of radioed deer for the winter 1976-77 was 180 ha. Weather and. Snow conditions were very mild and data were obtained only during the last month of winter when deer were beginning to move about prior to leaving the winter range. During this period three males had an average home range of 387 ha, which was signifi­ cantly greater (Pc.05) than 111 ha for nine females. During the 1976- 77 winter, the largest (700 ha) and the smallest (60 ha) home ranges recorded were for males. Average activity radii and standard diameters were also significantly greater (Pc.05) for males. The winter of 1977-78 was more severe than the previous winter (Table I). Twenty-two marked mule deer (16 females, 6 males) had an average home range"of 48 ha. Male and female neckbanded deer and female radio-collared deer had home ranges of 37, 44, and 58 ha respectively (Tables 9 and 11). Home ranges of radioed females were significantly smaller (Pc.05) during the 1977-78 winter than the 1976-77 winter. There was no significant difference (P>.05) between neckbanded males and females; however, neckbanded does had significantly (Pc.10) smaller winter home ranges than radio-collared does. This was probably due to the low ability to observe and identify non-radio marked deer. Home ranges among radio-collared females which remained on the Armstrong winter range averaged 109 ha for winter 1976-77 and 87 ha during the winter 1977-78 (Youmans in prep.). Yearling males had significantly (P<.05) larger home ranges (53 ha) than adult males (20 ha). Adult 69 males were relatively sedentary on the winter range while yearling ■ . males were still associated and travelled with, female-fawn groups. Schwarzkoph's (1973) data for the Armstrong area indicated smaller average activity radii and home ranges for two yearling males than for two adult males, contrary to the findings of this study. All radio-marked females returned in 1977-78 to. the same portion of the winter range that had been used the previous winter. However, three (chs 3-2, 3-9 and 4-1) showed home range shifts during the course of the winter (Fig. 8). All three initially used the same area south of Schafer Creek where slopes are 20-25° with a west-southwesterly exposure. When snow depth reached 35—45 cm and became somewhat compacted, between January 20 and February 5, they moved approximate­ ly 500 m northwest across Schafer Creek, where slopes were steeper, more southerly in exposure, and snow accumulation was less. Gilbert et al. (1970) reported snow depth of 45 cm or more precluded range use and caused concentrations of mule deer in Colorado. By March 17, when snow cover was beginning to break, all.returned to re-establish home' ranges with activity centers within 100 m of their early winter activity centers. These movements were typical of virtually all deer which used the area between Schafer and Middle Cottonwood Creeks in early winter. From December 21 to the end of January, 276 deer were observed south of Schafer Creek prior to heavy snow accumulation. From February I to LEGEND Channel 4-1 Channel 3-2 Channel 3 -9 EZI 1 -Dec21 — -Jan 31 2 - F eb l — -M a r l6 3 -Mar 17 '— - Apr 30 Figure 8. Shifts of activity centers by three female mule deer on the Schafer Creek winter range 1977-78. 71 March 16 only 68 deer were observed on the area, with no deer recorded from February 20 to March 13. After snow melt had begun (mid-March) from March 17 to April 27, 212 deer were observed. Mackie et al. (1976) also described movement to a "primary winter use area" for the Armstrong winter range. The primary winter use area of the Schafer winter range extended from Schafer Creek north along the mountain front to approximately 1.6 km north of Bostwick Creek (Fig. 4) and included approximately 334 ha . There may be differences in range use by mule deer between the Armstrong and Schafer areas. Winter home range size may tend to be smaller on the Schafer winter range. Also deer on the Armstrong area generally had larger winter activity radii (Appendix Table 26) and. standard diameters indicating greater mobility within their home ranges. The smaller home ranges, activity radii and standard diameters observed on the Schafer winter range may be due to deer entering the winter on a higher nutritional plane, to lower winter range densities, or to the range better meeting the life requirements and reducing energy costs of the deer and thereby allowing them to survive on a smaller area. Hamlin (1974) reported fawns and females of the Armstrong area to enter the winter with omental fat reserves close to a critical level, and Pac (1976) reported substantial fall losses of fawns. The general steady increase in fawn ratios from late summer to early winter (Fig. 9) could be an indication that fall ranges and holding areas, on the Fawns : 100 Adults Fawns : 100 Females Males : 100 Females J 77 F Figure 9 . Observed fawn and male ratios on the Schafer Creek study area January 1977 - April 1978. 73 Schafer study area, were allowing deer to arrive on the winter range in good physical condition. Densities on the Schafer winter range were lower, and over winter mortality also appeared to be lower as was indicated by Mackie et al. (1976). These factors would indicate the Schafer winter range can better carry deer through the winter. Sanderson (1966) and Robinette (1966) stated that with better habitat requirements at hand home ranges should be smaller. It would follow that survivability would be enhanced. Habitat Use Monthly and seasonal estimates of mule deer usage of various habitat types, based on all observations and relocations of deer on the Schafer Creek study area, are presented in Tables 12 and 13. Table 13 also lists seasonal preference indicies for habitat types. These data were calculated by dividing the percentage of all observations and/or relocations in a type by the percentage of the total winter range (winter) or the total study area (spring, summer and fall) on which that type occurred. The combined observational (aerial and ground, marked and unmarked animals) and relocation data appeared to provide the best estimate of habitat usage. Frequencies of use of the various types, calculated as the following groups: mule deer observed from the air, from the ground, radio-collared animals, for all other deer observed, and the total Table 12. Monthly frequencies of use of habitat types and series by mule deer associated with the Schafer Creek winter range. Habitat Type* Jan 77/ Feb 77/ Mar 77/ Apr 77/ May June July Aug Sept Oct Nov Dec Jan 78 Feb 78 Mar 78 Apr 78 77 77 • 77 77 77 77 77 77 Agriculture 0/12 0/1 7/1 0/0 FEID/AGSM 0/2 0/1 2 FEID/AGSP 5/3 8/Tr 3/3 2/0 4 6 ARTR/AGSP 0/12 39/10 6/20 29/19 Tr 22 ARTR/FEID 51/14 29/9 17/25 14/24 2 2 I 7 15 PSME/AGSP 0/24 16/31 22/25 10/15 5 2 I I 21 PSME/FEID 30/39 0/40 45/22 16/24 22 8 11 9 13 5 15 22 PSME/SYAL-A 14/2 8/5 1/2 11/4 4 2 3 I 2 2 I PSME/SYAL-C 0/3 0/3 0/1 18/14 43 20 13 12 18 25 16 10 PSME/CARU 3 5 11 I 2 I PSME/CAGE 7 5 2 Tr PSME/VAGL 0/Tr 7 11 2 I 4 20 ABLA/VAGL 9 9 10 17 13 16 27 ABLA/CAGE I 2 I 18 2 5 ABLA/CLPS 38 35 43 24 22 2 ABLA/ARCO I I PIFL/JUCO - 2 4 7 10 22 I FEID 5/5 8/1 3/3 2/0 4 8 ARTR 51/26 68/19 23/45 43/43 2 2 I 7 37 PSME 44/68 24/79 68/50 55/57 84 46 47 32 34 38 54 54 ABLA 10 50 46 61 55 40 34 PIFL 2 4 7 10 22 I 1 See Table 2 for h.t. abbreviations 2 Percent of all observations and/or relocations of mule deer for that month Table 13. Seasonal frequency of use of habitat types and series and preference indices (see text) for mule deer associated with the Schafer creek winter range. Habitat Type1 Winter % 77/Winter 78 Pref.2 Spring % Pref. Summer % Pref. Early Fall % Pref. Summer- Early fall % Pref. Late Fall % Pref. Agriculture 4/ld Tr FEID/AGSM 0/1 (0/.47) FEID/AGSP 4/2 (.4/.2) 4 (2.86) ARTR/AGSP 14/16 (3.18/3.64) I (1.67) ARTR/FEID 22/18 (1.57/1.29) 2 (1.0) I (.5) I (.50) 8 (4.0) PSME/AGSP 15/26 (1.83/3.17) 5 (1.72) I (.34) I (.34) PSME/FEID 31/32 (2.50/2.58) 22 (3.1) 10 (1.4) 12 (1.69) 10 (1.41) 18 (2.54) PSME/SYAL-A 6/3 (.83/.42) 4 (.85) 2 (.43) 2 (.43) 2 (.43) 4 (.85) PSME/SYAL-C 5/3 (.14/.08) 43 (2.81) 14 (.92) 18 (1.18) 16 (1.05) 16 (1.05) PSME/CARU 3 (.86) 7 (2.0) I (.29) 5 (1.43) I (.29) PSME/CAGE 5 (1.47) I (.29) 3 (.88) PSME/VAGL 0/Tr 7 (.37) 4 (.21) 3 (.16) 4 (.21) 23 (1.22) ABLA/VAGL 9 (.65) 11 (.8) 18 (1.30) 14 (1.01) 25 (1.81) ABLA/CAGE I (1.0) I (1.0) 14 (14.0) 6 (6.0) I (1.0) ABLA/CLPS 38 (2.66) 19 (1.33) 31 (2.17) 2 (.14) ABLA/ARCO I (2.0) Tr PIFL/JUCO 5 (1.72) 12 (4.14) 7 (2.41) I (.34) FEID 4/3 (.33/.25) 4 (2.35) ARTR 36/34 (1.96/1.85) 2 (.77) I (.39) I (.39) 9 (3.46) PSME 57/64 (.84/.94) 84 (1.36) 43 (.69) 37 (.60) 41 (.66) 62 (1.00) ABLA 10 (.33) 51 (1.66) 41 (1.66) 51 (1.66) 28 (.91) PIFL 5 (1.72) 12 (4.14) 7 (2.41) I (.34) 1 See Table 2 for h.t. abbreviations 2 Habitat preference Index: % use % area 3 Percent of all observations and/or relocations of mule deer for that season 76 combined data. These calculations differed considerably within and between seasons (Table 14). Aerial observations and relocations of radio-collared deer indicated use of more types as well as generally greater use of heavily forested habitat types and areas than ground observations. However, the concentration of effort on radio-marked deer undoubtedly also influenced the overall observed distribution of habitat usage, and may have introduced some bias. For example, the intensive monitoring of five radio-collared does, which were closely associated on both the winter and summer range, may have exaggerated the importance of habitat types frequented by these animals and resulted in underestimation of types used by other deer on the study area. In general, it appeared that mule deer used all of the habitat types available, at least to some extent and/or at some time of the year. During the course of field studies, evidence of current deer use (tracks, beds, pellet groups, and/or foraging) was noted in all habitat types and on all parts of the study area during the June-October periods. The only exception was the Pseudotsuga menzies-t-i/Physoaavpus matvaeeus (PSME/PHMA) h.t. along the lower portion of Middle Cottonwood Creek, where some past use, but only very limited current use was recorded. Mule deer fecal pellet frequencies (Table 15), recorded incidental to habitat typing, also indicated use of all types with the exception of PSME/PHMA and Philadelphus lewisii/Symphovieavpos albus (PHLE/SYAL) h.t.s. No deer were directly observed in either of these Table 14. Seasonal frequencies of use of habitat types for mule deer associated with the Schafer Creek winter range. Data are percentages in each type as determined by aerial/ground observations and by relocation of radio-marked deer/all other deer observed. Habitat Winter Early Summer- Late Winter Type1 1976-77 Spring2 Summer Fall Early Fall Fall 1977-78 Agricul. 0/4 0/4 0/1 0/Tr 0/Tr 0/Tr 1/1 1/1 FEID/AGSM Tc/1 Fr/l FEID/AGSP 4/4 0/4 4/- 4/4 3/2 0/3 ARTR/AGSP 17/13 16/14 I/-3 0/1 17/15 20/15 ARTR/FEID 17/23 7/22 2/- 0/4 2/1 1/2 1/0 I/Tr 1/1 8/- 5/9 16/19 9/19 PSME/FEID 10/36 13/33 22/- 15/26 8/11 3/16 13/0 9A5 11/10 4/15 18/- 11/20 26/33 33/31 PSME/AGSP 15/5 13/15 5/- 1/9 1/2 2/1 1/1 1/Tr 2V26 27/25 PSME/SYAB-A 12/4 9/6 4/- 1/6 1/3 1/3 2/4 2/2 1/3 1/3 4/- 2/4 4/2 Tr/3 PSME/SYAB-C 21/2 41/1 43/- 39/44 17/12 17/10 17/22 19/14 17/13 18/11 16/- 30/12 9/1 9/2 PSME/CARU 3/- 7/0 4/9 4/8 1/0 1/1 2/8 3/6 I/- 0/1 PSME/CAGE 3/6 5/5 1/4 0/2 2/6 3/3 PSME/VAGL 7/- 17/0 6/2 4/2 4/0 5/0 5/1 4/1 23/- 25/22 0/Tr 0/Tr ABLA/VAGL 9/- 10/7 17/6 18/7 19/13 16/21 18/6 17/10 25/- 18/27 ABLA/CLPS 1/0 36/41 37/42 16/39 22/13 26/40 30/30 2/- 5/1 ABLA/ARCO 1/0 1/0 1/0 1/0 Tr/0 ABLA/CAGE I/- 1/0 2/1 4/0 15/13 19/7 8/2 10/2 I/- 0/1 PIFL/JUCO 2/7 5/5 13/4 8A6 7/7 6/8 I/- 5/0 ^See Table 2 for h.t. abbreviations. 2Spring aerial only. 3Late fall, aerial and ground combined. 78 Table 15. Frequency of occurrence of mule deer fecal pellets among 2 x 5 decimeter plots and constancy among sites sampled as representative habitat types on the Schafer Creek study area. FEID/AGSM1 FEID/AGSP ARTR/FEID ARTR/AGSP PHLE/SYAL PSME/FEID (3) (3) (5) (I) (5) .10/.50 .63/1.0 .47/1.0 .44/1.0 0/0 .35/.80 PSME/AGSP PSME/CARU PSME/CAGE PSME/SYAL-A PSME/SYAL-C PSME/VAGL (6) (I) (2) (2) (7) (5) .45/1.0 .05/1.0 .10/1.0 .30/1.0 .17/71 .04/.60 PSME/PHMA ABLA/CLPS ABLA/CARU ABLA/CAGE ABLA/VAGL PIFL/JUCO (2) (2) (2) (I) (4) (2) 0/0 .30/.50 .12/1.0 .30/1.0 ,15/1.0 no data 1See Table 2 for habitat type identifiers. 2Number of sites. types or in the AMes lasiocappa/calamagrostis .vubesoens (ABLA/CARU) h.t. Seasonal Habitat Use Winter (Jan. I-Apr. 30, 1977 and Dec. 21-Apr. 30, 1978) Pseudotsuga menziesii/Festuoa idahoensis (PSME/FEID) was the most used h.t. and Artemisia tridentata/Agropyron spioatum (ARTR/AGSP) was the most preferred type for both winter periods. Together the PSME/ FEID, Pseudotsuga menziesii/Agropyron spioatum (PSME/AGSP), Artemisia tridentata/Festuoa idahoensis (ARTR/FEID) and ARTR/AGSP h.t.s were the most preferred and received the majority of use (82% in 1976-77 and 94% in 1977-78) by mule deer through the winter (Table 13). During the 79 relatively mild winter of 1976-77 recorded usage was highest in the PSME/FEID type followed by the ARTR/FEID, PSME/AGSP and ARTR/AGSP types. Artem-Lsia tridentata/Agropyvon spioatum appeared to be the most preferred h. t. followed by PSME/FEID, PSME/AGSP and ARTR/FEID. Aerial and telemetry observations (Table 14) indicated that the over­ all importance of PSME/FEID and ARTR/FEID may have been overestimated in ground observations; however,,the former were not made until the last month of winter. Also no deer were observed using the PSME/FEID type during February, a particularly snow-free period. The highest use of ARTR/FEID was recorded in January, after which usage steadily de­ clined through the remainder of the winter. Use of Festuoa idahoensis/ Agropyron spioatum (FEID/AGSP) was relatively minor and appeared to decrease after a peak in February. The winter of 1977-78 was more severe; though recorded snowfall was less, colder temperatures (Table I) resulted in greater snow accumulation on the winter range. Southerly and westerly slopes had 50 percent, or less snow cover for only a maximum of 69 days as com­ pared to 85 such days during 1976-77. As in 1976-77, PSME/FEID had the greatest recorded use followed by PSME/AGSP, ARTR/FEID and ARTR/ AGSP types (Table 13). Also like the previous winter, ARTR/AGSP appeared to be the most preferred type, followed by PSME/AGSP, PSME/ FEID and ARTR/FEID. The first three of these types received approxi­ mately equal use, while ARTR/FEID was used somewhat less, as deer were 80 arriving on the winter range in December. Recorded use of the ARTR types decreased in January and was the lowest of the winter period in February, the month of most persistent snow cover. This was followed by increased usage in March, as snow cover began to break. April use . was about the same as that for March. Observed use of PSME/FEID and PSME/AGSP increased in January and peaked in February, followed by a decrease in March. Usage of PSME/FEID remained relatively steady in April while PSME/AGSP further decreased. It appears that there was a compensating relationship between the ARTR and two PSME types. During both winters the Pseudotsuga mens-iesti/Symphor-iaappos dVbus- calamagrostis rubeseens phase (PSME/SYAL-C) h.t. was of relatively minor importance until April, when it received its greatest use. In 1976-77 importance of this type increased substantially with the ini­ tiation of aerial flights; however, earlier ground surveys indicated a moderate degree of use, for feeding and bedding,, had taken place. In 1978, prior to April, snow accumulation in this type was substantial, often 45 cm or more. Ground surveillance indicated that any use which occurred was primarily for bedding along the upper slopes, and travel from one south slope to another. The most notable difference in habitat usage between winters was the apparent shift in 1977-78 to greater usage of PSME/AGSP and less use of ARTR/FEID and greater preference of PSME/AGSP and less of PSME/ FEID than during the milder winter of 1976-77. Also minor habitats 81 received a greater amount of the total usage in 1976-77. The ARTR/ FEID h.t. occurs on sites which tend to hold snow, probably explaining the overall decline of use during 1977-78, particularly in January and February. On the other hand areas occupied by the PSME/AGSP type were among the first to lose snow cover and possibly were the warmest sites on the winter range, along with large rock outcrops, many of which were on PSME/AGSP sites. Although these factors may explain the observed difference in use between ARTR/FEID and PSME/AGSP,. they do not explain the increase in preference for PSME/AGSP compared with PSME/FEID, which supports more herbaceous vegetation (grass and forbs) as well as browse (Table 5). It is possible, however, that the PSME/ AGSP h.t., with less snow, provided greater availability of green succulent forage. Mackie et al. (1976) indicated habitat use may shift during winters due to availability of green grasses. Overall, snow cover appears to have been the major factor, influencing changes in habitat use between the two winters. Spring (May I-June 5) Dispersal from the winter range was under way and use of closed- canopy h.t.s (PSME/SYAL-C, Pseudotsuga menziesi-L/Vaeci-n-Lum globulave (PSME/VAGL) and Abies lasioearipa/Vaociniim globulave (ABLA/VAGL)) increased to a total of 59 percent of the recorded use. The most used h.t. was PSME/SYAL-C with PSME/FEID being the only other type of singu­ lar importance (Table 13). The two types with a Vaoainium understory 82 (PSME/VAGL and ABLA/VAGL) combined, ranked third in habitat use. The PSME/FEID h.t. had the highest recorded preference index, followed by FEID/AGSP, PSME/SYAL-C and PSME/AGSP. This indicated a preference for more open h.t.s on which "green-up" was more advanced, providing better foraging conditions for deer leaving the winter range. Bertram and Remple (1977) and Pac (1976) attributed use of open areas during spring mule deer migrations to advanced plant phenology and improved forage conditions. Summer (June 6-August 26) The AMes Ias-LoeaTpaZClematis pseudoaVpina (ABLA/CLPS) h.t. appeared to be the most used type, receiving twice the recorded use of any other habitat type for this time period. This may be an over­ estimation, due to distribution and concentration of marked animals in the head of Middle Cottonwood Creek (Fig. 4) where the majority of summer observations took place. Aerial and telemetry-aided observa­ tions also indicated that use of this type may have been overestimated (Table 14). Pseudotsuga menziesH/Symphovieorpos albus-Calamagvost-Ls vubescens phase and ABLA/VAGL h.t.s were ranked second and third in habitat use. Receiving highest preferential use was ABLA/CLPS followed by Pseudotsuga menziesi-L/Calamagrostis rubeseens (PSME/CARU), Pinus flexilis/Juhipeims eonmunus (PIFL/JUCO) and Pseudotsuga menziesiiZ Carex geyeri (PSME/CAGE); all of these were relatively open timber types with grass-forb understories. Use of PSME/SYAL-C decreased 83 substantially after June, possibly reflecting the general eastward and upward movement of the herd as it left the winter range. Use of PIFL/JUGO steadily increased through early summer and into October. Early Fall (Aug. 27-Nov. 14) A distinct shift in habitat usage was observed. Abies lasiocarpa/ Clematis pseudoalpina continued to be the most used h.t., but its usage decreased to half of that for summer. The ABLA/VAGL and Pseudotsuga mensiesii/Symphoriearpos albus - Agropyron spioatum phase (PSME/SYAL-A) h.t.s increased in importance to approximately the same level of use as ABLA/CLPS. A dramatic increase in the use of Abies lasioearpa/Carex geyeri (ABLA/CAGE) was observed; however, much of this was due to use by radio-collared animals (Table 14). The PIFL/. JUCO received its highest usage of the year. The ABLA/CAGE h.t. appeared to be highly preferred, while a high degree of preference was also recorded for PIFL/JUCO. The PSME/FEID h.t. ranked third in preference. Importance of ABLA/CLPS h.t. decreased steadily after August (Table 12).' Dramatic movements of deer out of this type onto ABLA / CAGE were observed following the first two snowstorms of the year, which resulted in a depth of 3 cm or more above 2130 m. This probably explains the high degree of use and preference of the minor ABLA / CAGE h.t. (I % of the study area) and may be an example of weather and/or behavior caused habitat selection rather than forage related selection. After snow had melted on.the 84 ABLA/CLPS h.t. deer returned to their "normal" home ranges and types used prior to the storms. Radio-collared deer summering be­ low 2072 m did not appear to experience this pattern of habitat usage.. Beirtram and Rempel (1977) documented the same pattern of movement for mule deer of the North Kings deer herd in California. The general increase in importance of forested types with shrub understories after August apparently reflected changes in plant phenology which resulted in a shift in food habits from succulent forbs to browse. Wilkins (1957) documented this shift in food habits, while Pac (1976). reported a similar pattern of habitat use for at least some deer during late summer-early fall (August-October). The general shift in early fall, after the first killing frost and snow, was recorded not only as a change in habitat use (Table 13), but also as changes in fawn:female and male:female ratios among deer observed on the study area (Fig. 9). The decrease in the male:female ratio after August may have been due both to decreased usage of the high elevation ABLA/CLPS h.t. by males and a slight increase in numbers of females with fawns using this type. In August, 72 percent of all males were recorded in the ABLA/CLPS h.t. This decreased to 27 percent in September. The actual number of males recorded in all h.t.s de­ creased by 24 percent while female numbers remained constant. Radio- collared females as well as males became less visable during this time period (Fig. 10); however, males decreased 72-85 percent in the ABLA/CLPS 85 Figure 10. FEMALES MALES 77-7876—77 Percent visual relocations for radio-collared mule deer on the Schafer Creek winter range 1977-78. 86 h.t. as females and fawns decreased only 26-33 percent. If radio- collared deer are representative of all deer on the area, this would suggest that movement of males off of this type (high elevation habitats) rather than a substantial increase in numbers of females with fawns in late summer-early fall, was the cause of the changes. Heavy use of high elevation habitats by male mule deer in the Bridger range has been reported by Schwarzkoph (1973), Hamlin (1974) and Pac (1976). Hamlin and Pac also documented a decrease in the percentage of males observed and an increase in the percentage of females with fawns observed in these habitats in late summer. They attributed the change to ,a movement of females with fawns onto these types from either the east side of the Bridgets (Hamlin 1974) or from lower elevations of the west slope (Pac 1976). Late Fall (Nov. 15-Dec. 20) Habitat use during this period reflected movement of deer from summering areas towards the winter range and the arrival of the first deer onto the winter range. The most important h.t.s for the period were timbered with Vaoeini-vm understories. The PSME/VAGL and ABLA/ VAGL h.t.s, together, accounted for 48 percent of the late fall usage. The PSME/FEID and PSME/SYAL-C types were also important (Table 13). Habitat types typical of the winter range began to receive use; for example, ARTR/FEID received eight percent of the total use. This was the highest recorded since the previous winter. Because of its minor 87 occurrence on the study area as a whole, ARTR/FEID received the highest preferential use of the season. Pseudotsuga menztes-Li/Festuea 'Lddhoensis3 which also received more use than at any time since spring, ranked second in preference. Closed-canopy habitats accounted for 64 percent of all observations and/or relocations. Bertram and Remple (1977) also reported high use and importance of heavy cover habitats during the fall period. Yearlong trends in use and preference of forest and shrubland series (Tables 12 and 13) indicated that h.t.s of the Pseudotsuga menzies-ii series were the most important for use by mule deer during the winter, spring and late fall periods and ranked second through the summer and early fall. It was the second most preferred habitat for summer, early fall and third in spring and late fall. Pac's (1976) observations indicated that Pseudotsuga Menz-Lesii types collec­ tively were most important for summer as well as spring and fall use on his study area on the north end of the Bridger range. The Pinus flexilis series was important only in summer and early fall when it ranked third in use and first in preference. Types of the Artemisia tridentata series were used to only a minor extent during all seasons except winter, when they ranked second in use and first in preference. Despite minor use ARTR was also the most highly preferred series for late fall. Due to its limited occurrence, the Festuoa idahoensis series was of minor importance in use over the entire year; however, it had 88 the highest habitat preference index for spring. Data were not adequate to indicate possible differential use of habitat types by sex and age of the animals generally. However, there was evidence that females with fawns may have occurred with greater frequency on certain h.t.s over others that were available during summer but not winter. Numbers of fawns:100 females (100 adults for winter) recorded by h.t. in which more than ten females and/or fawns were observed, during summer, fall and winter periods are shown in Table 16. To test for possible differences in fawn:female ratios among habitat types it was assumed that the number of fawns per 100 females would be the same for all h.t.s if deer were evenly distributed. Chi-square: tests were run for each season using the mean number of fawns:100 females for the season as the expected number of fawns:100 females for that season. Chi-square values (Table 16) show a significant deviation (P<.05) in fawn numbers observed among h.t.s for all seasons except winter 1977-78. The high value for the winter of 1976-77 may be partially due to observer inexperience in classifying deer and the open conditions which may have permited greater habitat selection than would normally occur in winter. Over the summer-early fall period substantially more fawns per female than average were observed in the PIFL/JUCO h.t. than in any other type. Greater than expected numbers of fawns:100 females were Table 16. Number of fawns per 100 females recorded by habitat type on the Schafer Creek study area. Habitat Early Summer- Late Winter Winter Type1 Summer fall early fall fall 1976-77 1977-78 Agric. (O)Z 3 (0) - - 34 4 FEID/AGSM — 36 FEID/AGSP — — — — 25 36 ARTR/AGSP _ — — (0) 36 31 ARTR/FEID (25) - (25) 67 24 42 PSME/FEID 22 25 23 40 27 31 PSME/AGSP (0) - CO) - 16 37 PSME/SYAL-A (133) - (100) (75) 10 23 PSME/SYAL-C 19 13 18 29 0 30 PSME/CARU 5 - 5 - - - PSME/CAGE 15 (0) 14 - - - PSME/VAGL (ID (0) 6 31 - (0) ABLA/VAGL 18 13 15 57 ABLA/CLPS 11 12 12 (0) - - ABLA/CAGE (0) 7 6 (0) - - PIFL/JUCO 42 29 34 - - - Season X X = 18.9 X = 16.5 X = 15 X = 44.8 X = 19.71 X = 33.33 Chi-square Values X2 3= 43.21 Xz= 26.41 X2= 47.74 X2= 24.66 X2= 43.7 X2= 6.98 n= 7 n= 6 n= 9 n= 5 n= 7 n= 9 1 See Table 2 for habitat type identifiers 2 Ten or fewer females and/or fawns observed, not included 3 No females or fawns observed 4 Fawn: 100 adults for winter 90 recorded for PSME/FEID. This may indicate a high preference by does with fawns for these types, or that does with home ranges on these types are more productive or successful in rearing fawns. Approximate­ ly the same number of fawns as the summer-fall means were observed in . ABLA/VAGL and PSME/SYAL-C, two h.t.s in which deer are hard to observe. These two types make up nearly 30 percent of the study area and prob­ ably contribute the major portion of the herd's annual fawn crop. The most used and preferred h.t. of the summer-early fall period, ABLA/CLPS, showed fewer fawns than the average observed. Hamlin (1974), as previously noted, reported that higher elevation habitats were used to a lesser extent by does with fawns than by females without fawns or by males. Fewer than the mean number of fawns were also recorded for the.ASIA/CAGE and PSME/CARU h.t.s. For late fall greater than average numbers of fawns were observed in the ABLA/VAGL and ARTR/FEID h.t.s, while about average numbers were recorded for PSME/FEID. For both winters, higher than average numbers of fawns were observed to use the ARTR/FEID type and lower than average used PSME/SYAL-A. The highest number of fawns:100 adults was recorded for ARTR/AGSP, a highly preferred h.t., and ARTR/FEID for the winters of 1976-77 and 1977-78, respectively, indicating this shrub series may be important to winter fawn survival. The even winter (1977-78) distri­ bution of fawn ratios among h.t.s may be due to mule deer behavior manifested as group home ranges. Though females with fawns may prefer 91 certain "better" h.t.s the high densities of mule deer on the winter range may preclude this. Food Habits Information on mule deer food habits during early summer was obtained by examination of seven feeding sites. Winter food habits were derived from 17 feeding sites and contents of five rumens. Per­ centages of plant use by taxon and forage class (grass, forbs, browse) and the relative availability (frequency of occurrence and canopy coverage) of plant taxa utilized in various habitat types are presented in Table 17 for early summer and Table 18 for winter 1977-78. Rumen sample data are listed in Table 19. These data indicate that utilization by general forage class was similar to that reported for deer associated with the Armstrong winter range during the same time periods (Wilkins 1957, Schwarzkoph 1973, Hamlin 1974, Pac 1976 and Morton 1976). Summer Forbs were the major forage class used during, early summer 19,77, comprising 88 percent of total utilization. Browse accounted for 11 percent and grass use was minor (Table 17). Utilization of 42 species was documented;. however, the two most important species at each of the sites comprised an average 67 percent of the total use at each. Only two species, northern bed straw and Nuttall violet (V-LoZa nuttaZZii- Table 11. Percent use (U)/frequency of occurrence (0)/canopy coverage (C) of plant taxa used at summer feeding sites by habitat type. ■ FORAGE CLASS I Species Habitat Typei Combined Total . U ofPfSME/C u/o/c PSME/FE1D u/o/c ABLA/CLPS U/O/C ABLA/VAGL U/O/C . P1FL/JUC0. U/O/C GRASS Agropyfon sps 2/.2/ T T Phleum alpina 2/.2/ 2 - T TOTAL GRASSES 4 I FORBS Agoseris gIauca 2/ T/ T - 12/.17/ 9 .4/ .1/ T 7/.5/10 .3 Aquilegia flavescens 12/ .2/ 7 8/.2/ 4 . 4 Arnica cordifolia 3/ .5/16 3/ .5/ I . I Arnica spp 17.2/ 3 T Aster conspicuus I/.05/ 2 ' T Astragalus spp 2/ .1/ I 2/ .7/14 , I/.3/10 , I Balsamorhiza sagitata 71/.2/10 9/ .4/12 I/ T/ T 8 Besseya Wyomingensis ■ 6/ .3/ 2 • T Castilleja miniata 54/ T/ T ■ I Comandra umbellata 15/.I/ 2 I Delphinium bicolor 2/ T/ T . T Epilobium angustifolium 2/ T/ T T Fragaria virginiana 13/.2/ 3 2 Galium boreal 27.25/ 7 8/i37/14 38/1.0/15 8 Geranium viscosissium 3/ .4/ 8 17/.3/3.3 3 Helianthella uniflora 18/ .3/12 . 4/ .8/21 3 Hieracium cynoglossoides 42/.25/ 8 12 Lupinus spp 6/.3/ 9 20/ .3/ 6 . 6 Osmorhiza chilehsis I/ .2/4 2/ .5/6 T Pedicularis paysoniana " 3/ T/ T T P. racemosa 11/.25/ 7 3 Solidago missouriensis 26/.15/ 4 8 Table 17. (continued) FORAGE CLASS Species Habitat Type1 Combined Total U ARTR/FEID U/O/C PSME/FEID U/O/C ABLA/CLPS . U/O/C ABLA/VAGL U/O/C PIFL/JUCO U/O/C FORBS (continued) Taraxacum officinale I/.8/19 ■ T Thalictrum occidentale 7/.34/25 2/.3/ 7 3 Townsendia parryi 6/ T/ T 16/ .6/ I I/ T/ T I Trifolium haydenii 44/ .4/18 ' 11 Valariana spp ■ 2/ T/ T T Viola nuttalli 26/ .4/ 4 27/ .5/ 2 6 Unidentified forbs 30/ .2/ T 2/ T/ T I TOTAL FORBS 100 99 90 93 . 54 88 BROWSE Ribes spp 2/.I/ 2. T Rubus parviflorus 9/ .3/ 3 I Shepherdia canadensis 37/.I/ 2 , - 5 Spiraea betulifolia 11 .9/11 I Symphoricarpos albus I/ T/ T 23/ .8/15 3 TOTAL BROWSE I 10 7 43 11 Instances of use 100 415 585 H O 201 1411 1See Table 2 for h.t. identifiers. Table 18. Monthly and total winter use In percent, by habitat type, of plant taxa which achieved greater than I percent uae In one or more months __________or habitat types. --------------------------------------------------------- ---------- - -------------------------------------------------- Habitat Type1 Month FOKAGE CLASS Number of sites Species Instances of use GRASS Agropyron splcatim Bromus carlnatus B. tectorum Bromus spp Carex geyerl Festuca ldahoensis Koeleria cristate Poa pratensia P. sandbergii Poa spp____ ""TOTAL GRASS ART*/AGSP J F M Wtr J PSME/AGSP F M Wtr PSME/FEID J F M Wtr ARTR / FEID F PSME / STAL-C F PSME / SYAL-A F Agricul­ tural J J Combined Totals2 F M Wtr 1 1 2 4 337 1491 904 2732 3 1364 I 2 546 1318 6 3228 I I I 656 1332 798 3 2686 I 1418 I 1201 I 1039 I 5 812 2357 6 5 16 6927 3020 12304 I 27 18 15 4 27 24 18 25 T3 13 13 9 2 10/.8" 11/.83 18/1.0 13/.88 8 3 I T I I 2 I T T 14 5 2/.17 I/. 06 46 15 8/.17 3/. 06 10 3 42 15 19 8 3 3 15/. 33 11/.6 9/. 31 3 I I T 4 I 3 215 I/.17 3/.6 I/.19 I/. 06 3 I 9 3 2 2 I 2 I/.2 T/.17 5/.6 2/. 31 3 I I I/.17 T/.2 I/.13 I 29 34 21 5 86 62 51 27 46 26 33 14 3 5 21 11/.8 30/1.0 41/1.0 27/.94 FORBS Achillea millofolium T T I T 2 T I I/.2 T/.17 T/.4 I/.25 2 I I/.4 T/.13 Balsamorhiza sagittata 9 T 2 4 15 I I 6 3 3 2 2 9/.8 I/.5 2/.8 4/. 69 Cerastium arvense I T T I I T T I/.6 T/.2 I/.25 Chrysopsis villosa 3 11 5 10 I 4 25 3 10 13 13/.8 I/. 17 7/.6 7/.5 Cirsium undulatum I I I I T/.2 T/.17 T/.2 T/.19 Lactuca serriola I T 3 T/.17 I/. 06 Lupinus spp 2 I I/.2 T/.06 Melilotus officinales 3 10 4 26 I/. 17 3/.2 I/.13 Tragopogon dubius 4 I T I 5 2 T T T/.4 T/.5 3/.6 I/.5 Urtica dioica 4 I/. 17 T/.06 Unidentified forb I T I T I _ I A 2 _ I/.6 I/.25 TOTAL FORBS 13 T 19 11 29 6 25 20 31 3 12 15 3 4 T 50 24/.6 3/1.0 19/1.0 15/.94 BROWSE Acer glabrum I T 48 7 T 9/. 33 T/.2 3/. 19 Amelanchier alnifolia T T 22 4/. 33 I/. 13 Artemisia tridentata 82 64 48 61 9 3 38 30 62 43 66 23 50 43/.8 39/.83 37/.4 40/.69 Juniperus scopulorum 4 I 49 6 18 14 5 18/.80 2/. 17 2/.2 7/.31 Philadelphus levisii 2 I 4 I 2 13 I/.2 3/. 33 I/.2 2/.25 Populus tremuloides I T T T/.20 T/.17 T/.13 Prunus virginiana I 5 2 T T 6 15 I/. 40 2/.33 I/.19 Pseudotsuga menziesii I T 5 2 15 T/.2 7/.33 3/.19 Ribes cereum 2 I 3 3 2 2 I 2/.6 T/.17 I/.2 I/. 31 Rosa spp 3 I T 3 I 5 T/.4 2/.5 I/.25 Symphoricarpos aIbus I T 2 I 4 T/.2 ■ 3/.5 I/.25 TOTAL BROWSE 86 69 48 68 65 8 l2 28 44 51 6i 52 85 54 96 20 65/1.0 6V/1.0 TITTS* 58/.88 MOSS I T I T/.2 T/.17 T/.13 1For habitat type names see Table 2. 2Does not include agricultural site. ^Trace amount <.5Z use. 4Percent of total use/frequency of use among feeding sites. 5Total of approximate equal uae of Poa pratenaie, Agropyron cristotisn, and Triticum aestiuum. VD 95 Table 19. Winter 1977-78 food habits determined from five rumen samples collected from predator involved mortalities. FORAGE CLASS February (3)1 March (I) April (I) Seadon Species % Vol/Freq2 % Vol % Vol % Vol/Freq GRASS .22 / 1.0 12 75 36 / 1.0 FORBS 3 / 1.0 4 5 4 / 1.0 BROWSE Artemisia tridentata 64 / 1.0 53 18 45 / 1.0 Berberis repens' I / .33 T3/ .2 Cornus stolonifera T / .33 T / .2 Juniperus scopulorum 8 / 1.0 28 I 12 / 1.0 Populus spp T T T / .4 Pseudotsuga menziesii 2 / 1.0 3 I 2 / 1.0 Pershia tridentata T / .33 T / .2 TOTAL BROWSE 75 / 1.0 • /CO 20 61 / 1.0 LICHEN . T / .33 T / .2 1Number of rumen samples. ^Frequency of occurrence among rumens. 3Trace5 less than .5 percent. Pursh.), were recorded twice as one of the two most important forages utilized at a site. This plus the fact that there was no evident correlation between the degree of use of a particular forage species and either frequency of occurrence (r=.10) or canopy coverage (r=.08), would indicate a wide range of individual forage preference among mule deer. Other plants frequently receiving major use were pale agoseris (Agosevis glauea (Pursh.) D. Dietr.), yellow columbine (Aquilegia flaveseens S. Wats.), arrowleaf balsamroot and lupine. 96 Winter During winter browse, grass and forbs received 58, 27 and 15 per­ cent, respectively, of the total utilization at feeding sites (Table 18) and comprised 61, 36 and 4 percent, respectively, of rumen contents (Table 19). Greater occurrence of grass and less of forbs in rumen contents as compared with feeding site data was also noted in data of Wilkins (1957) and Schwarzkoph (1973), and could be due to greater digestibility of forbs (Snider and Asplund 1974 and Short et al. 1974) and/or for the tendency of rumen analyses to emphasize the occurrence of coarse materials (Bergerud and Russel 1964). Green forage was available throughout the 1977-78 winter and may have influenced overall feeding habits. Succulent grasses and/or forbs made up 13, 16 and 56 percent of the total recorded utilization for January, February and March, respectively (Table 20). During both the 1976-77 and 1977-78 winters, many deer were observed feeding on and about large steep rock outcrops which appeared to absorb solar radiation, creating warm microclimates and allowing plant growth throughout the winter. Schwarzkoph (1973) reported deer using green grass on snow-free slopes as early as January on the Armstrong winter range. Big sagebrush was by far the major species.utilized in all h.t.s in which feeding sites were examined, except the PSME/AGSP h.t. where it was of limited availability. It provided approximately 40-45 percent of the diet and accounted for nearly 70 percent of the browse used. Table 20. Mean percent of all instances of use which were green grass. and/or forbs at feeding sites by habitat type. Habitat Typel . Combined Month Agric . PSME/AGSP. PSME/FEID ARTR/AGSP ARTR/FEID Totals Jan. 20 I 52 0 _2 13 Feb. 80 0 0 15 . 16 Mar. - 86 26 CM