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Habitat use patterns and associated movements of white-tailed deer in Southeastern British Columbia Smith, Christian Arthur 1977

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THE HABITAT USE PATTERNS AND ASSOCIATED MOVEMENTS OF WHITE-TAILED DEER IN SOUTHEASTERN BRITISH COLUMBIA by CHRISTIAN ARTHUR SMITH B.S., University of Alaska, 1973  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of Zoology)  We accept this thesis as conforming to the required standard  THE UNIVERSITY OF BRITISH COLUMBIA February, 1977 • © C H R I S T I A N A. SMITH, 1977  In  presenting  this  thesis  an a d v a n c e d d e g r e e a t the I  Library  f u r t h e r agree  for  scholarly  by h i s of  shall  this  written  in p a r t i a l  the U n i v e r s i t y  make  it  freely  that permission  thesis  for  It  financial  of  British  gain  Columbia  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  /£  ^J.  Columbia,  British  J ? ? ? -  for  the  requirements  reference copying of  I agree and this  shall  that  not  copying or  for  that  study. thesis  by t h e Head o f my D e p a r t m e n t  is understood  of  The U n i v e r s i t y  of  for extensive  permission.  Department  of  available  p u r p o s e s may be g r a n t e d  representatives.  fulfilment  or  publication  be a l l o w e d w i t h o u t my  ABSTRACT  This study of deer ecology was conducted from January, 1975, to May, 1976, in the East Kootenay region of British Columbia.  Plant  communities within the annual range of a population of white-tailed deer were classified into habitat types and analysed for species composition, productivity and forage utilization.  Relative levels of use of winter  habitat types were determined from track and pellet group counts.  Use  of summer habitats was determined subjectively from ground and aerial surveys.  Movements associated with changes in habitat use and season  were documented by observation and radio tracking of marked individuals. Forest succession over much of the winter range was found to have resulted in substantial decreases in availability of herbaceous and deciduous browse, causing quantitative and qualitative changes in the diet of this deer population.  To compensate for this situation deer  feeding activity was concentrated in the open habitat types which provide maximum quantities and qualities of forage.  However, snowdepth in one  winter was found to reach levels which prevented deer from exploiting these areas and concentrated them in areas where a maturing overstory reduced snowpack.  The impact of concentration in shelter types with  consequent reduction of available food, compared to a very mild winter, was an apparent 30% reduction in the juvenile:100 adults ratio the following spring. Summer distribution also appeared to be affected by large-scale forest succession which has produced a pattern of widely scattered, small openings.  These were found mainly along water courses or rocky  slopes at mid- elevations on the west side of the Rock Mountain Trench. Deer density was low throughout the summer range, but preference was observed for the open areas just described. Spring dispersal from the winter range was related to snow melt and green-up of vegetation, particularly cultivated alfalfa fields.  Summer  home ranges were relatively small and summer movements limited.  The  average distance travelled to summer range by nine deer was greater than that reported elsewhere in the literature and may be related to summer range habitat condition. lasting snowfall.  Fall movements were apparently stimulated by  Although density of deer on the winter range varied  greatly between years, home range loyalty was found to be relatively high.  ii  TABLE OF CONTENTS Abstract  . . .}  i  L i s t of Tables  iv  L i s t of Figures  v  Acknowledgements . . . . . . . .  vf  Introduction  1  Study Area .  4  Winter Range  4  Summer Range  6  Methods  8  Analysis of Winter Range Habitats  8  Analysis of Summer Range Habitats .  13  Weather  14  Use of Winter Range Habitats  15  Use of Summer Range Habitats  21  Individual Movements Results  . . . 21  .  23  Analysis of Winter Range Habitats  23  Analysis of Summer Range Habitats . .. . •  44  Weather .  54  Use of Winter Range Habitats  60  Use of Summer Range Habitats  77  Individual Movements  7 8  Discussion  $5  Conclusions  .110  Literature Cited Appendix I:  1  Individual Movements Accounts . . ii i  1  3  H8  List of Tables Table I II,  Page Proportions of habitat types on the Wolf Creek winter range  25  Species composition of the herbaceous layer: winter range  III IV  Available shrub browse:  VI VII VIII IX X  winter range  29  Annual production of browse in g/m by shrub 2  species: V  27  winter range  Shrub species utilization: Forest composition: Available f i r browse:  30 winter range  31  winter range  33  winter range  36  Species composition of shrub association: summer range Overstory composition of immature forest association: summer range . . . . Understory composition of immature forest association:  45 50  summer range  53  XI  Track count analysis:  winter 1974-75  XII  Track count analysis:  24 January, 1975  62  XIII  Track count analysis:  winter 1975-76  63  XIV XV XVI  .  .  .  . 61  Pellet group count analysis  64  Age ratios of deer observed in spring (15 April to 30 May) 1975 and 1976 Live-trapping results ,  69 79  List of Figures Figure  Page  1.  Study area  2.  Pellet group/vegetation transect design for sampling on the winter range  12  Vegetation macroplot layout for sampling on the summer range  12  Track count route and pellet group/vegetation transect locations  16  Distribution of habitat types on the Wolf Creek winter range  24  Shrub cover in the shrub association: summer range  46  Shrub cover in the immature forest association: summer range  52  3. 4. 5. 6. 7. 8. 9. 10. 11.  5  Three-day average minimum temperatures: December to A p r i l , 1974-75 and 1975-76 . • Average daily hours of direct sunlight: to A p r i l , 1974-75 and 1975-76 Total monthly snowfall: and 1975-76  • • • • 55  December  December to A p r i l , 1974-75 .  57 58  Depth of snow on the ground at month's end: December to A p r i l , 1974-75 and 1975-76  59  12.  Seasonal pattern of use of alfalfa fields, 1975  66  13.  Daily pattern of use of alfalfa fields, 1975  67  14.  Degree of selection of four winter habitat types: 1974-75 and 1975-76  71  15.  Sightings of marked deer, February 1975 to May, 1976 . . . . 82  16.  Relationship between canopy closure and annual production of preferred shrub forage Relationship between canopy closure and utilization of preferred shrub browse .  17.  v;  87 89  ACKNOWLEDGEMENTS  I would like to express my sincere appreciation to some of the many people who helped so much with this project.  Athough some names may be  omitted here for the sake of space, the memories of your support and friendship are firmly held. F i r s t , my thanks must go to Dr. Ian McTaggart Cowan for moral and financial support, direction and stimulation, and good company in the field.  I am glad to have had the opportunity to experience his supervision.  Next, I thank my committee members, in particular Dr. Fred Bunnell, for assistance in designing and carrying out the study and reviewing a previous draft of the thesis. The staff of the B.C. Fish and Wildlife Branch in Cranbrook, especially the "Godfather," Ray Demarchi, deserve very special mention.  Ray's  comments and criticism had a significant positive influence on the study and his Italian temper was an inspiration.  The provision of that fantastic  house at the Fish and Wildlife Research Station at Ta Ta Creek, and having use of many office f a c i l i t i e s were essential to the success of my research.  Conservation Officer Gerry Brunham's contributions to my  freezer were most welcome, and all the other direct and indirect support of folks like Bob Jamieson, John Russell, B i l l Warkenton and Dave Phelps are remembered.  Finally, though rumors in the Kootenays have i t that  l i t t l e good comes out of Victoria, B.C.F.&W. Research Coordinator, Don Eastman, is a notable exception whose time I gratefully acknowledge here.  vi  A great deal of the tedious botanical work that was part of this study was done by Miss Olivia Ng.  Her patience surpasses my own and I  appreciate her assistance and fine work. I thank Miss Judith Bradshaw whose artistic drafting speaks for  it-  self. Many friends and fellow students in both Vancouver and at the research station deserve recognition for their thoughts and actions, but none more than John and Ruth Foster who put me up (and put up with me) and assisted in production during those long days and nights cranking out the f i r s t draft of this thesis. Finally, I want to thank three generations of my family:  my parents  for encouragement, faith and financial support throughout my university years; my wife Michelle, for love and understanding as well as a lot of help with the actual study (like sitting on trapped deer even when she was eight months pregnant); and my children, Heather and Eryn, who, by representing the future, give my efforts to perpetuate natural systems meaning and purpose.  INTRODUCTION  The population of white-tailed deer {Odocoileus ockrourus,  vivginianus  Bailey) in the Premier Ridge region of the East Kootenays of  British Columbia suffered.a major die-off in the winter of 1964-65 (pers.. comm. R. Demarchi).  Since that time, the deer population has  remained well below the level that helped bring big game hunting fame to this area in the 1950's.  Local populations of other wild ungulates were  also on the decline about this time.  Although parasites played a major  role in the die-off of big horn sheep {Ovis canadensis  canadensis),  their ability to decimate the sheep was greatly enhanced as a result of poor winter range condition (Stelfox, 1971).  Russell (1967) examined  the possible role of parasitic infestation in the whitetail die-off, but found nothing conclusive.  Biologists in the B.C. Fish and Wildlife  Branch feel that the ultimate cause of the high mortality was the condition of the habitat on the, critical winter ranges.  Deterioration of the range is thought to have been the result of several concurrent events.  Most important was secondary succession of  forest types over large areas of the winter ranges (Demarchi, 1971). Forest succession has been shown to have generally negative influence on forage biomass and quality in such diverse ecological regions as the southern pine forests (Halls and Alcaniz, 1968), northcentral mixed woods (Wetzel et a l . 1974), northern B.C. (Cowan et a l . 1950) the coastal rain forests (Gates, 1968) and in the East Kootenays (Kemper, 1971).  On Premier Ridge, this factor has been complicated by Christmas  2.  tree farming ( c f .  Churchill, 1974) and the extensive use of some areas  by cattle which had led to modifications of the vegetative composition and productivity of some areas by the early 1960 s (D. Demarchi, pers. 1  comm.).  Although use of Premier Ridge by domestic.stock has been reduced  through the Fish and Wildlife Branch's land and grazing permit requisition program, the majority of the area supporting white-tails during winter has not yet recovered from the effects of overutilization (Demarchi, 1971; Barichello, 1975).  In view of the limited potential of the area for timber production (Kemper, 1971) and the detrimental effects of past grazing regimes, the B.C. Forest Service has agreed to cooperate with the Fish and Wildlife Branch in a Coordinated Land Use Planning program to improve the habitat for ungulates - - both domestic and wild.  In order for this coordinated  land use scheme to operate most efficiently with respect to white-tails, i t is necessary to increase our knowledge of their, habitat preferences and requirements, to document the present condition of these habitats and to formualte recommendations for future treatment of the habitats to improve their ability to support deer populations.  Recent studies have provided some of this information.  Barichello  (1975) and Farr (1975) have made detailed habitat assessments on property purchased by the Fish and Wildlife Branch, and a more general habitat analysis for the southern Rocky Mountain Trench was completed by the Environment Land Use Secretariat during 1975 (D. Demarchi, pers. comm).  3.  Previous work by Demarchi (1971) and Kemper (1971) also, gives insight into some of the vegetative aspects of deer ranges in this area.  Hudson et a l . (1974) examined the distribution and competitive relationships of cattle, big horn sheep, elk {Cervus canadensis  nelsoni),  mule deer [0. hemionus hemionus) and white-tails in the Premier Ridge area.  The species' general distributional elements were outlined, but  they failed to.make definitive comments on solutions to problems limiting ungulate populations here.  They did indicate, however, that white-tails  may be most seriously affected by forest succession and competition for cattle for both nutritional and distributional reasons (Hudson, et a l . , 1974.  pp. 39-40 and 62-63).  The study reported here was designed as an intensive investigation of a single population of white-tailed deer (based on their common use of a winter range area) to answer some basic questions about deer ecology in the East Kootenay.  This information is essential for the development  of proper land use plans. research were:  The objectives of the various phases of the  1) to identify the habitats used by the deer and determine  their species composition, productivity and physical structure as this relates to climatic influences;  2) to assess the relative levels of use  of these habitats and their forage resources by the deer; and 3) to document the movements of .individuals within the population.  This in-  formation would allow distributional and movement patterns to be related to the measured.habitat characteristics and climatic variables.  4.  STUDY AREA  Winter- Range  The winter range unit chosen for study lies on the lower slopes and terraces of Premier and Wasa Ridges in the Wolf Creek drainage about 48 km north of Cranbrook, B.C. (Fig. 1.)  These two ridges rise nearly 2000  and 1400 feet—'', respectively, above the east side of the floodplain of the Kootenay River on the floor of the Rocky Mountain Trench at 2500 feet above sea level.  The portions of these ridges used in most winters  by white-tailed deer l i e between the floodplain and 3500 feet.  Physically,  the region is characterized by a narrow floodplain that ends in steep slopes of 50 to 250 feet leading to rolling terraces with a complex series of slopes.  Above these rise the main bodies of the ridges with  moderate to steep angles and occasional bedrock outcrops.  The soils of  the flood plain are predominately thin, s i l t y gleysols on a thick gravel substrate.  The soils at higher elevations are primarily clay-podsols  and loess deposits on gravel substrates, or are thin veneers on bedrock. These soils are all deprived from limestone/dolomite parent material and are highly calcareous.  The soil pH depends on micro-climatic moisture  regime.  The area is located near the border of the Interior Douglas Fir zone and the Yellow Pine/Bunchgrass zone (Krajina, 1965).  The pre-1900  forest was a mature montane community of mixed stands of douglas f i r {Pseudoifgugct menziesii)  and yellow pine {Pinus ponderosa) with the  former being dominant in all cases except on drier, south aspects (Kemper, 1971). ]_/  All elevations given in feet (English units) for ease of reference to existing topographic maps.  5.  6. Only a few patches of mature timber remain now, as sporadic logging and fires prior to 1930, and range use by wild and domestic ungulates since then, have created a complex patter of dis-climax grass- and shrublands and serai forest communities with varying ages, species composition and stocking levels (Demarchi, 1971).  The relatively dry nature of the area (annual precipitation 30 to 60 cm, mostly snow) and extremely high temperatures in the mid- and late summer months effectively reduce the growing season for grasses, forbs and shrubs from the 180 day growing period to about 70 to 80 days in the spring and early summer.  A period of f a l l regrowth that varies annually  in length has been shown to be important to big horn sheep wintering on Premier Ridge (Hebert, 1973).  This period may also be important to  white-tails in some years.  Summer Range  The deer from the Wolf Creek drainage disperse over an immense area during summer, and i t is d i f f i c u l t to circumscribe a "summer range" as such.  A very small number of the deer remain on the "winter range", and  a few spend the summer along the Kootenay River.  However, the majority  of the deer migrate to the west to summer in the mid elevation valleys of Ta Ta, Lost Dog, Cherry and Skookumchuck Creeks.  The floor of the Trench on the west side of the river floodplain consists of ridges and terraces oriented along the line of the valley. These rise in gradual progression from about 2600 feet to 4500 feet.  7.  From 4500 feet upward rise the main slopes of the Purcell Mountains. The effects of recent glaciation are clearly evident on both landforms and soil development.  Morainal and fluvioglacial deposits f i l l the  depressions between bedrock ridges that are only partially covered with colluvial veneers.  The north-south stream courses and depressions in  places contain small, oblong lakes and occasional bog-like areas where organic matter has been accumulating, although not yet deep enough to warrant an organic soil classification (V. Hignet, pers. comm.).  The  soils are predominantly calcereous brunisols up to 4000 feet, but above that they are non-calcareous.  The summer range lies almost entirely within the Interior Douglas f i r zone, divided equally between the Upper and Lower sub-zones.  In a  few places at higher elevations the range reaches into the Engleman Spruce/Sub-alpine f i r zone and a natural grassland occupies part of the valley floor.  Extensive and severe burning of the area in the 1920 s 1  and 30's has all but totally removed the original forest cover. pine predominates the regenerating forest on the floodplain. levels, lodgepole pine {Pinus contorta)  Yellow  At higher  is most abundant, but micro-  climatic variations in soil moisture and temperature are reflected in species composition of the forest.  Occasional openings from more recent  spot fires or on slopes with too l i t t l e soil to afford major forest regeneration are covered with a shrub community.  Along most of the  stream courses a type of dense, deciduous brush community exists that varies from a few to almost 100 m. in width.  Due to elevation, aspect, and shading by the Purcells, the summer range is much cooler and more moist than the winter range.  Although  this precludes its use in the winter months when snow depth often exceeds 100 cm at 3000 feet, i t encourages a dense understory and prolonged plant growth that supplies lush vegetation until plant dormancy is forced by returning freezing weather in September or October.  METHODS  Analysis of Winter Range Habitats  Habitat types were outlined based on forest cover maps prepared by the B.C. Forest Service, aerial photographs, and personal reconnaissance of the area as suggested by Poulton and Tisdale (1961).  Forested types  were designated on the basis of age, species composition and/or canopy closure.  Nonforested types were classified by plant community (i.e.  grassland) or by soil moisture.  Except in cultivated fields where domestic grasses and alfalfa {Medioago sativa)  were assumed to be the only significant component,  species composition of the herbaceous layer was determined using Dau2 benmire's (1959) 1/10 m technique.  Frames were located at 1 m intervals  along two parallel transects 25 m in length, spaced 5 m apart. transects were randomly located within the habitat types.  These  9. Species composition of the shrub layer was determined using a modification of Passmore and Hepburn's (1955) technique based on counts of shrub stems in five circular plots of 10 m surface area at 25 m i n tervals along a 100 m transect (Fig. 2). type ranged from two to eight.  The number of transects per  A shrub stem was defined as any woody  part of a plant originating from the soil surface.  Since determination  of availability of forage in the previous winter was one of the major objectives of this survey, this definition which eliminated current year shoots seemed appropriate.  A subsample of two stems per species per  plot was measured for height prior to initiation of current year's growth to determine vertical distribution of forage.  The generally  stunted nature of the shrub layer ruled out the need for establishing an upper limit to the "available" zone used by most authors.  Use of shrubs by deer was based on the ratio of browsed to total number of twigs (expressed as percent) determined from the subsample of stems.  A twig was defined as a portion of a plant that ended in a ter-  minal bud or a browsed tip.  Measurements of diameter at point of  browsing (DPB) and diameter at annual growth (DAG) were also taken on the twigs of the subsampled stems to refine the percent browsed and estimate "utilization" as presented by Wetzel et a l . (1974).  Minimum production of browse by major shrub species was estimated 2 by multiplying stems/m by the mean number of unbrowsed twigs per stem, times the average oven-dried weight of at least ten annual growth segments collected at random from each species in each type.  Unbrowsed twigs  only were used here because twigs that had been previously browsed  10,.  would not produce, the same amount of growth.  Since they would probably  produce some browse, however, the values presented here should be considered minima.  Further, to be sure these values represent what is actually  availably to an overwintering deer, the twigs were collected in midJanuary.  Concentric with each center plot on each transect a circle of 100 2 m was sampled for overstory composition.  All trees within the circle  were measured for diameter at breast height (dbh) and their height was estimated.  Any tree with a dbh less than 5 cm was designated a sapling.  Demarchi and Demarchi (1967) have shown that douglas f i r provides nearly one-third of the volume of the winter diet and is universally consumed by white-tails in this area.  For this reason, i t was felt that  some estimate of available forage on f i r trees should be obtained.  I  used a regression estimation technique developed for conifers by King (1975) for this purpose.  This method uses stem diameter measurements to estimate browse.  I  defined a stem as any terminal or lateral branch originating from the main trunk of the tree.  I measured the diameter of 25 randomly selected  stems ranging from 2.99 mm to 29.52 mm to provide independent variables for the regression.  Then all leaves on the selected stems were removed  and collected, and all twig setments distal to the point where twig diameter equalled 3.5 mm (the maximum observed browsing diameter) were  n. clipped and collected.  This material was oven-dried for 36 hours at  65°C and the weights of twigs and leaves were obtained.  The natural  logarithm of the weights were regressed on the natural logs of the stem diameters to determine the least squares equation:  log Y = log a + b'(Vog X) e  e  e  where X = stem diameter, Y = predicted browse weight, and "a" and "b" are the regression constants.  I then sampled 10 saplings and 10 trees in each habitat where f i r occurs, counted a l l stems below 1.6 m above ground level, and measured the diameter of 10 of the stems.  Stem totals and average stem diameter  were then used to estimate the average amount of browse per sapling and per tree in each habitat.  With this information I could estimate f i r  forage biomass by the formula:  Biomass/ha = (average trees/ha)(average browse/tree) +. (average saplings/ha)(average browse/sapling)  Utilization was not quantified because the regression technique King (1975) demonstrated for determining use relied on the twig diameter at point of browsing to estimate biomass consumed. A large proportion of the fir.browse eaten by deer here is in the form of leaves removed from the twigs without taking woody tissue (Demarchi and Demarchi, 1967, and pers. obs.) so using DPB would have given an unrealistically low  13. estimate of consumption.  Subjective comments on intensity and characteristics  of browsing are made.  Analysis of Summer Range Habitats  The vegetative communities on the summer range are much more homogeneous than those on the winter range and consist primarily of open shrubland slopes, riparian brush, and dense immature stands of timber dominated by lodgepole pine.  The major objectives of the analysis of  this range were to determine the elevational trends in each of the three broad associations listed above and document present species composition. No attempt was made to assess the levels of utilization on any plant species due to the on-going nature of both plant growth and utilization through the summer survey period as well as a high degree of overlap of distribution of deer, cattle and elk on this range.  Transects with two to six square macroplots of 100 m area spaced at 100 m intervals were randomly established at various elevations in the shrub and forest associations between 2600 and 4100 feet above sea •p  level (Figure 3).  Daubenmire (1959) 1/10 m frames were placed at 1 m  intervals along the center line of the plot to determine composition of the herbaceous layer.  Shrub canopy coverage for each species was estimated  for the entire plot using Daubenmire's coverage classes.  All trees  within the plot were measured for dbh and their height was estimated. A subjective evaluation of species composition of the riparian brush was made at several elevations ranging from 2550 to 4000 feet.  14.  Weather  Many authors have investigated the relationships between deer behavior and the climate in an attempt to understand the role these interactions play in deer ecology.  Work has ranged from early observational  accounts (Hammerstrom and Blake, 1939; Cook and Hamilton, 1942; and Severinghaus and Cheatum, 1956) to the years of sophisticated energy flux studies summarized in Moen's (1973) excellent text.  Based on this  earlier work, I f e l t i t was sufficient for this study to examine the variations in the patterns of temperature, sunlight and snowfall during the winter months to help in interpretation of the observed patterns of habitat use and movements.  Weather data were collected from records at the Cranbrook Airport 32 km south of the study area.  Records of minimum, maximum and mean  daily temperatures, hours of sunlight and precipitation were collected throughout the study period.  Because of their relative positions in the  Rocky Mountain Trench, the winter range would be subject to slightly lower winter and higher summer temperatures with, somewhat less precipitation than the airport, while the summer range,would have generally lower temperatures and higher precipitation throughout the year.  The  airport data show clear trends and differences between seasons and years, and when coupled with limited data gathered on the study area, they provide information helpful in the understanding of habitat use and movement patterns.  15.  Use of Winter Range Habitats  Relative levels of use within the habitat types were determined by systematic track counts and pellet group counts.  The track counts, covering  seven types in 1974-75 and 10 types in 1975-76, were made 12 to 36 hours after the completion of a fresh snowfall of sufficient volume to effectively eliminate previous tracks.  A network of roads (Fig. 4) was either  walked, skied, or driven and all sets of tracks crossing the road were counted.  This method has been used successfully in the southern United  States (Harlow and Oliver, 1967) and in Minnesota (Irvin, 1975).  The proportion of the total tracks in each habitat was compared with the proportion of mileage surveyed in that type.  The hypothesis  that deer are uniformly distributed and therefore the proportions of tracks and mileage are not significantly different was subjected to a chi-squared test.  If a significant difference did occur (p$0.01) the  data were further tested using the method of Neu et a l . (1974) to deters  mine the degree to which each habitat contributed to the chi-squared value.  This technique places confidence limits on the proportion of  tracks in each habitat and preference was considered to be indicated for those types in which the confidence interval on the proportion of tracks was entirely above the proportion of miles.  For example, i f the propor-  tion of tracks in the Aspen type were 0.571 + ..645 and. the proportion of km censused were 0.365, selection i f the type would be inferred from the fact that the lower boundary of the confidence interval, 0.571 -.045 = 0.526, is greater than 0.365.  Inversely, avoidance was inferred from  FIGURE 4. Track count transect route (••—) locations (* ).  and pellet qroup transect  17.  confidence intervals that f e l l short of the proportion of km.  Use is  termed proportional (meaning neither avoidance nor selection) i f the confidence interval included the proportion of km.  Pellet groups were counted in each of the five circular plots used on the shrub survey (Fig. 2).  Each plot center was marked with a stake  and all groups within 1.78 m of the stake were counted..  A "group" was  arbitrarily defined as a collection of at least 12 pellets.  Border-line  groups were inlcuded only i f a majority of the pellets were inside the plot.  After counting, all pellets were removed from the plot with  minimum disturbance to the vegetation.  The small plot size was chosed based on a review by Neff (1968) and work by Smith (1968).  They felt that small plots were more efficient  and more precise than large ones because of the reduction of bias associated with missed groups.  Batcheler (1975) has shown, however, that the  inverse relationship between plot size and estimated,density of groups is mainly the result of bias due to border effect and.definition of pellet groups, not missed groups.  Thus small plots.may be more precise,  but they are not necessarily more accurate.  Although the data may not provide an accurate estimate of total deer numbers, i t seems valid to assume that any bias associated with using small, bounded plots will be approximately equal in each habitat. Since my main objective in counting pellet groups was to determine relative levels of use, any consistant bias is of no consequence.  18.  Each transect with, its five plots was treated as a. cluster sample, and the transects within each habitat type were grouped for analysis. Each cluster total, Y| , was the sum of the five individual elements (single plot counts), y-j  ..  The average number of groups per plot was  '3  estimated by:  -  Y,  y = 1. ITU  "•j  +Y  o  +...+Y  _n._  2. +m  9  . r  Y.  n  = i = 1  I1, .£  +.. .+m  n  '"'2 ' • • • '"'n  i m  ,  .  i = i m-j  where n = number of transects and m = number of elements (plots) per transect. The variance estimate i s :  S y = 2  1  . i = 1 yj -2y i =1 2  yjmj+y ^! mj 2  2  n(m)  2  n-1  The finite population correction factor, (N - n)/N, where N = total number of clusters in the population, has been dropped from the formula since the sampling fraction is unknown.  The sampling fraction is so  small, however, that (N - n)/N is nearly unity and the true sample variance is only slightly lower than the estimate presented here.  By  considering only the variance between grouped clusters and ignoring variance between individual elements of each cluster, the variance estimate is reduced using this treatment.  Mean pellet group density and  variance estimates were then used in selected t-test comparisons of habitat types,  t-tests were also used to evaluate the significance of  changes between years within each habitat type.  19.  The value of using both, track and pellet group counts cannot be overrated.  Each has an advantage over the other.  Track counts, for  instance, can be used to show temporal changes in habitat preference through the winter, a phenomenon impossible to detect from pellet group counts.  On the other hand, track counts are useless for estimating  absolute density or changes in absolute density--the most frequent objective of pellet group counts.  Perhaps even more important than  their specific advantages, however, these techniques have offsetting biases that gives their combined use greater strength.  Track counts are  biased toward active use areas and feeding behavior since deer obviously make more tracks while moving about in search of food.  Observations  also indicate that deer meander more in open habitats than in densely forested types, so multiple crossings of the survey road by one deer in the open could result in a higher count than a few deer in a forested area crossing once each.  This bias is countered by the tendency of  pellet groups to be concentrated near bedding areas (pers. obs. and V. Geist, pers. comm.) and the deers habitat types.  1  preference for bedding in forested  By using both techniques, one gets a more valid indica-  tion of temporal patterns of use, absolute density, and the behavior most prevalent within a given habitat.  Such information is essential in  proper evaluation of the role of various habitats in the ecology of deer.  In order to determine, what activities were being pursued in the various habitat types to help interpret the results of the track and pellet group counts, time was spent in each, habitat following the tracks  20  of individuals and groups noting feeding behavior and searching for bedding sites.  This indirect approach was necessary due to the great  flight distances and preferences for cover common to the whitetailed deer here.  Wetzel et a l . (1974) used tracking to quantify feeding  behavior, but the value of such an approach is questionable.  Infor-  mation of this type has only been used here in the interpretation of the quantitative data gathered in the manner already discussed.  The only significant use of cultivated fields, sown mainly in alfalfa and hay grasses, and the natural grasslands occurs in the late winter and spring.  The ability to observe deer on these areas at this  time was exploited to directly quantify use.  Although there are major  differences between the composition of plant species in these two areas, the response of the deer to them is the same, and their use is treated together in the remainder of the text.  The fields and grasslands were surveyed at irregular intervals as soon as they started to become snow free.  When deer use increased, the  censusing became regular and classified counts of the deer using individual fields were made several times each day on at least two days per week.  Counts were concentrated in the evening hours.  The most heavily  used fields were observed for periods of several hours at a time to detect daily patterns of use.  Observations after dark, in 1975 were made  with a "Startron" night vision device similar to the one used by Swanson and Sargeant (1972).  In 1976, a spotlight and binoculars were used.  21.  In addition to providing information on use of the fields and grasslands, these spring observations permitted estimation of the age ratios (juveniles:100 adults) in the population at the end of the two winter periods.  Use of'Summer Range Habitats  No satisfactory method of quantifying habitat use on the summer range was found.  Track counts were impossible; pellet group counts im-  practical due to low densities, lush vegetation and continuing deposition of groups until snowfall forced deer off the range; and the extremes in vegetation density would have biased census counts.  For these reasons,  the use of summer range was determined subjectively from observations of deer and deer sign and limited radio tracking.  Several hundred km of  roads, river and trails were covered through the summer of 1975 to try to outline the areas used by white-tailed deer in this section of the Rocky Mountain Trench.  Individual Movements  To supplement the information on relative levels of use, i t was desirable to obtain knowledge of the movement patterns of individual deer on the study area.  Deer were captured for individual marking in  three single-gate box traps (Clover, 1956) baited with alfalfa hay.  The  traps were generally set along trails in areas of relatively high deer density because success was too low in less-used areas.  I tied the  22.  traps in such a way as to permit quick release of the guy lines and folding of the trap with the deer inside.  Thus the trap became a  "squeeze shoot" which greatly facilitated handling the deer when alone. Once the trap was folded and laid on its side, the deer was blindfolded and manually restrained while beging fitted with marking devices.  No  drugs were administered to deer in this study.  Deer were marked with both material and radio-transmitter collars, ear tags and streamers.  The material collars and ear streamers were  made from an orange, vinyl-coated nylon material, "Armortite."  Each  collar was cut four inches by 27 inches and marked with four inch high letters and numbers with an indelible black ink.  The numbers could be  read from either side of the deer at up to 600 m with a 60x spotting scope.  The collar was placed around the neck of a trapped adult deer  and the ends overlapped until a loose, but secure f i t was made. ends were then fastened with three "pop" rivets.  The  "Ritchey" brand plastic  cattle ear tags with numbers corresponding to the collar were placed in the ears of each deer with a combination of up to three streamers of "Armortite."  This system provided sufficient variations in ear mark  pattern to permit individual recognition of deer even i f the collar was not visible or lost.  Records of sightings of marked deer were kept whenever they occurred on routine travels or while driving specific census routes in the study area.  Some were also identified from the air when I had the opportunity  to survey the area in mid-February, 1975 and 1976, with B. Warkenton (B.C.F.&W. Wildlife Technician) on his annual game flights.  23.  A total of ten deer was fitted with radiotransmitter collars. These collars were made by Wildlife Material Inc. of Carbondale, Illinois and ranged in frequency from 150.898 to 151.147 Mhz.  A three element  yagi antenna was used with a 12 channel AVM Instrument Co. receiver to determine directional bearings from known locations to the deer for triangulating their positions during the winter and spring.  Attempts  were made to radio-locate each deer weekly in winter and semiweekly during spring migration.  Four flights were made with a Cessna 172  aircraft during the summer of 1975 to locate the deer on their summer ranges.  RESULTS  Analysis of Winter Range Habitat  The plant communities on the winter range were divided into eleven different habitat types based on my interpretation of the deer's response to the physical structure or species composition of the type. are purely descriptive.  The names  This system was felt to be more relevant to the  practical approach I used than the more theoretical concepts used by Kemper (1971).  Figure 5. is a map of the distribution of the habitat  types on the winter range, and Table I lists their proportions.  Grassland:  On the winter range there are four grassland areas, all of which.  24.  YF RB MAF  FIGURE 5. Distribution of habitat types o n the winter range.  FO IFD SS MO Asp Cult *  Young Fir Riparian Brush Mixed Age Fir Grassland Immature Forested Open Immature Forested Dense Serai Shrub Mature Open Aspen Cultivated Riparian Mature Spruce Eastern edge of w h i t e tailed deer distribution  TABLE I PROPORTIONS OF HABITAT TYPES ON THE WOLF CREEK WINTER RANGE  Habitat  ha.  Grassland  330  6  Riparian Brush  55  1  Aspen  11  1  Serai Shrub  440  8  Young Fir  714  13  Immature Forested Open  1,319  24  Immature Forested Dense  1,374  25  Maturing Open  550  10  Mixed Age Fir  660  12  15  1  Riparian Mature Spruce  Total  % of the Area  5,468(21 mi ) 2  26.  show the influence of grazing by domestic stock.  Kemper (.1971) described  a "natural" grassland in this area as being dominated by bluebunch wheatgrass {Agropyron spioatum), junegrass {Koleria  cristata)  rough fescue (Fesiuca  and  scahrella),  with a combined ground cover of 80%; 10  species of forbs produced a ground cover of 6%.  In contrast, the grass-  land areas of the white-tail winter range are dominated by needle grass (Stipa richavdsoni,  cover = 13.2%) with some bluebunch wheatgrass (cover  = 6.6%), junegrass (cover = 4%) and bluegrasses {Poa spp., cover = 2.2%) giving a total of only 23% ground cover.  Twenty-one forb species were  found to cover approximately 10% of the soil surface with fleabane (Erigeron  pumi-lus, cover = 5.2%) and pussytoes [Antennaria  cover = 1.9%) being dominant. soil (See Table  miorophylla,  Nearly 20% of the ground surface is bare  II).  Riparian Brush Type:  This habitat was of limited distribution, and very d i f f i c u l t to sample, but it appears to be relatively important to the deer.  It con-  sists of the open brushy areas adjacent to Wolf Creek and differs from the serai shrub type in species composition due to greater soil moisture. In many places, this habitat consists only of narrow thickets of red osier dogwood (Comus stolonifeva) [Salix  or a marshy area with various willows  spp.) and a sedge ground cover.  of the measurements in these areas.  It was impossible to take many  I therefore concentrated on a few  spots where this type was dispersed over a drier substrate.  The species  composition of these areas differed somewhat from the dogwood thickets or willow marsh, but the trends in productivity and deer use are similar.  27.  TABLE II.  .  SPECIES COMPOSITION OF HERBACEOUS LAYER: WINTER RANGE Species*  Habitat Type Grass.  R.B.  Asp.  S.S.  tr/20  tr/10  2.8/52 1.0/40 1.1/14  Y.F.  I.F.O.  I.F.D.  M.A.F.  M.O.  Grasses: Agropyron splcatum Bromus tectorum Calamagrostls rubescens  6.6/23***  Carex condnnoldes C. rossil Festuca' scabrella Koleria cristata Poa spp. Stipa rlchardsonl  tr/16  Atragalus miser Castilleja minlata Chryopsis vllosa Collinsia grandiflora Crepis atrabarba Epilobium minatum Equisetum arvense Erigeron f i l i f o l i u m E. purailus  4.0/35 2.2/28 13.2/78  15.1/88  5.8/90  tr/20  1.9/18  tr/28 tr/24 1.5/20 5.8/62  7.9/42  tr/20 tr/22  tr/18  1.8/24  tr/10  tr/20  tr/10  1.3/52 1.3/12 tr/32 tr/16  5.7/58  4.4/65  6.4/33  4.6/64 1.8/14  2.3/30  tr/18  tr/20 8.3/76  tr/10  tr/10  1.8/20 tr/14  tr/20 3.4/28 tr/14  tr/14  3.0/20 2.6/30  tr/12  tr/16  tr/10  tr/31  tr/24  tr/18  tr/26 tr/11  tr/10  tr/26 tr/42  tr/20  5.2/48  Fragaria glauca Galium boreale Lathyrus nevadensis Lomatium trlternatum Kelilotus alba Penstemon confertus  tr/20  2.5/28  Forbs: Achilles millefolium Agroseris sp. Al 1ium cernuum Anemone patens Antennaria microphylla Aster sp.  2.9/30  2.2/52  1.3/20 tr/22  1.7/28  tr/20 3.1/66  1.2/32  tr/16 1.8/20  Phlox caespitosus Rubeckia hirta Taraxacum scopulorum  tr/12  Trifolium sp. Viola adunca  3.4/64  10.5/74 4.1/46  2.5/36  tr/10  1.6/34  tr/20  1.1/14  3.8/52  14.6/88  tr/28  1.3/33  1.2/18  1.6/42  tr.16  1.9/34  tr/10  1.7/28  Dwarf shrubs: Arctdstaphylos uva-urs1 Berberis nervosa Bare ground:  19.8/95  6.1/32  1.9/16  29.9/88  15.1/63 1.7/20  10.8/45  3.3/20  4.6/42  7.6/29  23.5/80  18.3/72  4.7/38  11.4/61  2.3/16  1.9/25  •Only species with at least 10% frequency of occurance in one type, shown. **Grass.= Grassland; R.B.= Riparian Brush; Asp.= Aspen; S.S.= Serai Shrub; Y.F.= Young F i r ; I.F.0.= Immature Forested Open; I.F.D. Immature Forested Dense; M,A.F.= Mixed Age F i r ; M.0.= Maturing Open ***% cover/% frequency of occurance  28.  Bluegrasses (cover = 15.1%) were the only abundant grasses, but Carex vossii  and bluebunch wheatgrass were also found in trace amounts  (Table II).  The relatively sparse forb component included twelve  species, but only strawberry (Fragaria introduced clover {Trifolivm  glauca,  cover = 1.3%). and an  sp., cover ~ 3.4%) were significant in  contribution to the approximately 5% ground cover.  The density of the  shrubs, however, left only 6% of the ground bare.  Seven shrub species were common (Table III),  with snowberry (Sym-  aVbus) and rose (Rosa woodsii and R. acioularis) contributing 2 most of the 248.3 twigs/m . Because of favorable sunlight conditions  phoricarpus  (no shading) and year-round availability of water, productivity is high, 2 and  the four most abundant species provide 21.64 g/m of forage annually  (Table IV). and  Utilization was low on snowberry (16.4%) due to its abundance  relatively low palatability, moderate on rose and willow (31.2 to  46.1%), and extremely high (105.5 to 110.0%) on the rare, but favored saskatoon {Amelanchier  alnifolia)  and choke cherry {Prunus  virginianus)  (Table V). Aspen Type:  In a few places along Wolf Creek and on the Kootenay floodplain, stands of maturing aspen (Populus tremuloides)  are found.  The herbaceous  layer in this habitat is relatively diverse containing at least 25 different species, but few of these provide ground cover in excess of 1%.  Most abundant are bluegrasses (cover = 5.8%), carex (cover = 2.9%),  29.  TABLE  III  AVAILABLE SHRUB BROWSE IN TWIGS/m : 2  Species  R.B." * Asp.S.S.  Purshia tridentata  Y.F.  R.M.S.  91.2  Amelanchier alnifolia Symphoricarpos albus Rosa woodsii R. acicularis  6.6  32.5  180.2  30.2  6.8  48.6  46.1  26.1  8.1  2.3  Spirea lucida Shepherdia canadensis Populus tremuloides  18.0  Salix glauca  0.3  36.9  27.8  32.8  4.6  0.9  0.6  9.8  20.1  14.7  8.4  0.5  1.3  0.3  0.2  1.1  1.7  0.2  0.6  2.0 0.6  2.4 0.2  Sambucus racemosa var. arborescens  0.5  Lonicera involucrata  0.2 248.3 156.0 101.6 88.5  :  20.8  3.5  Acer glabrum  R.B. S.S. = Asp. • • Y.F. = R.M.S.;  6.6  1.3  0.2  35.5  13.4  0.7  R. spectabilis  Mean:  4.0  13.5  3.0  Total:  21.1  2.0  Rubus parviflorus  6.0  I.F.D  7.4  0.7  Prunus virginianus  I.F.O.  9.2  Alnus sinuata  2.3  M.A.F.  45.4  1.7  Ribes irriguum  M.O.  14.8  Cornus stolonifera  *Habitat Type:  WINTER RANGE  26.0  33.8 11.1  Riparian Brush Serai Shrub Aspen Young Fir Riparian Mature Spruce  81.9  52.4  52.2  49.6  12.5  5.5  13.1  9.7  9.9  4.2  M.O. M.A.F. I.F.O. I.F.D.  = = = =  Maturing Open Mixed Age Fir Immature Forested Open Immature Forested Dense  30.  TABLE IV ANNUAL PRODUCTION OF BROWSE IN G»_2 /m BY SHRUB SPECIES:  WINTER RANGE Habitat Type  Species  R.B.*  Purshia tridentata  S. S.  0 .08  Symphoricarpos  19 .45  Rosa woodsii  0 .42  R. acicularis  1 .69  0. 30  R.IM.S.  Total:  21 .64  Mean:  5 .41 = = = = =  M,.0.  I.F.O.  M.,A.F.  I.I . D .  0,.97  0 .72  0 .18  0,.17  0,.93  0,.89  0 .46  0,.14  :  0 .07  1,.66  0 .67  0,.81  2 .03  0,.37  0 .07  0 .17  1 .01  0..63  0 .13  0,.05  0,.02  0 .17  1 .09  0..04  0 .96  0 .76  0..06  0 .01  3 .60  3.,20  3.13  2.,32  1,.70  0 .98  0..31  3. 72 0 .72  0.,64  0 .78  0.,58  0..43  0 .25  0..16  0. 18  Populus tremuloides  R.B. S.S. Asp. Y.F. R.M.S.  Y,.F.  10. 67  Amelanchier alnifolia  *Habitat Type:  Asp.  11. 15  Riparian Brush Serai Shrub Aspen Young Fir Riparian Mature Spruce  M.O. I.F.O. M.A.F. I.F.D.  = Maturing Open = Immature Forested Open = Mixed Age Fir = Immature Forested Dense  31.  TABLE V SHRUB SPECIES UTILIZATION*:  WINTER RANGE Habitat Type  R.B.** Asp.  Species  Y.F.  I.F.O.  I.F.D.  M.O.  M.A.F.  26.2  .30.9  29.5  18.3  52.3  45.7  25.3  36.5  49.6  50.4  36.1  18.1  15.6  24.3  54.1  48.7  28.7  45.9  40.0  23.1  Purshia tridentata 105.5  27.9  Symphoricarpos albus  16.4  58.7  Rosa woodsii  46.1  83.4  R. acicularis  31.2  42.5  Amelanchier alnifolia  S.S.  18.6  17.0  46.9  Populus tremuloides Salix glauca  44.7 44.5  27.1  48.0  Spirea lucida  36.7  28.8  34.6  Mean:  56.2 59.4  61.6  36.0 110.8  Cornus stolonifera Prunus virginianus  R.M.S.  110.0 59.0  61.9  82.8 48.9  19.6  56.6  38.7  30.3  28.2  31.6  54.2  •Utilization = (Diameter at Point of Browsing/Diameter at Annual Growth) x (% of .twigs browsed) **Habitat Type:  R.B. Asp. S.S. Y.F. I.F.O.  = = = = =  Riparian Brush Aspen Serai Shrub Young Fir Immature Forested Open  I.F.D. M.O. M.A.F. R.M.S.  = Immature Forested Dense = Maturing Open = Mixed Age Fir = Riparian Mature Spruce  32.  browneyed susan (Rubeckia  hirta,  cover - 10.5%), dandelion (Taraxacum  scopulorum, cover = 4.1%), purple pea (Lathyrus 3.1%), and violets (viola  nevadensis,  adunca, cover = 1.9%).  leaves only 1.9% of the soil surface bare (Table  cover =  This rich herb layer II).  The aspen type also has a well developed shrub layer in which five 2 major species provide 155.4 twigs/m (Table III). These species produce 2 3.6 g/m of browse annually (Table IV), of which nearly 50% is used (Table V).  The greatest use occurs on the most abundant and productive  species, Rosa  woodsii.  The overstory in this type is purely aspen; with a density of 4880 stems/ha.  The average dbh is 9.91 cm and mean height is 12 m.  Heavy  browsing by deer and cattle has affected the growth of the 3040 saplings/ ha (Table VI) and many of these were decadent.  Serai Shrub Type:  This habitat type occurs widely on Premier and Wasa Ridges, but a large portion of the shrubland habitat is at elevations unusued by white-tails in most winters.  Main grass species are junegrass (cover =  4.6%), bluebunch wheatgrass (cover - 2.8%) and fescue (cover = 2.5%) with some Bromus iectorum,  bluegrass and pinegrass  (Calamagrostis  rubescens) to total 13.8% grass ground cover (Table II).. are yarrow (Achilles  millefolium,  2.3%), phlox (Phlox caespitosus,  Common forbs  cover = 7.'9%), pussytoes (cover =* cover = 3.8%) and beardtongue (Pensiemon  33. TABLE VI FOREST COMPOSITION:  WINTER RANGE  Habitat Type  Species  Aspen  Populus tremuloides  4880 3040 7920  9.9 cm sapl.  Young Fir  Pseudotsuga menziesii  820 4120 20 20 4980  7.4 sapl. sapl. sapl.  6 3 3 3  117 217 1833 1317 567 100 4151  8.1 sapl. 9.1 sapl. 12.4 sapl.  6 2 8 3 10 3  950 300 450 4100 5800  10.2 sapl. 6.6 sapl.  10 3 6 5  Pinus contorta Larix occidentalis Immature Forested Open  Pseudotsuga menziesii Pinus ponderosa Pinus contorta  Stems/ha  Ave. dbh  Ave. 12 3  Immature Forested Dense  Pinus ponderosa  Mixed Age Fir  Pseudotsuga menziesii  175 725 1275 2175  29.2 9.9 sapl.  15 8 3  Maturing Open  Pseudotsuga menziesii  100 100 113 88 150 50 601  31.0 7.4 sapl. 25.3 11.2 sapl.  15 6 3 15 6 3  267 233 67 33 33 33 33 133 33 267 333 1465  36.6 9.9 29.5 sapl. 29.4 47.2 sapl. 9.1 sapl. 9.1 sapl.  16 6 15 3 16 19 6 6 5 6 3  Pseudotsuga menziesii  Pinus ponderosa  Riparian Mature Spruce  Picea glauca Pseudotsuga menziesii Larix occidentalis Populus tricocarpa P. tremuloides Betula papyrifera  34.  cover 2.5%), which, with 19 other species, cover 23% of the  oonfertus, ground.  Bearberry (Aratostatphylos  uva-ursi)  covers nearly 30% of the  ground surface, and 24% of the soil is bare reflecting both the dry nature of the areas and the influence of season long cattle grazing.  The dominant species in the shrub layer is bitterbrush  [Purshia  In an earlier study of this habitat Kemper (1971) gave  tridentata).  bitterbrush a coverage value of 25%.  I found that this species provides.  91.2 twigs/m , and saskatoon and rose [R. woodsii) 2  twigs/m , respectively (Table III). 2  provide 8.1 and 2.3  Annual production for these species  is 10.67, 0.30, and 0.18 g/m , respectively (Table IV), 2  of 11.15 g/m of forage produced in the type. 2  giving a total  Utilization of these  species was 23.1, 18.6, and 17.0%, respectively (Table V).  Since the  sampling period followed an unusually deep snow winter which reduced deer use of this open habitat, these values should be viewed as minimum levels of use for these species in this habitat.  Young Fir Type:  This habitat covers an area that was cleared and/or burned about 40 years ago on the northeast slope of Wasa Ridge and on the alluvial gravel deposits where Wasa Creek runs off the Hughes Range to join Wolf Creek.  It has been subject to human influence through grazing by cattle  and to Christmas tree farming.  These activities tend to retard maturation  of the overstory and most of the trees are less than 20 years old.  35.  Needlegrass (cover = 5.8%) and bluegrasses (cover - 1.5%) dominated pinegrass in the sampled area, but the reverse appeared true in some of the shaded areas on more northerly exposures.  Some 17 species of forbs  covered about 5% of the ground, with strawberry and violets, each with 1.7%  cover, most plentiful.  Bearberry was abundant and covered 15.1% of  the ground, and Oregon grape (Berber-is nervosa) covered another 1.7% (Table  II).  Saskatoon and flattop spirea {Spirea lueida)  dominated'the shrub  layer, but six other species were common and contributed to a total of 88.5  twigs/m (Table III).  Annual production was moderate, averaging  0.64  g/nr for each species and 3.2 g/nr for all species combined.  2  Utilization values are consistently high, averaging 56.6% (Table V).  The trees here are predominantly douglas f i r with 4120 saplings and 820  trees/ha.  Some lodgepole pine and larch (Larix  occur (Table VI).  Table VII  occidentalis)  also  shows that this overstory provides a total  of 3122 kg/ha of f i r browse, of which 72% is found on the saplings. Light use of browse was noted on the trees, but moderate to heavy use was common on saplings.  Some individuals were severely hedged.  Immature Forested Open Type:  Following removal of the original plant communities on the terraces or benchlands of Premier and Wasa Ridges, the i n i t i a l forest regeneration in the drier areas has been a mixture of yellow pine and lodgepole pine.  36.  TABLE VII AVAILABLE FIR BROWSE:  Habitat Type Young Fir  WINTER RANGE  kg/h (Sapl.)  kg/h (Trees)  kg/h (Total)  618 (T)* 1638 (N) ** 2256 (B)***  247 (T) 620 (N) 867 (B)  865 (T) 2258 (T) 3123 (B)  Immature Forested Dense  194 (T) 480 (N) 674 (B)  22 (T) 55 (N) 77 (B)  216 (T) 535 (N) 751 (B)  Mixed Age Fir  112 (T) 280 (N) 392 (B)  82 (T) 227 (N) 309 (B)  194 (J) 507 (N) 701 (B)  Immature Forested Open  23 (T) 56 (N) 79 (B)  17 (T) 42 (N) 59 (B)  40 (T) 98 (N) 138 (B)  Maturing Open  17 (T) 43 (N) 60 (B)  15 (T) 37 (N) 52 (B)  32 (T) 80 (N) 112 (B)  Based on the equations: (T)  log  e  (twig biomass) = -3.2534 +2.3442 (log  (N)  log  e  (needle biomass) = -2.7427 + 2.5913 (log  e  stem diameter); r  = .943  (B)  log  e  (browse biomass) = -2.0347 + 2.3593 (log  e  stem diameter); r  = .923  *(T) = twig biomass **(N) = needle biomass ***(B) = browse biomass  e  stem diameter); r  = .958  37.  These stands are relatively open and tree canopy coverage is about 40 to 50%.  Most of the trees are in the later part of the 30 to 50 year age  class.  Bluebunch wheatgrass is well distributed in the understory, but the plants are stunted and cover only 2.2% of the ground surface. grasses are only found In trace amounts. phlox (cover = 14.6%), Agvoseris  Other  The abundance of forbs like  sp. (cover = 3.4%) and pussytoes  (cover = 2.6%) point to an understory that is overutilized by grazing animals (Kemper, 1971).  In a l l , 22 forb species were found in the  understory and they cover about 25% of the surface.  Bearberry covers  10.8% of the ground and nearly 5% of the soil is bare.  The shrub layer here is much reduced from the open types that lack a significant conifer canopy.  Five species, dominated by saskatoon and  bitterbrush, provide 49.6 twigs/m ; a 50% reduction from the open serai 2  shrub areas with similar soil and moisture regimes (Table III).  Shade  and insufficient moisture are probably important in limiting annual production to 1.7 g/m , averaging only 0.43 g/m for each species (Table 2  IV).  2  Utilization is moderate, averaging 38.7% (Table V).  The overstory in this type consists of 25.7 trees and 1634 saplings/ha. The trees are 73% yellow pine, 23% lodgepole pine, and 4% douglas f i r . The sapling stands are composed of these three species in the proportions of 81%, 6%, and 13%, respectively (Table VI). 3.0 m t a l l , while the f i r are only 1.5 m t a l l .  The pine saplings: average The small fraction of  38.  f i r in this mixture results in only 138 kg/ha of conifer browse being available in this habitat (Table VI).  As in the Young Fir type, use of  f i r is light on trees and moderate to heavy on.saplings.  Immature Forested Dense Type:  This habitat is interspersed with the Immature Forested Open type and represents areas on the slopes and terraces that have a more moist micro-climate due, primarily, to exposure.  Yellow pine was the i n i t i a l  recolonizer following removal of the original montane forest, but douglas f i r is rapidly taking advantage of its greater tolerance for cooler, moister soils.  These stands originated about 40 to 50 years ago and  generally exceed 60% canopy closure.  Due to light conditions, the understory is sparse and the l i t t l e used pine grass covers only 8.3% of the ground.  Fourteen forb species  were found, but only one exceeds 1% cover and total forb cover is less than 2.5%.  Bearberry covers only 3.3% of the ground and 11.4% of the  soil is bare.  In addition to low light levels, the accumulation of  heavy l i t t e r (fallen dead branches) that covers only 80% of the ground (Table II)  is suppressing plant growth.  Only three species of shrub were found in the sample plots. Saskatoon, bitterbrush and flattop spirea supply only 12.5 twigs/m  2  (Table III)  and annual production of only 0.31 g/m for the two former  species combined (Table IV).  2  Utilization of shrubs is below average at  39.  30.3% (Table V), possibly reflecting lower palatability of shrubs growing in shaded conditions.  Sixty-eight percent of the 1400 trees/ha in habitat are yellow pine which are older and larger than the 32% douglas f i r .  However, 95%  of the 5850 saplings/ha here are f i r that are doing better than the pine (Table  VI).  It is the large number of young f i r trees that give this type its dense characteristic.  The degree of canopy closure has not only reduced  the understory, but most of the lower limbs on the "spindly" saplings have died.  Although this type provides more f i r browse per unit area  than is supplied by the Immature Forested Open or Maturing Open types, on a per sapling basis this habitat provides only one-third to onefourth the amount of the other types.  Quantitative differences may be  intensified by a qualitative difference since very l i t t l e of the available browse in the Immature Forested Dense type is current annual growth (see also Discussion).  Some saplings growing in small openings in this type were heavily browsed, but generally the sparsely-needled, dying branches within the truly dense stands were but l i t t l e used.  Mixed Age Fir Type:  This forest type represents areas that must have been selectively  40.  logged, or that were repeatedly subjected to "cool" fires and have developed a multiple-layered tree canopy composed almost exclusively of douglas f i r .  Canopy closure is highly variable, but'generally ranges  from 40 to 50% on south exposures and from 50 to 70% on north aspects.  Pinegrass (cover = 5.7%), carex (cover = 6.4%), fescue [cover = 1.8%), and traces of bluebunch wheatgrass provide a ground cover of about 14% in this habitat reflecting a good moisture regime and moderate grazing.  Wild strawberry (cover = 2.3%) and 20 other forbs cover about  4% of the ground.  Bearberry is common covering nearly 5% of the surface,  and only 2.3% of the soil is bare (Table  II).  Saskatoon is by far the most plentiful shrub with 27.8 twigs/m . 2  Snowberry and flattop spirea are common, providing 7.4 and 8.4 twigs/m , 2  respectively, and bitterbrush and rose also occur. is 52.2 twigs/m (Table III), 2  0.98 g/m annually (Table IV). 2  Total twig availability  and four of these species together produce Utilization of saskatoon, spirea and  rose is moderate, but snowberry and bitterbrush are ratherly lightly used (Table V).  The top layer of the overstory consists of f i r trees in excess of 60 years old. tall.  These have a mean dbh of 29.2 cm and average about 15.2 m  There are 175 of these trees/ha.  The next layer of younger  trees, aged 30 to 50 years have a mean dbh of 9.9 cm and average 8 m tall.  There are 725 of these trees/ha.  The bottom layer consists of  the sapling regeneration with an average height of 3 m and a density of 1275 stems/ha.  41.  The mature trees provide no browse for the deer, but the other two layers supply 702 kg/ha of f i r browse.  Use of browse on the trees is  light to moderate, but on the saplings is heavy to excessive.  Many very  small firs (less than 1 m tall) were browsed to the point where they were stunted and deformed.  Maturing Open Type:  This habitat is also a multiple layered type, but differs from the Mixed Age Fir in both density and species composition.  Canopy closure  rarely exceeds 40% and the drier nature of the soils in this type allow a mixture of pine and f i r .  Carex disappears from this drier type leaving pinegrass (cover 4.4%) the predominant species in the approximately 5% grass cover.  The  dry nature of the soil also prevented the 22 forb species found here from covering more than 4% of,the ground.  Bearberry which does well on  drier soils given sufficient sun covers 7.6% of the surface and only 1.9% of the soil was bare (Table  II).  Shrub biomass is slightly improved over the Mixed Age Fir type with both bitterbrush and saskatoon providing over 20 twigs/m . 2  Snowberry  and rose also contribute to the 52.4 twigs/m (Table III),  and all four  species produce a total of 2.32 g/m annually (Table IV).  The low  2  2  utilization values given tn Table V are most.likely a reflection of the influence of the unusually deep s'now during the winter prior to the sampling period.  42.  Both mature yellow pine and mature, douglas f i r are found here in densities of 88 and 100 trees/ha, respectively.  Younger pine and f i r  also occur with densities: of 150 and TOO stems/ha, respectively.  Fir  dominates the sapling layer with 113 stems/ha compared to only 50 for pine (Table VI).  Due to the openness of this type, the browse per tree is high, but total f i r browse is only 112 kg/ha (Table VII).  Level of use of the f i r  browse is inversely proportional to tree size:  the saplings are heavily  used, moderate to light use occurs on the larger saplings, and the trees are only lightly browsed.  Riparian Mature Spruce:  One small area in the Wolf Creek bottom is occupied by a habitat similar to the spruce communities found on the islands and banks of the Kootenay River.  Due to its limited distribution in Wolf Creek and also  to the level of agricultural  disturbance along the Kootenay floodplain  in this region, the type is of reduced importance on this winter range. It does appear to be a productive and preferred type where i t occurs though, and future investigations of white-tailed deer habitat in the Rocky Mountain Trench should place more emphasis on the study of this, and other riparian communities in undisturbed sections of the river floodplain.  Quantitative data were not gathered on the herbaceous layer, but i t  43.  was generally composed of lush, sedges with a wide variety of wetland plants such as:  skunk cabbage (Lysichiion  (Equisetum  avvense)i,  bedstraw (Galium  (Smilaoina  amplexicaulis),  kamtchaicense),  boreale)  false Solomon's seal  twisted stalk (Streptopus  various orchids (Cyprpedium  spp.].  horsetail  amplexifolius)  and  Oregon grape is common and covers  8.3% of the ground.  The shrub layer here is well developed and abundant, nonwoody growth of species like elderberry (Sambuous racemose and thimbleberry (Rubus parviflorus) story.  arbovescens)  var.  add to the lushness of the under-  Fifteen different shrub species here provide a total of 81.9  twigs/m (Table III). 2  Most important in providing browse are snowberry,  flattop spirea, rose, thimbleberry, and red osier dogwood.  Four of the  species were sampled for productivity and averaged .78 g/m , totaling 2  3.13 g/m (Table IV). 2  If all species were included, production might  approach 5.0 g/m in this type. 2  Utilization values are high on all  species except snowberry and rose (Table V).  Although some f i r did occur on the edges of this type in Wolf Creek (Table VI),  the model stand would lack this species.  tolerant white spruce (Picea  glauca)  is the major tree here having a  density of 267 mature trees and 233 younger trees/ha. wood (Populus stems/ha.  trichocarpa)  The more flood  Larch and cotton-  are also found as mature trees, each with 33  Younger cottomwobds, aspen, paper birch {Betula  pccpyvifera)  and choke cherry also contribute to a total density of almost 2000 stems/ha.  Many of the saplings of these, species are heavily browsed.  44.  Analysis of Summer Range Habitat  The plant communities of the summer range have been studied in much less detail than those of the winter range.  Accordingly I have chosen  to use the more general term "associations" rather than habitat types. The area has been divided into three plant associations: and Immature Forest.  Shrub, Brush  Sampling in each was designed only to document  variations and possibly identify trends in the associations.  Within the  Brush and Shrub associations, elevational trends were found, but the Immature Forest association was highly variable and trends were more d i f f i c u l t to ascertain.  Shrub Association:  Although the "shrubby" appearance of this association is consistent, Table VIII and Figure 6. illustrate the changes in the species composition of the shrubs with increasing altitude.  At lower levels the association  is dominated by bitterbrush and saskatoon; a situation not too unlike the shrublands of the winter range.  However, bitterbrush quickly  disappears upslope and is replaced by species indicative of more moist areas:  evergreen ceanothus [Ceanothus velutinus)  and snowberry.  Saskatoon and rose vary about high (16.6%) and low (1.6%) mean coverage values, respectively, and no definite trend is expressed.  Incomplete  data prevent analysis of elevati.0n.9V changes in the herbaceous layer. However, i t is expected that total herb cover would increase slightly with more favorable conditions (i.e.  greater moisture and reduced cattle  45. TABLE V I I I SPECIES COMPOSITION OF SHRUB ASSOCIATION:  Species  2650'  SUMMER RANGE  Elevation 2800'" 3300'  3600'  Shrubs: Amelanchier a l m ' f o l i a Arctostaphylos u v a - u r s i Acer glabrum Ceanothus v e l u t i n u s Populus tremuloides Purshia t r i d e n t a t a Rosa woodsii S a l i x glauca  30.0/100 tr/10  6.7/100 **  1.7/67  5.0/33  6.7/100  15.0/100 tr/33  tr/33 2.5/100 2.5/100  17.5/100* 1.5/10  5.8/67 1.7/67 1.7/67  Shepherdia canadensis Spirea l u c i d a Symphoricarpos albus Total shrub cover Herbs:  35.7  49.2  2.0/50 tr/33 tr/33  3.0/50 2.0/50  1.7/67 5.0/33 5.8/67  15.0/100  25.9  Achillea millefolium Agropyron spicatum Antennaria m i c r o p h y l l a Arabis h o l b o e l l i  5.5/57 11.4/83  Astragalus miser Balsamorhiza s a g i t t a t a Bromus tectorum Calamagrostis rubescens  13.3/67 1.8/10 tr/33 tr/10  Galium boreale Koleria cristata Penstemon confertus Poa pratens  tr/37  2.4/43 1.2/17 5.1/40 tr/10 tr/10 tr/17  tr/10 1.0/40 tr/10 tr/10 2.6/53 1.3/20 3.2/17  Solidago s p a t h u l a t a Tragopogon dubius  8.0/100 56.0 **  ***  Campanula r o t u n d i f o l i a Chryopsis v i l c s a Epilobium minutum Erigeron pumilus  12.0/100 ** 2.0/50 12.0/100  1.2/13 7.3/60 7.1/57 3.2/27  tr/27  Total herb cover  38.3  Other: Bare s o i l Litter  17.8/83 57.9/100  27.5  26.1/73 40.3/97  *% ground cover/% frequency of occurrance ( t r = l e s s than 1%) **data not a v a i l a b l e ***only species w i t h a t l e a s t 10% frequency i n a t l e a s t 1 s i t e shown.  46. 30  A  r  /  \  / 25  \  \  Amelanchier  \  \  Purshia  \  tridentata  Ceanothus  \  Rosa  alnifolia  velutinus  woods ii  \  20  \  \ \  or UJ  \  >  \  o  \  \  10  0  2600  2800  3000  !  ELEVATION  3200 IN  3400  3600  FEET  FIGURE 6. Elevational trends in coverage values of four shrub species in the Snrub Association on the summer range.  47.  grazing) at higher elevations.  Species composition should also change as  dry soil ^"plants like bluebunch wheatgrass, pussytoes, and  Astragalus  miser disappear and are replaced by more moisture dependent species.  Brush Association:  This riparian association occurs on islands in the Kootenay River and along streams at various elevations. consists of two sub-associations. a willow-dogwood-rose  On the river islands, i t  Along the banks and in old channels,  sub-association with a Carex-Elymus  understory  provides a dense thicket two to four meters high of varying width that supplies both forage and cover.  Local, moderate use of the leaves of  these shrub species was noted in late summer and f a l l .  The other sub-  association occurs on interior, drier portions of the islands and is characterized by large clumps of water birch {Betula occidentalis) 8 m tall spaced at about 10 to 15 meter intervals. almost completely covered with a layer of Votentilla approximately 1 m high. are sedges.  Beneath the Potentilla,  up to  The ground is fruticosa  that is  the only common plants  Deer were found to bed in this sub-association, but did not  feed much on the plants here.  These sub-associations have been, and are, greatly affected by annual flooding.  Recently, their productivity was seriously hampered  due to the excessive silting that occurred during the record flood of June, 1974.  The source of the s i l t was a section of the floodplain  directly upstream that had not developed a ground cover following clearing a few years before.  48.  Along the creeks at elevations above the floodplain, this association increases in species diversity, and composition of any particular section is influenced by surrounding vegetation.  Where the creeks run  through narrow channels In open areas at lower levels, the brush is dominated by aspens which occasionally reach tree size.  Red osier  dogwood is common, as are willows, rose, silverbush (Eleagnus mountain maple {Acer glabrum) and water birch. sedge and forb.  commutata),  The herb layer is mixed  These areas often show abuse from cattle grazing and  trampling.  At several elevations, the wet areas spread out and form sloughs or bogs.  In some places the process has been encouraged through damming by  beaver (Castor canadensis). (Salix  glauca,  S. barrattiana,  Equisetum understory.  Here, the brush is predominantly willows and S. bebbiana) with a sedge-rush-  At the marshy margins, alders (Alnus sinuata)  and  some birch and aspen may form a dense wall of'woody vegetation, with a sparse sedge-forb understory.  These areas are less affected by cattle.  Where the creeks flow through established forest communities, especially at higher elevations, the Brush association becomes much less dense and willows and birches are highly reduced.  Red osier dogwood,  rose and some alder and maple are common, as are shade tolerant species like twinberry (Lonicera  involucrata)i,  Cattle rarely disturb, these areas.  thimbleberry and snowberry.  49.  Immature Forest Association:  The majority of the area considered in the "summer range" was subjected to extensive severe fires In the early 1900's.  These fires  created ranges that supported deer, elk and "hundreds of wild horses" (H. Campsall, pers. comm.) about 30 to 40 years ago.  Now, however, the  range has regenerated into relatively dense immature forests.  Once tree growth began, the more favorable moisture regime west of the floodplain promoted rapid growth.  Table IX l i s t s the composition of  the overstory at the forested sites sampled west of the Kootenay River. On the floodplain, immature yellow pine stands are found that are similar to the Immature Forested Open type on the winter range, except for more intense cattle use.  The high degree of variation above the  floodplain is the result of micro-climatic factors.  As a result of more  favorable growing conditions, the average dbh here is greater than on the winter range despite similar stand age.  Dominance of lodgepole pine  and the presence of larch, aspen and birch reflect the more severe fire history and increased moisture, respectively, relative to the winter range.  The wide variation in the overstory at the sample sites introduces another element in analysis of understory with respect to elevation. For this reason, caution must be used in evaluation of the data. types of changes were noted In shrub cover:  Two  density and species composition.  Figure 7. shows that the total shrub cover at sites above 2800 feet was  50. TABLE IX 0VERST0RY COMPOSITION OF IMMATURE FOREST ASSOCIATION:  Site  Elev.  Species  2600'  Pinus ponderosa Pseudotsuga menziesii  2800'  Pinus ponderosa Pseudotsuga menziesii Pinus contorta  3000'  Pinus ponderosa Pinus contorta Pseudotsuga menziesii Larix occidentalis Populus tremuloides  3300'  Pinus contorta Larix occidentalis Populus tremuloides Pseudotsuga menziesii Betula papyrifera  3300'  Pseudotsuga menziesii Populus tremuloides Larix occidentalis Pinus contorta Pinus ponderosa  Stems/ha 840 400 400 10 1650 370 240 10 50 670  SUMMER RANGE  Ave.dbh 10.7 cm sapl.* 10.2 sapl.  Ave. ht 5.5 m 2.4 5.5 1.8  11.2 sapl. sapl. 14.7  6.4 3.0 3.0 11.0  90 90 300 440 • 50 10 20 70 1070  12.4 15.7 8.4 sapl. 11.4 sapl. sapl.  8.2 12.2 7.3 3.0 10.4 4.3 6.1 3.0  1433 67 733 567 167 167 33 33 67 3264  12.2 sapl. 9.9 sapl. 10.9 sapl. sapl. 5.6 sapl.  13.7 6.1 11.9 3.4 13.7 4.6 4.6 7.6 4.6  340 260 40 150 10 10 10  11.4 sapl. 13.2 sapl. sapl. sapl. sapl.  8.2 3.4 10.1 2.7 6.1 2.4 6.1  .6.4  TABLE IX (cont.) Site  Elev.  Species  3300'  Pinus contorta Pseudotsuga menziesii Larix occidental is Betula papyrifers  3500'  Pinus contorta Larix occidental is Pseudotsuga menziesii  3700'  Pseudotsuga menziesii Larix occidental is Pinus contorta Thuja plicata  4100'  Pinus contorta Larix occidental is Populus tremuloides Populus trichocarpa  *sapl. = sapling  Stems/ha  Ave.dbh  Ave. ht.  11.4 cm sapl. 12.7 sapl. 16.0 8.4 sapl.  12.2 m 6.1 12.2 5.2 12.2 10.1 4.6  190 30 30 10 20 280  10.9 sapl. 13.2 sapl. sapl.  12.2 3.0 13.7 1.5 1.5  733 800 533 100 167 567 2900  11.7 sapl. 18.0 sapl. 17.8 sapl.  9.1 4.0 16.2 4.0 12.2 1.5  1720 550 40 20 10 2340"  9.1 sapl. sapl. sapl. sapl.  12.2 4.6 4.6 3.0 3.0  440 80 90 90 60 120 410 1290  Total Shrub Cover oo in  co tn —  p 3» 1 to  -to  D  Purshia  Arctostaphylos  Rosa  Linnea  Vaccinium  Shepherdio  tridentata  uva-ursi  woodsii  borealis  spp.  canadensis  n  1 1 1  •A.:v-  m  Frf>  I I I I I ELEVATION  IN  -Fr  FEET  ^o'oUo'o'o oo o o o o o  1  CN CN P") CO O CO T  FIGURE 7. Elevational trends in coverage values of shrub species in the Immature Forest Association on the summer range.  53. TABLE X UNDERSTORY COMPOSITION OK IMMATURE FOREST ASSOCIATION: SUMMER RANGE  Species*  2600  2800  3000  Elevation)feet) 3300 " 3300  3300  3500  3700  4100  Shrubs: Acer glabrum Alnus alnifolia Amelanchier alnifolia Arctostaphylos uva-ursl Berberis nervosa Ceanothus velutinus Cornus stolonifera Juniperus communis J . scopulorum Llnnaea boreal is Lonicera involucrata L. utahensis Henziesia ferruginea Populus tremuloides Prunus virginianus Purshia tridentata R1bes irriguum Rosa acicularis R. woodsii  2.1/83  6.6/100  22.5/100  11.7/62  7.8/43  3.9/40 5.6/43  Achillea millefolium Agropyron spicatum Agroseris sp. Allium cernuum Arnica cordiflora Astragalus miser Calamagrostis rubescens Carex concinoides Cornus canadensis Festuca scabrella Fragarla Glauca Gentlana acuta Hedysarum sulphurescens Heuchera cylindrica Hleracium albiflorum Koleria cristata Melampyrum lineare Penstemom confertus Pyrola asarifolia  Other: Bare soil Litter  1.5/10 7.8/60  8.5/37  10.9/63  tr/33 tr/33 2.5/17 2.1/83  2.5/100 15.0/100  tr/33  2.5/100  10.0/67 2.5/100  1.7/67  2.5/100  1.7/67  tr/33  tr/33 2.5/20  tr/10 3.3/33  6.7/100  5.8/67  8.2/37  9.6/50  5.7/63 1.3/50  2.5/100  7.5/50 7.5/50  2.5/100  1.3/50  tr/33  10.0/67  1.7/67 tr/33  tr/33 1.3/50  tr/33  tr/17 tr/17  tr/13 1.2/30 tr/10 tr/10 2.2/15 2.1/13  tr/10  10.8/100  2.5/100 tr/33  5.0/33  30.0/100  54.2/100  10.0/67  30.0/100 17.5/67  .15.0/100 10.8/100  17.5/100  29.2/67 15.0/100 25.0/67  tr/3  8.6/80  16.0/90  20.1/90  14.1/90  1.0/23  37.5/100  26.3/100  15.0/100  37.5/100 5.8/67 5.0/33 2.4/17  8.8/100 1.3/50 23.2/70  11.4/43  7.4/70  tr/17  19.7/37  1.8/17 1.1/10  tr/14 1.1/10 tr/10 1.3/20  tr/18  6.6/32 66.3/98  25.8/100  3.4/23  1.3/20  1.5/10  13.0  6.7/100  3.8/33  1.5/10  tr/6  tr/33  tr/10  5.3/47 7.9/57 1.6/13  1.3/50 1.3/50  tr/33 18.3/100  tr/33  P. secunda Total herb cover  1.6/13 10.3/37  tr/33  S. glauca S. mackenziana Sambucus racemosa  Spirea lucida Symphoricarpos albus Vaccinium caespitosum V. scoparium Herbs:  2.5/100  5.0/33  Rubus parvlflorus R. spectabilis Salix agrophylla S. bebbiana  Shepherdia canadensis  tr/33* 1.7/67 tr/33  1.3/14  tr/27  1.3/25  tr/20 tr/10  19.0  10.0  26.0  22.0  18.0  22.0  14.0  6.0/33 64.3/100  1.6/13 83.4/100  71.2/100  72.1/100  77.7/100  69.1/100  73.0/100  * Only species with at least 10% frequency of occurrence in at least one macroplot shown. '* t ground cover/% frequency of occurrence (tr • less than 1%).  1.8/23 13.0  63.2/100  54.  greater than that at or below this level (see also Table X).  The d i f -  ference is probably due to increases soil moisture resulting from greater precipitation and greater water holding capacity due to nearsurface bedrock instead of a thick gravel substrate beneath the s o i l . Reduced cattle grazing may also be a factor.  Changes in moisture and  temperature are also reflected in the species composition of the shrub layer as bitterbrush and bearberry disappear and soapolallie canadensis),  {shepherdia  huckleberries [Vaccinium scoparum and V. caespitosum),  twinflower (Linnaea borealis)  and  increase at higher levels.  At a l l elevations, herbaceous plant cover was limited due to the overstory.  No distinct trends were found except for an increase in  pinegrass at mid-elevations that declined farther up slope, possibly because of the dense overstory of both trees and shrubs (see Table X).  Deer use of available forage in the Immature Forest association was impossible to estimate.  At lower elevations, cattle use, which in some  areas was excessive, compounded deer use.  At higher elevations, the  understory was so dense and deer were so widely distributed that i t was impractical to attempt any type of forage consumption survey.  Weather  Temperature:  Figure 8. is a graph of the averages of the minimum temperatures  ••  5°r  Winter  1975-76  Winter  1974-75  P e r i o d s of  1  I DECEMBER  JANUARY  I FEBRUARY  temperature  in winter  of  1  I MARCH  APRIL  MONTH  FIGURE 8.  warmer  minimum  Three-day average minimum temperatures, December to April, 1974-75 and 1975-75.  1974-75  56.  for three day periods from 1 December to 3Q A p r i l , 1974-75, and 1975-76. The three day average was chosen because ft smoothed out the curves while maintaining clear evidence of the temperature patterns.  From this  i t can be seen that the winter of 1975-76 was not as cold as 1974-75. The only notable exceptions to this were the month of December, and the f i r s t two weeks of March which were warmer during the f i r s t winter.  Sunshine:  Figure 9. illustrates the average daily amount of direct sunlight, during the winter months in both years.  It demonstrates that the winter  of 1975-76 was sunnier from mid-January to late March, the period of maximum thermal stress.  Snow:  Thermal stress is one aspect of winter severity; the others are snowfall and the characteristics of the snowpack.  Snow conditions are  important because they influence the deer's ability to deal with thermal stress by 1.) increasing the energy cost of moving about, 2.)  restricting  the use of some habitat types and, 3.) burying certain portions of the understory making them unavailable for feeding. vulnerability to certain predators.  Snow may also increase  Although a crusted snow that will  support a deer may raise the upper limits of the available forage zone, such conditions were not observed on the study area.  1974 -75 1975-76 to rCC  uj<  iorr 3  LL LiJ  LU  rr !Z  1  ° =>5 < o rr x - i LU z  > u. 3  <Oy)  DECEM8ER  JANUARY  i  FEBRUARY  MARCH  APRIL  MONTH  FIGURE 9. Average hours of direct sunlight, December to April, 1974-75 and 1975-76.  r  58.  80 r-  E o  <  60  1974-1975  <= o |  x lz o  j  1975-1976  40  Q  UJ  <  20  o o < DECEMBER  JANUARY  FEBRUARY  MARCH  MONTH  FIGURE 10.  Total snowfall, December to March, 1974-75 and 1975-76.  10  S N O W ON T H E  I—I  AT  cn cr I  TO  cr>  m  MONTH'S  GROUND END  (Cm) OI  o m o m  a ro •a  ro  o  to  OI  o  2  m oo 3 O  >  cu ct-  C > -<  O 3  fD  a  tt> O  o  m  CD  c >  -<  ro 3  cr fD -s  3 >  o 5=  X  •a  > I  73  cu Q.  to  cn  i  •si  '69  CO •Nl  J> i -si  60.  Figure 10. shows the pattern of snowfall and Figure 11. the depth of snow on the ground at the Cranbrook Airport for both winters.  It can  be seen that not only was there substantially more snow in 1974-75, but the cold weather during that winter prevented melting and settling of the snow, and the snowpack lasted longer.  The same patterns were displayed  on the study area, but the different degrees of canopy closure and aspect resulted in habitat-specific depths and melting rates.  Although density was not quantified, i t is intuitively obvious that the prolonged cold weather in 1974-75 kept the deep" snow very soft.  Use of Winter Range Habitats:  Tables XI, XII and XIII l i s t the results of the systematic track counts and their statistical treatment.  In all cases the chi-squared  test showed highly significant differences from a "uniform" distribution (p^O.Ol), so I have only tabulated the information on degree of habitat selection.  Table XIV l i s t s the results of the pellet group counts for both winter periods.  The estimated mean number of groups per plot and the  sample variance in each year as well as the percent change between years are presented for each type.  Two habitats were found to occupy more than one aspect, and i t was felt that better understanding of their use might be gained by sub-  61.  TABLE XI TRACK COUNT ANALYSIS: Transect P r o p o r t i o n Length of Total (km) Transect Length  Habitat Grassland  1.13  .146  WINTER 1974-75  Track °c °d 2 d  R i p a r i a n Brush  Y . F . (west a s p e c t )  e  M.O.  .000 .000 .024  Confidence Degree Interval Snow of fa=0.05) S e l e c t i o n Depth 3  .OOOSpJ.063  -  30 cm 50 cm 40 cm  —  0.32  .041  3 3 20  .042 .103 .244  .000{p*.097 .000jp$.235 .133*p$.355  0 0 +  30 cm 40 cm 30 cm  0.64  .083  7 3 15  .097 .103 .183  . 014?p*.180 .000.<p<.235 .173jp<r.l93  0 0 +  25 cm 40 cm 30 cm  3.06  .396  42 4 15  .583 .138 .183  .445*p<.719 .000$p<.287 •173?p*.193  +  -  20 cm 50 cm 40 cm  20 19 30 .  .278 .655 .366  .152$p*.402 .449*p$.861 .242cp<.490  0 + 0  15 cm 25 cm 20 cm  M.A.F. (south aspect)  2.58  Total  7.73  a. b. c. d. e.  Proportion of Track  .333  72 29 82  Degree o f S e l e c t i o n : .. - = avoidance; 0 = p r o p o r t i o n a l ; + = s e l e c t i v e . Count conducted on 18 January, 1975. Count conducted on 16 February, 1975. Count conducted on 10 March, 1975. Y . F . = Young F i r ; M.O. = Maturing Open; M.A.F. = Mixed Age F i r .  62  TABLE XII  Habitat  TRACK COUNT ANALYSIS:  24 JANUARY, 1975  Proportion Transect Length of Total (km) Transect Length  Proportion of Track  Track  Confidence Degree Interval of . (<*-0.05) S e l e c t i o n a  Snow Depth  Grassland  1.61  .251  39  .115  .109Sp<.201  _  30 cm  Serai Shrub  0.16  .026  19  .056  .049<p<.063  +  25 cm  •0.48  .077  19  .056  .049<p<.063  -  25 cm  I.F.O.  0.32  .051  10  .029  .008<p$.050  -  20 cm  M.A.F. (north aspect)  3.70  .590  252  .743  .688«p$.798  +  15 cm  Total  6.27  Y.F. (north a s p e c t )  a. b. c.  0  339  Confidence i n t e r v a l on p r o p o r t i o n o f t r a c k s (ex= 0 . 0 5 ) . Degree o f S e l e c t i o n : - = avoidance; 0 = p r o p o r t i o n a l ; + = s e l e c t i v e . Y . F . = Young F i r ; I . F . O . = Immature Forested Open; M.A.F. = Mixed Age F i r .  63 TABLE X I I I TRACK COUNT ANALYSIS:  Transect P r o p o r t i o n Length o f Total (km) Transect Length  Habitat  WINTER 1975-76  Track  Proportion of Track  Confidence Degree Interval of (cx=0.05) S e l e c t i o n a  -  0  Snow Depth  - '' 0  2 cm 15 cm 15 cm  0.32 0.32 0.32  .010 .009 .011  0 d 11  .000 .001 .019  .000* p< .004 .003< p< .035  1.61 2.25 0.32  .048 .066 .011  12 90 45  .022 .055 .080  .0051 p{ .039 .0381 p< .072 .0485 p5 .112  2.74 3.06 2.25  .082 .090 .075  7 27 3  .013 .016 .005  .000* p i .027 .007$ p< .025 .000$ p< .013  1.93 1.77 1.77  .058 .052 .059  80 203 23  .147 .123 .041  .1041 p i .190 .100$ pi .146 .017* p< .065  + 0  2 cm 15 cm 15 cm  2.25 2.25 2.25  .067 .066 .075  10 120 40  .018 .073 .071  .0025- p i .034 .055$ p i .091 .0411 pi .101  0 0  2 cm 15 cm 15 cm  Y.F. (west aspect)  1.29 1.29 0.64  .039 .038 .021  18 150 60  .033 .091 .106  .0121 pi .054 .071 5 pi .111 .070* p f .142 .  0  2 cm 15 cm 5 cm  I.F.D.  5.47 3.86 4.18  .163 .114 .139  79 115 31  .145 .070 .055  .102< p< .192 .053< p i .087 .028i pi .082  0  2 cm 15 cm 15 cm  I.F.O.  8.37 8.53 8.37  .250 .251 .278  131 220 75  .240 .133 .133  .188* p i .292 • 110J p i .156 .0941 p i .172  0 -  2 cm 15 cm 15 cm  5.15 5.63 4.83  .154 .165 .161  146 305 176  .267 .185 .312  .2141 p i .316 .1581 p i .212 .257£ p$ .367  -+  2 cm 15 cm 5 cm  4.35 4.99 5.15  .130 .147 .171  63 416 101  .115 .253 .179  .0781 p i .152 .219* p& .277 • 1 3 3 < p l .225  0  Aspen  R i p a r i a n Brush  Grassland  S e r a i Shrub  Y.F. (north  M.O.  M.A.F.  Total  aspect)  f  33.48 33.95 30.08  C  2  e  0 +  +  + +  -  0 +  +  0  546 1648 565  a. b. c. d.  Confidence i n t e r v a l on the proportion of tracks (<*.= 0.05). Degree o f S e l e c t i o n : - = avoidance; 0 = p r o p o r t i o n a l ; + = s e l e c t i o n . Count conducted on 31 December, 1975. Count conducted on 12 January, 1976.  e; f.  Count conducted on 27 February, 1976. Y.F. = Young F i r ; I.F.D. = Immature Forested Dense; I.F.O. = Immature Forested Open; M.O. = Maturing Open; M.A. F. = Mixed Age F i r .  2 cm 15 cm 15 cm 4 cm 15 cm 15 cm  2 cm 15 cm 15 cm  TABLE XIV PELLET GROUP COUNT ANALYSIS 1976  1975 Habitat type  Mean  Var.  Aspen Grassland Riparian Brush Serai Shrub  0.20 1.15 2.00 1.00  0.00 1.00 1.00 0.07  Y . F . (North aspect) Y.F. (West aspect) I.F.O. I.F.D. M.O. M.A.F. (North aspect) M.A.F. (South aspect)  0.80 5.40 0.77 1.50 1.23 0.90 4.35  0.04 0.80 0.08 0.11 0.05 0.25 0.16  Overall average  1.59  0.05  b  0  Rating  6 1* 7 3* 4 5 2*  Mean  Var.  a 0.15 2.40 0.55  0.01 4.00 0.02  0.53 1.60 0.37 0.73 1.18 0.60 0.85  0.14 0.45 0.06 0.06 0.04 0.20 0.05  0.79  0.02  Rating  % Change -87* +20 -45*  6* 1* 7 4 2*  5 3  -34* -70* -48* -51* - 4 -33 -80* -50*  a. uncountable due to flooding b. Y.F.. = Young F i r ; I.F.O. = Immature Forested Open; I.F.D. .= Immature Forested Dense; M.O. ^.Maturing Open; M.A.F. = Mixed Age F i r c. total number of groups/total number of plots (excluding Aspen) * Significant at the 0.05% level.  65.  dividing them.  Thus the Young Fir type is split into north and west  aspect, and Mixed Age Fir type is split into north and south aspect.  Aspen:  This minor component of the Wolf Creek winter range was not surveyed for tracks during the 1974-75 winter, but pellet groups were counted here in the early summer of 1975 to determine previous winter use. The reverse was true the next year as tracks were counted, but flooding prevented counting of pellet groups.  Both Table XIII and Table XIV reveal l i t t l e winter use of the Aspen habitat type.  The track counts indicate deer avoid this type through  much of the winter, and the pellet group density was lowest here.  Grassland and Cultivated Fields:  The track count data (Tables XI, XII and XIII) demonstrate that throughout the winter deer avoid the Grasslands and Cultivated areas regardless of snow depth.  The levels of use indicated by the pellet  group densities are the result of spring grazing of these areas by the deer.  The temporal pattern of use of these areas in the spring has two aspects:  seasonal and daily.  These are generally similar for both  types, but the Grasslands receive much less use.  Data from several  Bradford  t  1  o  o  Jenning's Upper  *  *  Jenning's Lower  DATE (SO DAY PERIODS) FIGURE 12.  Seasonal pattern of use of four a l f a l f a f i e l d s on the study area, March to May, 1375.  CM CTl  FIGURE 13.  •  • Bradford (no traffic)  A  A Jennlng's Lower (grave! road)  A  A Dr.Green  (Hwy 9 3 - 9 5 )  Daily pattern of use of three alfalfa fields on the study area, March to May, 1975.  68  fields are presented here to demonstrate the characteristics of use. Figure 12. shows the average number of deer seen in four fields for 10 day periods in the spring of 1975. There is a definite rise and fall in numbers over a period of several weeks in each f i e l d .  Figure 13. illustrates  the daily pattern of use for three fields combining several days' data for major use periods in the spring of 1975.  Data from the "tails" of  the lines in Figure 12. were omitted, as were counts on unseasonably cold or rainy days.  The influence of distrubance can be seen by examining  the diel patterns of use of the three fields.  In a field with no road  past i t (Bradford's), use may begin early in the day and rise steadily to a peak in the evening.  Increasing levels of traffic on a gravel road  (Jenning's Lower) and a major highway (Dr. Green's) act to suppress diurnal use of the fields.  There are insufficient data to carry the  comparison beyond 22:30 hours.  Deer use of the fields and Grasslands in spring made i t possible to obtain composition counts to estimate the adult:juvenile ratio in the population at the end of the two winters.  Table XV l i s t s the results of  these counts and indicates a significant difference in juvenile in the two winters.  survival  In the spring of 1975, there were only 42 juveniles  per 100 adults whereas in 1976, there were 60 per 100 adults.  Riparian Brush Type:  The Riparian Brush type along Wolf Creek is thought to be important to the deer on this winter range.  With the exception of the Serai Shrub  TABLE XV AGE RATIOS OF WHITE-TAILED DEER: APRIL AND MAY, 1975 & 1976 Year  Number Classified  Juveniles: 100 Adults  1975  1226  42  1976  421  60  70.  habitat, this is the only type that supplies large quantities of deciduous browse.  It is highly productive (Table IV),  the browse is well used  (Table V) and the extended growing season due to lack of moisture limitations may lead to more nutritious forage.  Pellet group densities in this type were high, but so variable (Table XIV) that i t is impossible to make clear statements about its relative standing based on this statistic alone.  The track data in  Tables XI and XIII and Figure 14a. show that there is a distinct seasonal trend in use.  In both years, intensity of use increased as winter  progressed.  The only significant use of this type was for feeding at night and an obvious daily movement of deer down from the terraces into the brush at dusk was observed.  When use intensified in late winter (particularly  in 1974-75) deer were found in some parts of this type during the day, but generally they moved back up onto the terraces by sun-up.  This  movement can be demonstrated by analysing the proportions of tracks going up or down the slope from the creek to the terraces.  A count of  58 sets of tracks that were made between 1:00 AM and 9:00 AM on 18 January, 1975, indicated that 90% of the morning movement is upslope. By contrast, a count of 329 sets of tracks representing 24 hours of movement on 23 and 24 January, 1975, indicated 51% downslope and 49% upslope movement.  71.  FIGURE 14. Degree of Selection for four habitat types, December to March, 1974-75 and 1975-76. (D.S.= proportion of tracks minum proportion of transect.)  72.  Serai Shrub Type:  This type has received a great deal of attention in the East Kootenay.  It is generally felt that the carrying capacity of ungulate  winter ranges in the East Kootenays is directly related to the proportion of Serai Shrub type on the area.  Kemper's (1971) thesis clearly explains  why this should be the case.  Whitetailed deer use of this habitat was found to be strongly influenced by snow depth during the study period.  In late January,  1975, deer were found to be selecting this type despite 20 cm of light snow on the ground (Table XII).  It was obvious at that time that they  were feeding heavily on the remaining leaves and the tips of the leaders of bitterbrush.  Although no other systematic track count covered this  type in 1974-75, use is known to have decreased dramatically in early February when snow depth quickly passed the 40 cm mark.  Only occasional  tracks were seen here, usually at the ecotone between this and a forested type, until very late in the winter when snow density increased.  Even  in late winter, however, use did not reach earlier levels as the only exposed vegetation was the tops of the larger bitterbrush plants.  In 1975-76 with different nival conditions, this pattern of use changed markedly.  Table XIII shows that this habitat was being selected  throughout early and midwinter.  Although use was estimated to be  proportional in late winter, i t is believed that the track count on 27 February, 1976, underestimated use of this type (see Discussion, p. 93.).  73.  The pellet group data may underestimate actual use of the serai shrublands, but increased selection of this habitat in 1975-76 relative to 1974-75 seems to be indicated by its rise in relative standing. Another possible indicator of greater use during the second winter is the cange in estimated pellet group density between years, 45%, which is somewhat less than the drop in the overall average density (50%).  Young Fir Type:  This habitat type was divided into two aspect classes, west and north, on the basis of the 1975 pellet group data.  The f i r s t winter's  track count data was insufficient for drawing conclusions about the importance of aspect from this measure, but Table XIII clearly supports the concept that use based on track counts varies with aspect.  Further-  more, use of this type was found to increase through the winter in both years on the west aspect (Figure 14b.), and in at least one year on the north.  The intense use of the west facing portion of this type in mid to late winter led to its having the highest pellet group density of all forested types in 1975. This was also true in 1976, although the 70% decline in density between years is above average and indicates relatively less use during the second winter.  The densities on the north slope,  however, were less than anywhere else except the Immature Forested Open, which was lower, and the north facing Mixed Age Fir which was not significantly higher (Table IV).  The below average drop in density  74.  between years indicates relatively more, use in 1975-76 than in the previous winter.  It may be that the frequency of bedding is somewhat  lower on the cooler north slope than in other areas, so these pellet group densities may underestimate actual value of the habitat.  Immature Forested Open Type:  This habitat type was covered only once in the track counts in the 1974-75 winter (Table XII) 1975.  and was found to be avoided in late January,  Snow depths here reached 30 to 35 cm by mid-February, 1975, and  use appeared to decline further.  A similar pattern of decreasing use  was seen in 1975-76, even with very l i t t l e snow (Table XIII).  The lack of use of this habitat is further substantiated by the pellet group counts.  The estimated densities here following both  winters were significantly lower than any other forested type (Table IV).  Immature Forested Dense Type:  Unfortunately, this type was not sampled by track counts in 197475, so the temporal pattern of use cannot be determined in that winter of deep snow.  However, the pellet group density here was l i t t l e below  the overall average and ranked third among the forested types in 1975. Although i t slipped to fourth, in 1976, the difference between this type and the one above i t , south facing Mixed Age Fir, was not statistically  75.  significant.  Careful interpretation of these densities must be made  though, since the track data in Table XIII indicate that deer avoid this type under certain conditions.  Maturing Open Type:  Use of this type was also markedly affected by snow depth.  Although  the deer were selecting the Maturing Open type in early winter 1974-75, the excessive snow in January and February caused the deer to avoid this type in mid and late winter (Table XI).  Heavy use of this type during  the transitional period between winter and spring helped elevate its pellet group density somewhat, but the habitat s t i l l ranked fourth and had a below average density in 1975.  In the winter of 1975-76 with less  snow, use decline only briefly in mid-winter (Table XIII).  The decline  in pellet group density between years was statistically insignificant (Table XIV),  indicating a large increase in relative use.  facing Young Fir type ranked higher in the second year.  Only the west These trends  are shown in Figure 14c.  Mixed Age Fir Type:  This habitat occupies sections on both sides of the Wolf Creek drainage, and thus has both north and south aspects.  Although track  data were largely missing for the north facing slope in 1974-75, the pellet group data indicated that splitting the type was justified (Table IV).  However, the expected difference in tracks the following winter  76.  did not appear, as neither slope was preferred over the other in the f i r s t count, the north facing slope was preferred in the second count, and the south facing slope was more heavily used at the time of the third count.  As a result, the data have been pooled in Table XIII and  Figure 14d.  During both winters, use of this type based on track counts displayed a "peaked" pattern, being proportional at both ends of the winter, but selective in mid-winter (Figure 14d).  The degree of selection was  greater in the snowy winter of 1974-75 as indicated in Figure 14d.  Intense use of this habitat during the 1974-75 winter when snow depths were significantly lower (p^0.05) than in any other habitat led to high pellet group densities on the south facing slope (second only to the Young Fir west aspect) while the north facing slope could be grouped with the Maturing Open (not significantly higher) and the Young Fir north slope (not significantly lower).  During the second winter, there  was an 80% decrease in use as measured by pellet groups on the south facing slope, while on the north slope, use was down 33% (Table  IV).  These changes in density eliminated statistical significance between densities with respect to aspect indicating more uniform use of this habitat in a year when critical snow relief was not needed.  Riparian Mature Spruce:  Use of this habitat was not quantified for evaluation in relation  77.  to the other forested types on the winter range due to its limited distribution and more complex flooding pattern.  However, use was f e l t  to be light to moderate in 1974-75 and moderate to heavy in 1975-76. There also appeared to be two types of use of this habitat.  A small  number of deer spent most of their time here, and others used this type only when crossing from one slope to the other or for cover between feeding periods in the open brush along the creek.  Use of Summer Range Habitats  For the reasons already mentioned, statements about summer range use must be subjective and cannot be based on quantified data.  There is  value however in documenting what was observed during the study.  Use of the three associations on the summer range can be categorized in the following way.  Brush:  is predominantly, for feeding.  use is consistent and the nature of use Shrub:  use is inconsistent and varies  with location of the association and species composition, with most use being for feeding.  Immature Forest:  use is extremely variable being  practically nil in the dense stands, moderate in the more open areas and highest in areas near small meadows, stream courses and the Brush association where time is spent between feeding periods.  In no place  did densities approach those common on the winter range.  Of the two deer with radio-collars that were monitored through the summer, both had seasonal ranges that included the open associations.  78.  Adult male #1B centered bis activities around a series of sloughs, creeks and beaver ponds in a small tributary drainage of Lost Dog Creek. Adult doe #R2 spent most of the summer months just north of Lost Dog Lake in an area composed of Brush association, stream bottom and a slope that was divided about equally between an open Immature Forest association and Shrub association dominated by Ceanothus  velutinus.  individual Movements:  The study of individual movements was based on 33 deer live trapped between 21 January and 18 A p r i l , 1975, and between 21 December, 1975, and 22 February, 1976.  During the f i r s t winter 6 adult males, 19 adult  females, 2 male fawns and 3 female fawns (Table XVI) were captured.  Of  these, 1 adult male and 5 adult females were fitted with radio-transmitter collars, the remainder with material markings.  In the second winter, 3  adult males and 1 adult female were trapped and radio collared, and one female fawn was ear tagged.  (Table XVI).  A total of 84 visual sightings of 21 positively identified deer provided information on individual movements during the study period. Another 17 sightings of unidentified marked deer gave futher details for the population.  Some of these unidentified deer were reported to me by  local people who knew of my work.  Others were deer I saw but was not  able to positively identify due to insufficient sighting time or due to loss of markers.  79.  TABLE XVI LIVE-TRAPPING RESULTS Dates  Trap nights  21 January to 18 A p r i l , 1975 21 December,' 1975 to 22 February, 1976  108 59  Male adults 6(1)** 3***  Female adults  Male fawns  19(7)  2  3  38  35  0  1  6  10  1  Female % fawns Total Succ*  * % Success = total number of captures/total number of trap nights x 100 ** ( ) indicates number of recaptures. ***Note: Two of these were recaptures of deer tagged the previous winter.  80. Rate and quantity of tag loss for all deer is unknown.  However,  one deer tore its collar off within .23 days while crossing a barbed-wire fence, and a few lost one streamer within 39 days.  Adult doe #1A lost  her ear tag between 57 and 60 days after capture, and stepped through her collar at some time between. 72 and 146 days.  At 160 days, the  collar was s t i l l around her neck, over her left shoulder and under the left foreleg and she was seen favoring this leg (B. Jamieson, pers. comm.).  She was not seen again and i t is unknown how long the collar  remained on this deer.  Of the six deer marked in the winter of 1974-75  and positively identified again in 1975-76, two does retained all markers (totalling 2 collars, 2 tags and 4 streamers), the fawn already mentioned had lost 1 of 2 streamers, one doe had lost a streamer by 357 days and its collar by 397 days (retaining 1 tag and 2 streamers) and two bucks each lost their collars and two streamers within 327 and 337 days respectively, retaining 1 tag each and one streamer on one.  This 20%  sample of the marked deer implies that, after about one'year, a minimum of 40 to 60% of the collars were shed, 14% of the tags were lost, and 43% of the streamers were missing.  This high level of loss of markings  suggests that estimates of the proportion of the deer returning to this winter range in 1975-76 based on sightings of deer marked in 1974-75 are probably low.  How much too low is unknown.  The high collar losses are thought to be the result of using the loose-fitting material collars which could catch on the brush or numerous wire fences and tear off.  Subsequent studies should find alternatives.  Ear tag losses were similar to those experienced in cattle using the  81.  same type of tag (H. Campsall, pers. comm.).  The 14% could possible be  cut by using metal tags, but these would not provide identification at a distance like the tags used here.  Streamer loss might be lowered by  shortening the running ends thus reducing the chance of tagling in brush and being ripped out.  The duration of contact with deer fitted with radio-transmitter collars varied from 33 days for 2 does to 427 days for two other does, and averaged 135 days.  The number of radio-locations varied from five  to 19 and averaged 10.  It was found that i t generally took almost an  hour to locate a deer with precision.  The varied topography deflected  the signals and it was often necessary to obtain more than two bearings before a true location could be determined.  Figure 15 is a map of the 84 sightings of marked deer made between February 1975, and April 1976, divided into winter, spring, and summerf a l l periods to show the seasonal distribution of the population.  Of  the 33 deer captured, 17 with material collars and 7 with radio collars were relocated and identified a sufficient number of times to provide useful movement data.  The details of these movements are mapped and  described in Appendix I;  they can be summarized as follows.  Throughout the winter of 1974-75 the track counts and general observations indicated a definite shift of the population to the eastern portion of the winter range as snow depth reached unusual levels in the study area.  This shift was exemplified by the movements of 7 deer  82.  FIGURE  deer.  15.  Winter (o), Soring (A), and Summer-Fall (•) observations of marked  83.  mapped in Figures Al to A3.  In late winter, as the snow began settling  and south facing slopes became snow free, the deer began to expand back to the west.  These movements resulted in several relocations in areas  used in early winter (see Fig. A l , A3 and A4), and eventually deer were seen in the fields on the benches to the east of the Kootenay River.  As spring progressed and the westward procession of green-up continued, the deer kept moving west.  The movements of 7 deer illustrated  in Figures A4, A7 and A8 typify the spring redistribution that took place from late april through early June 1975.  Figure 15 also shows  that most of the spring relocations were made on the floodplain and valley floor.  Many of these were associated with intensive use of  cultivated fields in late April and May.  Following mid-June, relatively few deer were seen along the river or at elevations below 3000 feet west of the river.  The majority of the  summer-fall observations were made in the Lost Dog Canyon-McNair Lakes area.  The movements of 5 marked deer illustrated in Figures A5, A7 and  A8 coincided with this general pattern and i t was felt that the majority of the population did move to mid-elevation levels on the west side of the Trench.  However, two extreme movements were noted.  One adult male  (believed to be #4B) was seen on 24 July, 1975 by a forestry-wildlife graduate student, Brian Churchill, near the confluence of Mary Anne Creek and the Kootenay River. site.  That is over 65 km northeast of the trap  Another buck (believed to be #6B) was reported at the confluence  of Find!ay and Lavington Creeks, 40 km northwest of the trap site in the  84.  f i r s t week of September, 1975.  These animals were recaptured in January  and February, 1976 in the Wolf Creek drainage indicating that these extremes are probably not dispersals, but seasonal movements.  Although the data are limited, the summer home ranges of the deer appear to be relatively small.  No two summer relocations for one deer  were more than 3 km apart and the home ranges for two radio tagged deer were less than 200 ha in total area.  Detailed information regarding timing of f a l l movements was not obtained from marked animals.  No signals were received from any of the  six radio collared deer from August to December.  Other information on f a l l movements is very limited.  Tracks in the  snow indicated that deer were s t i l l west of the river using the area from 3800 to 4300 feet on 27 November, 1975, although heavier use was occurring at the 3300 to 3800 foot level.  However, by 10 December snow  depths had surpassed 20 cm at 3500 feet and no deer tracks could be found above that level.  At that time deer were beginning to be seen on  the floodplain and on Premier and Wasa Ridges.  Prebaiting for the second trapping period was initiated on Premier Ridge on 12 December, 1975, and limited use of the hay occurred prior to 21 December.  However, use increased rapidly after that date and the  f i r s t capture of the winter was made on 27 December.  Of the four adult  deer captured in the 1975-76 winter, two were recaptures from the previous  85.  winter (see Fig. All and A12 and accompanying text).  This indicated  that some white-tailed deer in this area return to the same winter home range in consecutive years.  In contrast, the lower density of deer on the ridges and higher density of the floodplain in 1975-76 compared to the previous winter indicates that not a l l deer return to exactly the same area each winter. This was found to be the case for doe #4A (see Fig. A8).  No eastward shift of the population was noted in the low snowfall winter of 1975-76.  The home ranges of four radio-tracked adult deer  (Fig. A9 to A12) showed continued use of western portions of the winter range.  These home ranges averaged 279 ha and had a mean value of 4.9 km  for the maximum distance between extreme relocations.  Spring movements in 1976 followed the same pattern as in 1975 (see Fig. 15, All and A12).  However, the milder winter weather resulted in  an earlier greenup, and the redistribution occurred approximately two weeks sooner in 1976.  DISCUSSION  The process of forest succession and its detrimental effects on ungulate winter ranges throughout the East Kootenays has been well documented.  The scale of this change was shown by Demarchi (1971) who  stated that there had been " . . . a reduction of 58% of the potential big  86.  game winter ranges in 40 years."  This statement applied to the loss of  open, serai shrub communities to young forests on all the winter range areas in the southern Rocky Mountain Trench.  Considering the d i s t r i -  bution of white-tailed deer and habitat types on Premier and Wasa Ridges (Fig. 5 and Table IA), has been even greater.  the proportional loss of this productive habitat Less than 10% of the area remains in serai shrub  habitat and over 50% of the area is now covered by unproductive habitat types.  The limiting nature of this tremendous reduction in unforested habitat becomes clear when changes in forage production are analyzed. Kemper (1971) found that the productivity of the herbaceous layer declines to 25% of the level in the open, and the shrubs lose over 70% of their productivity by the time a forest on Premier Ridge reaches 50 years of age or 50% canopy coverage.  My results concur with these.  Figure 16 illustrates the change in shrub productivity with increasing canopy coverage.  The same general trend has been reported by numerous  other authors (Cowan et a l . 1950; Kelbenow, 1965; Halls and Alcaniz, 1968; Blair and Enghardt, 1976; see Kemper, 1971, for others.)  Kemper has also discussed the influence of changes in species composition within the herbaceous and shrub layers and indicated that many preferred species decrease and non-preferred species increase (i.e. soapollalie) under forest canopies.  In addition to this, my data  indicate that increasing canopy closure influences the intensity of utilization, possibly reflecting lower palatability of the preferred  87.  25  r  20 CM  E  o 1-  o  15  Q O  or  < 10  10  SS  C/>  YF  A A  RMS  MO  IFO" MAF #  0-10  II - 4 0  J_  > 60  41-60  CANOPY C L O S U R E  N IFD  (%)  FIGURE 16. Relationship between forage production by shrubs and percent canopy closure on the winter range.  88.  species of shrubs (Figure 17).  Klebenow (1965) found a similar trend on  a Montana mule deer winter range.  These relationships are not surprising  in view of the facts that palatability is at least partially a function of the nutritive quality of a plant (Longhurst et a l . 1968) and nutritional quality of forage plants is inversely related to canopy coverage (Cowan et a l . 1950; Lay, 1956).  Another major element in the successional influence on the nutrition of the deer is the composition of the diet.  Demarchi and Demarchi  (1967) have shown that, in general, the winter diet of deer on the Wolf Creek winter range consists of 60% browse, 25% forbs, 5% grass and sedges and 10% miscellaneous other items.  These percentages are similar  to those reported by Martinka (1968) who found that white-tails in Montana have winter diets composed of 74% browse, 24% forbs and 2% grasses.  However there are major differences in the species composition  of the browse segments in the two diets.  The major species for the  populaton that Martinka studied are aspen (32%) and saskatoon (22%); for the Wolf Creek deer they are douglas f i r (33%), bearberry (19%) and other deciduous shrubs dominated by bitterbrush  (8%).  Martinka's population was assumed to be expanding slowly, and showed no signs of intra-.or inter-specific competition.  The high use  of aspen (mainly dead leaves) is probably significant to their well being since this browse averages about 10% crude protein in winter (Tew, 1970).  The other important species, saskatoon, has been found to be  lower in protein in late f a l l at about 6.2% (Diets'et a l . 1958'in_ Ullrey  RB YF  X  RMS  y = 60.55 - .401x ^  r = -.829  s  MAF  \ \  ,  F  0  '  \  \  IFD*  MO  SS  I O-IO  I 11-40  I > 60  41-60  CANOPY CLOSURE  {%)  FIGURE 17. Relationship between average utilization of shrub species (Table V) and percent canopy closure on the winter range. *These values may be depressed due to deep snow in the winter prior to sampling, and they were deleted in the calculation of the line equation.  90.  et a l . 1967), but protein content on a dry matter basis would increase slightly through the winter (1)11 rey et a l . 1967).  Thus their diet  appears to be satsifactory in both quantity and quality.  I do not  believe this is the case in the Wolf Creek area.  Various authors have reported that douglas f i r has crude protein levels between 6 and 9% in the current annual growth and 4 to 6% in old needles (Longhurst et a l . 1968; Oh et a l . 1970).  However, douglas f i r  also contains various essential oils that can inhibit rumen function (Oh et a l . 1967 and 1970) and the concentration of these oils increases with age of the twigs and needles.  The ability of a deer to obtain  sufficient value from ingested f i r browse is a function of the balance between nutrient and essential oil content (Longhurst et a l . 1969). Among black-tailed deer {0. h. colvmbianus) is  in California, this balance  such that use of f i r is limited to new growth and only occurs for a  short time in spring when protein is at its peak, and the essential oils are lowest (Oh et a l . 1970).  Deer use of f i r in the northwestern states, however, is almost entirely limited to winter during periods when snowfall makes other forage less available (Crouch, 1966 and Miller, 1968).  The same pattern  of use generally holds true in the East Kootenays (Demarchi and Demarchi, 1967), but succession and competition have led to a situation in which other forages are quantitatively limited regardless of snow depth. As a result, the deer are forced to rely heavily on this browse. Although there are substantial quantities of f i r browse available in the  91.  Wolf Creek drainage (Table VI),  only a very small percentage of this is  current annual growth except in the Young Fir type.  The browsing survey  showed that on certain trees the deer are using much more than the current annual growth, particularly in the areas where they are concentrated during severe winters.  As a result, they must be consuming  browse with high levels of inhibiting oils and low nutrient quality. Unless they can obtain sufficient quantities of other forages to compensate for this, they are bound to suffer from poor nutrition through the winter months.  Interpretation of the distribution and movements of the whitetailed deer provides insight into their attempts to compensate for the reduced forage availability and quanity on this winter range.  Use of the unforested habitats, which permits exploitation of the maximum observed forage production, was found to be selective much of the time.  However, their ability to use these areas can be strongly  influenced by nival conditions.  Within the Riparian Brush type in the  winter of 1974-75, deep, soft snow made i t very d i f f i c u l t for the deer to move about and use was only proportional to availability in early and mid-winter.  It late winter, with warmer daytime temperatures and noctur-  nal freezing, the density of the snow increased enough to prevent a deer's sinking in more than 10 to 15 cm and the deer responded by selecti this type at that time.  Jones (1975) observed similar use of open areas  by black-tailed deer on Vancouver Island when snow density in clearcuts reached a level that would support a deer.  92.  Snow depth cannot be the only factor involved in the temporal pattern of use of this type, however.  In the winter of 1975-76 snow  depth never surpassed the 25 cm level Hepburn (1959) called "restrictive" for whitetails.  In spite of this lack of snow, early winter use  was even lower than before as deer were found to be avoiding this type at that time.  As before, use increased as the season passed (Fig. 14a).  It seems likely that use in early and mid-winter 1975-76 may have been low due to lower densities on the winter range and the deers' abilities to obtain sufficient food in other types.  As forage in the  other types was reduced through browsing during early and mid-winter; as nonbrowse foods became less available due to increasing snow and less nutritious due to aging; and as body fat reserves were depleted, i t became increasingly necessary to concentrate feeding activity in the Riparian Brush type.  The relative importance of the browse in this type  due to forage reduction in less productive areas would increase even more through a winter with a heavy snowpack since the deer rapidly deplete food when concentrated in the sheltered areas.  This may, in  fact, be indicated by the higher use observed in the deep snow winter of 1974-75 (Fig. 14a).  Use of the Serai Shrub type was also found to vary inversely with snow depth.  The selection of this type at times of low snow is thought  to be a function of forage preference for the components of the herbaceous layer and especially for the productive and well-used shrubs found here. The decrease in use in mid and late Winter, 1974-75, can be simply  93.  related to increasing snow depth, but the drop in late winter, 1975-76, requires more explanation.  The apparent drop in use in late winter is thought to be a result of a biased sample rather than a true decrease in use.  The unseasonably  warm weather in February led to incomplete snow cover on the winter range at this time.  This weather and lack of snow brought about a shift  in the deers' distribution.  Part of the population moved to higher  levels on the southwest slopes of both ridges and continued to use the Serai Shrub type up to nearly 4000 feet elevation.  However, their  tracks were not counted on the survey route that Only covered portions of this type at lower levels.  The track count technique could not be  applied at the higher elevation because of the presence of mule deer with the white-tails there.  The high selection of this type based on track counts indicates that i t is a favored feeding area for the deer throughout a winter of low to moderate snow depth.  Coupled with the relatively high producti-  vity of this type and its potential for covering large acreages, this information further emphasizes the importance of this habitat to the wellbeing of wild ungulate populations in the southern Rocky Mountain Trench.  The deer were also found to make selective use of the Riparian  Brush type during late winter critical periods in both years.  Such use  implies that they do respond to the availability of concentrated quantities of higher quality forage in the open habitat types.  94.  Of a l l the forested types, the Young Fir habitat is especially valuable.  It is favored because i t has the most productive shrub layer  of the conifer types, the super abundant browse on the healthy young f i r trees is probably more nutritious than elsewhere because a greater proportion of i t is represented by current annual growth, and the trees provide excellent cover except under severe snow conditions.  The large  increase in use of the west slope in both late winter periods was also related to the deers' response to the warm afternoon sun.  The lack of a  closed canopy provided numerous sunny bedding sites with sufficient hedge effect to break any wind and isolate the deer from disturbance. The increase in use began later in 1974-75 (Fig. 14b) due to the deep, soft snow in February which did not settle and become dense enough to support deer until March.  The reduced snowpack in 1975-76 permitted  exploitation of this type throughout the winter.  It can also be seen from Figure 14b. and the pellet group data that late winter use of this type was substantially higher in 1974-75 than in the following year. related to the deers  As with the Riparian Brush type, this may have been 1  greater need to seek forage in types that were  less used during the times of confinement by snow conditions.  It was  probably also the result of the concentration of deer in the eastern portion of the winter range during that f i r s t winter.  Excessive snow depths in the more open types on the western half of the area (Immature Forested Open, Maturing Open, Serai Shrub, Cultivated, Aspen, and Riparian Brush) forced the deer onto the lee side of the  95.  ridges and into the relief of the Mixed Age Fir habitat.  The movements  of several of the marked deer which were seen at this time from 0.5 to over 4 km to the east of their trap sites exemplified this redistribution of the deer (Figs. Al-A3).  During mid-February, 1975, when snow depth in the open were exceeding 40 to 50 cm, much of the ground in the Mixed Age Fir type was covered with less than 30 cm.  Snow depth here was significantly less than in  other types (p^O.05) and, in fact, within this type the areas used by the deer as indicated by the presence of tracks and beds had a mean snow depth of 24 cm which was significantly less than the 37 cm mean depth in unused areas.  Thus the deer were not only heavily concentrated  in this type, but within this habitat they selected the low snow areas and especially for bedding, they selected the warmer south aspect.  This  shelter seeking movement was the cause of the temporal pattern of use of the Mixed Fir type displayed in Figure 14d.  During the winter of 1975-76, conditions were much milder and snow relief was unnecessary.  No eastward shift occurred; in fact, many deer,  like doe #4A (Fig. A8) remained west of the Kootenay River for the second winter.  Although the basic shape of the temporal use curse of  this type (Fig. 14d) was the same, the degree of selection was much lower since deer were s t i l l able to use other, more productive types such as the Young Fir (west aspect) and the Maturing Open.  The pellet  group counts also indicated higher levels of use for this habitat in the severe winter with preference for the south aspect, and more uniform  96.  distribution of fewer deer in this type in the mild year.  These results  lend further support to the interpretation of the value of this type as a snow relief habitat.  It is important to note, however, that at no time was observed use less than expected (which would indicate avoidance).  Thus use is dedi-  cated to this habitat regardless of the need for snow relief.  This is  related to its provision of good cover, a moderately productive understory, and the presence of quantities of f i r browse that are intermediate in quality to those in the Young Fir and Immature Forested Dense types.  Canopy coverage in the Maturing Open type is relatively low, and as with the other open types, forage productivity and availability are improved relative to the densely forested types.  The deer respond by  selecting this type whenever snow depth permits.  However, the open  nature of the canopy provides for relatively l i t t l e snow being intercepted by the trees.  In 1974-75, this led to mid-February snow depths  that averaged over 40 cm.  As a result, use, which had been high in the  early winter f e l l off sharply (Fig. 14c).  The failure of use to increase  significantly in early March when snow density increased allowing exploitation of other open types (i.e. related to two factors.  Riparian Brush and Young Fir),  is  These are the mid-winter shift of the population  to the east away from this type, and past grazing pressure which causes the growth of the understory to be vertically stunted.  Thus very l i t t l e  browse including f i r which is relatively scarce in the Maturing Open type was available above the dense snow.  97.  The open canopy and the generally south facing aspect of the Maturing Open type results in an extremely early melt off of snow in late winter.  Almost the entire surface of the ground here was snow free  in 1975 long before patches began to open up in any other type.  The  deer were seen to respond quickly to snow melt and the initiation of forb growth which commenced in late March, 1975.  New growth attracted deer to the Maturing Open type, initiating expansion of the deer back toward the summer range.  Although no track  counts could be made at this time, direct observation indicated very heavy use of this type in A p r i l , 1975. marked deer (Figs. A1-A3).  This was demonstrated by several  Hudson et a l . (1974) reported the same  population shift to southwest slopes in the late winter-spring period.  With the change in snow conditions in 1975-76, several things were altered in the pattern of use in the Maturing Open habitat.  First, as  Table XIII and Figure 14c indicate, use remained high throughout the winter since snow depth was never excessive.  Second, the pellet group  density here in 1976 ranked second and was only 4% lower than in 1975. Since the overall average dropped by 50% and the average drop by types was 43%, this slight decrease in the Maturing Open indicates a substantial increase in relative use in the mild winter.  Finally, a large  influx of deer to this type in late winter-spring was not observed in the second year. 76.  This is also a result of the reduced snowfall in 1975-  Because there was so l i t t l e snow everywhere that year, the Maturing  Open did not represent the only habitat with bare ground and a source of  98.  new growth of forbs.  In fact, the period that had been late winter in  the f i r s t year (25 March to 10 A p r i l , 1975) was spring in 1976, and use of fields and grasslands was beginning.  The Immature Forested habitats, both Open and Dense represent areas that are generally avoided by white-tailed deer.  In the Immature  Forested Open the explanation for this low use in 1974-75 is simplified by snow depth.  The relatively open pine canopy intercepted but l i t t l e  snow, and the deer could not exploit this type without excessive energy costs in traveling about.  The deers' requirement for relief from deep  snow in dense stands in that cold, snowy weather led them to seek other forested habitats.  The repetition of this pattern of low use in a relatively mild winter (1975-76) indicated, however, that snow depth was not alone in discouraging use of this type.  The understory is poorly developed and  locally affected by heavy grazing so available forage is low regardless of snow pack and the limited amount of f i r in the overstory provides very l i t t l e of this important browse.  Hout (1974) found that white-  tails in Quebec not only selected types that provided cover, but bedded near the larger trees in those types.  I generally noted the same here  and Dr. R. Ream (pers. comm.) found this to hold true for elk in Montana. Moen (1968 a. and b.) has given energetic evidence to show why this selection occurs.  Given this selection and its energetic basis, the  relatively open nature of this type composed almost entirely of small trees would give i t a very low shelter preference rating.  Lacking both  99.  food and shelter, this habitat is of l i t t l e value to the deer, and they respond accordingly.  Interpretation of the use and value of the Immature Forested Dense type is not as simple.  There appears to be a discrepancy in the data  since the track counts indicate avoidance of this type through much of the winter, but the pellet group densities indicate that this type receives only slightly less than average use.  The explanation for this discrepancy lies in the fact that the structure of the habitat is such that its use intensified the two biases in the sampling techniques.  It has already been shown that this type  virtually lacks an understory and that although the total quantity of f i r browse is high, the growth form of the individual trees is such that they produce very l i t t l e , and probably very low quality forage. fore, the deer spend l i t t l e time feeding here.  There-  On the other hand, the  high degree of canopy closure results in the interception of much of the snowfall and the density of saplings makes i t ideal cover.  It is  used heavily for bedding and relief from the occasional severe storms in winter.  Since track counts are biased toward feeding sites while pellet  groups are highest where there is a concentration of bedding, the two use estimates are drawn in opposite directions in a habitat like this that offers shelter, but l i t t l e food.  Since this habitat is used primarily for relief from climatic stresses, the influence of the proportion of the winter range covered by  100,  this type must be evaluated in relation to the general winter conditions and the distribution and productivity of surrounding habitats that f u l f i l l other l i f e support requisites.  There are good indications that  too much of the Wolf Creek winter range is allocated to the Immature Forested Dense type for the maximum benefit of the whitetailed deer in view of typical winter weather and the potential productivity of the unforested types.  The remaining two forested types, Aspen and Riparian Mature Spruce, were relatively minor and apparently unimportant components of this winter range.  In the former habitat, forage production is relatively  high and more use might be expected in this type.  Martinka (1968) and  Allen (1968) both found white-tails to use a similar habitat extensively in Montana.  Along Wolf Creek, however, the patches of Aspen habitat are  very small and offer no cover whatsoever.  They are also located near a  road so diurnal use is restricted as the deer here do not generally tolerate vehicular and human disturbance.  Finally, the lack of a conifer  overstory means that snow builds up here almost as quickly as in the open types and in a winter like 1974-75, use would be limited by nival conditions.  The Riparian Mature Spruce type was also found to be relatively productive, and the spruce canopy intercepted much of the snowfall. However, the level of use was found to be light to moderate in 1974-75 and moderate to high in 1975-76.  This pattern of use could be related  101.  to the habitat's being uncomfortable in cold winters due to its shaded and humid nature.  Both the Aspen and Spruce habitats are more significant components of the riparian deer ranges found north of Skookumchuck and south of Fort Steel.  The present study may not provide a complete analysis of  the value of these types and future research on white- tailed deer range in the Rocky Mountain Trench should include a more thorough examination of these areas.  The grasslands and fields in the Wolf Creek drainage are virtually untouched in the winter months.  This is probably related to their lack  of cover and the influence of heavy grazing or the mowing of a l f a l f a . On the natural grasslands, cattle grazing has altered the community by eliminating the bunch grasses that can withstand f a l l rains and snow and that remain nutritious in winter (Kemper, 1971).  The increased forb and  annual grass cover is decadent, prostrate and of l i t t l e value following the typically hot, dry summer weather in this area.  On the cultivated fields, two cuttings of hay are the rule and many ranchers then fall-graze the stubble.  This practice severely reduces  the quantity, quality and height of the available forage regardless of snow cover.  That deer would use these fields under different treatments  was demonstrated by several deer feeding for a few weeks from late November, 1974, to January, 1975, in a section of one field that was not cut a second time nor grazed in the f a l l .  The deer were even "cratering"  102.  in 24 cm of snow to feed on the a l f a l f a .  (Cratering observed in natural  habitats occurred in snow no deeper than 17 cm with Balsamovhiza and bearberry as target species.)  saggittata  Similar behavior in other single cut  fields was reported to me by G. Seaton, a local conservation officer.  The very conditions that eliminate usefulness of these habitats during the winter enhance their value in spring.  The almost complete  removal of above ground material creates a situation in which new growth begins early and is immediately available to spring-grazing deer.  The  animals respond quickly to this high quality forage and spend many hours in the spring feeding in these areas.  The advanced availability of  these types in spring, however, should not be viewed as justification for continuation of present treatment of these areas.  Continued complete  removal of annual production on cultivated fields may be of minor importance in terms of overall winter deer forage availability and the negative soil effects can be offset to some degree by heavy f e r t i l i z i n g  (which is  not generally done), but continued abuse of natural grasslands will lead to severe depletion of soil nutrients and will eventually reduce theirvalue as spring range.  The 20% bare ground in the grasslands (Table  may indicate that some nutrient depletion has already occurred.  I)  (It  should be noted here that season- long grazing by domestic livestock which lead to this deterioration was replaced in 1976 with a system of deferred and rest rotations oh the area north of Wolf Creek.  R. Demarchi,  pers. comm.).  The i n i t i a l use of these areas in spring is relatively low and  103.  probably represents pioneering use by those individuals whose winter home ranges border on the f i e l d . be bare of snow.  At this time, only a few patches may  As more snow melts and the alfalfa initiates growth,  other deer are attracted to the field and shift their activity to the field and the cover at its edges.  This shift was demonstrated by marked  deer #2C (Fig. A4), and several authors have reported similar behavior in white-tailed deer in eastern North America (Marchinton, 1968; Downing et a l . , 1969; and Byford,. 1969).  As forage green-up occurs in other  areas, deer become less dependent upon alfalfa and shift their away from the fields.  activity  A similar rise and f a l l pattern was reported by  Bartmann (1974) for mule deer on alfalfa fields in Colorado.  To some extent, the d r i f t of the population toward the summer range was responsible for the observed series of peaks of use in the fields illustrated in Figure 9.  These peaks occur in an east to west fashion  in relation to the rate of snowmelt (earliest in the east) and distribution of the population as indicated by observations of marked deer.  Thus  although use of the easternmost field dropped off after 9 A p r i l , some . (and probably most) of these deer had just moved to the west and were using other fields.  It can be seen then that these fields, and to a  lesser extent the grasslands, are a highly used type throughout April and May.  Hall (1973) has shown that the availability of high quality forage during this period is critical to late term fetal development and neonatal survival in white-tailed deer.  Availability, however, is not  104.  just a function of what is actually growing in the field as determined by the arrival of spring weather.  The timing of initiation and amount  of use of these areas is controlled by several factors.  Detailed studies of deer use of alfalfa fields have indicated that weather exerts an important influence on deer use of f i e l d s .  Boyd  (1960) and Progulske and Duerre (1964) found that cold, rainy weather reduced the numbers of deer using study fields.  This was also noted in  my observations, but i t was not sufficiently quantified for statistical evaluation.  An especially cold or rainy spring could be detrimental by  reducing both rate of plant growth and exploitation of the new growth by the deer.  Another important factor is human disturbance, and the influence of this on the daily pattern of use.  I have shown that human dis turbance  can influence the initiation of daily use of fields (Fig. 10).  The data  are too incomplete to carry the comparisoon on through the night, but the trend in all fields was for a consistent, gradual decline resulting in the fields being empty at sunup. (1960) and Bartmann (1974).  This pattern was also found by Boyd  It would appear then that traffic and  associated human activity have the effect of reducing deer use of the fields.  Although the "late starters" might be able to compensate by  feeding in the fields for longer periods through the night, I have no data to indicate that this occurs..  Also, their ability to do this would  be limited somewhat by the fact that the peak of use on Dr. Green's field occurs two weeks later than on Bradford's and the night is almost an hour shorter at that time.  105.  Thus the potential value of the open, non-shrubby habitats is a function of their location, their treatment, the weather and the proximity of human disturbance.  Verme (1968) has stated that winter severity is a function of two things:  the physical restraints of the snowpack and.cold weather.  The  different amounts of snowfall on the study area in the two winters have been shown to have had different influences on the winter distribution of the deer and their ability to exploit the forage resources on the winter range.  The identicle phenomenon was documented in New Brunswick  by Drolet (1976).  In general, the deep snow of 1974-75 confined the  deer at high densities to the nonproductive, closed canopy forested types, whereas the minimal snows of 1975-76 did not physically use of any area.  restrict  As a result, the deer not only had to expend relatively  more energy to move about in the f i r s t winter, but also suffered from intraspecific competition and the inability to utilize the most productive forage areas.  From this standpoint then, the f i r s t winter was more  severe.  The intensity of cold weather is a function of several climatic variables.  Verme (1968) found that the most import ant of these is  temperature and i t is obvious that the colder the weater, the greater the energetic cost of maintaining homeothermy.  Once again, the winter  of 1974-75 rates more severe since average temperatures then were lower than in 1975-76 (Fig. 8).  106.  Another environmental factor that is involved in the thermal regime of a deer is direct solar radiation.  Although Verme (1968) found poor  correlation between this and air c h i l l , his "chillometers" were shaded from direct sunlight for the majority of the time, and their shiny metal surfaces reflected much of any light that struck them.  On the contrary,  a deer has the option of moving to place itself in direct sunlight and its surface is not as reflective as metal.  On many occasions, deer were  found bedding in sunny areas, and the previously discussed differences in track counts and pellet group densities indicate obvious selection for aspects that receive direct solar radiation.  Loveless (1964) has  discussed similar behavior in mule deer in Colorado.  It has been shown that the winter of 1975-76 was sunnier from midJanuary to late March.  This means that the deer had the option of  exposing themselves to more solar radiation in the second winter. Whether or not they exercised this option—or indeed i f they needed to exercise it—during this warmer winter is unknown.  The weather data I gathered indicated that the winter of 1974-75 was much more severe in all aspects than the following year.  I believe  that this severity is an important agent in the different age ratios found in the spring.  There is no indication that anything other than  greater forage availability due to decreased snow and reduced energy costs due to milder weather influenced the population in such  a way as  to bring about a 50% increase in the proportion of yearlings in the population in the second spring.  107.  The fact that white-tails are found in many areas where they experience longer, more severe winters does not necessarily imply that weather is not important in this population, or that, the differences between years is of no significance.  As Moen (1968 a.) has emphasized,  the dietary level is the regulating element in the deers' ability to withstand adverse climatic conditions.  Secondary succession has been  shown to have reduced the quantity and quality of the forage available to this deer population.  It seems logical that restrictive snow condi-  tions—such as those observed, in 1974-75--may be sufficient to reduce nutritional levels and raise energy costs to the point where thermal stress can exert a significant effect on survival.  Forest succession has also affected the summer range of this deer population.  The total quantity and average quality of the forage is  undoubtedly less than what was. found during the peak deer population years.  However, nutritious forage that is lightly used relative to what  is available is s t i l l found in small openings in.the Immature Forest .association and throughout the Brush and Shrub associations.  Deer densities are so low on the summer range that each individual has the opportunity of selecting highly nutritious forage in quantities unlimited by competition or environmental factors such as snow depth. My analysis of summer range use indicates that individuals in this population do take advantage of this opportunity by distributing themselves in relation to these open areas. North America,  summer distribution of white-tailed deer has also been found to be  108.  primarily related to availability of preferred foods (Kohn and Mooty, 1971).  Some authors have discussed the limitations of summer range for northern deer populations (i.e. McCaffery and Creed, 1974) and relate its importance to the need for deer to obtain body reserves for use in the winter period when available food is limited by climatic factors. Summer range can become a limiting factor when deer are no longer able to build up these required reserves.  All the deer I had the opportunity to examine in the f a l l of 1975, from both hunter k i l l s and accidental road k i l l s , were in excellent condition with large deposits of subcutaneous and mesenteric fat.  These  deer are apparently able to obtain sufficient excess energy and nutrients in summer to permit storage of these resources for use in the coming winter.  It is not known to what degree the present level of energy and  nutrient storage approaches the maximum physiologically possible for white-tailed deer.  The greatest value of information gathered from individually marked deer is that i t verifies the movements associated with the observed shifts in distribution of the deer resulting from changes in relative levels of use of the habitat types in response to environmental factors.  Unfortunately, the data are insufficient and the sample size  too small to allow critical analysis of the influence of specific factors such as snow depth or succession on individual home ranges.  109.  However, general observations indicated that home range size and distances moved daily varied inversely with snow depth.  Such a relation-  ship should be expected and.Drolet (1976) and several others authors have clearly demonstrated that this occurs in other white-tail populations.  The average distance from the trap site to summer relocation for nine deer sighted between the end of June and the end of November, 1975, was 24.0 km.  Because those relocations made in October and November may  have been of deer already moving back to the winter range from their point of maximum summer dispersal, 24 km should be considered a minimum estimate.  Even so, this value is well above all of the average dis-  persals for white-tailed deer found in numerous studies listed by Verme (1973).  The highest average he found in the literature was 15.6 km  reported by Carl sen and Farmes (1957) for the prairie-deciduous biome.  There appears to be an inverse relationship between habitat diversity and average distance from winter to summer range (Verme, 1973).  It may  be that forest succession on the summer range of the Wolf Creek deer has progressed to the point where the patches of preferred forage are becoming more widely dispersed, and as a result, the deer must move greater distances from winter to summer range.  Although the sample size is extremely small (4) i t is interesting to note that 50% of the adult deer trapped in 1975-76 were recaptures from the previous year (Table XIV).  These were the two males believed  no. to have traveled a minimum of 40 and 65 km and were both recaptured within 100 m of their 1974-75 trap site.  At least five other marked  deer returned to the Wolf Creek area and there is no way of knowing how many others came back but had either lost their tages or were not seen. Winter range fidelity appears to be as high in the East Kootenay region as in any other area even though the deer here may range farther between seasons.  CONCLUSION  Forty years of forest succession on the terraces of Premier and Wasa Ridges aided by at least 25 years of intensive fire control has resulted in major changes in the winter range used by the white-tailed deer in the Wolf Creek drainage.  Serai Shrub habitat, which once  covered the majority of this area, is now limited to less than 10%. Various forested habitats in which forage production is less than half that in the Serai Shrub type now cover over 80%. of the winter range.  The quantitive reduction of forage has been intensified by a  generally, negative change in forage quality.  As a result of these changes in the shrub and herb vegetation, the deer have come to rely heavily on douglas f i r as a winter food.  Infor-  mation in the literature indicates that the timing and intensity of these animals' browsing on douglas f i r is such that they are obtaining very l i t t l e nutritional value from this food, and must have alternate sources of energy and nutrients to survive the winter.  This becomes  m. more important each year as the trees mature and the ratio of current annual growth to older needles gets smaller.  In a winter when snow depth does not reach a confining level, distribution of the deer appears to be controlled by the availability of forage in the understory as the deer selectively use the more productive, open-canopied or unforested habitats. animals is high.  In such years., survival of young  During winters when snow depths in the open exceed 30  to 40 cm these deer change their distributional patterns by selecting the habitats which afford the greatest amount of relief from the physical restraints of the snow pack.  These types have closed forest canopies,  however, and available forage is seriously reduced.  As a result, over-  winter survival is lowered,, especially for juveniles.  In late winter and spring the population undergoes a redistribution as individuals concentrate on.those areas where forage greenup has begun.  I n i t i a l l y , these are the southfacing, open slopes and later the  natural grasslands and cultivated fields.  Use of these fields is  influenced by the level of human disturbance and their position on the gradient from winter to summer range as well as climatic factors.  Movement of the deer from winter to summer range begins in late April and continues into May.  The majority of the deer from Wolf Creek  spend the summer months on the west side of the Kootenay River on the benchlands that extend,to the Puree!! Mountains.  Their distribution  here is primarily controlled by the availability, of.preferred forage species.  112.  Forest  succession has also influenced the habitat.on the summer  range and resulted in wide dispersal of the remaining areas of nutritious forage.  The deer seem to be responding to this by moving a greater  average distance between seasonal home ranges than is true of other populations of white-tailed deer.  The deer are s t i l l able to obtain  sufficient excess energy and nutrients to permit storage for the f o l lowing winter, so summer range is not judged to be a limiting factor.  Little information was obtained on f a l l movements back to the winter range, but the deer appeared to move only as they were forced to by snow depth.  The lower densities observed on the winter range in the  second year were the result of wider dispersal of the deer indicating that they remain off the winter range as long as possible.  Despite this  change in the two winters, loyalty to the winter home range was high and several deer were found to use the same area in both years.  113  LITERATURE CITED  Allen, E.O. 1968. Range use, foods, condition and productivity of whitetailed deer in Montana. Jour. Wild!. Manage. 32(1): 130-141. Barichello, N. 1975. Management proposals for the 3-Sons Property. Career '75 Project. B.C. Fish & Wildl. Branch. 92pp. mimeo. Bartmann, R.M. 1974. Guidelines for estimating deer numbers in connection with claims of damage to growing crops. Outdoor Facts. Colo. D.N.R. Game Info. Leaflet #97. 3pp. Batcheler, C L . 1975. Development of a distance method for deer census from pellet groups. Jour. Wild!. Manage. 39(4):641-652. Blair, R.M. & H.G. Enghardt. 1976. Deer forage and overstory dynamics in a loblolly pine plantation. Jour. Range Manage. 29(2):104-108. Boyd, R.J. 1960. An evaluation of spring grazing on alfalfa by deer in western Colorado. Proc. West. Assn. Game & Fish Comm. 40:130-147. Byford, J.L. 1969. Movement responses of white-tailed deer to changing food supplies. Proc. Ann. Conf. S.E. Assn. G.&F. Comm. 23:63-78. Carl sen, J.C. and R.E. Farmes. 1957. Movements of white-tailed deer tagged in Minnesota. Jour. Wildl. Manage. 21(4):397-401. Churchill, B.P. 1974. Christmas tree permits and their relation to ungulate forage in the Sprinbrook-Canal Flat region of southeastern B.C. Career '74 Project. B.C. Fish & Wildl. Branch. 25pp. Mimeo. Clover, M.R. 1956. Single-gate deer trap. Calif. Fish & Game. 42(3):199-201. Cook, D.B. & W.J. Hamilton. 1942. Winter habits of white-tailed deer in central New York. Jour. Wildl. Manage. 6:287-291. Cowan, I.M., W.S. Hoar & J . Hatter. 1950. The effect of forest succession upon the quantity and upon the nutritive values of woody plants used as food by moose. Can. Jour. Res. 28 Sect. D(5):249-271. Crouch, G.L. 1966. Preferences of black-tailed deer for native forage and douglas f i r seedlings. Jour. Wildl. Manage. 30(3):471-475. Daubermire, R.F. 1959. A canopy coverage method of vegetation analysis. Northwest Sci. 33(l):43-64. Demarchi, D.A. 1971. Ecology of big game winter ranges in the southern Rocky Mountain Trench, East Kootenay Region. Wildl. Mgt. Div. B.C. Fish & Wildl. Branch, Victoria, B.C. 28pp.  114.  Demarchi, R.A. & D.A. Demarchi. 1967. Winter and spring food habits of white-tailed deer in the East Kootenay. paper presented to 4th Ann. Mtng. of B.C. Fish & Wildl. Branch, Kamloops, B.C. A p r i l , 1967. 10pp. Dietz, D.R., R.H. Udall, H.R. Shepherd & L.E. Yeager. 1958. Seasonal progression in chemical content of five key browse species in Colorado. Proc. Soc. Amer. For. 1958:117-122. Downing, R.L., B.S. McGinnes, R.L. Petcher & J.L. Sandt. 1969. Seasonal changes in movements of white-tailed deer. Proc. of White-tailed Deer Symp. March 25-26. Nacogdoches, Texas. Drolet, C A . 1976. Distribution and movements of white-tailed deer in southern New Brunswick in relation to environmental factors. Can. Fid.-Nat. 90(2):123-136. Farr, A. 1975. Management alternatives for the Gordon Earl Ranch. Career '75 Project. B.C. Fish & Wildl. Branch, (untyped) Gates, B.R. 1968. Deer food production in certain serai stages of the coast forest. M.Sc. Thesis. U.B.C. Vancouver, Canada. 105pp. Hall, W.K. 1973. Natality and mortality of white-tailed deer, (Odoooileus vivginiccnus daootensis Goldman and Kellogg) in Camp Wainwright, Alberta. M.Sc. Thesis. U. Calgary. 117pp. Halls, L.K. & R. Alcaniz. 1968. Browse plants yield best in forest openings. Jour. Wildl. Manage. 32(1):185-186. Hammerstrom, F.N., Jr. and J . Blake, 1939. Winter.movements and winter foods of white-tailed deer in central Wise. Jour. Mammal. 20:206-215. Harlow, R.F. .& W.F. Oliver, Jr. 1967. Natural factors affecting deer movement. Quart. Jour. Fla. Aead. Sci. 30(3):221-226. Hebert, D.M. 1973. Altitudinal migration as a factor in the nutrition of bighorn sheep. Ph.D. Thesis. U.B.C. Vancouver, Canada. 357pp. Hepburn, R.L. 1959. Effects of snow cover on mobility and local distribution of deer in Algonquin Park. M.Sc. Thesis. Univ. Toronto, Ont. Canada. 55pp. Hout, J . 1974. Winter habitat.of white-tailed deer at Thirty-one Mile Lake, Quebec. Can. Fid. Nat. 88:293-301. Hudson,.R.J., V.'.C. Brink & W.D. Kitts. 1974. The grazing question in B.C., an investigation of habitat utilization by wildlife and livestock in the southern Rocky Mountain Trench. Final Report Vol.1. Irvin, L.L. 1975. Deer-moose relationships on a burn in northeastern Minnesota. Jour. Wildl. Manage. 39(4):653-662.  115.  Jones, G.W. 1975. Aspects of.the winter ecology of black-tailed deer (Odocoileus hemionus oolumbianus Richardson) on northern Vancouver Island. M.Sc. Thesis U.B.C. Vancouver, Canada. 78pp. Kemper, B.J. 1971. Secondary autogenic succession on Premier Ridge; B.C. M.Sc. Thesis. U.B.C. Vancouver, Canada. King, D.R. 1975. Estimation of yew and cedar availability and utilization. Jour. Wildl. Manage. 39(1):101-107. Klebenow, D.A. 1965. A montane forest winter deer habitat in western Montana. Jour. Wildl. Manage. 29(1):27-33. Klein, D.R. 1970. Food selection by North American deer and their response to overutilization of preferred plant species. In:A. Watson (ed) Animal Populations in Relation to their Food Resources. Oxford, pp.25-46. Kohn, B.E. & J . J . Mooty. 1971. Summer habitat of white-tailed deer in northcentral Minnesota. Jour. Wildl. Manage. 35(3):476-487. Krajina, V.J. 1969. Ecology of forest trees in B.C. In.: V.J. Krajina (ed.) Ecology of western North Amer. 2(1):1-147. Dept. of Bot.. U.B.C., Vancouver. Lay, D.W. 1956. Some nutritional problems of deer in the southern pine type. Proc. Conf. S.E. Assn. Game & Fish Comm. 10:53-58. Longhurst, W.M.,. H.K. Oh, M.B. Jones, & R.E. Kepner. 1968. A basis for the palatability of deer forage plants. Trans. N.Amer. Wildl. Conf. 33:181-192. Longhurst, W.M. J.H.K. Oh, M.B. Jones & R.E. Kepner. 1969. Deer forage palatability, digestibility and management implications. Paper presented at Calif.-Nev. Sec. Wildl. S o c , Berkeley, Calif. Loveless, C M . 1964. Relationships between wintering muld deer and the physical environment. Trans. N. Amer. Wildl. Conf. 29:415-431. Martinka, C J . 1968. Habitat relationships of white-tailed and mule deer in northern Montana. Jour..Wildl. Manage. 32(3):558-565. McCaffery, K.R. and W.A. Creed. 1969. Significance of forest openings to deer in northern Wisconsin. Wise. Dept. Nat. Resources Tech. Bull. 44. 104pp. Miller, F.L. 1968. Observed use of forage and plant communities by black-tailed deer. Jour. Wildl. Manage. 32(1):142-148. Moen, A.N. 1968a. Energy exchange of white-tailed deer, western Minnesota. Ecol. 49(2):676-682.  116.  Moen, A.N. 1968b. Thermal energy exchange of a birch tree and a spruce tree at night. Ecol. 49(1):145-147. Moen, A.N. 1973. Wildlife Ecology. Freeman & Co., San Fran. 458pp. Neff, D.J. 1968. The pellet-group count technique for big game trend, census and distributions review. Jour. Wildl. Manage. 32(3):597-614. Neu,  C.W., C R . Byers & J.M. Peek. 1974. A technique for analysis of utilization-availability data. Jour. Wildl. Manage. 38(3):541-545.  Oh, H.K., T. Sakai, M.B. Jones & W.M. Longhurst. 1967. Effect of various i essential oils isolated from douglas f i r needles upon sheep and deer rumen microbial activity. Jour. Applied Microbiol.... 15(4) :777-784. Oh, J . H . , M.B. Jones, W.M. Longhurst & G.E. Connolly. 1970. Deer browsing and rumen microbial fermentation of douglas f i r as affected by fertilization and growth stage. For. Sci. 16(1):21-27. Passmore, R.C. & R.L. Hepburn. 1955. A method for, appraisal of winter range of deer. Ont. Dept. Lands & For. Res., Rept. #29. 7pp. Poulton, C.E. and E.W. Tisdale. 1961. A quantitative mthod for the description and classification of range vegetation. Jour. Range Manage. 14(1) :13-21. Progulske, D.R. & D.C. Duerre. 1964., Factors influencing spotlighting counts of deer. Jour. Wildl. Manage. 28(1):27-34. Russell, L.J. 1967. Parasites of the white-tailed deer, Odoooileus vivginianus ochrourus of B.C. M.Sc. Thesis. U.B.C. Vancouver, Canada. Severinghaus, C.W. & E.L. Cheatum. 1956. Life and times of the whitetailed deer. Inj W.P. Taylor (ed.) The Deer of North Amer. The Stackpole Co. Harrisburg, PA. pp.57-186. Smith, R.H. 1968. A comparison of several, sizes of circular plots for estimating deer pellet-group density. Jour. Wildl. Manage. 32(3):585-591. Stelfox, J. 1971. Bighorn sheep in the Canadian Rockies: A History 1800-1970. Can. Fid.-Nat. ,85(2):101-122. Swanson, G.A. & A.B. Sargeant. 1972. Observation of nightime feeding behavior of ducks. Jour. Wildl. Manage. 36(3):959-961. Tew, R.K. 1970. Seasonal variation in the nutrient content of aspen foliage. Jour. Wildl. Manage. 34(2):475-478. Ullrey, D.E. W.G. Youatt, H.E. Johnson, L.D. Fay & B.L. Bradley. 1967. Protein requirements of white-tailed deer fawns. Jour. Wildl. Manage. 31(4):679-685.  117.  Verme, L.J. 1968. An index of winter weather severity for northern deer. Jour. Wildl. Manage. 32(3):566-574. Verme, L.J. 1973. Movements of white-tailed deer in upper Michigan. Jour. Wildl. Manage. 37(4):545-552. Wetzel, J . F . , J.R. Wambaugh and J.M. Peek. 1975. Appraisal, of white-tailed deer winter habitats in northeastern Minnesota Jour. Wildl. Manage. 39(1).-59-66.  118.  APPENDIX I INDIVIDUAL MOVEMENT ACCOUNTS  Adult Females #5A, #7A and #2B:  Figure Al illustrates the known movements of three adult females in the winter of 1974-75. 1975.  Doe #7A was captured along Wolf Creek 2 February,  She was next seen just west of Lazy Lake on an aerial survey  flight, 18 February.  She was not seen again until late March when she  was found feeding twice daily on both 27 and 29 March on a southfacing slope just west of the trap site.  (This slope, in Maturing Open habitat,  was heavily used in late winter, 1975.)  Her i n i t i a l  "transitional" or  spring movement was detected the next day when she was located in an alfalfa field in lot 8103 on the afternoon of 30 March 1975.  Doe #5A was captured on 31 January, 1975, and recaptured 200 m to the east on 16 February.  She was next seen on the morning of 6 March  feeding on an open, southeast facing slope with doe #2B which had also been trapped 1.9 km to the west on 19 February.  This association may  have lasted some time, as both deer were seen together again on the evening of 2 April about 0.5 km west of their original trap site. Although no spring movement locations were determined for these does, #2B was seen several times in the late summer-fall period (see Fig. A5).  119.  FIGURE A l . Known movements of adult females 7A (A-*), 2B ( « - - ) , and 5A (••'••) in the winter of 1974-75. 0 indicates deer seen together and dotted arrow indicates direction to summer range of 2B (see text and Fig. A5.).  120.  Adult Female #9B and Female Fawns #2F and #4F:  Figure A2 is a map of the known movements of an adult doe, #9B, and two female fawns, #2F and #4F, both believed to be her offspring.  The  f i r s t one of the three captured was fawn #2F on 16 February 1975. The other fawn and the doe were captured together about 2 km. to the east on 3 March.  This pair was seen again by B. Jamieson another 2 km to the  east feeding with 7 to 10 other deer in the brush along Wolf Creek on 19 March at 3:00 p.m.  The three deer were seen together in the same general  area on 27 March at 12:05 p.m.; #4F was seen here with 6 unmarked does and 4 unmarked fawns on 29 March at 5:30 p.m.; and the three (#9B, #2F, and #4F) were again together feeding in the brush on 3 April at 10:04 a.m. with 15 other deer. April at 5:35 p.m.  The trio was next seen 1 km to the west on 8  Although all three were not seen together again, #9B  and #2F were seen 3 km to the west on the previously mentioned heavily used southfacing slope on 14 April at 5:00 p.m.  The next sighting was  of #9B with #4F in an alfalfa field at 4:15 p.m. on 22 April.  Fawn #2F  was not noticed among the 20 other deer in the field at that time. was repeated on 24 April at the same time.  This  #9B was seen again in that  field at 11:30 a.m. on 7 May with two unmarked deer.  None of these three deer were seen again until the following year on 31 March, 1976, when #9B and #2F were seen in that same field at 7:30 p.m. feeding with 31 other deer.  121.  FIGURE A2. Known movements of adult female 9B (A) and female fawns 2F ( « ) and 4F (o) in winter, 1974-75. — * indicates movement of 2 F ; — i n d i c a t e s movements of 9B and 4F; and • > • indicates movements of all three. j|| i n dicates deer seen together.  122.  Adult Females #9A and #7B:  Figure A3 is a map of the known movements of two adult females, #9A and #7B.  The former was captured on 17 February, 1975.  One month later  on 15 March, she was seen 2.4 km to the northeast in the Mixed Age Fir type.  After that, she was seen six times between 20 March and 9 April  feeding on the heavily used Maturing Open slope.  She was usually seen  in the company of three or four other deer including #1A once and #7A twice.  The single point west of Highway 95 in lot 338 represents a  sighting a year later on 19 A p r i l , 1976.  Doe #7B was i n i t i a l l y captured in the Mixed Age Fir type on 1 March, 1975.  She was seen 12 days later 1.2 km to the southeast moving  toward the southfacing slope.  On 2 A p r i l , she was seen another 1 km to  the southwest fedding on the heavily-used slope at 6:30 p.m. with six unmarked deer.  The other two points for this deer were locations on 19  February and 31 March, 1976, indicating use of the same general area in both winters.  Adult Females #1A and #2A and Adult Males #2C and #3C:  Figure A4 is a map of the known locations of four adult deer and is provided to show spring movements.  Doe #1A was captured on 24 January,  1975, and was not seen again for two months.  Like many other deer in  the late winter of 1975, however, she was frequently seen in the Maturing Open habitat on the south facing slope northwest of her trap site.  She  123.  FIGURE A3. Known movements of adult females 9A ( « - » • ) and 7B A. • • 1975-76). -  1974-75;  124.  FIGURE A4. Known movements of adult females IA (A) and 2A (o) and adult males 2C (©) and 3C (A) in the spring of 1975. indicates direction to lA's summer range-  125.  was seen here five times between 22 March and 9 A p r i l , once in the company of #9A.  She was not seen again during spring, but Figure A5  shows several relocations during the early summer of 1975.  Doe #2A was i n i t i a l l y captured on 30 January, 1975, and recaptured about 200 m to the east on 17 February.  She was relocated twice during  spring on the evenings of 24 and 26 April at the edge of Skookumchuck Prairie in lot 338.  Buck #2C was trapped on 16 February.  He was not seen again until  6:00 p.m. on 7 April when he was found feeding in an alfalfa field 1 km to the west.  (He could have made this move earlier i f he were the  "collared deer" reported in the same field by M. Jamieson on 27 March.) For the next month, he was seen a total of six times feeding on two neighboring fields and the adjacent grassland.  Buck #3C was captured on 25 February, 1975.  He was next seen on  the open shrublands west of Highway 95, just south of Skookumchuck in late June by several men who work at the nearby pulp mill. until late August he was occasionally found in that area.  From then He was killed  by a hunter at the final mark on the map at 6:30 p.m. on 14 September, 1975.  Adult Females #1A and #2B:  Figure A5 is a map of the known movements of does #1A and #2B  126.  FIGURE A5. Known movements of adult females IA ( O ) and 2B ( • ) during the summer of 1975.  127. during the 1975 summer and f a l l , respectively. June, 6 and 8 July at the locations shown. a large cleared "grassland."  Doe #1A was seen on 22  These are all very close to  Several deer were seen feeding in this  clearing each night in early summer and i t is thought that this-doe's movements were centered here at that time.  Doe #2B was seen four times between 3 and 27 November, 1975, as shown in Fig.A5.  This doe's movements at this time were observed in the  clearing mentioned above, and on the slope up to 1 km to the west.  It  is not known whether she used this area throughout the summer or not.  Adult Females #6A and #R3:  Figure A6 is a map of the sighting and radiolocations of two adult females, #6A and #R3.  Doe #6A was i n i t i a l l y captured on 1 February,  1975, sighted about 0.4 km to the northwest on 20 March, and recaptured in the same trap and radio-tagged on 2 April.  She was next located 2.4  km to the south on Wasa Ridge on 18 April and then made a move to the west side of the Skookumchuck Prairie where she was found six times between 24 April and 12 May, 1975. Here her spring range (the area circumscribed by a line connecting the outer locations) was about 13 ha in area and included Grassland, Immature Forest and Brush areas around a small lake.  No signal could be heard in this area from 19 May to 28  May, and i t was not until I made an aerial search of the area on 4 June that I found her again.  At that time, she had returned to Wasa Ridge.  I spent most of 5 June in that area on the ground trying to relocate her, but no other signal was ever received from this deer.  128.  FIGURE A6. Late winter - sprina range of adult female R3 ( « ) and known movements and spring range of adult female 6A (A) in 1975.  129.  Doe #R3 was trapped on 19 April and thereafter was only radiolocated five times in an area about 1 km to the north between 5 and 22 May, 1975.  No more signals were received after that date.  The total  area in the late winter-spring range used by this deer was about 40 ha, predominantly in the Immature Forested Open habitat type.  Adult Female #R1 and Adult Male #1B:  Figure A7 is a map of the relocations of adult doe #R1 and adult male #1B.  The doe was captured and radio-tagged on 3 A p r i l , 1975.  All  subsequent locations from 18 April to 21 May were to the south, with the final location being near some alfalfa fields south of Lazy Lake.  Total  area for her late winter-spring range was approximately 65 ha.  Buck #1B was i n i t i a l l y caught on 19 February, 1975, and seen on 6 March 4 km to the northeast with does #5A and #2B.  He returned to the  area northwest of the capture site by 20 March, and was seen twice (20 March and 2 April) and recaptured and radio-tagged (9 April) on the heavily used southfacing slope.  Four radio-locations from then until 12  May provided the outline of his late winter range that totaled 31 ha. His i n i t i a l movement toward summer range occurred between 12 and 19 May when he was found on the Kootenay floodplain.  No signal could be heard  from the floodplain from 20 May on, and i t is believed that he continued on toward his summer range on the night of 19 May.  Five relocations  between 4 June and 5 August indicated a minimum summer range of 195 ha for this buck.  No signal could be detected after 5 August, 1975.  130.  FIGURE A7. Late winter - spring range of adult female Rl (A) and winter movements- ( o — l a t e winter - spring range, spring movements (•--*) and summer range of adult male IB.  131.  Adult Females #4A and #R2:  Figure A8 is a map of the sightings and radio-locations of two adult females, #4A and #R2.  Radio contact was maintained for the  greatest time with these deer.  Doe #4A was i n i t i a l l y captured on 31 January, 1975.  She was  recaptured 200 m to the east on 21 February and again on 29 March at which time she was fitted with a radio-collar.  She was seen crossing  the road, moving upslope above the trap site to the north of Wolf Creek at 9:45 a.m. on 3 April and once again trapped at the same spot on 8 April.  Two subsequent radio-locations and a sighting (with #R2 and #4B)  between 9 April and 7 May, along with frequent radio contact that could not provide exact locations, indicated this doe used a relatively small late winter range of about 7 ha.  This range included both Grassland and  Mixed Age Fir habitat.  Contact was lost from 7 to 19 May during which time she crossed to the west side of the floodplain.  No clear signal could again be heard  from 20 May until a flight on 4 June found her another 2 km to the norhtwest.  On 3 July, she was again found from the air another 3 km  northwest.  Flights throughout the Trench in early August and September  failed to pick up a signal and none was heard from the ground.  On a  final flight on 10 December, 1975, I received a good signal in the same area where she had been on 19 May, and intermittent contact was maintained with this doe in this area from then until 20 March, 1976.  Here her 10  132.  FIGURE A8. Known locations and movements of adult females 4A in 1975 (A—>) and 1976 ( A ) and R2 in the winter of 1974-75 (•--*) and the summer of 1975 (o). R2 returned to the same winter range in 1975-76.  133.  ha range was located in a small section of what would have been classified Maturing Open habitat had i t been on the terraces of Premier or Wasa Ridges where detailed habitat analysis was conducted.  Doe #R2 provided complete information on the annual cycle of movement.  First trapped on 9 A p r i l , 1975, she was radio-located three times  and sighted once between then and 7 May.  The,next signal was picked up  on the west side of the floodplain on 19 May where she was s t i l l with #4A (as just described).  At this point, however, the association broke  up at #4A continued to the northwest and #R2 had returned to Premier Ridge between 22 and 28 May.  She then moved to her summer range (by 4  June) and was relocated there several times between 4 June and 5 August. No signals were picked up from the ground after that, nor from the air on 10 December.  However, on 14 January, 1976, a good signal was received  on Premier Ridge in the vicinity of her previous winter home range. From then until mid-March, 1976, she was relocated within the same home range area although she moved very l i t t l e .  After 15 March, the bearings  to the radio failed to change direction and i t is thought the collar was either shed or the deer was dead.  A search for her body was unsuc-  cessful as the area inhabited at the time was in the Riparian Mature Spruce habitat with a dense underbrush and complex series of beaver ponds.  Adult Male #R5:  Figure A9 is a map of the trap site and radio locations for adult  FIGURE A9. 1975-76 winter home range of adult male R5 based on trap location (x) and radio-locations (o).  135.  male #R5.  He was captured on 19 February, 1976, and seven subsequent  locations between then and 16 April indicated a minimum home range of 233 ha located primarily within the Young Fir and Mixed Age Fir habitats. No signal was received after 16 A p r i l , but i t is unknown whether this is the result of radio failure or the deer's leaving the study area.  Adult Female #R6:  Figure Al0 is a map of the trap site and radio-locations of an adult female, #R6.  After her capture on 22 February, 1976, she was  found to occupy a home range area of approximately 190 ha mainly in the Mixed Age Fir habitat to the northeast until contact with her was lost on 7 May, 1976.  Adult Male #6B:  Figure All is a map of the movements of adult male #6B.  He was  i n i t i a l l y captured on 21 February, 1975. Although not positively identified again until his recapture 200 m to the northeast of the f i r s t trap site a year later on 18 February, 1976, it is believed from the observer's description that this is the buck reported from the Lavington and Findlay Creek confluence area in September, 1975.  If so this would  indicate a minimum of 40 km between his winter and summer range.  This buck was the most mobile of any deer that was radio-tracked. Extensive movements from the Wolf Creek drainage to the westfacing  136.  FIGURE A10. 1975-76 winter home range of adult female R6 based on trap location (x) and radio-locations (•).  137.  FIGURE A l l . 1975-76 winter movements and home range of adult male 6B based on trap location (x) and radio-locations {&). Dotted line indicates exact location unknown.  138.  slopes of Premier and Wasa Ridges indicated that this animal ranged over 670 ha between 18 February and 7 May, 1976.  He began to move toward a  summer range between 2 and 24 May when he was located on the bank of the Kootenay River.  He continued from there 4 km farther to the northwest  by 28 May, 1976.  Adult Male #4B:  Figure A12 is a map of the known locations of adult male #4B. He was captured at the same spot on 20 February, 1975, and 13 January, 1976.  Two resightings in 1975 f e l l just outside of his 122 ha 1976  winter home range indicating use of the same general area in both winters. This home range loyalty is especially interesting since this is almost certainly the deer reported over 65 km to the north during the intervening summer.  This buck's i n i t i a l spring movement between 16 April and 8 May,  1976, indicated that he might be heading back to the upper Kootenay Valley.  No signal could be heard in the study area as far north as  Skookumchuck between 9 and 28 May, 1976.  139.  FIGURE Al2. 1975-76 winter home range and spring movement of adult male 4B based on trap location (x) and radio-locations (©). o indicate sightings in the previous winter.  


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