Open Collections

UBC Theses and Dissertations

UBC Theses Logo

UBC Theses and Dissertations

Habitat utilization by mule deer in relation to cattle and California bighorn sheep in the Ashnola River… Morrison, Douglas Charles 1972

Your browser doesn't seem to have a PDF viewer, please download the PDF to view this item.

Item Metadata

Download

Media
831-UBC_1972_A6_7 M67.pdf [ 13.27MB ]
Metadata
JSON: 831-1.0101708.json
JSON-LD: 831-1.0101708-ld.json
RDF/XML (Pretty): 831-1.0101708-rdf.xml
RDF/JSON: 831-1.0101708-rdf.json
Turtle: 831-1.0101708-turtle.txt
N-Triples: 831-1.0101708-rdf-ntriples.txt
Original Record: 831-1.0101708-source.json
Full Text
831-1.0101708-fulltext.txt
Citation
831-1.0101708.ris

Full Text

HABITAT UTILIZATION BY MULE DEER IN RELATION TO CATTLE AND CALIFORNIA BIGHORN SHEEP IN THE ASHNOLA RIVER VALLEY, ' BRITISH COLUMBIA by DOUGLAS CHARLES MORRISON B.S.A. , University of British Columbia, 1967 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of PLANT SCIENCE We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA September, 1972 In present ing th i s thes is in pa r t i a l f u l f i lmen t o f the requiremehts for an advanced degree at the Un ive rs i t y of B r i t i s h Columbia, I agree that the L ib ra ry sha l l make it f r ee l y ava i l ab le for reference and study. I f u r the r agree that permission for extensive copying of th is thes is fo r s cho l a r l y purposes may be granted by the Head of my Department or by h is representat ives . It is understood that copying or pub l i c a t i on o f th i s thes is fo r f i nanc ia l gain sha l l not be allowed without my wr i t ten permiss ion. Department of P L A N T SCIENCE The Un ive rs i t y of B r i t i s h Columbia Vancouver 8, Canada Date September 25. 1972 1 ABSTRACT Habitat use by mule deer, particularly in relation to-use by cattle and by California bighorn sheep on the bighorn winter-spring ranges of Flatiron Mountain- was studied from January 1968 through November 1969. Observations were made of (1) food habits, (2) forage production and utilization, (3) the effect of spring and summer utilization on subsequent forage production and (4) spatial and temporal distribution of range use. The results indicate that competition for forage between the native ungulates, deer and sheep, is•largely obviated by differential habitat use. This may point to long term evolutionary ecological niche specialization. Some competition for forage occurs for a short period in the early spring when both ungulate species seek succulent new grass, the supply of which is at f i r s t limited. Cattle use of the winter-spring ranges was excessive and the diets of cattle and the native ungulates are similar, with the exception that utilization of grass by deer was less. Range use by cattle contributed to intra-specific cattle-deer competition on the grasslands in the spring and cattle-bighorn competition on the grasslands during the winter. The study of spring range utilization indicated that deer use was not detrimental to the 1969 annual forage production in areas used by deer. Spring range utilization by bighorn or bighorn in combination with deer reduced the standing crop Of: forage produced on the Agropyron spicatum dominated winter-spring ranges. Sheep 2 utilization on South Slope during the summer, when forage growth was declining, further reduced the amount of forage available to the wintering bighorn population. 3 TABLE OF CONTENTS Page ABSTRACT , .• • ....... 1 TABLE OF CONTENTS ... ..... .' ....... 3 LIST OF TABLES • .. 6 LIST OF FIGURES . .. .• • • • • • 9 LIST OF APPENDICES ..... • ...... ' 10 ACKNOWLEDGEMENTS • •. . .. •, •• .. 12 1. INTRODUCTION • 14 2. THE STUDY AREA • • .... • ... 17 2.1 Description • • ... 17 2.2 Past and present use • •.. ....... 24 3. LITERATURE REVIEW . . .• 31 3.1 Competition and ecological adaptation • 31 3.2 Niches occupied by wild and domestic ungulates 33 3.3 Forms and patterns of competition • 37 3.4 Review of bighorn sheep-mule deer-cattle range relations •, 44 4. METHODS AND PROCEDURES • .• ..48 4.1 Food habit analysis ...... 48 4.1.1 Feeding observations - 48 4.1.2 Grazed plant loop-transects • 49 4.1.3 Rumen sample analysis .... 4.1.4 Forage preference index • ^ 4 4.2 Range forage appraisal • 53 4.2.1 Site selection .• 53 4.2.2 Forage measurement 54 4.2.3 Community description .; • :.. • 53 4.2.4 Total nitrogen analysis • 59 4.3 Data analysis .... • 59 4.3.1 Food habits • ...... 59 4.3.2 Forage production and utilization . .• gg 5. OBSERVATIONS AND RESULTS : 61 5.1 Food habits 61 5.1.1 Cattle spring food habits > • 61 5"; 1.2 Bighorn spring food habits •—. . 65 5.1.3 Mule deer seasonal food habits — .. 71 , 5.1.3.1 Fall diet 72 • 5.1.3.2 Winter diet ....... • • 76 5.1.3.3 Spring diet . .... 77 5.2 Range forage appraisal .... t 86 5.2.1 Site locations 86 5.2.2 Community descriptions ......... 88 5.2.3 Forage measurements 6. DISCUSSION ' • .... .'105 6.1 Food habits ... .• ... 105 6.1.1 Comparison of food habit studies • 108 6.1.1.1 Cattle • !Q8 6.1.1.2 Bighorn sheep •. jgg 6.1.1.3 Mule deer • -yL2 6.1.2 Potential dietary competition and animal equivalence 120 6.1.2.1 Potential competition in the spring ... 120 6.1.2.2 Potential competition in winter .. •. ]_24 5 6.1.3 Animal equivalence • •. 126 6.1.4 Limitations of and recommendations for food habit studies 131 6.1.4.1 Rumen sample analysis 131 6.1.4.2 Grazed plant loop-transect 133 6.1.4.3 Trailing feeding animals 1 3 4 6.2 Range, forage appraisal 134 6.2.1 Forage measurements •. 135 6.2.2 Forage measurements on the alpine summer range • • 144 6.2.3 Limitations of range forage appraisal 2.4-4 6.2.3.1 Forage production and utilization ...... 7. RECOMMENDATIONS 146 8. SUMMARY ; 150 9. LITERATURE CITED 156 6 LIST OF TABLES Page 2.2.1 Cattle stocking rates on Flatiron Mountain, including Joe Lake range 25 2.2.2 Numbers of mule deer observed in the Ashnola, 1957 to 1959 , 30 5.1.1 Cattle, bighorn and mule deer spring food habits. Relative proportions by oven dry weight, expressed as a percentage, for each taxon and forage class from rumen sample analysis.. 62 £ 63 5.1.2 Statistical analysis of cattle, bighorn and deer spring food habits as indicated from rumen sample analysis. Relative proportions by oven dry weight, expressed as percentages, of identified forage only 64 5.1.3 Forage class frequency of grazing and preference indices for bighorn and deer during the spring, 1968 - 67 5.1.4 Bighorn sheep forage species preference index as indi-cated from South Slope, Agropyron community, grazed plant loop-transect data, July 7, 1969 68 5.1.5 Bighorn sheep forage species preference index as indi-cated from South Slope, Agropyron-Artemesia community, grazed plant loop-transect data, July 7, 1969 69 5.1.6 Seasonal mule deer food habits. Relative proportions by oven dry weights, expressed as a percentage, for each identified taxon and forage class from rumen sample analysis Vv .... 73 S 74 5.1.7 Statistical analysis of mule deer seasonal food habits as indicated from rumen sample analysis. Relative proportions of the forage classes are expressed as percentages, from identified forage only. 75 5.1.8 Statistical analysis of within-spring mule deer rumen sample analysis. Relative proportions by oven dry weight, expressed as percentages, of identified forage only .... 80 7 5.1.9 Mule deer forage species preference index as indicated from Juniper Slope, Poa community, grazed plant loop-transect data • 82 5.1.10 Mule deer forage species preference index as indicated from Juniper Slope, Artemesia community, grazed plant loop-transect data . ... 83 5.2.1 Topographic and soil information for exclosure plot sites on Flatiron • Mountain and Joe Lake -.- 87 5.2.2 Species composition of the four winter-spring range communities studied, Flatiron Mountain, 1969 89, 90 S 91 5.2.3 Species 'composition of the two alpine (summer range) communities studied, Joe Lake, 1969 92 g 93 5.2.4- Fall forage utilization - by cattle on Flatiron Moun-tain, 1968 96 5.2.5 Over winter forage weathering and shattering losses on South Slope and Juniper Slope 1968-69 and on Juniper Slope 1969-70 - 97 5.2.6 Fall and over winter forage utilization by sheep on Flatiron Mountain, 1968-69 99 5.2.7 Forage production, utilization and the effect of wildlife spring grazing on forage production, shown as differences in mean dry matter yields between protected and grazed areas, Flatiron Mountain, 1969 100 5.2.8 Soil moisture at subsequent dates in 1969, for six inch soil horizons from 0 to 24 inch soil depth, at two study sites on Flatiron Mountain 102 5.2.9 Forage production and utilization by wildlife on two alpine communities, Joe Lake, 1969 104 6.1.1 Food habits of cattle on North American ranges 110 6.1.2 Food habits of bighorn sheep populations I l l 6.1.3 Food habits of mule deer populations 115 8 6.1.4 Animal equivalence calculations using seasonal ungulate food habit information from Flatiron Mountain ranges, 1968-69 130 6.2.1 Utilization of forage produced in the Poa pratensis community, Juniper Slope, 1968, by cattle and wild-l i f e • • • ' 136 9 LIST OF FIGURES Page 2.1.1 Location map of the Ashnola Study Area ............. , 18 2.1.2 Aerial-photograph showing locations of slopes and winter-spring range study sites on Flatiron Mountain • 20 2.1.3 Aerial photograph showing location of summer range study sites,.Joe Lake Area....; 22 4.2.1 One-meter square quadrat and hand clippers used in measurement of forage production and utilization ... 55 4.2.2 Fenced plot on the Poa. pratensis community, Juniper Slope..... ........ .. 55 5.1.1 Highlined and.decadent condition of Douglas f i r regeneration between Starvation and Flatiron Slopes, Flatiron Mountain, 1968 • • 78 6.1.1 Percent distribution of grass, forbs and browse utilization in relation to their availability factor value during the spring; from grazed plant loop-transect data ,]_07 6.1.2 Seasonal•variation in oven dry weights of grass, forbs and browse, expressed as percentages, found in mule deer rumen samples • .............. nu 6.1.3 Relative forage class - use by mule deer on Juniper Slope and bighorn on South Slope in the spring.' Data from grazed plant loop-transects .122 6.1.4 Variation in grass, forbs and browse utilization in the spring food habits of cattle, bighorn and deer, from rumen sample analysis.................................. 123 6.1.5 Comparison of winter food habits of mule deer and bighorn... 127 6.2.1 Forage remaining on Juniper Slope in.the f a l l of 1968 after combined cattle and wildlife spring and summer use.. • 138 6.2.2 Forage remaining on Juniper Slope in the f a l l of 1969 after spring grazing by wildlife only • •.• 138 6.2.3 Effect of spring grazing on subsequent winter-spring range• forage growth rates, Flatiron Mountain, 1969 • . 141 10 LIST OF APPENDICES Page 4.2.1 Paired plot design for measurement of forage utiliza-tion by cattle in the f a l l of 1968, and forage u t i l i -zation by sheep during the winter of 1968-69 166 4.2.2 Field plot design for measurement of alpine forage production and utilization, 1969 167 4.2.3 Field plot design for forage production and utiliza-tion measurements , 1969 • 168 5.1.1 Hereford cattle spring food habits, as indicated from rumen sample analysis, in gms .... 169 5.1.2 Bighorn sheep spring food habits as indicated by rumen sample analysis, in gms .. ....... 5.1.3 Forage species occurrence and utilization by big-horn sheep on South Slope, Agropyron community, as indicated from grazed plant loop-transect data. July 7, 1969 , 1 7 1 g ± 7 2 5.1.4 Forage species occurrence and utilization by big-horn sheep on South Slope, Agropyron-Artemesia community, as indicated from grazed plant loop-transect data. July 7, 1969 .• • 1 7 3 g 1 7 4 5.1.5 Mule deer f a l l food habits as indicated from rumen sample analysis, in gms .• 175 5.1.6 Mule deer winter food habits as indicated from rumen sample analysis, in gms 1 7 5 5.1.7 Mule deer spring food habits, as indicated from rumen sample analysis, in gms 177 g JJQ 5.1.8 Forage species occurrence and utilization by mule deer on Juniper Slope, Poa community, as indi-cated from grazed plant loop-transect data. June 13, 1969 179 g iso 11 5.1.9 Forage species occurrence and utilization by mule deer on Juniper Slope, Artemesia community, as indi-cated from grazed plant loop-transect data. June 15, 1969 • - 181- g 182 5.2.1 Scientific and popular names for plant species collected on the winter-spring ranges, Flatiron Mountain, 1968-69 ......... 183,4,5,6,7 5.2.2 Scientific and popular names for plant species collected on the summer ranges, Joe Lake area, 1968-69 . 188 g 189 12 ACKNOWLEDGEMENTS Field, work was financed by research grants from the Canada Department of Agriculture and National Research Council of Canada (N.R.C.), held by Dr. V.C. Brink. Financial assistance was provided by an N.R.C. Bursary in 1967-68, and N.R.C. Postgraduate Scholarships from 1968 through 1971. The Department of Plant Science provided laboratory facilities and office space. The use of Mr. Monks's cabin in the Ashnola River Valley was greatly appreciated, as was the use of the University cabin in-the study area. I am especially grateful to Dr. V.C. Brink of the Department of Plant Science, Faculty, of Agriculture, University of British Columbia, who supervised the study. Fellow students Eike Scheffler and Fred Harper also gave many helpful suggestions in the early stages of the study. Dr. I. McT. Cowan, Dean of Graduate Studies; Dr. P.J. Bandy, Head, Research Division, Messrs. D. Hum, Regional Supervisor, and D. Spalding. Regional Biologist, of the British Columbia Fish and Wildlife Branch, and Mr. D. Pearce, Department of Plant Science, provided valuable advice and assistance during the study. Dr. G.W. Eaton, Department of Plant Science, and Dr. A. Kozak, Faculty of Forestry, provided assistance with statistical analysis of data. Thanks are due D. Roger, E. Storness-Kress, L. Stellick, M. Neible, D. O'Dell and summer assistants C. Shaw and G. Barr who 13 provided excellent field assistance. I am indebted to the Wabnegger and especially the Quaedvlieg families of the Barrington Ranch, Keremeos, and gratefully acknowledge their sincere hospitality and invaluable assistance. To these people, and many others whose cooperation and help made the study possible, I extend sincere thanks. -14 1. INTRODUCTION An ecological study of ungulate habitat utilization was initiated to evaluate biotic factors affecting the quantity of available forage on the California bighorn, sheep (Ovis canadensis califbrniana)^ winter range of the Ashnola. Previous investigators, Blood (1961, 1967) and Demarchi (1965) have suggested that retrogression has occurred on the Ashnola winter ranges due to the misuse of open rangelands and that the lack of available winter forage limits the bighorn population. They evaluated forage utilization by cattle and wildlife and stated that competition was occurring. The extent of range utilization by mule deer, Odocoileus hemionus hemionus Rafinesque) however, was not studied as a discrete factor. This, then is the f i r s t intensive study of mule deer on Interior ranges, particularly as they may compete for forage with bighorn and cattle in the northern portion of mule deer distrib-ution. The forage production and1 utilization phase of the study was concentrated on the grasslands rather than the forested range since (1) i t was reported (Blood, op.cit.) and confirmed by this study that bighorn preferred and-relied upon the grasslands rather than the forested range, and (2) interspecific competition can only be evaluated on areas grazed in common. In discussing the Ashnola range competition problem, Blood (op.cifr~.') stated (it) "... is basically one of spring and f a l l u t i l -ization by cattle of sheep winter-spring range". This, however, should be expanded to include spring utilization by sheep and deer of sheep "Sfaxnraalian nomenclature after Cowan and Guiguet (1965). 15 winter-spring ranges. The three ungulate'species use the grasslands synchronously during the spring months with competition possibly arising at that time and/or later during the winter months as a result of spring and summer utilization, (a sequential use pattern). Factors affecting range utilization, and hence an evaluation of intra- and inter-species range competition, were studied.from January 1958 through November 1969 on the Flatiron Mountain section of the Ashnola winter ranges. In the west, winter ranges are cr i t i c a l for the survival of big game populations (Stoddart and Rasmussen, 1945). Forage competition on winter ranges i s , as a result, a very important problem. In the Ashnola, competition on the winter ranges is subtle, for i t may be the spring and summer grazing on the winter ranges that removes forage important for the wintering sheep.population. The focal points of the study then were two; f i r s t , to measure spring and early summer utilization of the bighorn winter range by each ungulate species, hereford cattle, bighorn and mule deer, and to evaluate the effect of that utilization on the standing forage crop at the time of. maximum production; second, to evaluate winter use of the bighorn winter range.. Of the various, factors affecting the standing crop of useable forage available to overwintering animals the major abiotic ones are the length of the growing season as determined primarily by soil moisture (Harper, 1969) and the depth of the winter snow pack. The'major biotic factor is the extent of grazing during the forage growing season. The following variables were evaluated to describe ungulate habitat utilization, and to evaluate the presence or severity of range 16 competition: 1. The duration and season that each ungulate class utilized the bighorn winter range, 2. The spatial distribution of the three ungulate classes in relation to habitat types in the study area, 3. The numbers of each ungulate species involved, 4. The winter and spring forage preferences of each ungulate species, 5. The quantity of the forage utilized by cattle, deer and sheep in the spring, early summer and winter while on the bighorn winter ranges, and the effect of early use on subsequent forage production.' In addition biological data were collected on Interior mule deer, and the collection of bighorn population dynamics data was continued. For the purpose of the dissertation two sections of the total study are presented; they are: 1. Food habits, and 2. Range forage appraisal. Ultimately, management of the ranges may necessitate limiting the numbers - of each ungulate species to achieve a balance of range utilization. To obtain this, information concerning the seasonal food habits of each species, the effect of range use by each species and a more ecologically based system of animal equivalence is essential. The1 sections of the total study presented herein provide data on which management decisions may be based. 17 2. THE STUDY AREA 4 2.1 Description The Ashnola study area (Figure 2.1.1) is immediately north of the 49th Parallel in south central. British Columbia, approximately 150 miles each of Vancouver. The intensive study area on Flatiron Mountain lies approximately ten and one-half miles southeast of Keremeos on California bighorn,sheep and mule deer winter ranges (Figure 2.1.2), and on the Joe Lake section of the bighorn.summer range (Figure 2.1.3). It is within the 35,000 acre Ashnola Bighorn Sheep Management Area. Physiographically, the area lies within the mountainous southern portion of the Interior Plateau (Brink and Farstad, 1949). It has also been described as lying within the transition region between the Plateau and the Cascade Mountain system which lies to the south, east and west (Holland, 1964). A detailed description of the study area's physiography was undertaken by Blood (1961) and Scheffler (1972). The•general ecology of the Interior Inter-Mountain Region, its climate, soils and vegetational zones have been described by Spilsbury and Tisdale (1944),.Tisdale (1947), Tisdale and McLean (1957), Van Ryswyk et a l . (1966), and McLean and Marchand (1968). Climatolog-ically the area is characterized by the "rain shadow" effect of the Cascade Mountains, and two seasonal maxima for precipitation (Tisdale and McLean, op.cit.).. The major rainfall maximum in spring, May through July, is .important in determining forage quantity; the lesser rrBximum of precipitation in December and January is important in recharging soil moisture, and developing the winter snow pack. It is during the latter Figure 2.1.1. Location map of the Ashnola Study Area 19 Figure 2.1.2. Aerial photograph showing locations of slopes and winter-spring range study sites.on Flatiron Mountain. Legend: 1. Poa pratensis community - deer only grazing. 2. Stipa-Arternesia community - deer only grazing. 3. Agropyron spicatum community - sheep only grazing. 4. Agropyron spicatum community - deer and sheep grazing. Scale: 40 chains = 1 21 Figure 2.1.3:'. Aerial photograph showing location of summer range study sites, Joe Lake area. Legend: 1. Carex community. 2. Danthonia-Carex community. Scale: 20 chaiLns = 1" 23 period that snow accumulates in the study area, occasionally to a depth of several feet, which restricts winter range use by bighorn, and mule deer in the area. However, the ridges on the south and south-west facing grasslands characteristically blow relatively free of snow and provide forage for the wintering bighorns. The basic climatic variables in the Ashnola basin were studied by Scheffler (op.cit.). Harper (1969) described the climatic variables affecting forage production on Flatiron Mountain. The soils and vegetational types occurring in-the area have been locally described by Blood (op.cit.), and Demarchi (1965), whereas Scheffler (op.cit.) inventoried them on a more extensive basis. The altitudinal zonation effect of climate, soils and,vegetation as des-cribed by.Spilsbury and Tisdale (op.cit.)•is evident. The higher mountains in the southern part of the area provide summer range while the grass and forest ranges at lower elevations in the northern part of the area, provide winter range for the'ungulate population. At the lower elevations, north, east and west slopes support forests of Douglas 2 f i r (Pseudotsuga menziesii var. glauca) (Beissn.) Franco., or Lodgepole pine (Pinus- contorta) Dougl. in burnt over areas. The Douglas f i r range has received light cattle use and is in good to excellent condition over most of Flatiron Mountain. This potentially valuable range, under proper grazing management, could provide forage beneficial to cattle in the early summer (McLean, 1967; McLean et a l . , 1969; Freyman-and Van Ryswyk, 1969; and Harper, 1969). The lower elevation grasslands 'Nomenclature after Hitchcock et .al. (1969). 24 on south and southwest to west.slopes between 2500 and 6000 feet have received intensive livestock use and were described (Blood op.cit and Demarchi op.cit.) as .retrogressive and below optimum in production over much of their acreage. At higher elevations the forest is dominated by alpine fir'(Abies lasiocarpa) (Hook.) Nutt. and Engleman. spruce (Picea engelmannij) Parry ex Engelm. Open areas are of two types: (a) subalpine meadows dominated by grass and forbs and (b) alpine tundra dominated by sedges. The latter is the summer range of the bighorns, while the former, together with the sub-alpine forest, is the summer-fall range for deer. The. meadows and sub-alpine forest edge is the main summer range for cattle. 2.2 Past and present use In the Ashnola.region cattle, domestic sheep, horses, mountain goats (Oreamnos americanus Blainville), mule deer and California bighorn sheep have been the ungulates utilizing the range. Domestic stock have been grazed in the Similkameen and Ashnola valleys since the 1880's. Prior to 1950 or 1951 as many as 600 head of cattle were carried on the Flatiron and Crater Mountain ranges, along with 1000 head of domestic sheep, which grazed for seven seasons between 1913 and 1923. However, even .as late as the early 1950's Brink and Cowan (pers. comm.) observed a band.of domestic sheep utilizing the Ashnola ranges. The ranges also received year round use by at least 250 horses prior to 1919, when most.were removed leaving less than 80 on the ranges until the 1930's. In more•recent times range use by horses, used for 25 recreational purposes has been increasing. Use of bighorn winter ranges by mountain goats in the past was undoubtedly light. Present use is negligible due to the low density of the goat population. Specific cattle grazing data, stocking rates by area, exist since 1952 when separate grazing permits were issued for Crater and Flatiron Mountains. Light to moderate grazing occurred between 1952 and 1957. Since 1957 Crater and Flatiron Mountains, the latter including Joe Lake range, have been grazed by 195 and 150 cattle respectively between May 1 and the middle of October. Cattle use the lower elevation ranges in spring, early summer and f a l l . Past use by domestic stock, especially sheep and horses, was in a l l probability the important.factor causing the present retro-gressive condition of the Ashnola ranges. To the further detriment of the range, a recent series of short duration grazing permittees in the area did not help to secure proper grazing management (Table 2.2.1). Table 2.2.1. Cattle stocking rates on. Flatiron Mountain, including Joe Lake Range, 1965-1970 Date Number Duration Permittee 1965 150 cattle May l-Oct. 31 E S M Cattle Co. Ltd. 1966 150 cattle May l-Oct. 31 Christa Ranch Ltd. 1967 150 cattle • May l-Oct. 31 Rimson Equipment Ltd. 1968 150 cattle May l-Oct. 31 Rimson Equipment Ltd. 1969 Grazing rights purchased by Fish and Wildlife Branch. 26 In 1968 the British Columbia Fish and Wildlife Branch purchased the grazing permit for 35,000 acres of the Ashnola range, including the Crater Mountain, and Flatiron Mountain winter ranges and a portion of the sheep summer range at the head waters of Juniper, Gilanders and' Susap Creeks. Cattle use on these ranges was terminated following the 1968 grazing season to enhance the preservation of the bighorns. A more complete discussion of past domestic stock use in the Ashnola may be found in Blood (1961) and Demarchi (1965). The numers of California bighorn in the area in pristine times is not known. It is believed to have been a substantial number and this is supported by Buechner's (1960) discussion of bighorn status in adjacent Washington, and Blood's (1961) historical account of the Similkameen herds. The'numbers.of sheep are believed to have declined in the late 1880's. In 1909 a strict sheep hunting closure was imposed and the population began to make a comeback (Williams, 1925). However, after the f i r s t World War, their numbers remained relatively constant, probably as a result of poaching, the retrogressive range condition and.the alienation of low elevation ranges cri t i c a l for the survival of a large population in severe.winters. In more recent times there appears to have been an increase in the total British Columbia population. In 1960 Sugden (1960) estimated the British Columbia herds totaled 1,235 individuals. A more recent estimate by Spalding and Mitchell (1970). .placed the population at 1,879 individuals. In the Ashnola, Blood (1961) observed 70 and 170 sheep, on Crater and Flatiron Mountain winter ranges respectively, for a total of 240 sheep. Demarchi (1965) 27 observed 75 arid 175 sheep in 1964 for a total of 250 in the Ashnola. On June 29, 1967, Scheffler and-Harper (pers. comm.) observed 182 ewes, lambs and young rams on South slope. In addition they stated there were approximately 35 legal rams, for a total of 217 individuals on Flatiron Mountain. D. Spalding (pers. comm.) estimated that in 1970 there were 230 sheep on Flatiron,Mountain and an additional 120 on Crater Mountain (including Paul Creek) for a total of 350 bighorn in the Ashnola. Big-horn, use the Flatiron Mountain ranges.in late f a l l , winter and spring. A small segment of the population, primarily ewes, lambs and young rams, remain on lower elevation ranges throughout the year. "v"' The numbers of mule deer using the range in the past is unknown since they received less attention in the Ashnola than the bighorn, and they are more difficult to, observe and to enumerate. It might be inferred from discussions with local residents and a review of past accounts that the population was once larger than that presently using the ranges. (Due to time limitations a review of possible material contained in the Provincial Archives, Victoria was not undertaken during this study). Adam Monks, Ashnola River Valley (pers. comm.) stated that in the past one could see 300 deer in one day's travel over the Ashnola winter ranges. "Garney" Willis, Cawston, B.C. (pers. comm.) stated that in April, 1916, he observed between 400 and 500 head of deer and-sheep on South Slope. If the average''maximum number of big-horns using South Slope in the spring i s , and likely was in the past, about 170 individuals then possibly as many as 200 to 300 deer could have been observed on South Slope. Trout (1886) and Phillipps-Wooley (1894) stated that mule deer and mountain sheep were in abundance in the SiLmilkameen area. Game abundance is also indicated by the number; of ] 28 deer slaughtered for the hide markets. L.L. Dyche's collecting exped-ition to Mount Chapaca and adjacent B.C. encountered hide hunters in this region, and remarked "... the piles of deer and sheep skins which lay about the camp made i t evident that they were skin hunters. It was later learned that these two men killed over 200 deer during that summer" (Edwords, 1893). In 1887, Phillipps-Wooley (1888) made a hunting excursion into the Ashnola region where he found game in abundance on what must have been Crater Mountain, according to the description in the party's records. In describing the numbers of deer in this area he stated, "This forest seemed f u l l of mule deer", and further "In the morning a l l the downs are alive with them, in big bands". Williams (1925) in discussing the general decline in mule deer populations suggested that "The decrease was brought about in many ways, but the start began when market hunters slew them solely for their hides. At one time there was an almost constant stream of wagons passing from the Okanagan Valley across the line to the United States, and their sole freight consisted of mule deer skins". Spalding (1968) in describing the historical background of deer numbers in the "Boundary" region of southern B.C. discusses the influence of forest succession following fires, on the extent and quality of deer range. Large fires in the early years of this century produced success-ional range conditions suitable for an increase of deer populations in many areas of southern British Columbia,, including the Ashnola-Simirkameen. At present, the Fish and Wildlife Branch does not know the 29 precise size of the Ashnola mule deer population (Harper, 1969). The results of ground and aerial counts carried out in the Ashnola by-personnel of the B.C. Fish and Wildlife Branch and others indicate only the numbers of observable deer (Table 2.2.2). Dave Spalding (pers. comm.) estimates that within the Ashnola drainage there may be 1800 to 2500 deer. Table 2.2.2. Numbers of mule deer observed in the Ashnola, 1957 to 1959 . Date Deer Observed South Slope Juniper Starvation Flatiron Mountain Crater Mountain Total Method January 24/57 112 April 20/58 109 January 3/59 10 May 17/59 January 22/60 3 March 12/60 4 May 1960 February 3/61 May 21/61 January 11/62 April 1962 April 29/62 15 Apr. 16-17/64 March 22/65 March 23/66 February 10/66 March 15/67 February 15/68 March 28/68 66 January 8/69 January 21/69 8 91 12 74 13 3 144 100 5 108 Total Ashnola 13 94 70 155 19 101 40 20 152 149 165 205 130 355 41 175 60 27 313 18 144 66 293 15 224 100 152 149 165 98 478 414 91 Aerial?" Census Ground Aerial Ground" Aerial 1 Aerial Ground: Aerial_ Ground^ Aerial-^ Aerial-^ Ground^ Ground^ Ground,, Aerial Aerial^ Ground^ Ground L Ground £ Aerial Aerial^ Aerial^ 4 Stated values indicate distribution of deer tallied in the /'total" value: " Values indicate number of deer. 2 E. Taylor. Monthly reports to B.C. Fish and Wildlife Branch. ~ Keremos and Cawston Rod and Gun Club. J Blood (1961) M.Sc. Thesis. D. Spalding, Regional Biologist, B.C. Fish and Wildlife Branch. Letter dated October 30/70. C O o 31 3. LITERATURE REVIEW 3.1 Competition and ecological adaptation Ecological niches of different species may overlap and as a result competition may occur (De Vos, 1969). Dice (1952) defines "niche" as a l l of the features of the ecosystem that a species utilizes and states that the term does not include,. except indirectly, any consideration of the functions that a species serves in the community. Deer and mountain sheep largely utilize.different ecological niches as defined by Dice (ibid) but use the same niche in "terms of Elton's (1927) and Miller's (1967). "functional" interpretation of the word. Miller (ibid) states that competition by direct exploitation is a primitive and unstable form of interaction. He believes that interference mechanisms such as territoriality represent evolutionary advances toward more effective biotic control of population interactions and numerical stability. Wild and domestic ungulate populations have not been interacting long enough nor freely enough on North American ranges to have developed such behavioral mechanisms, hence competition can and often does occur. Skovlin and Harris (1970, p. 77) stated "Forage use by big game alone was very small when compared with that of cattle and big game". Further " . . . i t is apparent that cattle management had a decided impact on availa-b i l i t y of forage for game". Few studies have been conducted to investigate interference mechanisms within mule deer populations (Copperider, 1967) and between mule deer and bighorn sheep populations (Buechner, 1960). They indicate that Miller's interference mechanisms do not exist here either. 32 It is the degree to which ecological adaptation has developed which functions to reduce actual competition in areas of overlap between wildlife species occupying the same functional niche, as defined by Miller (1967). In this respect different habitat preferences, that is evolutionary adaptation to separate ecological niches, reduces competition between deer and sheep. This is especially true when availability of preferred habitat is not restricted due to alienation from ranges, competition from domestic stock, severity of weather con-ditions and successional changes in vegetation. In discussing the role of physiology in the evolution of adaptation and competition between animals Beament (1961) emphasizes the development of behavioural physiology (behavioural habits). These-habits, he points out, limit an animal to an environment far narrower than that in which i t can survive; hence by this means behavioural characteristics largely eliminate interspecific competition. In discussing behavioural adapta-tions to competition, Brian (1956) employed the term "selective segregation" which involved "...the selection by instinctive behavior of totally different habitats (that is resources- and modifying factors combined)". Taber and Hoffman (1963) discussed the behavioural res-ponses of mammals in mountainous habitats. They noted that there is "pronounced restriction of some species to specific habitats in portions of their ranges in response to the presence of closely related species". Bighorn sheep have evolved with and occupy bunchgrass ranges over most of their geographical distribution. They have also evolved with mule deer as a coinhabitor of the ecosystem. Actual competition 33 is reduced by adaptation to this co-existence, especially on winter ranges,through the utilization of separate'ecological niches. Talbot and Talbot (1963a and 19.63b) and Vesey-Fitzgerald (1965) have documented examples of. ecological niche adaptation in the plains ecosystem of East Africa. They observed complementary co-existence and non overlapping preferred diets'of various native plains herbivores, thus allowing a larger animal biomass and efficient utilization of available forage. 3.2 Niches occupied by. wild and domestic ungulates Bighorn sheep and mule deer inhabit different ecological niches, probably as a result of evolution in the same geographical area at about the same time as described by Cowan (1940, 1956) and Buechner (1960) . Throughout their range bighorn sheep (Ovis canadensis) are known to inhabit open ranges, either natural or created through distur-bance of climax vegetation, which are interspersed with rough escape terrain (Buechner, ibid; Flook, 1964; Capp, 1968). However, Sugden (1961) pointed out that the open ranges of the California bighorn were not always associated with mountainous terrain; probably they once made extensive use of arid grasslands in the major B.C. Interior river valleys. Desert bighorn being the exception, .all other populations are. closely associated with two.general communities of grasslands: (1) Agropyron spicatum-Poa secunda, and (2) Festuca idahoensis- Agropyron spicatum (Buechner, on, c i t . ) . Seasonal migrations take sheep, into extensive alpine and sub-alpine ranges in the summer. They return in the autumn to their usually 34 restricted winter ranges after the f i r s t snow storms blanket the summer range (Capp, op.cit.). However, in certain areas, non-migratory seg-ments of normally migratory populations have been observed: Smith (1954) and'Morgan (1968) noted this in Rocky Mountain bighorn herds in Idaho, Sugden (1961) in a California bighorn herd in the Churn Creek region of British Columbia, Reade Brown (pers. comm.) in the transplant California bighorn sheep herd reintroduced into the Sinlahekin area, Washington. Sugden also reports that the California bighorn populations at Riske Creek, and Dog Creek undergo no marked seasonal movements. Bighorn sheep are generally considered to be grazers, mainly utilizing herbs; some browse may be important during the winter. (Sugden, ibid; Blood, 1967; McCullough and Schneigas, 1966; Demarchi, 1965; Cooperider, op.cit.; Capp, op.cit.). Mule deer inhabit a wider range of habitats, extending from the valley bottoms to slopes of moderate to steep gradient in the mountainous shrub and. coniferous'forest ranges of the west (Cowan, 1956). In a review of literature dealing with ungulate-range relationships, Capp (op.cit.), summarizes studies that indicate mule deer prefer forest and shrub communities and their edges rather than grassland communities, except for a short period in the spring. This habitat prefernce was substantiated on summer range by Reynolds (1966a, 1966b) i n Arizona, Skovlin et al . (1968) in Oregon and on winter ranges by Cowan (1947) in Jasper National Park. Mackie (1970) studying seasonal habitat utilization in Montana found that mule deer preferred forested range for feeding and bedding except in the winter and spring when, due to forage 35 availability, they fed in the open Arternesia-Agropyron community. Populations, other than those of the grasslands at the eastern limits of their range, on the plains, and in desert shrub communities, exhibit seasonal migrations from alpine and subalpine summer range to restricted lower elevation forested winter ranges (Einarsen, 1965; H i l l , 1965). Weather conditions, primarily snow depth, dictate.when deer are forced to return to the winter ranges (Shepherd, 1960; Dorrance, 1966; Richens, .1967). Mule deer winter ranges, like those of the bighorn which they often overlap geographically, are characterized by low precipitation. Mule deer are considered to be browsers throughout their range. Winter forage consists predominantly of browse; however, during the summer when forbs are.readily available, browse utilization becomes secondary. It appears that only for a short period in the spring, when the new growth.of grass provides succulent forage, does grass.form a significant portion of the deer's diet (Yeager, 1960; Lovass, 1958; Wilkins, 1957; Smith, 1952; H i l l , op.cit.; Capp, op.cit.). Cattle differ from wild Northern American ungulates primarily in that they were introduced to North American ranges in relatively recent times. Also, their seasonal use of range, movements and habitat u t i l i - . zation, is or can be controlled by man. A review of the literature indicated that there•is a sparse literature on cattle range preferences, and on cattle distribution in relation to various range conditions and available habitat types. Generally cattle prefer open range when left unherded, rather than open-ings created in forests, or forested range. This preference for open 36 range was demonstrated in studies conducted by Wagnon (1963) in , California, Reynolds (1966a and 1966b) in Arizona, and Mackie (1970) in Montana. On the Starkey Experimental range in Oregon, Skovlin et al. (1968), and Skovlin and Harris (1970) found that cattle use was equally distributed between forest and grassland.. However, this distribution pattern likely occurred as a result of efforts, following 1948, to obtain better livestock distribution and more efficient use of the herbage on the Starkey Experimental range (Harris, 1954). Harris (ibid) further stated that prior to 1948 cattle use was more intensive on the grasslands than on the timbered range. A review of the literature failed to produce any published data from studies designed to evaluate cattle habitat preferences on British Columbian rangelands. Sugden (1961) however, does mention that cattle used exposed gentle slopes and benches, the aspen-willow forest type and a few sedge meadows in the Churn Creek area of the Cariboo. Another, rather indirect, reference to the habitat preferences of cattle can be interpreted from McLean's (1967) work on cattle growth rates in central British Columbia. He found that when cattle were confined, by fencing, to forested range generally consisting of a pine grass (Calamagrostis rubescens) dominated under-story, they were able to attain promising daily weight gains. Cattle prefer forage consisting primarily of grass and grass-like plants (Skovlin, et a l . , op_. c i t . ; Mackie, op_. c i t . ; Cook et al. , 1967), especially when forage quality and variety has not been reduced by prior heavy grazing. Smith and.Doell (1968) and Lesperance et al. (1970) demonstrated that cattle w i l l utilize browse species when the more succulent - forage has been consumed. No reference to cattle food 37 habits on B.C. ranges was found in a review of the literature. 3.3. Forms and patterns of competition Competition is an often inaccurately used term, its interpreta-tions are varied depending upon what factors are emphasized. For the purpose of this discussion competition may- be separated into two major divisions depending whether the actions .and effects of competition are being considered either at the animal population interaction level, or at the habitat (range) utilization level. ' It has become necessary, as w i l l be seen below, to state what form of competition is being con-sidered. The two divisions can be further subdivided into interspecific (between species) pr intra-specific (within species) competition. Odum (1959) discusses competition as one of eight basic types of interactions between two species at the animal population level. The•system is based on the positive or negative -results of the inter-actions on population growth equations. This system is applicable for intraspecific as well as interspecific competition because the "self-crowding effects" in a single species population situation can be applied in the same terms to its population growth equation. In his discussion, Odum (ibid) states that "competition" is an interaction in which each population adversely affects the other in the struggle for food, nutrients, living space, or other common need. He does not stipu-late that the supply of materials has to be limited. Gross. (1970) like Odum. (op. cit.) interprets competition by way of the results that population interactions have on reproduction, 38 mortality, immigration and emigration. Thus, he converts subjective evaluation of competition to objective quantitative values for the purpose of computer modeling of population data. However, Milne (1961) disagrees with this approach.. Milne (ibid) also criticizes Odum's (op. cit.) definition and "result" rather than "cause" style of inter-preting competition. This is based on the observation that "most of the common results of both intra-specific and, interspecific competition, such as debility, retarded development, stunting, reduced natality, increased mortality and emigration can equally well be produced by predators or parasites or pathogens or weather". Certain causal.factors related to habitat and not directly associated with the supply of forage per se may also be included in this l i s t . These include, changes in forage species composition, changes in nutritive value of available forage due to retrogression or plant•succession, or alienation from traditional ranges. This criticism is also applicable to the approach taken by Gross (op_.. cit.) in analyzing for or quantitatively measuring the existence of competition. Milne (op_., cit.) believes that the forms and results (of competition) are irrelevant to its definition, hence he propounds a strict definition for animal competition along the same lines as Clements and Shelford (1939). "Competition is the endeavour of two (or more) animals to gain the same particular thing, or to gain the measure each wants from the supply of a thing when that supply is not sufficient for both (or a l l ) " . Miller (1967) allows more latitude in the definition, "biological competition is the active demand .... for a common resource or.requirement that is actually or potentially limiting". Competition interpreted from the habitat approach involves 39 understanding and measuring factors affecting the cause and severity of competition, rather than observing the reusIts of competition upon the animal population. Smith and Julander (1953) differentiated two forms of habitat competition. The f i r s t "land use competition" they associated with common use of forage species which were adequate to meet the .require-ments of both populations, but where the reduction of one population would permit an increase in the numbers of the other.. The second "forage competition", occurs when the supply of a forage species used in common is inadequate. They stated further that i f the combined use of a forage species does not go beyond the point which is considered "proper", there is no competition, because up to that point there is forage available for each ungulate population to take as i t chooses. Cole (1958 cited in Capp 1968) l i s t s four components necessary for competition; the fourth of which is "The forage plants being used are in a limited supply or are deteriorating in production as a result of combined use". This is an example of Miller's (1967) "potentially limiting" competition factor. Similarly '-Sugden (1961) defines competition in "actual" or "potential" terms in relation to'the habitat. He stated that "competition for forage occurs between individuals of the same species or between different species whenever there is insufficient food or whenever the soil or plants are damaged, with subsequent reduction of available forage". He explained that "in the latter case, animals may not suffer harmful effects of competition until-a year or more after i t occurs". Another more subtle form of land use competition is that possibly resulting from range "fouling"'or "contamination" by livestock on an 40 area prior to f a l l and/or winter game use. Few references refer to this form of competition. Sugden (1961) .reported that overwintering bighorns prefer bunchgrass plants possessing ungrazed stalks after cattle use. He also mentions the . possibility that wild sheep do not tolerate "contamination" of the range by domestic sheep. Skovlin et a l . (1968) stated that even light cattle use on the Starkey Experimental Forest and Range, Oregon, affected elk use in the area as much as moderate or heavy use did; deer use however, was not affected. The effect on elk distribution was attributed to the presence or absence of ungrazed forage (Skovlin and Harris, 1970)., Range "fouling" generally does not appear to be a simple cause and effect relationship on range use. Several references indicate that i t is the effect of grazing on species composition, or on the quality of the standing crop of range forage that influences subsequent use. Farrell (pers. comm.) found that elk, as shown by distribution patterns in Grant County, Oregon, preferred to use range maintained in a grazed or serai condition by cattle.. This is in opposition to the results, of Skovlin et a l . (1969), but may be explained by differences in i n i t i a l and/or grazing induced species composition of the range communities involved. Deer also seem to prefer range areas caused to be i n , or maintained in, a forb serai stage by livestock (Julander, 1962; Lesperance et a l . , 1970). Cook et a l . (1953) in Utah have shown that forage remaining after grazing is reduced in nutritive value, and that the nutritive value is further decreased as grazing intensity is increased, hence possibly affecting subsequent grazing by game in the area. 41 In context with the above discussion the present study inves-tigated factors contributing.to range (forage) competition. Several authors have listed factors pertinent to an evaluation of range compet-ition. Stoddard and Rasmussen (1945) calculated competition by evaluating differences both in areas used and in food habits. Smith and Julander (1953) proposed that the supply and utilization of forages in "key areas" (important feed-grounds for both species in question) should be evaluated in relation to the "proper" degree of utilization, and secondly that the extent of overlapping use on preferred species should be studied. Julander (1958) stated that, where forage is limiting, forage competition could be evaluated by studying two contributing factors: 1. The extent to which game.and livestock graze the same area, and 2. The extent to which they prefer the same species. Buechner (1960) lis t s information necessary for a precise and.detailed analysis of competition: 1. data on the diets of the species involved, 2. numbers of each kind of animal, 3. comprehensive and statistically adequate sampling of the vegetation, 4. knowledge of seasonal and annual,variation in-forage production, 5. the distributional habits of the animals, and 6. the past history of the area with special reference to grazing and fire. Range competition exists in a variety of patterns, each requiring a different approach to evaluate its presence and severity. Most com-petition studies are conducted on winter ranges, because as stated by Stoddart and Rasmussen (1945), "the winter range almost everywhere in the west is the area of limited, feed for game". In a review of 42 game-range literature Capp (1968) summarized that "the c r i t i c a l period, or period when competition is most active is during winter". Julander et a l . (1961), and Jones et al . (1956) however, have shown the importance of quality of summer range to the reproductive proficiency of ungulate populations, and hence the importance of competition on summer ranges. The simplest competition pattern is intraspecies competition in game or livestock areas grazed by a single ungulate species. In this case i t is a relatively simple process to measure and delineate areas receiving excessive use. The approach and investigational techniques become more involved when two species utilize the same range. Here one has to evaluate interspecific or combined intraspecific and inter-specific competition. In this case several patterns of competition may exist, some more difficult to evaluate than others. Difficulties arise with the occurrence of overlap in seasonal range-use and whether i t is domestic and wildlife species or only wildlife species being studied. The simple two-species competition patterns occurs when livestock occupy the area during one•season and game species in another, i.e. a sequential pattern. Examples of competition studies with this pattern are Julander (1955, 1958), Sugden (1961, p. 35), Skovlin et a l . (1968), Dasman (1949). A more complicated pattern occurs when the season of range use overlaps forming a pattern of synchronous use. Dorn (1970) for example, was not able to collect data when moose and cattle were in the area at the same time.; Due to synchronous range use, Lesperance et a l . (1970) studied only food habits and hence "potential dietary competition" between mule deer and cattle in Nevada. In Utah, Smith and Julander (1953) were able to measure forage utilization between 43 domestic sheep and mule deer, during synchronous range use, by con-trolling sheep movements. Buechner (1947) studied only food habits to evaluate competition between cattle and antelope in western Texas. However, as mentioned previously, competition in a synchronous range use pattern may be alleviated, especially between wildlife species, by utilization of separate habitat types or ecological niches within the same range area. In this case the measurement of competition factors is much less complicated. Sugden (1961) observed this at Churn Creek and, especially, at Riske Creek with mule deer and bighorn on their winter range. Cooperrider (1969) in Montana made similar observations. Evaluation of competition between three or more species can be very difficult due to the occurrence of synchronous range use. In such cases i t is almost impossible to conduct a complete evaluation of com-petition for each species as outlined by Buechner (1960). Examples of the problems encountered are illustrated by the following studies: Morris and Schwarty (1957) in studying competition between elk, mule deer and bison found that they could use only rumen analysis to study forage consumption because of concurrent range use. Mackie (1970) in a competition study between mule deer, elk and cattle on the Missouri River Breaks was only able to' measure animal distribution, and the food habits of each ungulate class. He was not able to quantitatively determine forage utilization by cattle and/or elk. Cowan (1947) like Mackie (op_. cit.) did not quantitatively determine utilization by moose, deer, elk and bighorn in his evaluation of range competition in Jasper Park. 44 3.4 Review of bighorn sheep-mule deer-cattle range relations Comprehensive reviews of literature pertinent to ungulate range utilization and competition have been published by Buechner (1960) and Capp (1968). A review of the literature indicates that studies of bighorn-cattle range relations are fewer than those of deer-cattle and deer-bighorn range relations. In the United States Buechner (ibid) stated that on bighorn winter ranges in Colorado, Montana and Wyoming bighorn populations are suffering as a result of changes in range condition induced by domestic stock grazing. Basically this is true on many Canadian bighorn winter ranges as well. Sugden (1961) studying California bighorn winter ranges at Churn Creek and Riske Creek, B.C., found indications of competition. Excessive forage utilization by cattle on the Chum Creek bighorn winter ranges indicated intraspecific competition.• However, competition with bighorn was not considered to be significant due to the,small number of sheep involved, and the negligible amount of sheep grazing occurring on areas of excessive cattle use. Although forage competition was not considered significant, "land use competition" was, in that the area could support five times as many wintering sheep in the absence of cattle grazing. In the Ashnola Blood (1961) found only slight forage competition in the summer of 1960. Summer cattle grazing on South Slope resulted in forage utilization, ranging from proper to excessive, on only 20% of this major wintering range. Competition was further alleviated since winter sheep distribution only marginally overlapped the areas of intensive cattle use. However, Blood (ibid) states that in 45 the past cattle grazed South Slope in the spring or summer and again in the f a l l . This more intensive use would probably force cattle to utilize forage on areas important for overwintering bighorn. Demarchi (1965) concluded that summer use by cattle caused severe forage compet-ition on the South Slope winter range, and that the quantity as well as the quality of available forage was limting the bighorn population. Relatively few studies have been conducted on deer-cattle range relations; the majority of these have concentrated on the results of summer and f a l l foraging by cattle on deer winter ranges. Smith and Doell (1968) found early cattle grazing beneficial because the removal of herbaceous forage allowed increased bitter brush production on Utah winter ranges. However, they stated that competition could occur i f cattle use was allowed in late summer or f a l l , as cattle consumed browse at that time. Similarly, Lesperance et al. (1970) found that potential dietary competition occurred when cattle were forced to consume signif-icant amounts of browse as a result of insufficient herbaceous forage. They stated that (1) dual range use-was necessary because the elimination of livestock from western ranges would eventually reduce deer populations, and (2) they observed that in Nevada and Utah, land misused by cattle tends to create range conditions advantageous for deer populations through changes.in species composition in favor of browse. This observation may be true to a point, but one must realize that prolonged range misuse w i l l result in reduced forage productivity and hence reduced secondary productivity. This problem was.observed by Julander (1962) on winter-spring ranges in Utah. Julander (1955) discussed factors affecting deer-cattle range distribution and forage competition on the summer and 46 winter ranges of a range unit in Utah. • Range relations between deer, elk and two systems and three intensities of cattle stocking were studied in.Northeastern Oregon by Skovlin et al . (1968). They observed no significant effect on deer range use patterns as a result of cattle stocking, although deer tended to use grasslands more where cattle stocking was heavier. They suggested conservative rather than heavy cattle stocking to inaximize productivity on the ranges, even though no direct game competition was observed. Cattle-deer summer range relations were studied by Mackie (1970) in Montana. He observed that areas of spring deer use coincided with range affected by past cattle use. Deer preference for serai stage grassland communities in the spring was also observed in the Ashnola by Demarchi (1965). Many studies of competition for forage between mule deer and bighorn sheep have found l i t t l e i f any competition of,'; significance between these two species. Such"studies include: Cowan, (1947) in Jasper National Park; Flook (1964) in Banff and Jasper National Parks; Sugden (1961) studying California bighorn in B.C. stated that competition was not observed at Riske Creek nor Churn Creek. However,, in the future, competition may be observed at Churn Creek i f either the sheep or deer populations were to increase. Competition may then occur as a result of deer use of pasture sage, an important sheep winter forage. Cooperrider (1969), working in Montana, found l i t t l e competition for grass but observed that "deer use browse species which in small quantities may be importnat in the winter diet of sheep". In the Ashnola Valley, B.C., both Blood (1961) and Demarchi (1965) found l i t t l e evidence of competition on the winter ranges but stated that early spring range use by deer may 47 be in part responsible for maintaining the retrogressive condition of the Ashnola bighorn winter ranges. Sheppard (1960) and Wishart (1958), cited in Sheppard (1960), found l i t t l e evidence of competition in Alberta except-in the spring when "both were feeding on new grass, the supply of which i s , at f i r s t , limited". Sheppard (op_. cit.) believed that spring forage competition may have been severe enough to be responsible for the death of some animals. Couey (1955) believed that certain of the Montana bighorn,herds were encountering competition from mule deer.on restricted winter ranges. Competition for forage between bighorn and mule deer appears to have been avoided in most studies by differential habitat preferences and/or differences in winter food habits. Where competition was observed, preferred winter deer forage was limited in availability, hence forcing deer to consume a larger volume of grass important for overwintering sheep. Buechner (1960) concluded that where vegetation has deteriorated and high density mule deer popula-tions exist on bighorn winter ranges, efforts should be made to reduce the deer herds to lessen the chance of deer-sheep forage competition occurring. In.summary, i t appears that forage utilization by deer and sheep generally has l i t t l e effect on forage available for cattle use, as a result of differences in food habits or range areas preferred. However, cattle management can and does affect the availability of forage for game species. Many authors have suggested that dual range use is not only more productive in terms of•secondary productivity but.that i t also allows more efficient use of the range. This is true provided that the total effect of combined use is not detrimental to the range. 48 4. METHODS AND PROCEDURES Methods of a practical nature were employed to obtain data for the evaluation of synchronous range utilization by three ungulate species. Time was limiting and, in some cases, sampling would not satisfy rigid statistical demands. 4.1 Food habit analysis In addition to the winter, spring and f a l l food habits of mule deer, the spring food habits of bighorn and cattle were concurrently investigated on the same ranges. The food habits of mule deer and bighorn sheep were determined by a combination of techniques; the analysis of ingested forage, and observations of feeding animals and feeding sites. Cattle food habits were determined only by rumen sample analysis, as cattle grazing on the Flatiron and Crater Mountain portions of the Ashnola range was terminated following the 1968 grazing season. The techniques used were similar to those described by Saunders (1955), Wilkins (1957), Blood (1961) and.Demarchi (1965). In presenting the food habrhdata the chamaephyte, pasture sage (Artemesia frigida) and buckwheat (Eriogonum..heracleoides), were classified as browse species. The native ungulates consumed both species as browse during periods of the year when only the woody portions of the plants were available. 4.1.1 Feeding observations Observations on foraging deer and sheep were recorded to sub-stantiate and expand information on food habits obtained by more 49 detailed methods. The observational techniques employed included: 1. direct observation on browsing deer with the aid of binoculars, and 2. the examination of vegetation selected by grazing animals. This was conducted by back-tracking on.the trails of deer or sheep, when fresh snow covered the ground to a depth of not .more than a few inches. Utilization of one plant was recorded as one observation. Signs of browsing on forage species were recorded when encountered in "deer only" or "sheep only" areas after cattle grazing was terminated on the Flat-iron ranges. 4.1.2 Grazed plant loop-transects Grazed plant loop-transects were conducted to substantiate spring food habit information obtained from analysis of deer and big-horn rumen samples. The transects were made on the grasslands used in common by deer and sheep during the spring.• Transects were subjectively located, following the departure of ungulates from the Flatiron Mountain winter-spring ranges in 1969 in areas observed to have been used by deer only or sheep only. Sites used only by deer in the Artemesia  frigida - Stipa columbiana and Poa pratensis dominated communities on Juniper Slope were studied from Jule 13 to 15. Sites grazed only by sheep in the Agropyron spicatum-Artemesia frigida and Agropyron  spicatum dominated communities on South Slope were studied from July 5 to 7. Three 100 foot transects were located in each, community and observations, in a 4.5 inch diameter loop, were made at one foot intervals along a steel tape. Data recorded at each observation point included the species present, their percent coverage and the presence 50 or absence of grazing on each species. The presence of a species or forage class , either grass, forb or browse, was considered to be a single observation in each loop, even though many individuals of the species or representatives of a forage class may have been present. From the transect data, frequency of utilization on forage classes and individual taxon was observed in relation to abundance. Frequency of use values were computed two ways. That i s , grazing on a given species or forage class as a percentage of: 1. the occurrence for that species or forage class, and 2. the total grazing observed in the community. In addition, botanical composition of the range was described by the frequency and cover data obtained from the transects, as outlined by Brown (1954) and by M.A.S.-N.R.C. (1962). 4.1.3 Rumen sample analysis Rumen samples were collected from mule deer in the winter, spring and f a l l for quantitative botanical analysis. Bighorn and cattle samples were collected.for food habit.analysis during the spring of 1968 because range'use-overlap between the three ungulate species was observed only during this .season. Dietary competition during the spring was investigated by both rumen sample analysis and grazed plant survey techniques. One to three mule deer were collected each month for a l l months that deer utilized the winter-spring ranges on Flatiron Mountain. Additional mule deer samples were obtained from hunter and winter k i l l s . Bighorn sheep were collected on South Slope during the spring only, 51 because winter food habits had been investigated previously (Blood, 1961; Demarchi, 1965). However, one winter-kill sample from Juniper slope was analyzed. Investigation of the spring food habits of Hereford cattle was made possible through the purchase of two short yearlings, a steer and a heifer, from the Flatiron Mountain grazing permit holder, Rimson Equipment .Company. The study animals were allowed to graze normally with the rest of the herd on Flatiron Mountain. They were slaughtered while grazing on the range, one on Juniper Slope, the other on Juniper Top in June, 1968. The rumen sample analyses employed modifications of those discussed by Medin, 1970. Rumen•samples of one.litre volume were collected from several parts.of' the less digested portion of the rumen contents. Samples taken in spring were preserved in the field by addition of 10% formalin and then frozen for storage. Winter samples were allowed to freeze in the field and were stored frozen. A one pint sub-sample was chosen randomly for botanical analysis. Partial segregation of the sample was achieved by washing the material through a series of four sieves: No. V' (6.35 mm); No. 3Jg (5.66 mm); No. 5 (3.96 mm); No. 8 (2.0 mm). The identifiable material remaining on the f i r s t three sieves was combined. The unsegregated residue was combined with the material remaining on the fine sieve and composed the unidentifiable "fiber" portion of the sample. Plant particles of sufficient size to allow identification were segregated, then oven dried and weighed by genus and species. Particles that were not identified to species could usually be placed in one of three forage class categories: grass, forb or browse. It . 52 was not possible, using standard techniques and reasonable time expenditure, to separate even to generic level the various grasses occurring in the rumen samples. To' facilitate species identification a representative plant collection was made and identified; in addition the collection of E. Scheffler was made available. The- percent by weight of each taxon and each forage class was calculated as a percentage of only the total segregated material. This was done on a seasonal basis using the "aggregate" method (Martin et a l . , 1946), modified to a weight basis. Food habits were thus calculated for 3 seasons: winter, spring and f a l l . The unidentifiable residual fiber material for each sample was dried and weighed. The unidentifiable fibrous portion is presented as a percentage, for each season, of the aggregate total weight of a l l samples analyzed from a given season. 4.1.4 Forage preference index A forage class preference index was .calculated from the analysis of the spring rumen samples of deer, sheep and cattle and data on the availability and use of spring forage determined from the grazed plant loop-transects. The following formula was employed: Forage class % of forage class in rumens x % frequency of use on preference = the forage class  index Forage class availability where the "forage class availability" factor % canopy cover x % frequency of occurrence 100 53 This factor is similar to the "forage quantity" value of Buechner (1960), which is calculated from cover x frequency index. Spring forage preference for individual species within a forage class, for deer only and sheep only, was determined by analyzing the grazed plant loop-transect data. These transects were conducted in deer only and sheep only grazing areas,. in the absence of cattle grazing in the spring of 1969. The following formula was employed: Species preference = % frequency of grazing on a species ; L n^ e x species availability where the "species availability" = % canopy cover x % frequency of occurrence factor 100 1.2 Range forage appraisal 4.2.1 Site Selection Reasonably homogeneous sites were chosen for evaluating forage production and utilization after general ungulate distribution patterns were determined. Within the sites, plots were established in the summer of 1968 in areas grazed by a single ungulate.species as determined by observed range use during the preceding winter and spring. On these plots forage utilization by a given ungulate species, forage production and the effect of grazing on productivity were measured. As • cattle grazing, however, was terminated on Flatiron Mountain after the 1968 grazing season, new plots were established prior to spring growth in 1969 to take advantage of the absence of cattle on the wildlife 54 spring ranges. Three of the four plots were s t i l l established in areas used by.a single ungulate species. Two deer-use plots were located on Juniper Slope in the Poa pratensis and Artemesia-Stipa communities. On South Slope a sheep-use plot was located in the upper portion of "sheep basin", and a deer and sheep use plot was located in the lower portion of the basin; both in an Agropyron spicatum community. Cattle use of the range in the f a l l of 1968 was measured in an Agropyron-Stipa community on the lower portion of South Slope, in a Bromus-Stipa community on the flats of "sheep basin", and in a Poa  pratensis community on Juniper Slope. Production,and utilization of alpine forage on deer and sheep summer range was measured at'two sites near joe Lake in 1969, using fenced exclosure plots. During the winter of 1968-69 forage use by sheep was measured in the Agropyron spicatum community in sheep basin, South Slope, and in association with f a l l cattle use of the Poa pratensis community on Juniper Slope. Over-winter forage losses due to forage "weathering and shattering" were measured at both sites. In addition, total forage losses during the winter of 1969-70 were measured at both deer use study sites on Juniper Slope. 4.2.2 Forage measurement Forage production and forage utilization were measured by hand clipping, to ground level, the total forage present in eight one-meter-square quatrats per treatment (Figure 4.2.1). Sample quadrats were clipped and separated into forage classes, viz. grass, forb and browse, Figure 4.2.1. One-meter square quadrat and hand clippers used i n measurement of forage production and u t i l i z a t i o n . ma. 11 Figure 4 .2.2. Fenced plot on the Poa pratensis community, Juniper Slope. 56 but this was discontinued due to the high between-sample-variance observed. Quadrats at all- sites were laid out and marked with spikes using a square meter frame...They were spaced one foot apart to avoid common border effects and to facilitate clipping. The samples were air dried in. the fi e l d , oven dried within two weeks and weighed to the nearest tenth gram. Forage use by cattle in the f a l l of 1968 and by sheep during the winter of 1968-69 were measured by "before and after use" forage clippings of unexclosed sites' (Appendix 4.2.1) Weathering losses of forage over the winter were measured by clipping in the f a l l and spring within fenced areas. In 1969, a l l areas used for .forage clipping measurements were preclipped to ground level prior to spring growth to remove the old vegetation. In the alpine plots the old forage was removed, but not to ground level as the,Carex species over-winter in a green condition. Alpine forage production and utilization by deer and sheep were measured at two locations on the summer range by clipping "fenced" and "unfenced" areas, as illustrated in Appendix 4.2.2, in late August, 1969. The fenced plots designed to facilitate measurement of forage production and utilization during 1969 are illustrated in Figure 4.2.2 and Appendix 4.2.3. They were centrally located within one acre game observation areas. Six foot high, seven strand barbed wire fences were erected during the grazing period to provide total ungulate exclosure when desired. This system of plot location and sequential fencing was necessary to evaluate forage use by individual ungulate species and the 57 subsequent effect of that use. The plots were designed to exclude only ungulates, thus forage use by invertebrates and rodents was not greatly influenced and was considered to be an unmeasured but constant factor on a l l quadrats. Field plot design was the same at three sites, but at the fourth, the sheep and deer use plot, plot section "B", was lacking. The gener-alized plot design is shown in Appendix 4.2.3. Section "A" of the plot was fenced at the time of construction as a total exclosure"area. The remainder of the plot was fenced at intervals to permit: 1. measurement of forage growth and forage use by a single ungulate species during a three week spring grazing period (Martin, 1970; Brown, 1954). This was accomplished by clipping the treatment L:,quadrats, erecting a fence around section "B" and then clipping the quadrats for treatments 2 and 3 at the end of the short period. This combines "before" and "after" grazing and "grazed" and "ungrazed" area clipping techniques (Shepherd, 1962). Animal numbers in the one acre "macroplot" surrounding the exclosures were observed at intervals during the day to assess forage use per animal; This was attempted so that forage use data obtained from areas of single ungulate species grazing could be utilized to calculate forage use on range areas synchronously used by more than one ungulate species. 2. measurement of forage use during the entire spring grazing period. Quadrats for treatments 4 and 5 were clipped and area "C" was fenced when the deer and/or sheep departed from the winter spring ranges. 3. measurement of the effect of spring grazing on annual forage production. Treatments 6 and 7 were undertaken at the time of l 58 maximum forage production as indicated by soil moisture depletion. (Harper, 1969, p. 33). 4. calculation of forage carry-over in the f a l l , measured by clipping treatment 8. The percent forage.loss from time of maximum forage production t i l l late f a l l was also calculated. Cattle grazing was suddenly' and unexpectedly terminated on Flatiron and Crater Mountains, as mentioned earlier, after the 1968 grazing season when the British Columbia Fish and Wildlife Branch purchased the grazing permits. As a result, i t was no longer possible to measure.forage use by cattle or to measure the effect of use on forage production in the study area as originally planned. In .place of this a forage clipping study was initiated to measure the effect of various cattle turn out dates on forage production in the Poa pratensis community on Juniper Slope. 4.2.3 Community description Two 100 foot line-point transects were laid out at each exclosure site to describe the botanical composition and to measure the frequency contribution of each major grass, forb and shrub to the forage production and utilization measurements. Observations were taken at six inch intervals along the transects by vertically lowering a pin to ground level. As the pin descended through the forage canopy the last "aerial hit" and "basal hit" were recorded. In the event that the observation point f e l l in interstices between the canopy, observations were recorded as a "non-vegetative hit" and the basal observation as one of l i t t e r , s o i l , rock, moss or lichen. Plants were collected and identified for both the winter-spring and summer ranges; identifications were verified by 59 Dr. V.C. Brink. Hitchcock et a l . (1969) was used as the taxonomic authority for a l l species, except for grasses where Hitchcock.(1950) was employed. 4.2.4 Total nitrogen analysis Total nitrogen analyses were conducted on f a l l clipped forage from the two alpine communities. The macro-Kjeldahl technique was modified to include.nitrates (Chapman and Pratt, 1961; p. 150-152). 4.3 Data analysis Field data are presented in appendices, whereas summarized data are presented in tables in the text.• The data have been statistically analyzed, with the assistance of Dr. A. Kozak and Dr. G.W. Eaton, using the IBM 360 computer and programs located at the University of British Columbia. 4.3.1 Food habits Forage class consumption values, on an oven dry weight basis, for cattle, sheep and deer were obtained from individual rumen sample analysis results using the aggregate.technique of Martin et a l . (1946), modified-to a weight basis. Differences in the forage class weights from rumens of mule deer were tested for significance within and between seasons using the multi-factor analysis of variance (MFAV) program which incor-porates Duncan's New Multiple Range test. Variation within and among forage class use in the spring by the three ungulate species was similarly analyzed. 60 4.3.2 Forage production and utilization Eight replicates of one meter-square quadrats were clipped per treatment, and the dates of forage clipping were considered to be the treatments for the analyses. Differences in oven dry weights of the total forage clipped from meter square quadrats at each treatment date were tested for sig-nificance with the MFAV analysis of variance program. The field designs used to collect.the forage data included a randomized split plot design on the "before" and "after" f a l l cattle and over-winter sheep grazing clippings : and a partially randomized plot design on the "fenced" and "unfenced" deer and sheep spring and summer grazing studies. 61 5. OBSERVATIONS AND RESULTS 5.1 Food Habits 5.1.1 Cattle spring food habits Analytical results of rumen samples collected in June, 1968, are presented in grams in Appendix 5.1.1, and in percent in Table 5.1.1. Grasses comprised 97.5% of the identifiable material in the samples, forbs 1.7% and browse species only 0.8%. Consumption of grass was found to be significantly greater (p < 0.01) than that of forbs and browse. (Table 5.1.2). It was found that from the total rumen subsample mass analyzed, only 30.7% of the material was identifiable to species or forage class, while the remainder, 69.3%, consisted of masticated and partially digested unidentifiable grass-like fibrous material. The three forage classes were found in both samples. Only two species of forbs were identified, and consumption of forbs was small. Spring use of browse by cattle in the Ashnola appears to be even less important than that of forbs and consisted of the ingestion of dead pine needles, probably picked up while grazing. Table 5.1.1. Cattle, bighorn and mule deer spring food habits. Relative proportions by oven dry weight, expressed as a percentage", for each taxon and forage class from rumen sample analysis. Ungulate species Sample size Cattle Bighorn Deer 2 3 11 % " Freq. Rank % Freq. Rank % Freq. Ran} GRASS 1. Mature 19.2 2 2 46.2 3 1 17.3 11 2 2. Green 78.3 2 1 44.8 3 2 20.7 11 1 Subtotals 97.5 2 1 91.0 3 1 38.0 11 •2 FORBS 1. Achillea millefolium Tr 1 2 0.7 1 4 Tr 3 6 2. Allium cernuum - - - - - 0.2 1 5 3. Antennaria sp. - - - - - 0.9 5 3 4. Arenaria capillaris - - - - - 0.2 1 5 5. Erigeron compositus - - - - - Tr 2 6 6. 'Geranium viscosissium - - - - - - Tr 1 6 7. Heuchera cylindricata - - - - - 0.5 4 4 8. Lupinus (sericeus) - - 2.1 1 2 - - -9. Microserius alpestris - - - - - Tr 1 6 10. Oxytropus campestris - - - - - 1.2 3 2 .11. Penstemon (procerus) - - - - - 0.5 2 . . 4 12. Phacelia (hastata) - - 1.4 1 3 - - -13-. Potentilla gracilis - - • - - - 0.9 4 3 14. Ranunculus sp. - - - - - 0.2 4 5 15. Sisymbrium altissimum - - - - - - 0.2 1 5 16. Stellaria sp. Tr 2 6 "Percentages based on weight of identified material only Table 5.1.1., continued. Ungulate species Cattle Bighorn Deer Sample size 2 3 11 % Freq. Rank % Freq. Rank % Freq. Rani 17. Taraxicum officinale Tr 1 6 18. Viola sp. - - - Tr 1 5 Tr 2 6 19. Zygadenus venenosus Tr . 1 2 - - - ' - - -20. Unknown sp. 1.7 2 1 •2.8 3 1 ' 2.8 9 I Subtotals 1.7 2 7.0 2 7.7 3 TREES AND SHRUBS 1. Arctostaphylos uva-ursi - - - - - - 2.3 4 • 2 2. Artemesia frigida - - - - - - 2.1 5 3 3. Berberis repens - - - - - - Tr 1 7 4. Chrysothamnus nauseosus - - - - - - 2.1 1 3 5. Eriogonum heracleoides - - - 2.1 1 1 1.6 9 4 6. Juniperus scopulorum - - - - . - -' 0.5 1 6 7. Pinus contorta 0.8 1 1 - - - 0.9 2 5 8. Pseudotsuga menzlesii - - - - - - 43.2 11 1 9. Ribes sp. - - - - - 0.9 2 5 10. Unknown sp. Tr 1 2 Tr 1 2 - - -Subtotals 0.8 3 2.1 3 53.8 1 LICHENS - - - - - - 0.5 - 4 TOTALS 100 100 100 UNIDENTIFIED MATERIAL 69.3 2 72.5 3 84.6 11 cr>: G O Table 5.1.2. Statistical analysis of cattle, bighorn and' deer spring food habitats as indicated from rumen sample analysis. Relative proportions by oven dry weight, expressed .as percentages, of identified forage only. Cattle Bighorn Deer Ungulate Species Percent Signifi-cance within species Signifi-cance between species Percent Signifi-cance within species Signifi-cance between species Percent Signifi-cance within species Signifi-cance between species Grass 97.5 N.S. 91.0 N.S. 38.0 N.S. Forbs 1.7 N.S. N.S. 7.0 N.S. N.S. 7.7 ** N.S. Browse 0.8 N.S. N.S. 2.1 N;S. N.S. 53.8 N.S. N.S. Lichens 0.0 t N.S.. 0.0 + N.S. 0.5 + N.S. TOTALS 100 100 100 Sample Size 2 2 11 N.S. = not statistically significant * = statistically significant at .05 level ** = statistically significant at .01 level t = Lichens not included in statistical analysis between forage classes within ungulate species due to lack of homogeneity of variance. 65 5.1.2 Bighorn spring food habits Three bighorn specimens, two ewes and a young ram, were c o l l -ected for food habit study while grazing on South Slope during late May and early June, 1968. In addition, the rumen contents of a winter k i l l found on Juniper Slope was analyzed. Results, in grams oven dry weight, of individual rumen sample analyses are presented in Appendix 5.1.2, while the forage class percentage distribution for the spring diet is presented in Table 5.1.1. Grasses, the most important component of the bighorn's spring diet, comprised 91% of the samples while forbs and browse contributed only 7.0%, and 2.1% respectively. Four species of forbs were identified, the most important of which was silky lupine (Lupinus sericeus) and white leaf phacelia (Phacelia hastata). Each of these species occurred• in only one of the samples (Appendix 5.1.2). Buckwheat, the browse consumed in the spring, occurred in only one of the three samples. Browse formed 10% of the winter rumen sample, made up primarily of buckwheat and pasture sage with some Douglas f i r (Appendix 5.1.2). Statistically analysis indicates that spring u t i l i -zation of grass by sheep, like that of cattle, was significantly greater than forbs or browse (p < 0.01). No 'significant difference between forb or browse utilization was measured (Table 5.1.2). Only 27.5% of the rumen subsample was.identifiable, the other 72.5% consisted primarily of unidentifiable grass-like fibrous material. Bighorn grazing observations, from grazed plant loop-transects, conducted on the Agropyron and Agropyron-Artemesia communities of South Slope are presented in Appendices 5.1.3 and 5.1.4. These data are summarized by forage class in Table 5.1.3 and to the more important species in Tables 5.1.4 and 5.1.5. , Forage class- and species-preference 66 index evaluations are presented in these tables. Average sheep use on South Slope indicated that forage was grazed and "preferred" in relation to its availability. Grass, the most frequently grazed and apparently "preferred" forage class, was the most available. Forbs.and browse, being the second and third most available forage respectively, were the second and third most frequently grazed and "preferred" forage classes. In the Agropyron spicatum community, grass was the most frequently grazed forage class, accounting for 63.6% of a l l observed grazing, but only 32.6% of the grass plants observed were utilized (Appendix 5.1.3). Three of the five grass species occurring on the site were grazed by sheep. Bluebunch wheatgrass was the most abundant species, but was second to Junegrass (Koeleria cristata) in percent frequency of a l l observed grazing, viz. 26% compared to 27.9% for Junegrass. Idaho fescue (Festuca idahoensis) was the most preferred grass species as revealed by the relatively high frequency of use in relation to its low abundance (Table 5.1.4). Forbs were the second most frequently used forage class. Only 24.4% of the forbs observed were grazed,.but use on forbs formed 30.5% of a l l grazing observed in the community. Nine of the sixteen forb species observed were grazed; however, only four were of any importance. Yarrow (Achillea millefolium), silky lupine, wild onion (Allium cernuum) and.timber milkvetch (Astragalus miser) each accounted for 5.8 to 7.8% of a l l observed grazing. Silky lupine, the most abundant forb encountered in rumen samples ranked third on the preference index, after wild onion and timber milkvetch (Table 5.1.4). The consumption of browse by sheep in spring was slight, in that i t formed only 5.9% of a l l observed utilization. Only 20.5% of a l l Table 5.1.3. Forage class frequency of grazing and preference indices for bighorn and deer during the- Spring, 1968„ Ungulate Species Forage Class Percent in Rumens Percent Frequency of Grazing* Communities Forage Class Availability Factor* Communities Forage Class Preference Index Communities 1. Cattle Poa Art Aver. Poa Art Aver. Poa Art Aver. Grass 97.5 22.3 6.1 14.2 Forb 1.7 - - - 3.9 1.1 2.5 - - -Browse 0.8 - - - 0.003** 5.2 2.6 — • _ — Lichens 0 2. Bighorn Ag spic Ag-Art Aver. Ag spic Ag-Art Aver. Ag spic Ag-Art Aver. Grass 91.0 63.6 73.5 68.6 11.5 10.2 •• 10.9 503.3 655.7 572.7 Forb 7.0 30.5 17.0 23.8 3.4 0.6 2.0 62.8 198.3 83.3 Browse 2.1 5.9 9.5 7.7 0.3 1.1 • 0.7 41.3 18.1 23.1 Lichens 0 - - - - - - • - - -3. Deer Poa Art Aver. Poa Art Aver. Poa Art Aver. Grass 38.0 69.4 63.9 • 66.7 22.3 6.1 14.2 118.3 398.1 178.5 Forb 7.7 30.1 19.9 25.0 3.9. 1.1 2.5 59.4 139.3 77.0 Browse 53.8 0.5** 16.4 8.5 • 0.003** 5.2 2.6 8966.7** 169.7 175.9 Lichens 0.5 * Percentage values derived from grazed plant loop-transects conducted on Juniper Slope (Poa and Artemesia communities) in June 1969 for deer and on South Slope (Agropyron and Agropyron-Arternesia communities) in July 1969 for bighorn. ** Inadequate browse observation sample size. cn -J 68 Table 5.1.4. Bighorn sheep forage species preference index as indicated from South Slope, Agropyron community, grazed plant loop-transect data. Availability Frequency Preference Rank Rank Species Factor n °. ,. Index A B Grazing" GRASSES 1. Agropyron spicatum 5.70 26.0 4.6 3 9 2. Festuca idahoensis 0.42 9.7 23.1 1 5 3. Koeleria cristata 1.81 27.9 15.4 2 6 4. Poa secunda 0.01 0 0 0 0 FORBS 1. Achillia millefolium 0.98 7.8 8.0 4 8 2. Allium cernuum 0.01 5.8 617.0 1 1 3. Antennaria rosea 0.05 0 0 0 0 4. Arabis;. sp. 0.01 0 0 0 0 5. Astragalus miser 0.04 5.8 134.9 2 2 6. Lappula redowskii 0.10 0 0 0 0 7. Lupinus sericeus 0.10 7.1 70.5 3 4 BROWSE 1. Artemesia frigida 0.26 2.6 10.1 2 7 2. Eriogonum heracleoides 0.04 3.3 85.3 1 3 * Calculated as the percentage of a l l plants grazed. Rank A = preference rank within a forage class. Rank B = preference rank between forage classes. 69 Table 5.1.5. Bighorn sheep forage species preference index as indicated from South Slope, Agropyron-Artemesia community, grazed plant loop-transect data. July 7, 1969. Availability Frequency preference Rank Rank Species r w c ^ x x x c y f Factor n . Index A B Grazing GRASSES 1. Agropyron spicatum 2... .Festuca idahoensis 3. Koeleria cristata 4. Poa secunda 3.75 34.0 0.0.5 4.1 1.53 29.9 0.01 4.8 9.1 4 . 8 91.1 2 4 19.5 3. 7 363.6 1 2 FORBS 1. Achillia millefolium 0.03 2.0 77.2 3 5 2. Allium cernuum 0.01 5.4 500.0 1 1 3. Antennaria rosea; 0.02 0.7 35.4 4 6 4. Lappula redowskii 0.02 :ti. :o 0 0 5. Lupinus sericeus 0.01 4*1 297.1 2 3 BROWSE 1. Artemesia frigida 1.10 8.2 7.4 70 plants of the two browse species observed were browsed. Buckwheat was preferred as spring browse over pasture sage (Table 5.1.4). In the Agropyron-Artemisia community on South Slope, grass was again the most frequently used and available forage class (Table 5.1.3). It formed 73.5% of a l l the grazing observed in this community. At the same time the observed frequency of use was greater, 43.7% (Appendix 5.1.4). Five of the six species were.grazed. Bluebunch wheatgrass was . s t i l l the most.abundant species and in this community the most, frequently grazed, accounting for 34% of a l l grazing. Junegrass ranked second in utilization. However, Sandberg bluegrass (Poa secunda) was the most "preferred" species encountered and Idaho fescue ranked second (Table 5.1.5). Forbs were again the second most frequently used forage class. In this community, the observed use was reduced to 17.0%, but the frequency of utilization of. forbs increased to 37.9%. This may be accounted for by the relatively low abundance of forbs in this community (Table 5.1.3). Seven of the twelve species encountered were grazed, but only two, wild onion and silky lupine contributed frequently to the total observed grazing (Table 5.1.5). They ranked f i r s t and second respectively in the forb preference index. In this community the increased availability of the two browse species is reflected in the greater frequency of browse use (Table 5.1.3). Use of browse formed 9.5% of a l l grazing observed, and 21.5% of a l l plants of the two browse species observed were browsed. It appears that buckwheat was again the preferred species, but sufficient observations were not obtained to, confirm this. However, in terms of availability and amount contributed to the spring diet of bighorn in the Ashnola, pasture sage is the more important species. Sheep use of pasture sage in this 71 community was the highest observed in the spring; i t formed 8.2% of a l l grazing with 20% of the plants observed grazed. In November and December, 1968, the feed of bighorn trailed in the shallow snow pack was, by .number of.observations: kinnikinnick 4; Vaccinium sp. 1; buckwheat 1; pasture sage 2 ; pine 1; chickweed 1; lupine 1; fescue 3; bluegrass 16; pinegrass 24; bluebunch wheat-grass 52. In April, 1969, as a result of f a l l and spring use by sheep (winter use was precluded due to the snowpack) bluebunch wheatgrass in Sheep Basin, South Slope, was grazed to a stubble height of 4 to 7 inches. During the summer of 1969 a significant amount of bluebunch wheatgrass and fescue on the ridges in and around the basin on South Slope was consumed by the herd of bighorn summering on the winter-spring ranges. 5.1.3 Mule deer seasonal food habits A total of 22 deer- rumen samples were collected and analyzed from Starvation Slope, Juniper Slope, South Slope and the adjacent forest ranges, from January to June, 1968, and September to December, 1968. Two of the samples, included in the above, were from a winter k i l l (no. 69-1) and a f a l l hunter k i l l (No. 69-5) in 1969. The analyses of the individual rumen samples are presented in Appendices 5.1.5, 5.1.6 and 5.1.7, on the basis of three arbitrarily delineated seasons. The " f a l l " sampling period October and November began with the arrival of the occasional deer on the ranges and termin-ated with the arrival of the main herd. The "winter" sampling period began in December and continued until the growth-of new herbaceous 72 forage was initiated in the' spring, viz. December through February. The "spring" sampling period began after this time and continued until the deer migrated from the spring ranges, viz. March through June. Data obtained for the three seasons are summarized on the basis of a l l identified taxa within the three forage classes, in Table 5.1.6. 5.1.3.1 Fall diet The f a l l diet of mule deer was determined from three animals collected on the north side of Joe Camp Ridge. Browse species pre-dominated, forming 59.6% of the material identified, while grass.and forbs contributed 17% and 23.4% respectively. Only 15.6% of the rumen sample material was identifiable; the other 84.4% consisted of finely masticated and digested unidentifiable fibrous.material. Analysis of the data presented in Appendix 5.1.5 indicates that there were no stat-istical l y significant differences between amounts of browse, forbs and grass use by deer in the f a l l (Table 5.1.7). Use was recorded on nine species of forbs, the most important of which in order of weight observed are: lupine (Lupinus sp.) , yarrow and clover (Trifolium repens). In the browse class use of seven species including lodgepole pine (Pinus contorta), kinnikinnick (Arctostaphylos  uva-ursi) and sagebrush (Arternesia tridentata) was observed (Table 5.1.6). Sagebrush was present in each of the f a l l deer samples, whereas the other two species were present in only two out of the three samples. Use of grass at this time included some succulent f a l l regrowth. Table 5.1.6. Seasonal mule deer food habits. Relative proportions by oven dry weights, expressed as a percentage*, for each taxon -and forage class from rumen sample analysis. Season Sample size Fall Freq. Winter Rank Freq. Rank Spring 11 Freq. Rank 1. 2. GRASS Mature Green Subtotals 17.0 12.9 17.3 20.7 38.0 11 11 11 2 1 2 FORBS 1. Achillea millefolium 2. Allium cernuum 3. Antennaria sp. 4. Arenaria capillaris 5. Aster sp. 6. Capsella bursa-pastoris 7. Equisetum arvense 8. Erigeron compositus 9. Geranium viscosissimum 10. Heuchera cylindricata 11. Lupinus sp. 12. Microserus alpestris 13. Oxytropus campestris 14. Penstemon procerus 15. Potentilla gracilis 16. Ranunculus sp. 17. Silene sp. 18. Sisymbrium altissimum 19.. Stellaria sp. 2.9 1.-2 0.6 0.6 6.4 0.6 Tr. 1 1 1 2 1 1 4 5 5 1 0.2. 7.8 4.-4 1 3 4 1 1.5 Tr 0.2 0.9 0.2 Tr Tr 0.5 Tr 1.2 .0.5 0.9 0.2 0.2 Tr 3 1 5 1 2 1 4 1 .'3 2' 4 4 1 2 4 2 4 1 3 2 4 -Percentages based on weight of identified material only Table 5.1.6. , continued. Season Fall Winter Spring Sample size 3 8 11. % Freq. Rank Freq. Rank % Freq. Rank 20. Taraxicum officinale • 0.6 1 5 Tr 1 21. Trifolium (repens) 1.6 1 3 - - - - - -22. Viola sp. - - - - - - . Tr 2 -23. Unknown forb sp. 8.8 3 - 5.8 8 - 2.8 9 -Subtotals 23.4 2 19.7 2 7.7 3 TREES AND SHRUBS 1. Arctostaphylos uva-ursi 14.0 2 2 - - - 2.3 4 2 2. Artemesia frigida - - - 0.8 5 4 2.1 '.. 5 • 3 3. Artemesia tridentata 12.3 3 3 - - - — — — 4. Berberis repens - - - - - - Tr ' • 1 -5. Chrysothamnus nauseosus • - - - - - 2.1 1 3 6. Eriogonum heracleoides 1.8 1 . 4 8.3 4 2 1.6 9 4 7. Juniperus scopulorum - - - - - - 0.5 1 6 8. Pinus contortus 22.8 2 1 3.9 2 3 0.9 2 • 5 9. Pseudotsuga menziesii 0.6 1 ' 5 52.9 6 1 43.2 ' 11 1 10. Ribes sp. - - - - - - 0.9 2 5 11. Salix sp. 1.-8 •2 4, - - - - - -12. Shepherdia canadensis - - - 0.2 1 5 . ; • — - — 13. Vaccinium scoparium 1.8 2 4 - - - - - -14. Unknown browse sp. 4.7 2 - 0.5 2 - - - -Subtotals 59.6 " 1 66.8 1 53.8 1 LICHENS - - - 0.7 - 4 0.5- - 4 TOTALS 100 100 100 UNIDENTIFIED MATERIAL 84.4 3 - 61.6 8 - 84.6 11 . -Table 5.1.7. Statistical analysis of mule deer seasonal food habits as indicated from rumen sample analysis. Relative proportions of the forage classes are expressed as percentages, from identified forage only. Season . Fall Significance Significance Percent within between species species Winter  Significance Significance Percent within between species species Spring  Significance Significance Percent within between species species Grass 17.0 Forbs 23.H Browse 59.6 Lichens 0.0 TOTALS 100 N.S. N.S. N.S. t N.S. N.S. N.S. N.S.. 12.9 19.7 66.8 0.7 100 N.S. N.S. N.S. N.S. N.S. N.S. 38.0 7.7 53.8 0.5 100 N.S. N.S. t N.S. N.S. N.S. Sample Size 11 N.S. = not significantly different " = statistically significant at .05 level ** = statistically significant at .01 level t = Lichens not included in statistical analysis between forage classes within seasons due to lack.of• homogeneity of variance with other forage classes. cn 76 5.1.3.2 Winter diet The "winter" deer diet was determined from eight samples: two from the open lodgepole pine forest at the east end of Juniper Slope in December, three in the Douglas f i r forest between Juniper and.Star-vation Slopes, one in December and two in January, and three in the open Douglas f i r forest forming the western boundary of South Slope in February. Use of browse species increased from 59.6% in the f a l l to 66.8% of the winter diet, while forbs and grasses contributed 19.7 and 12.9% respectively. Lichens formed the remaining 0.7%. Of the rumen sample material analyzed, 38.4-% was identifiable; the-balance, 61.6%, consisted of unidentifiable fibrous material, i.e. considerably less than that found in the f a l l samples. Statistical .analysis, of the individual rumen sample data presented in Appendix 5.1.6, indicated that there was.significantly more use of browse (p < 0.01), than of forbs or grass (Table 5.1.7). Use of only four forb species, the most important of which was pussytoes (Antennaria sp.), was recorded from the deer collected to the west of South Slope. Shepherd's purse (Capsella bursa-pastoris) was found in one deer collected in the forest between Juniper and Starvation Slopes. In the browse class, use of five species was recorded. The most important wasrtdpsyan'd?"needp.es.--pf^ Bpuglas f i r , often boughs or bough tips blown to the ground by winter storms. Douglas f i r was identified in six of the eight deer sampled and formed 52.9% of the winter diet (Table 5.1.6). The second most important browse.species, found in four of the samples, was buckwheat. It should also be noted that pasture sage, although not plentiful, was in five of the samples. 77 Deer utilizing the last two species were those collected in the open forest to the west of South Slope and one of those collected in the forest between Juniper and Starvation Slopes. The use of lodgepole pine by deer collected in the pine forest to the east of Juniper Slope, and of soapberry (Shepherdia canadensis) by one of the deer collected between Juniper and Starvation Slopes is unusual. Supplementary information on winter food habits was obtained by trailing feeding deer in the snow. In January and December, 1968, in the Douglas f i r community between Juniper and Starvation Slopes the following species and frequencies of use were observed: Douglas f i r -numerous blown down bough tips5'!; pine 2; lupine 19; strawberry 1; chickweed 1; sedge 2 ; bluegrass 3; fescue 9; pinegrass 15; and wheatgrass 4. In January, 1969, on the steep westerly exposed Douglas f i r community below Starvation Slope use on the following species was observed: Oregon grape 2; ceanothus 2 (both plants heavily browsed); Douglas f i r 2; rabbit bush 2. Browsing on Douglas f i r was observed to the very heavy in the area west of South Slope and north of, and east of, Starvation Slope where young trees were highlined and regenerating f i r was severly hedged (Figure 5.1.1). 5.1.3.3. Spring diet The "spring" mule deer diet was determined from eleven rumen samples, five from the open Douglas f i r forest (three to the south and "Difficult to quantify as the tips just l i e on the snow; may number into the thousands. Figure 5.1.1. Highlined and decadent condition of Douglas f i r regeneration between Starvation and Flatiron Slopes, Flatiron Mountain, 1968. 79 east of Starvation Slope, two from the west of South Slope flats); five from the open grasslands, (two from Starvation Slope and three from Juniper Slope); and one from the Ashnola Valley bottom below Flat-iron Slope. In the "spring" period use of grass increased to 38% while that of forbs and browse decreased to 7.7 and 53.8% respectively. The residual, 0.5%, was lichens (Table 5.1.6). Only 15.4% of the rumen sample material was identifiable, while 84.6% was unidentifiable fibrous material. Analysis of data presented in Appendix 5.1.7 indicated that there was not a significant difference between grass and browse utilization in the spring. Use of forbs was significantly below that of the other two forage classes (p < 0.01), Table 5.1.7. It was possible to observe trends in the spring diet by statistically analyzing the four "early" spring, March, observations and the seven "late" spring, April-June, observations. In the results presented in Table 5.1.8, within the.sample periods, the only significant difference is the greater browse utilization in the "early" spring (p < 0.01). Between period comparisons showed that only the utilization of forbs varied significantly, increasing in the "late" spring period (p < 0.01). Use of grass increased and of browse• decreased from the "early" to the "late" spring periods, but the changes were not statistically significant. Green forage in the grass forage class, primarily leaf blades, formed only a l i t t l e more than half of the total spring grass use observed in the rumen samples between March and.June.' The- diversity of species utilized in the spring exceeds that for the other periods studied. Sixteen species of forbs were used, the largest number for any Table-5.1.8. Statistical analysis of within Spring mule deer rumen sample analysis. Relative proportions by oven dry weight, expressed as percentages, of identified forage only. Percent Early Spring Significance within periods Significance between periods Percent Late Spring Significance within periods Significance between periods Grass Forbs Browse 27.9 3.6 68.5 TOTALS 100.0 Sample Size N.S. N.S. N.S. N.S. 58.1 15.5 26.4 100.0 N.S. N.S. N.S. N.S. N.S. N.S. = not statistically significant * = statistically significant at .05 level &ft = statistically significant at .01 level CO o 81 given season or ungulate species. Only five of the above species contributed appreciable amounts to the diet, and none occurred in more than five of the 11 samples (Table 5.1..6). During this season, u t i l i z -ation of nine browse species was observed, the most importnat of whcih was again the needles and bough tips of Douglas f i r . This browse occurred in each of the samples analyzed and formed 43.2% of the spring diet.. Kimikinnick was the second most important species. Spring use of pasture sage increased while that on buckwheat decreased from the observed winter use (Table 5.1.6). Buckwheat was observed in rumens from a l l areas sampled, while pasture sage was observed in samples taken to the west of South Slope flats, and on Starvation and Juniper Slopes proper. The most important "miscellaneous items" observed in the rumens were juniper (Juniperus scopulorum), not generally considered to be a palatable species. It was found in only two samples, one of which was a winter k i l l . Other items included lichens, mainly arboreal species, in winter and'spring, and balls of deer hair during the spring. Additional information on the spring food of mule deer relating . frequency of•use to relative availability, by forage class and'individual taxon, was obtained from the grazed plant loop-transects in deer-only feeding areas on Juniper Slope in the spring of 1969. . The results from the two communities, Poa pratensis, sodgrass-forb swale areas, and Artemesia frigida sparsely vegetated ridge areas, are presented in Appendices.5.1.7 and 5.1.8. The deer spring grazing transect data are summarized in Table 5.1.3, on a forage class basis, and in Tables 5.1.9 and 5.1.10 for the more important species only. Forage class and species 82 Table 5.1.9. Mule deer forage species preference index as indicated from Juniper Slope, Poa community, grazed plant loop-transect data. Species Availability Factor Frequency of Grazing* Preference Index Rank A Rank B GRASSES 1. Agropyron smithii. 0.02 1.4 63.6 1 4 2. Koeleria cristata 0.17 5.7 33.6 2 7 3. Poa pratensis 26.49 57.9 2.2 4 11 4. Stipa sp. 1.47 ' 4.3 2.9 3 10 FORBS 1. Achellia millefolium 0.07 4.8 65.8 3 3 2. Arabis sp. 0.007 2.4 363.6 1 1 3. Lappula redowskii 0.003 0.5 166.7 2 2 4. Penstemon procerus 0.01 0.5 34.2 5 6 5. Phacelia linearis 1.63 12.9 7.9 7 9 6. Taxacicum officinale 0.34 3.8 11.3 6 8 7. Viola adunca 0.05 1.9 38.8 4 5 * Calculated as a percentage of a l l plants grazed. Rank A = importance rank within a forage class. Rank B = importance rank between forage classes. 83 Table 5.1.10. Mule deer forage species preference index as indicated from Juniper Slope, Artemesia community, grazed plant loop-transect data. Species Availability Frequency Preference Rank Rank . Factor „ . Index A B Grazing GRASSES 1. Agropyron smithii 0.04 1.2 32.4 1 4 2. Festuca idahoensis 0.60 13.7 23.0 4 7 3. Poa pratensis 0.11 2.5 23.2 2.5 5.5 4. P. secunda 1.21 27.9 22.1 5 8 5. Stipa sp. 0.75 17.4 23.2 2.5 5.5 FORBS 1. Allium cernuum 0.01 5.0 555.6 1 1 2. Antennaria rosea. 0.95 - 0 0 0 0 3. Arabis sp. 0.03 6.8 198.8 2 2 4. Oxytropis campestris 0.05 3.1 67.4 3 3 BROWSE 1. Artemesia frigida 7.89 16.0 2.0 1 9 84 preference index evaluations are presented in these tables. Studies of spring deer use on Juniper Slope indicate that grasses were used in relation to their availability (Table 5.1.3). The higher preference value for browse than forbs, when forbs on the average were more frequently utilized, can be explained by deer feeding in the forest habitat for part of each day. This resulted in greater browse consumption and consequently more browse in the rumen samples than was indicated as available from the availability data which was • collected on the grasslands;• This discrepancy is reflected in the use of the "preference" index formula. In the Poa pratensis community grass was the most frequently used forage, accounting for 69.4% of a l l observed grazing, while only 33.4% of the grass observed was utilized (Appendix 5.1.8). Four of the seven species were grazed. Kentucky bluegrass was the most important in terms of availability and frequency of use, however, i t ranked last in terms of preference. The most "preferred" grass in this community was western wheatgrass (Agropyron smithii) (Table 5.1.9). Forbs were the second most frequently used forage class. Only 27.3% of the forbs observed were .utilized,:but forb use formed 30.1% of a l l grazing observed in the community. Eleven of the 19 forbs observed were used, however, only five were taken frequently. The most frequently used species was phacelia (Phacelia linearis), yet i t appeared to be the least "preferred". The most preferred species was rockcress (Arabis sp.) and i t contributed only 2.4% to a l l the observed gracing. Two'of the most important species in the rumen samples, loco weed (Oxytropis monticola) and cinquefoil 85 (Potentilia gracilis) were not sampled with great enough frequency by the transect technique for preference evaluation in the bluegrass community. Deer use of browse in the bluegrass community was very low, forming only 0.5% of a l l observed grazing. This was attributed to scarcity of browse in this community, rather than to dislike. In the second community studied on Juniper Slope, the Artemesia community, grass was again the most available and most frequently used forage class (Table 5.1.3). Of the grasses observed, 44.8% had been utilized and this formed 63.9% of a l l grazing observed. The mule deer's increased preference for grass in the spring is illustrated in this community by the increased percent frequency of grass use in response to a reduction in its relative availability compared to the Poa pratensis community. Use was made of seven of the eight species observed. Sandberg bluegrass was the most available and frequently used grass, followed by needle and thread (Stipa sp.) and'Idaho fescue. Once again the most available and frequently used species was the least preferred, whereas needle and thread, and Kentucky bluegrass were less frequently used due to reduced availability and ranked second on the preference index. Idaho fescue was only slightly less preferred and third on the l i s t . The most preferred species, western wheatgrass, was-the least available and hence cannot be a major component of the spring diet (Table 5.1.10). Forbs were again the second most.frequently used forage class. The reduced availability of forbs in this community is reflected in the reduced- use of the forb forage class in relation to a l l grazing observed in this community, viz. 19.9%. However, a greater percentage (35.6) of the forbs observed were grazed. Of the 14 forb species encountered, ten were observed to have been utilized. Rockcress was taken more 86 frequently than wild onion, but ranked second to the latter in terms of preference (Table 5.1.10). The availability of browse, in the form of pasture sage, was greater in the Artemesia community. The result was an increase in the frequency of browse utilization to 16.4% of a l l grazing observed in the community. This was a l l on pasture sage, however, only 16% of the pasture sage plants observed had been browsed. Pasture sage was the main browse species available, and dominated the observed browse utilization. Supplementary information on spring food habits was obtained by direct feeding observations on deer in the field. In April and May, 1968, deer were seen feeding in the Douglas f i r communities on the western slopes below Juniper Slope and South Slope. The following species and frequencies of use were observed: grass, primarily pinegrass, the occasional forb and kinnikinnick 45; Douglas f i r 31; currant 13. In March and May, 1969, deer were observed feeding in the Douglas f i r community between Juniper Slope and.Starvation Slope, to the west of South Slope, and.to the east of Flatiron Slope. The following species and frequencies of use were recorded: pinegrass and the occasional forb 9; currant 6;. Douglas f i r saplings and bough tips that had been blown to the ground 27. 5.2 Range forage appraisal 5.2.1 Site locations The elevation, slope, aspect and soil type for the four winter-spring range and two summer range plots are presented in Table 5.2.1; site locations are illustrated in Figures 2.1.2 and 2.1.3. Table 5.2.1. Topographic and soil information for enclosure plot sites.. Ungulate Species Location Elevation Slope % Aspect Soil Type Mule deer 1. Poa pratensis 2. Stipa-Artemesia WINTER-SPRING RANGE 5600 5600 5 19 SW S Orthic Black Orthic Bark Gray Bighorn Sheep South Slope 3. Agropyron spicatum 5700 55 Rego Dark Gray-Sheep and Deer South Slope 4. Agropyron spicatum. 5500 35 Rego Dark Gray SUMMER RANGE Sheep and Deer Joe Lake 1. Carex 2. Eanthonia-Carex 7400 7100 4 28 S SSE CO 88 5.2.2 Community descriptions' The. species composition of the four winter-spring range study sites are given in Table 5.2.2 and of the two alpine summer range study sites in Table 5.2.3. The scientific and popular names, for plants collected on the winter-spring and summer range areas are presented.in Appendices 5.2.1 and 5.2.2 respectively. The percent frequency of occurrence values for each species were calculated using the number of vegetative observations as a total, whereas those for the sundries were based upon the total 400 observation points. The community f l o r i s t i c data are not comparable" due to differences in the accuracy and efficiency of line^point sampling on "sodgrass", "bunchgrass" and alpine tundra ranges. In the Poa pratensis community on Juniper Slope the major associa-ted grass species were Stipa sp. , primarily S_. columbiana; grass formed-76.6% of the aerial and 88.2% of the basal observations. Of the ten forb species present Oxytropus campestris and Potentilla gracilis were the most common; forbs formed 23.6% .of the aerial and 12% of the . basal observations. Shrubs were absent in this community. In the Stipa columbiana community on Juniper Slope the major associated grasses were Koeleria cristata and Poa secunda. Grass formed 76.6% of the aerial and 77.9% of the basal observations. Achillea  millefolium and Aster sp. , primarily A. compositus, were the major forb species. Forbs formed only 3.7%.of the aerial and 7.0% of the basal observations. Shrubs, actually the chamaephytes Artemesia  frigida and Eriogonum heracleoides,. formed 19.7% of the aerial and Table 5.2.2. Species composition of the four Winter-Spring range oommunities studied, Flatiron Mt., 1969 Percent Frequency of Occurrence by Community Last Aerial Hits Basal Hits Kind of Cover Poa prat. Stipa Ag. Spic. Ag_. Spic. Poa prat. Stipa Ag. Spic. Ag. Spic. GRASS and SEDGE 1. Agropyron spicatum - - 54.8 ' 30.3 - - 67.6 45.0 2. Agropyron smithii 1.0 0.6 — - - - - -3. Carex sp. 1.3 0.3 0.3 3.1 3.2 2.9 10.0 4. Festuca idahoensis - 1.8 6.9 - — - 5.0 5. Koeleria cristata - 14.7 11.5 6.9 1.1 8.1 8.8 10.0 6. Poa ampla - - 0.3 0.8 - - - -7. Poa pratensis 70.8 1.1 - - 82.8 - - -8. Poa secunda - 9.6 - - - 44.2 - -9. Stipa columbiana 3.5 50.3 - 14.4 1.1 25.6 - 10.0 . SUBTOTAL 76.6 76.6 68.7 62.4 88.2 77.9 79.3 80.0 FORBS 1. Achillea millefolium 0.8 1.4 10.0 7.5 2.2 2.3 2.9 2.5 2. Agoseris,'sp. - 0.3 - 1.1 - - - -3. Antennaria sp. - 0.5 0.3 1.7 1.1 2.3 8.8 -4. Arabis sp. 0.8 0.6 - 1.4 1.1 1.2 - -5. Arenaria capillaris - - 3.6 1.1 - - 7.5 6. Artemesia michauxiana - - 1.8 3.1 - - - -7. Aster sp. 2.0 1.1 - 0.6 - - - -8. Astragalus serotinus - - 2.4 3.1 - - - -9. Cirsium sp. - - 2.4 0.3 - - -10. Collinsia parviflora - - 0.3 - - - - -11. Fragaria virginiana 0.3 - - - - - - -12. •"Geum ""mcrbphyllum- "• ' - - 0.8 - - - -cont... Table 5.2.2., oontinued. Percent Frequency of Occurrence by Community Last Aerial Hits Basal Hits Kind of Cover Poa prat. Stipa Ag. Spic 1 . 2 Ag. Spic. Poa prat. Stipa Ag. Spic1 . 2 Ag. Spic. 13. Heuchera ovalifolia 1.5 0.3 14. Lappula redowskii - - 1.5 - - -15. Lupinus sericeus - - 0.3 5.3 - - -16. Oxytropus campestris 4.5 - - - 4.3 1.2 -17. Phacelia hastata - - 1.5 - - -18. Potentilla gracilis 11.6 0.3 - 0.8 2.2 - - -19. Silene acaulis - - 1.2 0.8 _ _ _ 7.5 20. Taraxacum officinale 2.5 - - 0.3 - - - - -21. Verbascum thapsus 0.3 - - - _ - -22. Viola adunca and nutt a l l i i 0.3 - - - - - . - -23. Zygadenus venenosus 0.5 - - 0.3 - -SUBTOTAL 23.6 3.7 23.2 31.0 12.0 7.0 11.7 17.5 SHRUB 1. Artemesia frigida _ 16.9 5.8 3.1 14.0 2.9 2.5 2. Eriogonum heracleoides - 2.5 2.1 3.6 1.2 5.9 -3. Rosa sp. - 0.3 • — - - - - -SUBTOTAL - 19.7 7.9 6.7 - 15.2 8.8 2.5 Table 5.2.2. , continued. Percent Frequency of Occurrence by Oonmunity Last Aerial Hits Basal Hits 1 2 1 2 Kind of Cover Poa prat. Stipa Ag.. Spic. Ag. Spic, Poa prat. Stipa Ag. Spic. Ag. Spic. NON-VEGETATIVE Soil - - 52.0 31.0 22.5 Rock - - - - - - 36.0 38.8 Litter - - - 76. 8 26.5 24.5 28.8 Non-vegetative observation 0.8 11.5 17.5 9.8 1. Area grazed by sheep only. 2. Area accessible to cattle in past; grazed by sheep and deer. Table 5.2.3. Species composition of the two alpine (summer range) communities studied, Joe Lake, 1969. Percenta^ ie frequency of occurrence by community Last Aerial Hits Basal Hits Species Carex Danthonia-Carex Carex Danthonia-Carex GRASSES AND SEDGES 1. Agropyron saunderii - 0.5 - -2. Carex alborigra ) 3. C. phaeocephala ) 40.6 26.4 11.3 18.3 4. C. praticola ) 5. C. scirpoidea ) 6. Danthonia intermedia - 27.4 - 15.9 7. Festuca ovina 7.8 0.3 3.2 1.2 8. Koeleria cristata 0.3 7.3 - 7.3 9. Luzula spicata 2.2 1.3 3.2 2.4 10. Poa alpina - 0.5 - 1.2 11. P. pratensis - 5.7 - 2.4 12. P. rupicola 10.6 - 4.0 -13. P. secunda - 0.8 - 1.2 14. Trisetum spicatum . 3.0 0.8 0.8 1.2 SUBTOTAL 64.5 71.0 22.5 51.1 FORBS 1. Achillea millefolium 1.1 7.6 0.8 1.2 2. Agoseris glauca 0.6 1.0 - -3. Antennaria anaphaloides - 0.5 - -4. A.lanata and umbrinella - 13.7 -5. Antennaria rosea - - 1.2 6. Arenaria capillaris - 5.0 - 7.3 7. A. obtusiloba 2.2 0.3 23.4 1.2 8. Aster sp. 0.3 - - -Cont; Table 5.2.3. , continued. Percentage frequency of occurrence by community Last Aerial Hits Basal Hits Species Carex Danthonia-Carex Carex Danthonia-Carex 9. Delphinium menziesii 0.3 10. Geum triflorum 0.8 - 0.8 -11. Oxytropis campestris 9.2 3.4 5.6 2.4 12. Polemonium pulcherrimum 0.8 0.3 - -• 13. Potentilla diversifolia 9.7 5.0 1.6 4.9 14. Selaginella sp. - - - 17.7 2.4 15. Silene acaulis - 1.8 9.7 -16. Solidago multiradiata 6.7 0.3 3.2 -17. Stellaria longipes 3.6 2.3 0.8 4.9 18. Taraxacum sp. - 0.3 - -SUBTOTAL 35.0 28.1 77.3 25.5 SHRUBS 1. Arctostaphylos uva-ursi - 0.3 - 1.2 2. Potentilla fruticosa 0.3 - - -3. Eriogonum umbellatum - 0.8 - 14.6 SUBTOTAL 0.3 1.1 - 15.8 SUNDRIES Moss _ _ 9.5 24.8 Lichens - - 24.3 6.8 Litter - - 30.8 43.0 Soil - - 0.8 4.5 Rock - - 3.8 0.8 Non-veg. observations 10.5 4.3 - • -94 15.2% of the basal observations. In the two Agropyron spicatum sites on South Slope the major associated grasses were Festuca idahoensis and Koeleria cristata. The frequency of Stipa columbiana observations increased in the deer and sheep use study area, an area accessible to and used by cattle in the past. In the two sites''studied,grasses formed 68.7 and 62.4% of the aerial and 79.3 and 80.0% of the basal observations. • The major forb species observed were Achillea millefolium and Lupinus sericeus. Forbs formed 23.2 and 31.0% of the aerial and 11.7 .and 17.5% of the basal observations in the two sites. Shrubs, Artemesia frigida and Eriogonum  heracleoides, formed 7.9 and 6.7% of the aerial and 8.8 and 2.5% of the basal observations in the, two sites. In the Carex community studied on the summer range the major associated grasses were Poa sp. and Festuca ovina. The sedges and grasses combined formed 64.5% of the aerial and 22.5% of the basal observations. Forbs contributed 35.0% of the aerial and 77.3% of the basal observations, with many species being frequently observed (Table 5.2.3). Shrubs were virtually absent in this community, fo:rming less than 1% of the observations. In the Danthonia-Carex community, Danthonia intermedia was observed more frequently than the combined observations on the associated Carex sp. In this community grasses and sedges formed 71.0% of the aerial and 51.1% of the basal observations. Forbs formed 28.1% of the aerial and-25.5% of the basal observations, several species being observed with equal frequency. Shrubs formed only 1.1% of the aerial 95 but 15.8% of the basal observations as a result of their "cushion" growth form. 5.2.3 Forage measurements Forage utilization by cattle in the f a l l of 1968 as measured in Agropyron-Stipa and Stipa-Bromus sites on South Slope and in a Poa  pratensis site on Juniper Slope are presented in Table 5.2.4. In-a two week grazing period between September 18 and October 9, 1968, more intensive grazing occurred on the South Slope communities. The loss of forage quantity by leaching and weathering over the winters of 1968-69 and 1969-70 are presented in Table 5.2.5. Forage loss for 1968-69 was measured in an Agropyron spicatum community on South Slope and in a Poa pratensis community on Juniper Slope. Cattle grazing on these areas had reduced the amount ofz. forage. present at the start, of the winter and weathering losses were 41.8% and 16.0% for the two comm-unities respectively. Since cattle did not graze this range during 1969 the standing crop of usable forage was much larger at the beginning of the 1969-70 winter and forage weathering losses were increased. However, the losses measured include over winter wildlife use (possibly 9-10%) as the areas clipped were not fenced. In the Poa pratensis community the loss of the standing forage crop was 90.5%; forage pressed to the ground accounted for 36.1% of this, while the other 54.4% was "lost", from the site by shattering, leaching and "weathering". In -the Stipa-Artemesia community only a combined forage loss of 72.9% was measured. Data, on f a l l and over-winter forage use by sheep in 1968-69 is presented 96 Table 5.2.4. Fall forage utilization by cattle on Flatiron Mountain, 1968. .. Forage yield g/m Difference immunity & J toBefore After g/m2 Sept. 18/68 Oct. 9/68 Agropyron-Stipa 58. 3 32.3 26.0* 44.6 Stipa-Bromus 158. 5 63.8 94.6* 59.8 Poa pratensis 44. 3 37.5 6.8* 15.6 * Significant difference at 5% level 97 Table 5.2.5. Over winter forage weathering losses on South Slope and Juniper Slope 1968-1969 and on Juniper Slope 1969-1970. Community Forage Yield gm/m Fall Spring Weathering Losses % t gm/m Poa pratensis 1968-69 Oct. 9 44.3 April 22 37.2 16.0 7.1* Agropyron spicatum Oct. 11 96.8 April 20 56.3 41.8 40.5'' Winter Use and Weathering Losses % gm/m^  1969-1970A Poa pratensis standing forage pressed to ground Total forage Oct. 6 96.7 May 14 9.2 34.9 44.1 90.5 36.1 54.4 87.5* 52.6* Stipa-Artemesia Oct. 6 63.1 May 14 17.1 72.9 46.0* to Percentages based on f a l l forage yields. *'.;;'Differences significant at the 5% level. r ) A ' 'Clipping conducted on unfenced areas. 98 in Table 5.2.6. Fall to spring.utilization in two communities was measured and calculated as a percentage of the•forage available in the f a l l : A higher utilization percentage was' measured in the Poa pratensis community on Juniper Slope (37%) than in the Agropyron spicatum site on South Slope (25%). However, South Slope provided more forage in terms of grams per square meter. The mean dry matter forage production, utilization and affect of wildlife spring grazing on the standing crop of usable forage produced on the winter-spring ranges is presented in Table 5.2.7. The values presented are either increases, positive percentages, or decreases, negative percentages, between the protected and unprotected areas clipped on a given date. Differences in forage yields at the end of the three week grazing t r i a l at June 9, 1969, indicated 21% and 16% forage utilization on the sheep, grazed Agropyron spicatum community and the deer grazed Poa pratensis community, respectively. In the Stipa-Artemesia community on Juniper Slope the "deer-only" grazed area yielded 19% more forage than the ungrazed or protected area. The con-i • • founding effect of forage growth on utilization measurements during the grazing interval was taken into consideration. This was. accomplish-ed by calculating forage growth during the grazing t r i a l by "before and after" clippings in a protected area. At approximately June 12, when the deer, and June 24, when the sheep, departed from the winter-spring range a second series of clippings was conducted. In the Stipa- Arternesia and Poa pratensis communities grazed only by deer during the spring.there was a non-significant 12% and 4% increase in forage pro-duction on the grazed over the ungrazed areas. In the Agropyron Table 5.2.6..^  Fall and over Winter forage utilization by sheep on Flatiron Mountain, 1968-1969. Difference Forage Losses Fall to Spring Community F a l l yield Spring Yield 2 Weathering Utilization' gm/m^  gm/m^  gm/m % % %t gm/m^  Poa pratensis Oct. 9 April 22 37.5 21.0 16.5* 44.0 .16.0 37.0 13.9 Agropyron spicatum Oct. 11 April 20 62.5 35.7 26.8* 42.9 41.8 25.0 15.6 * Difference significant at the 5% level. t Utilization percentage based on forage available in the f a l l . CD CD Table B-J2.U). Forage production, utilization and the effect of wildlife spring grazing on forage production shown as differences in. mean.'- dry/ matter yield between/protected and grazed areas, Flatiron Mountain, 1969, Difference in mean yields between protected and grazed areas Community Type Grazing t r i a l June 9 Animals leave Spring range Deer Sheep June 12 June 24 Effect at time of irBximum production July 30 Maximum summer yield g/m2 July 30 Fall yield g/m2 Oct 6 Difference g/m2 Aug-Oct Stipa-Artemes i a 5.5 + 19% 8.3 + 12% 1 +1% 85.4 63.1 22.3* 27% Poa pratensis 6.8 - 16% 4.9 + 4% 10 - 7% 136.7 96.7 40.0* 24% Agropyron spicatum-1 10.2 - 21% '-35.6* - 30% 22.9* -15% 141.9 108.9 33.0 8% Agropyron spicatum2 -31.0* - 21% 25 -13% 187.9 109.3 78.6* 33% * Differences significant at the 5% level. 1 Sheep only grazing 2 Deer and sheep grazing 101 spicatum community grazed by sheep and deer in the spring there was a significant 21% reduction in forage.production on the grazed area. In the Agropyron spicatum area. grazed by sheep only there was a significant 30% reduction in forage production. Forage growth continued from the dates the wildlife species left the winter-spring ranges until late July when the permanent wilting point extended to sufficient'depth to promote summer dormancy in the Poa and Agropyron communities (Table 5.2.8). Another set of forage clippings was completed between July 30 and August 1, on grazed and ungrazed areas at each of the four sites. At this time only in the Stipa-Artemesia community was more forage harvested from the grazed area, but the yield difference (1%) between the grazed and ungrazed areas was not significant. In the Poa pratensis. community a nonsignifi-cant 7% reduction in forage yield was measured on the area grazed only by deer. Significant decreases in forage yields were measured in the "deer and sheep" (13%) and "sheep only" (16%) grazed areas of the Agropyron spicatum community. The loss in standing crop of usable forage from the time of JTaximum production to October was significant at three of the four sites; i t was not significant at the Agropyron spicatum sheep only grazing site (Table 5.2.7). It ranged from a low of 8% in the relative-ly climax Agropyron spicatum site to 33% in the second Agropyron  spicatum site which has substantial amounts of Koeleria, Stipa and Carex present in the canopy cover (Table 5.2.2). Forage losses in the Poa pratensis .and Stipa-Artemesia communities were intermediate between the f i r s t two. Table 5.2.8. Soil moisture at subsequent dates in 1969, for six inch soil horizons from 0 to 24 inch soil depth, at two study sites on Flatiron Mountain* •Site Percent available moisture Depth June 8 June 15 June 24 July 3 July 18 July 27 Oct 5 0-6 85 66 <20t 52 49 <20 26 6-12 88 78 24 <20 29 <20 <20 12-18 86 84 45 43 26 <20 <20 18-24 89 89 72 62 48 -.26 <20 Bouyoucos moisture readings (ohms resistance) June 7 June 10 June 15 June 24 July 5 July 19 July 29 0-6 895 3,675 9,525 10,000+ 778 4,825 10,000+ 6-12 650 10,000+ 8,125 10,000+ 3,430 5,550 10,000+ 12-18 677 8,475 10,000+ 10,000+ 10,000+ 10,000+ 10,000+ 18-24 "650 . 5,650 10,000+ 10,000+ 10,000+ 10,000+ 10 ,0.00+ Poa pratensis mesic Agropyron spicatum mesic * Sites established by F. Harper. t <20 percent available moisture or 10,000 ohms is below the permanent wilting point. 103 The mean dry matter forage production and utilization at two sites on the alpine summer range, near Joe Lake, is presented in Table 5.2.9. Summer grazing, to August 21, primarily by mule deer did not significantly reduce forage yield outside the fenced plot at either site, although 14% and 6| utilization were measured in the Carex and Danthonia-Carex communities respectively. Forage production, measured within the fenced areas, was significantly greater in the Danthonia- Carex community. The total nitrogen and crude protein levels were 1.25% and 7.8% in the Carex, and 1.39% and 8.7% in the Danthonia-Carex communities respectively (Table 5.2.9).' The levels of crude protein, in grams per square meter, was higher at the Danthonia-Carex than the Carex community. Table 5.2,9. Forage production and utilization by wildlife on two alpine communities, Joe Lake,- 1969. Community Type Mean Yieldt Aug 21 Protected Grazed area area % Difference 9-Nitrogen % Protein Carex.sp. 138.0* +14.3SD 119.4 ± 9.9SD 14 1.25 7.8 Danthonia-Carex sp. 176.8* +32.5SD 166.5 +11.7SD 6 1.39 8.7 * Difference significant at the 5% level. t Oven dried mean yields derived from 8 one square meter quadrats. r—' O -P 105 6. DISCUSSION 6.1 Food habits The observed variation of individual forage species in food habit and range forage studies indicated that a more inclusive grouping of plants was necessary for the evaluation of potential dietary competition between ungulate species. The main objective in food habit studies should not be analysis of rumen samples to forage species. Smith (1952) and Ghamrad and Box (1968) illustrated that detailed food habit findings are only of local value due to regional differences in forage species and their relative availability. They stressed that the most useful food habit character with general application or merit for comparative purposes is the.proportion of,browse, forbs and grass utilized during a given season. To facilitate comparisons,with other studies and to evaluate poten-t i a l dietary competition between the three ungulate species, individual forage species have been grouped into forage classes: grasses, forbs and browse. However, analysis of rumen samples and range forage utilization were conducted to the species level. This was necessary to evaluate potential dietary competition resulting from statistically significant use of.the same forage class by different ungulate species. In this regard, comparison of the major forage species contributing to the overlap in use of each forage class was as essential for evaluating potential dietary competition as are 1) food habits, 2) percent forage utilization and 3) the area or ecological.niche used by each ungulate-106 species for evaluating range competition. , • •. Cook and Stoddart (1953)., Heady (1964) and Kelsall (1968) stress the necessity of relating forage use to availability. This is also true for evaluating forage species or forage class preferences as pre-ferences vary in relation to regional availability, and other factors discussed by Heady (op.. c i t . ) . The preference index formulas employed by Chamrad and Box (1968) are based upon rumen analysis and in this "way are comparable to the present forage class preference formula. Those presented.by Drawe and Box (1968) are based on field observations and are comparable to the present forage species preference index. Preference indices however do not reflect the volume each forage species contributes to an animal's diet. The grazed plant observations indicated that the least available species often receive a high frequency of use and as such receive a high preference value. They may not however, contribute greatly to the bulk of the diet due to their limited availability. Conversely the abundant and hence important species receive a lower preference index value due to the frequency of their observance. Preference calculations and frequency of use observations at the forage class level, relating use to availability are, as a result, better for the purpose of comparing food habits. The relationship between ~" class use and the availability value, presented in Figure 6.1.1, i l l u s -trates that forage use by deer and sheep is related to relative forage availability. The same relationship was observed for spring and early summer cattle grazing on South Slope by Blood (1961); however his observations may be confounded by spring deer grazing as deer tend to Figure 6.1.1. Percent distribution of grass, forb and browse utilization in relation to their availability factor value during the spring; from grazed plant loop-transect data. 100 DEER SHEEP DEER SHEEP DEER SHEEP GRASS FORB ' BROWSE 108 use areas which receive heavy cattle use. Food habits, frequency of use and relative forage availability information was collected from the grassland communities in the spring. This habitat type is used in common by the three ungulates and this is where and.when interspecific and intraspecific range competition between cattle, sheep and deer may be originating. No attempt was made to evaluate winter or spring browse use in the forest habitat because i t was observed that cattle and bighorn made only limited use of the forested range; cattle could in a l l probability make some use of i t , however, i f .encouraged to do so. 6.1.1 Comparison of food habit studies Comparison of ungulate food habits from various areas is most validly conducted on a forage class basis as a result of regional vari-ation in forage species and their availability. 6.1.1.1 Cattle ' Data presenting the spring diet of cattle on Juniper Slope, Flatiron Mountain (Section 5.1.1) have shown that the cattle select 97.5% grass. Intensive use was also evident in the summer of 1968 when the grass on Juniper Slope was uniformly grazed to a height of two inches or less. (Compare Figures 6.2.1 and 6.2.2 of range after combined cattle and wildlife use in 1968 and after only wildlife use in 1969). Blood (1961), studying the spring and summer food habits of cattle on South Slope, a big game winter range, observed that grass formed 96.5% 109 of the diet. The predoirrinant species grazed were bluebunch.. wheat-grass, junegrass and bluegrasses. Food habits of cattle in the Ashnola are comparable to those of other areas where grass formed between 64 and 83% of cattle diets (Table 6.1.1). Lesperance et a l . (1970). and Smith and Doell (1968) found that cattle preferred to use herbaceous forage, especially grass, but would consume browse i f the supply of herbaceous forage was depleted. In the Ashnola the general lack of shrubs, especially on Flatiron Mountain, restricts cattle to a diet of grass and forbs. Chamaephytes, such as pasture sage, were moderately abundant on the range but were not used by cattle during the spring. 6.1.1.2 Bighorn • sheep_ Data presented in Section 5.1.2 indicate that the California Bighorn on Flatiron Mountain utilize grass as the major constituent of their spring diet (91%), whereas browse formed less than 1% of the spring diet. This'..is true for bighorn populations in general (Capp, 1968, Table 5); however, only two of the 16 papers he reviewed present data on spring food habits. ' The results presented earlier compare favorably with those of Blood (1961) (Table 6.1.2). Although pasture sage is an important winter forage (Sugden, 1961; Blood op_. ext.; and Demarchi, 1965) i t was not observed in the May and June rumen samples. Blood (op. cit.) observed that use of pasture sage and buckwheat declined from 15% to trace and 12% to '2% respectively from March through May, and that grass formed 95% of the observed spring grazing. The three most frequently used grass species Table 6.1.1. Food habits of cattle on North American ranges. Location of Study Technique Season Percent of diet Grass and Grass-like Plants Forbs Browse Authority South Slope, Ashnola, B.C. Missouri,River Breaks, Montana Grazed stem counts Feeding site exarnination Spring and summer Spring Summer Fall 96.5 82 64 76 1.6 Tr 3 7 1.7 18 33 17 Blood, 1961 Mackie, 1970 Starkey Exper. For. and Range. Oregon Nevada Occular estimate by plots June to October May-Nov. 90 71-83 7-17 0-22 Skovlin et al. 1968 Lesperance et al. 1970 Nevada Arizona Rumen fistula samples General diet-73 September to December 82 13 19 Lesperance and Tueller, 1969 Gait et a l . , 1969 • H o Table 6.1.2. Food habits of Bighorn sheep populations«. Location of Study Technique Season Percent of diet Author Grass and Grass-like Plants Forbs Browse Ashnola, B.C. Rumen samples Spring* 93 2 4 Grazed stem counts Spring* 94 3 3 Trailing in snow Winter 44 7 49 Grazed stem counts Winter 64 4 32. 0 Annual Means* 74 (72)t 4 (4) 22 (24) Ashnola, B.C. Rumen samples Winter 88. .1 . 5. 9 6. 0 Grazed stem counts and trailing in snow Winter 53. .5 9. 7 34. 9 Churn Creek, B.C. Trailing feeding animals in snow Winter 29 9 62 Yellowstone Examining feeding National Park sites after fresh snowfall Winter 61. ,4 17. 2 21. 5 Blood,.1967 Demarchi, 1965 Sueden, 1961 Oldemeyer et a l . , 1971. * Data re-evaluated from Blood (1961) to include only spring rumen samples, t (Mean values) are those from Blood (1967). 112 were bluebunch wheatgrass, junegrass and bluegrasses. Observations of sheep grazing by the loop-transect technique conducted in two communities early in July, 1969, in areas not grazed by either cattle or deer, indicate that grass formed only 68.6% of the observed grazing. Bluebunch wheatgrass and junegrass were the most important species in terms of frequency of use. In the Agropyron community Idaho fescue was the most preferred species and no utilization was observed of the small amount of bluegrass present. In the Agropyron-Artemesia community however, bluegrass was the most preferred species, but ranked third in frequency of use behind bluebunch wheatgrass and junegrass. Browse formed 7.7% of the observed grazing; the majority (5.4%) occurred on pasture sage. However, only 18% of the pasture sage observed was grazed, so winter availability should not be affected.. Bighorn spring food habit results obtained by the two methods of study are similar. Observations from the grazed plant loop-transects, in comparison with the rumen sample analysis, indicate a lesser use of grass, with a • corresponding increase in use of both forbs.and browse. However, grazing observations indicate only the frequency of use in relation to a l l grazing observed, not the quantity used. Also , the , grazing observations being restricted to current annual growth omitted use of mature forage produced -in the previous season and this formed over 50% of the increased grass use observed by rumen analysis. 6.1.1.3 Mule deer Data presented in Section 5;1.3 indicates that the diet of mule deer on Flatiron Mountain is dominated by browse during the f a l l , winter and spring. Trends in the seasonal utilization for forage classes 113 were observed (Figure 6.1.2), although there was considerable sample to sample variation in any given season. Forb use was greatest during the f a l l , but not significantly greater than winter use. It decreased significantly during the spring compared to f a l l and winter, but increased significantly from "early" to "late" spring as forbs became more available. Maximum browse use was observed in winter and minimum use in spring, but the seasonal differences were not statistically significant. Grass use decreased to its lowest level in winter and increased.to a maximum in spring,, but again seasonal differences were not statistically significant. This seasonal pattern in the food habits of mule deer is similar to that presented by Taylor (1956) and in studies reviewed by Capp (1968) and Cooperrrider (1969). Additional information on the food habits of mule deer is presented in Table 6.1.3 from studies not reviewed by the two authors (supra vide). Mule deer winter ranges on Flatiron Mountain differ markedly from the optimum habitat described by Dietz et a l . (1962) in the lack of deciduous shrubs available for winter use; this was also observed by Blood (1961). It -can be surmised, from the reviews referred to above, that deer are basically browsers especially during the winter. Mule deer in the study area were maintaining the normally high browse•intake by.foraging on conifers. In the f a l l , deer inhabited the lodgepole pine community on Joe Camp Ridge and used the lodgepole pine as forage. The major f a l l forages used were lodgepole pine, kinnikinnick and big sagebrush. Use of the latter two species is common and is recorded by other workers. However, use of pine has been previously recorded only by Blood (1961) Figure 6.1.2. Seasonal variation in oven dry weights of grass, forb and browse, expressed as percentages, found in mule deer rumen samples-t—• FALL WINTER SPRING * Table 6.1.3. Food habits of mule deer populations. Location of Technique Season Percent of diet Author Study Grass, and Grass-like Plants Forbs Browse Utah - - Spring 66 12 22 Smith and Julander, 1953. Nevada - - Year long average 6 13 80 Lesperance and Tueller, 1969. Nevada - -Year long Average Summer Winter 1-12 5 3 9 46 3 62-97 49 94 Lesperance et a l . , 1970. Montana Rumen sample Spring Summer Fall • Winter 26 Tr 29 11 30 36 32 27 36 59 39 73 Mackie, 1970. Rock Creek, Montana Rumen samples Winter Spring 4-12 49 8-43 29 45-88 ' 22 Cooperrider, 1969. New Mexico Rumen samples Year long average 2.2 16 75 Boeker et al. , 1972 Chum Creek, B.C. Observation and trailing Winter 4.7 5 90.3 . Sugden, 1961. Ashnola, B.C. Rumen samples Winter 24 15 61 Blood, 1961. H cn 116 and Sugden (1961) for mule deer collected.on Flatiron Mountain in the Ashnola and in Big Basin,.Churn Creek respectively. Nellis and Ross (1969) and Lovass (1958) found that mule deer used Ponderosa pine in Montana when occupying the pine community in winter. As a result of increased availability from f a l l regrowth, use of grass in the f a l l exceeded that observed in winter. In the winter deer maintained a 67% browse intake by consuming bough tips and needles of- Douglas f i r . It was the most available species in the Douglas f i r community which they inhabited at that season. Deer inhabiting and collected in a lodgepole pine community in December 1968 were found to be eating pine. Observations on the highlined and decadent condition of Douglas f i r regeneration compared to the lightly browsed condition of soapberry and currant shrubs indicates that the latter two are not preferred species. Hedged willows in the open lodgepole pine forest to the east of Juniper Slope have been heavily browsed. This cannot be attributed to deer use only* however, because cattle use the area in the f a l l when herbaceous forage is not abundant, on Juniper Slope. Douglas f i r has been recorded, in small.amounts, as a winter deer forage by.other authors, Lovass (1958), Mackie (1970), and Cboperrider (1969). Sugden (1961), however, reported that i t was the most important winter deer food, together with kinnikinnick, in the Big Basin area of Churn Creek. Nellis and Ross (1969) found that con-ifers, both. Douglas f i r and Ponderosa pine, were heavily utilized during winter in response to a scarcity of deciduous shrubs on the National Bison Range in Montana, and that pasture sage was used when other green 117 forage was scarce. Sugden (1961) reported that deer use of pasture sage on Sheep flats, Churn Creek, during winter was 63% of a l l deer grazing observed by trailing feeding animals. In the Ashnola Blood (1961) observed deer using pasture sage and he found that i t averaged 20% in the 3 rumen samples he analyzed. However, two of his "winter" samples were collected in March and April. In the present study very light use of pasture sage was.found during the winter (< 1%), increasing only in the latter part of the spring grazing period, and then only to 2%. Grass is apparently the last important forage class for wintering deer. Cooperrider (1969) studying the winter food habits of mule deer at Rock Creek, Alberta, reported that grass formed less than 10% of the diet. Sugden (1961) found winter utilization of grass to be less than 5%. An-exception to the apparently reduced use of grass during winter is the data presented by Blood (1961). On Flatiron Mountain, Blood (op_. cit.) found that deer use of grass exceeded that of forbs during the winter, averaging 24% of the diet as determined from the three "winter" rumen samples. Present observations indicate that deer use new grass as soon as i t is available which was .in March in the spring of 1968. In his study Blood (op_. cit.) did not differentiate use of new growth from that of matured grass and this may account for the high winter use reported. In the present study, grass formed only 10% of the diet in January and February but this increased to 38% by March and April, 1968. Deer are doubtless to quite a degree opportunist in their foraging habits, which could explain why-Blood (ojp_. cit.) observed heavy use of grass during the "winter". Flatiron Mountain. 118 ranges do not provide -an abundance of deciduous shrubs for winter deer foraging, as discussed earlier; hence during winters of a light'snow pack deer may take advantage of the increased accessibility of grass. This relationship between use of herbaceous forage and snowpack is discussed by Coblentz (19.70). The opportunity to use grass existed during the f i r s t winter of the present study, 1967-68, and deer made some use of the open grasslands from January through spring. However, the use of grass •> as indicated by rumen sample analysis 5 averaged only 10% in January and February, with one January sample containing as much as 43% grass. Blood (1961) suggested that further reduction of browse availability in the Ashnola could force deer to utilize more grass and pasture sage thus increasing winter competition with bighorn. Sugden (1961) thought that deer on the Churn Creek ranges were utilizing large quantities of pasture sage out of necessity because the taller browse species were lacking. Cowan (1947) considered mule deer winter use of grass in Jasper Park above normal when he observed 15% winter use and thought i t was the result of a lack of available browse. This suggests that under certain conditions deer modify their winter diet to include unusual amounts of grass or pasture sage. Food habit studies indicate a marked (but non-statistically signifi-cant) increase in use of succulent new grass during the spring. The increase is accentuated because grass is available before the new growth of forbs or shrubs and .hence is sought early in the spring (McMahan, 1964; Leckenby, 1967; Yeager, 1960). This is supported by the results of rumen sample analysis and grazed plant loop-transects. However, the agreement between the two techniques for mule deer spring 119 food habits was not as good as that for the bighorn. Rumen samples indicated less use of grass and forbs and more use of browse than the grazed plant transect observations. Two factors contributed to the observed differences. In the spring deer use the open grasslands primarily in the morning and evening, spending the remainder of the day on the forested ranges where browse species are more available and apparently utilized, as shown from rumen sample analysis. Secondly, the grazing transects were conducted only on the open grasslands, as this is where sheep and deer range use overlaps, and only two of the nine browse species used by deer in the spring occur on the grasslands. This sampling problem influenced the mule deer forage class preference values. The mule deer spring rumen samples contain a forest forage influence, mainly browse species, the availability of which was not accounted for by studying relative spring forage availability on the grasslands. In the preference index calculations this has reduced the mule deer's apparent preference for grass in the spring, while exaggerating the importance of browse (Table 5.1.3). In the early spring grass use reached a seasonal maximum• whereas use of forbs increased during the spring.- Mackie (1970), Lovass (1958), Smith and Julander (1953) and Julander (1952) observed similar trends in spring use of forage classes. The greatest number of forb species was used at this time but only 11 out of the 28 species used were important. Increasing use of herbaceous forage affected corresponding reductions in the use of browse. Use of Douglas f i r progressively decreased from March and April through May and June. • Use of buck-wheat, although•light, only 3% of the spring browse or 1.6% of the 120 spring diet, was frequent in that i t was present in 9 of the 10 rumens analyzed. Pasture sage was present in 5 of the 10 samples but use was light and i t formed only 3.9% of the spring browse use and 2.1% of the spring diet. The. observed frequency of use by the grazed plant transects was also relatively light, reaching a maximum of 16% on the Juniper Slope Ridge or Artemesia community. In contrast, Blood (1961) found 21% pasture sage in two deer rumen samples collected in March and April as discussed earlier. This indicates that deer may have made greater use of the grasslands in the spring of 1961 than during the spring of the present study, 1968. 6.1.2 Potential dietary competition and.animal equivalence Competition discussed in this section has been qualified as "potential" because i t is based only upon food habit studies. This information relates season of use and.class of forage used by the ungulate species, but one cannot discuss range competition until the areas grazed by each species are known and the adequacy of supply is related to the observed-use. 6.1.2.1 Potential competition in the spring Food habit studies indicate that during the early spring use of grass by mule deer, when the supply of the succulent new forage is at first.limited, may result in competition between deer and.sheep. The extent depends upon the intensity of use and the amount of range used in common in the early spring. The grazed plant observations (Table 5.1.3), which represent frequency of forage class-use, not the 121 amount contributed to the diet, indicate that the spring food habits of deer.and sheep, while on the grasslands, are quite similar (Figure 6.1.3). However, the bias in.the mule deer spring grazing observations has been discussed in Section 6.1.1.3, mule deer spring food habits. Briefly stated, the bias i s : 1) deer use the grasslands for only part of their daily foraging and as such the grazed plant observations, conducted only on the grasslands, do not represent the mule deer's total spring diet; 2) observations of spring grazing by deer were conducted on Juniper Slope (Poa dominated) whereas observations of spring grazing by sheep were conducted on South Slope (Agropyron  spicatum dominated). This disparity was necessary in order to observe areas of single species grazing. This may result in use being observed on different plant species but the relative proportions of forage classes used should not be greatly altered. Since Sugden (1961) found that deer in winter would utilize A. spicatum when wintering on a range dominated by that species i t may be questioned that deer using South Slope in spring would.use the A. spicatum and Festuca available there. Analysis of spring rumen samples indicated that deer relied heavily upon browse whereas sheep and cattle used grass almost exclusively. The relative spring use of the three forage classes by cattle, bighorn and deer is presented in Figure 6.1.4. • Statistical analysis of cattle, bighorn and deer spring food habits, as indicated from rumen analysis results, are presented in Table 5.1.2. ' The analyses show that bighorn and cattle diets were not significantly different; both were utilizing grass very heavily during the spring. Deer use of grass in the spring, although at a seasonal peak, was significantly less than that of either Figure 6.1.3. Relative forage class use by mule deer on Juniper Slope, and bighorn on South Slope in the spring. Data from grazed plant loop-transects. Figure 6.1.4. Variation in grass, forb and browse utilization in the spring food habits of cattle, bighorn and deer, from rumen sample analysis. rO CO CATTLE BIGHORN DEER UNGULATE SPECIES 124 cattle or bighorn. Deer use of browse remained high during the spring but i t was not significantly greater than that of cattle or bighorn. There was no significant difference observed in the use of forbs during the spring between the three ungulates. Cattle grazing on the winter-spring ranges while sheep and deer are s t i l l utilizing them in the spring could present a forage competi-tion problem. Some degree of potential competition is possible between cattle and deer in the late spring as cattle use - especially on Juniper and Starvation Slopes - - reduces forb availability. It has been shown earlier that deer use of forbs increases significantly from early to late spring. Relatively light spring use of browse species occurring on the grasslands, viz. buckwheat and pasture sage, suggests that use and hence competition for these species is not important during the spring. 6.1.2.2 Potential competition in winter Potential interspecific and intraspecific competition during the winter can arise in two ways: 1. If the diet of an ungulate species in the spring or spring and early summer, is similar to its winter diet, or the winter diet of another ungulate species, and 2. If there is similar forage class use within or between ungulate species during the winter. As in case one above interspecific potential dietary competition is possible during the winter as a result of spring and early summer-cattle use of grass and sheep use of grass in winter. In addition, and 125 for the same reason, competition during winter could exist, but to a lesser extent, between deer and sheep. In the same way intraspecific potential dietary, competition could arise among sheep during the winter as a consequence of grass use by sheep during the spring and early summer. Light spring use by cattle, deer and sheep of browse species occurring on the grasslands should reduce the possibility of competi-tion occurring for this forage class during the winter. However, Blood (1961) suggested that sheep utilized a l l the available pasture sage early in the winter of 1960-61, and as a result had to use other forage. If this is their normal pattern then .any • .< ; : " .. use on pasture sage which reduces the availability of this forage for sheep use during winter w i l l be increasing competition for- this- forage. The competition w i l l be (a) intraspecific sheep competition either bet-ween seasons of use from spring use to winter use or just during the winter, and/or (b) interspecific competition between sheep in winter and deer in the spring, as a result of deer use reducing the potential production of browse on the grasslands and hence its availability for winter use. The effect of spring forage use on the potential forage production of the grasslands w i l l be evaluated in the range forage section. It is possible that potential intraspecific dietary competition is occurring as a result of deer browse use. Deer use browse as a major .component of their diet during their occupancy of the winter-spring range.. Therefore, competition may occur either between seasons of use, from spring use to f a l l and winter .use (several browse species .126 used in spring are also important during the winter) or as a result of excessive use during the winter period only. In relation to case two above, interspecific potential dietary competition in winter can only occur between deer and sheep as cattle are removed from the wildlife winter ranges in f a l l . The possibility of competition between deer and sheep on Flatiron Mountain during winter is reduced as a result of the difference in their winter foraging habits (Figure 6.1.5) and their use of habitat types. Intraspecific competition may also occur during the winter i f forage use is excessive by an ungulate species. However, since this has to be evaluated in terms of quantity of forage used in relation to availability, intraspecific competition w i l l be discussed in the range forage section. Potential competition for forbs either among or between ungulate species between seasons is not as strong as that for grass because forbs were only.a minor part of the seasonal food habits. In addition although spring use may have some effect on forb productivity and hence winter availability this w i l l be minor compared to that of late summer drying and disintegration of forbs. 6.1.3 Animal equivalence .Computation of animal equivalence from an ecological basis is important. Ultimately decisions concerning the numbers of each ungulate species allowed on the. range to achieve the optimum balance of forage utilization w i l l have to be made. Percentage distribution of forage class use r-o o o CD O 00 o o o C D C D TO M o I?1 4 O & r!-OP CD f—1 cn s? CD 3 cn O O ft rt> 4 H' (t Hi cn o S N O r—1 CL CO CD S ? 3 r+ fl) 4 r+ cn tr o 3 CD CD 8 CD 3" r+ cn s a r-t) ^ 3 P £ 2 1 128 Blood (1961) equated range use and land use competition between ungulate species on the basis of sheep unit months; where "one sheep unit month (S.U.M.) can be defined as the use made of any particular range by one sheep of average weight, for one month". Deer and cattle use was equated to a S.U.M. by calculating five bighorn sheep or three mule deer equal to one cow on the basis of dry matter intake. Demarchi (1965) also compared sheep.use to that of cattle and deer on a total dry.matter intake basis as above. .. Ecologically speaking, however, one cannot equate the three ungulate species directly on biomass or metabolic .energy requirements for several reasons: (a) the food habit differences between each ungulate species; (b) the season of the year the range is used, actively growing versus dormant; (c) the ecological impact of range use by ungulate species upon product-ivity, vigor and succession of plant species; (d) the region of range or ecological niche used to obtain the total forage intake in relation to the other ungulate species. It is suggested, for these reasons, that animal equivalence should- be calculated taking as many of the above factors into account as possible rather than just metabolic requirements. Considering only the food habits variable, in addition to the dry matter intake, the equivalence ratios between deer, bighorn and cattle are different than those employed by Blood (1961) and Demarchi (1965) or those recommended by the American Society of Range Management (1964), Rasmussen and Doman (1943), and Stoddart and Smith (1955). Deer, bighorn and cattle use of the grassland could be equated on a forage class basis using only grass, as i t is the major forage class 129 utilized and produced on the ranges. . Results of rumen sample analysis presented, earlier indicate that when a l l three- ungulate species are using the grasslands in the spring, grass forms 97.5% of the cattle diet, 91% of the sheep diet and 38% of the deer diet. Using these data and knowing that 660 pounds of forage or one animal unit month (A.U.M.) is equivalent to one cow with calf or five sheep or five deer, as defined by the American Society of Range Management (1964), then one can calculate modified equivalence values for the three ungulate species. Food habits considered at a forage class level vary with the seasons, hence animal equivalence ratios, w i l l also vary seasonally. Consider, as an example, cattle-deer equivalence in the spring: one cow and calf requires 660 pounds of forage per month, 98% of which is grass. This means 646 pounds of grass per A.U.M. One deer would use one-fifth of the total forage.of a cow and calf or 132 pounds of forage, of which 38% or 50 pounds would be grass. Thus i f 50 pounds of grass maintains one deer for one month then the 646 pounds of grass (the equivalent of one cow A.U.M.) would support 12.9 deer for a month in the spring. This means that 12.9 deer on the grasslands in the spring would consume as much grass as one cow with calf. However, the 12.9 deer and one cow may s t i l l not have equal range competition values because the two ungu-late species may (1) use different species of grass, (2) obtain the grass from different areas and (3) not have the same ecological impact on the range as a result of spring use. Using calculations similar to that above, other animal equivalence for spring and winter are presented in Table 6.1.4. 130 Table 6.1.4. Animal equivalence calculations using seasonal ungulate food habit information from Flatiron Mountain ranges 1968-69. Season Species % grass in diet Equivalent to Equivalent number Spring deer sheep, deer cattle Winter deer 38 91 38 97 13 1 cow AIM .. 1 cow AUM 1 sheep AUM 1 sheep AUM 1 sheep AUM 12.9 5.4 2.4 0.18 6.7 131 6.1.4 Limitations,of and recommendations for food habit studies 6.1.4.1 Rumen sample analysis Description of ungulate food habits by.rumen sample analysis techniques has.certain shortcomings. _ These include: (a) Variation in ease of identifying fragments of various forages and percent of sample identificaiton. Bergerud and Russell (1964) found that they could only identify 10 to 18% of screened caribou rumen samples'. In the present study the identifiable portion ranged from 15.4 to 38.4% for seasonal mule deer samples, and was 27.5% for bighorn and 30.7% for cattle during the spring. (b) Norris (1953), Bergerud and Russell (1964) and Scotter (1967) have noted variation in the rate of passage for different forages in rnjminants. Differences in digestion rates also reflect upon the validity of identifying only set fragment sizes in rumen samples to accurately represent the ungulates total diet (Scotter, 1966). This suggests that rumen sample analysis is more qualitative than quantitative. • The authors listed above, reported that herbaceous material passes through the rumen more rapidly than does coarse forage. Differential rates of passage have been partially explained by the presence of oils in Douglas f i r (Ho et.al.,1967) and in sagebrush (Nagy et a l , , 1964) which inhibit rumen bacteria. It is generally agreed, as a result, that rumen sample analysis tends to overestimate the importance of browse and underestimate the importance of herbaceous forage in-the diet. 132 (c) High variability.in the use of forage species has been observed between individual rumen samples. This means that large sample sizes are required to accurately represent use of even the major , forage species. Medin (1970) in a review of stomach content analysis reported that to provide an adequate estimate of major food items within 15% of the true mean at the 95% confidence level would require a sample of 9 3 mule deer rumens. A large sample such as this is difficult to obtain and requires a great deal of time to analyze. Rumen sample analysis to the level of forage class is recommended, when acceptable to the study, because: between:' sample varia-b i l i t y is reduced; the data collected is comparable between ranges with different forage species availability; analysis time can be reduced because fewer rumens would be required and the analysis would require less time per sample.-(d) A definite limitation to rumen sample analysis is the time involved in separating and identifying forage species encountered. Conifers and most browse species can be readily identified, forb species are more difficult and the majority of food habit studies reviewed did not attempt separation of grasses to species or even genera. Attempts to reduce time requirements and quantify rumen•sample analysis have achieved varying degrees of success using modified techniques (Chamrad and Box, 1964; Gait et a l . , 1968; and Robel and Watt, 1970). 133 6.1.4.2 Grazed plant loop-transect (a) The forage species occurrence and frequency of use observations recorded on the deer and bighorn grazed plant loop-transects appear to be adequate for only the major forage species. Fewer observation sites are required with "loops" than with "points" to represent the forage species present, but Smith (1962) found that the loop method is deficient in detecting the less frequently occurring species. Time limitations pre-vent more comprehensive sampling. (b) The grazed plant loop-transects, conducted only on the grass-lands ,.did not provide data on the availability or frequency of use of browse in the forest range. However, deer.use of browse from the forest.range was reflected' in the results of rumen sample analysis. Consequently, the preference index, computed from data of use in relation to availability, overestimated the preference for browse' by deer. The deer grazing-transects were not conducted on range used by both deer and sheep as.it was not possible to distinguish between deer and bighorn grazing. In addition, i t was not possible to locate a "deer only" grazing area on an Agropyron  spicatum range such as South Slope and consequently the "deer use" transects were located on a Poa pratensis dominated area, Juniper Slope. 134 6.1.4.3 Trailing feeding animals Additional deer and bighorn forage use observations obtained by trailing feeding animals while snow blanketed the range are not quantitative measures of forage use. Use of this technique was further complicated in that snow in excess of approximately three inches caused the animals to paw through the snow to.obtain forage. The pawing resulted in fragmentation of plants occurring in the "snow crater" making i t difficult to deterniine which species had been fed upon. 6.2 Range forage appraisal Measurement of browse use by deer on the forested winter ranges was deleted from the study. Inasmuch as other browse species are absent, Douglas f i r is perforce the key winter browse for deer on Flatiron Mountain. Quantitative measurements of its utilization and availability are difficult to obtain because i t is utilized by rodents as well as deer and winter storms affect its availability. Bough tips blown to the ground from the forest crown, by the more severe winter storms, remain available only until the next snow f a l l . 135 5.2.1 Forage measurements The unexpected termination of cattle grazing on Flatiron Mountain at the end of the 1968 grazing season prevented evaluation"of cattle and wildlife forage utilization during 1969. However, using the 1968 Poa pratensis maximum production figure (Harper, 1969), i t was. possible to calculate seasonal forage utilization and carryover values for Juniper Slope in 1968 (Table 6.2.1). Lack of maximum forage production values for the Agropyron spicatum and Stipa-bromus communities on South Slope necessitated calculation of percentage estimates for f a l l cattle use, overwinter sheep use and overwinter forage weathering loss from f a l l forage values. Total forage losses, including late summer weathering losses (33.1%, Harper op_. cit.) and use by cattle and wildlife from the spring until September 18, amounted to 71.3% of the annual maximum production on Juniper Slope. Further losses by f a l l cattle grazing, over-winter sheep grazing and weathering losses contributed to the 89.3% removal of forage produced in 1968. This rate of utilization allowed only 28.7% forage.carry-over at the end of the summer and 10.7% carry-over the following spring. Thus the combined cattle and wildlife range use exceeds the recommended f a l l carry-over value of "at least half of the total season's growth" (McLean and Marchand, 1968). In 1969 only wildlife grazed the Ashnola ranges on Juniper Slope, primarily during the spring, and this resulted in removal of 29.3% of the annual Kentucky bluegrass production on Juniper Slope. This permitted 70.7% forage carry-over in the f a l l of 1969, an increase of 46% over that of Table 6.2.1. Comparison of seasonal forage.utilization in the Poa pratensis eommunity, Juniper Slope in 1968, by cattle and wildlife and in 1969 by wildlife only. Cattle and Wildlife Use Accumulative Seasonal Forage Utilization 1968-1969 Cattle-Deer Use Spring to Sept. 18/68 Fall Cattle Use Oct 9/68 Winter Sheep Use Oct to April 22 Weathering Loss Spring Carry-over April 22/69 Percent of annual production removed Percent forage carry-over 71.3 28.7 75.7 24.3 80.3 19.7 89.3 10.7 Wildlife Use Only 1969-70 Spring Wildlife use only Oct 6/69 Winter use and weathering loss Spring Carry-over May 14/70 Percent of annual production removed• Percent forage carry-over - Standing forage only - Total forage carry-over 29.3 70.7 67.7 6.7 32.3 137 1968. Winter1 weathering loss and over-wintering sheep use, measured on May 14, 1970, totalled 67.7% of the 1969 annual production. The over-winter weathering losses were possibly increased due to the amount of forage remaining on Juniper Slope in the f a l l . Of the 1969 annual production 6..7% remained in May as standing forage; whereas the total forage carryover was 32.3%. Figures 6.2.1 and 6.2.2 illustrate the difference in 1968 and 1969 forage carryover in the f a l l on Juniper Slope. Wildlife grazing alone did not make efficient use of the forage produced during the 1969 growing season, especially on Juniper Slope. The forage remaining in the Poa pratensis community in the f a l l has•a low nutritive value for wildlife winter use and muchcf i t , 54%, was not available due to collapse and other over-winter weathering losses. In the spring the depressed forage could be'nutritionally detrimental as i t makes the succulent new forage more difficult to obtain. However, the solution to this problem does not l i e in returning cattle grazing to the range, even at a reduced rate, as any degree of utilization would likely further retard significant changes in species composition of the deteriorated portions of the range (McLean and Tisdale, 1972). Results of the spring and summer forage clipping studies at the four exclosure plots on the Flatiron Mountain winter-spring ranges was presented in Table 5.2.8. Forage clippings conducted on June 12, shortly after the deer had departed from the winter-spring ranges, indicated that spring forage utilization by deer on the Poa pratensis and Stipa-Artemesia communities was less than the increase in forage Figure 6.2.2. Forage remaining on Juniper Slope i n the f a l l of 1969 after spring grazing by w i l d l i f e only. 139 production due to the stimulatory affect of light early spring grazing. The increase in forage productivity made i t impossible to estimate the average forage consumption by deer during the spring. Forage clippings conducted on June 24, shortly after the major portion of the sheep population departed from the winter-spring ranges, indicated that the rate of forage utilization by sheep alone or in combination with mule deer on the Agropyron spicatum community was intensive enough to exceed the rate of forage production and hence reduce the standing crop of forage. It would be possible to estimate average utilization by sheep from the numbers observed in the one acre census plot and the forage clippings conducted during the short early spring grazing t r i a l . However, this was not done in view of the .' interaction between utilization and the stimulatory affect of light spring grazing indicated from1 results: of'" the;:mule: deer'grazing, t r i a l . After departure of the native ungulates from the winter-spring ranges forage production continued until soil.moisture declined and remained below the permanent wilting point. This occurred at a depth of 10 cm. by July 27 on the Poa pratensis community and at a depth of 20 cm. by July 29, 1969, on the Agropyron spicatum community. Forage clippings conducted at the four sites at this time were used to measure the effect of spring grazing by deer only, sheep only and deer and sheep combined on subsequent forage production. Spring use could affect the amount of forage potentially available for the sheep popula-tion wintering on Flatiron Mountain. The effect of grazing has been illustrated on the basis of differences in growth.rates between the grazed and ungrazed areas subsequent to spring grazing and differences 140 in forage yields at the time of .iraximum forage production (Figure 6.2.3). Deer-only spring grazing on the S.tipa-Artemes ia community has no significant affects on the forage growth rates or the forage yields at the time of maximum forage production. Whereas, on the Poa  pratensis community early spring grazing by deer-only significantly reduced the subsequent forage growth rate in relation to that of the protected or ungrazed area, but there was no measurable effect on the amount of forage produced at the time of maximum forage production. This indicates: (1) that the stimulatory affect of early spring grazing on forage production was offset by a subsequent compensating reduction in productivity, and (2) that forage removal in the early spring by deer did not significantly affect the amount of forage potentially available for the wintering sheep population on Flatiron Mountain. The lack of significant reductions in the observed amount of forage remaining on the grazed areas at the time of maximum production indicates that the grazed areas actually produced more forage than the protected areas on Juniper Slope. Blood (1961, p. 60-61) in reference to an observation of 144 deer grazing on Juniper Slope in May 1961 states: "Unfortunately this grazing (by deer) occurs during the early growth.stages when plants are damaged most easily. Such heavy spring use may be in part responsible for the advanced successional stages of such ranges as Juniper Slope and Starvation Flats, and certainly cannot be ignored as a long term factor in bighorn sheep range compe-tition". Figure 6.2.3. Effect of spring grazing on subsequent winter-spring range forage growth rates, Flatiron Mountain, 1969. 200 150 * 100 •9 50 Growth rates DEER Poa pratensis grazed area; 200 protected area 126.6 J ^ June 12 CD bO July 30 200 0) 150 i c 8J o 100 50 DEER Stipa-Artemesia June 12 July 30 150 100 50 200 SHEEP Agropyron spicatum 18% 141.9 120 o 18%-^ < O 119.0 J June 23 150 100 50 August 1 T87.9 128.4 DEER and SHEEP . Agropyron spicatum -p June 23 DATE OF FORAGE CLIPPING August 1 142 Data from the deer grazing.study discussed above refute Blood's supposition and indicate that light spring grazing by deer on Juniper Slope may increase forage production. This is further supported by results of the "simulated cattle grazing" clipping study which showed no significant reduction in forage production or plant vigor in a Poa  pratensis community subjected to a much heavier rate of forage removal during the spring and.summer. On the Agropyron spicatum community grazed by sheep,, only ." the* ' .. growth rates of the grazed and ungrazed areas were not signifi-cantly different; there was a significant reduction in the amount of forage remaining on the grazed areas at the time of maximum production. Similar results were observed in the Agropyron spicatum community grazed in common by deer and sheep. Thus, early spring grazing by sheep, or sheep in combination with deer, significantly reduces the amount of forage remaining in the Agropyron spicatum community for use by sheep wintering on Flatiron Mountain. In reference to grazing on the bluebunch wheatgrass range Blood (1961, p. 49) states that: "The short period of grazing by sheep on spring growth on South Slope should not materially affect the competition relationship". This statement appears to be incorrect on the basis of observed effects of spring grazing by sheep, and deer or sheep only on forage production, that is the standing crop of usable forage, available for winter sheep use in the Agropyron spicatum community, South Slope. Demarchi (1965, p. 74) states that: 143 "It is likely that grazing by mule deer is partly responsible for the maintenance of serai communities as the heaviest deer use occurs while the forage is emerging from winter dormancy". This may be true in part and cannot be concluded definitely from results of the present study in that i t was not possible to locate an area of deer-only spring grazing to study the effect of that use on the productivity of an Agropyron spicatum community on South Slope. However, spring grazing of the Agropyron spicatum community by deer does not appear to be more detrimental than that by sheep because the area grazed in common by sheep and deer was not more severely affected than that grazed, by sheep alone (Table 5.2.8). The-apparent difference in the effect of spring grazing by sheep and that by deer may not occur as a result of the ungulate species involved but rather the physiology and growth form of the species of forage grazed. Deer tend to utilize areas dominated by bluegrass and needle grass rather than bluebunch wheatgrass, as indicated by studies of animal distribution and community preferences not presented herein in detail. The frequency of deer use of sod grass versus bunch grass domin-ated areas on various slopes was found to be: Percentage by community type Location Animals observed Sod grass Bunch grass Starvation Slope Juniper Slope South Slope 617 787 389 80 68 26 32 20 144 6.2.2 Forage measurements on the alpine summer range Mean dry matter production and utilization measured in the Carex  Danthonia-Carex alpine summer range communities is presented in Table 5.2.9. Forage production measured in the Carex community, at 7400 f t . elevation, was greater than that measured on the Stipa-. Artemesia. and Poa pratensis communities at approximately 5600 f t . elevation on the Flatiron Mountain winter-spring ranges. Forage pro-duction in the Danthonia-Carex community, at 7100 f t . was greater than that of the two winter-spring range communities mentioned above and one of the two Agropyron spicatum community sites studied. The productivity of the two alpine sites is•comparable to other alpine communities studied in B.C., but the nitrogen levels were generally lower than those observed for B.C. .coastal alpine (Brink et a l . , 1972). However, the crude protein levels of the two alpine sites exceed the levels observed from three communities on the winter-spring ranges of the Ashnola on August 30, 1967. (Harper, 1969). 6.2.3 Limitations of range forage appraisal 6.2.3.1 Forage. production and Utilization ' Martin (1970) suggests that there is a poor correlation between measurements of forage production and utilization on the range and.the forage consumed by the animals, especially when forage•pro-duction and utilization are low and the utilization occurs over a long grazing period. Forage use and the effect of use by rodents and invertebrates, was not measured independently and was assumed to'be equal inside and outside of the fenced areas. 145 The annual forage losses due to maturation and erosion . (weather-ing) from the time of maximum production to the f a l l , and.from f a l l to the spring were measured at some sites and the losses were found to vary with the community and quantity of forage involved. Temporal and spatial overlap, that is synchronous use of the range, by the three ungulate species prohibited the use of standard range use evaluation techniques. Forage growth and the effects of grazing on growth during the spring and summer grazing period complicates utilization estimates. To minimize this, forage growth during the grazing interval and the effect of grazing on forage production were measured. However, during what is believed to be a c r i t i c a l period in the early spring the amount of new green forage.is relatively very small and use is complicated by rapid growth. 146 7. RECOMMENDATIONS Acquisition of the bighorn winter-spring ranges of Crater and Flatiron Mountains by the Provincial Government (B.C. Fish and Wild-l i f e Branch) was a positive management step to maintain or improve the present herd of California bighorn in the Ashnola River Valley. In -that the geographical distribution of the bighorn is more restricted than that of the mule deer in B.C., i t is sound practice to manage the Ashnola ranges primarily for bighorn. Habitat management and acquis-ition is the key to long term maintenance of viable populations and i t should be based on sound fundamental information. For sound management a better understanding of habitat require-ments must be achieved. Extensive knowledge of the use, by domestic and wildlife species, of the niches they occupy and of the effect of their use on the habitat and its use by other species must be achieved. In the future i t may be possible and necessary to limit the numbers of specific ungulates using a range to attain optimal ecologically balanced range use. As discussed in Section 6.1.3, ecological animal equivalence values should be based upon more than the metabolic equiva-lence of species concerned. If the Ashnola ranges are to be managed for the production of bighorn then they should be managed to promote high production of the bighorn's staple winter forage, bluebunch wheatgrass. Such a program may possibly reduce spring use by mule deer of the grassland range as they seek serai grasses and forbs in spring. Restoration of the over-grazed portions of the winter ranges, as -suggested by Demarchi (1965), 147 may .....result . in an increase in the bighorn herd. The present forage studies indicate that optimum use of the annual forage production was not obtained by wildlife grazing only. However, use of cattle grazing, as previously allowed, as a management tool to improve range quality for wildlife by reduction of the standing crop of "less valuable" forages would probably be detrimental to the regeneration of bluebunch wheatgrass. Cattle grazing in the 1968 season included spring (May 24 to June 15-18) and f a l l (September 20 to 29 or October 2) use of South Slope, and excessive use on the other slopes as measured on Juniper Slope, between May 8 and July 8. Due to the remote nature of the Flatiron Mountain ranges use of short duration intensive cattle grazing during specific seasons is not economically feasible. To improve the productivity of bluebunch wheatgrass in a reasonable time span on at least some of the present range communities, may require implementation of range improvement techniques, not just cessation of cattle grazing. South Slope,. Juniper Slope, Starvation Slope and Flatiron Slope function as the major bighorn winter ranges during the "average" winter. They are not, however, the "critical winter" ranges in that a heavy snow pack, such as that of 1968-69, reduced the number of bighorn wintering on South Slope from 157 individuals on December 15, 1968 to 52 by early January and to 49 individuals by February 22, 1969. The majority were forced to move down the valley-to the area of Gillanders Greek and beyond. For this reason more serious management consideration should be given to the South and Southwest facing grass-lands in the lower, portions of the Ashnola Valley. 148. Following the removal of cattle grazing from the major winter range, South Slope, i t appears as though grazing by bighorn during the spring and summer is the major biotic factor affecting the amount of forage available to wintering bighorn. This is especially true of the. bighorn which spend the, majority of the summer on South Slope and utilize the bunchgrass from the ridge areas important for wintering bighorn. Removal of animals summering on the winter ranges should be considered after determining their distinction as an entity within a larger migratory population. In part, innate environmental factors are responsible for the limitations on the Flatiron Mountain wildlife populations. The effect of the habitat, as i t does or does not satisfy the requirements of the animals, and innate environmental factors such as snowpack and spring weather conditions should be evaluated. The absence of many preferred browse species on the Flatiron Mountain winter ranges may be limiting the deer population. However, direct eivdence of this, such as an increase in winter k i l l following the moderately severe 1968-69 winter, was not observed. In the case of the bighorn, possibly a change in wintering and lambing range elevations has occurred as a result of alienation of lower elevation ranges. A change in elevation could adversely affect sheep reproduction (1) directly through poor weather conditions during the early lambing period, as was observed on May 13, 1970, when up to 5 inches of snow covered the ranges, or (2) indirectly through timing of the availability of nutritious spring forage in relation to the increased nutrient requirements during the 149 latter one third of the gestation period. Lack of high quality forage at this cr i t i c a l time may reduce the number- of live births or delay milk production at the time of birth. The effect of environmental factors should be investigated as they may help to explain the apparently low spring lamb:ewe ratios observed on South Slope in relation to the 100% pregnancy rates observed in ewes retained at the Okanagan Game Farm, Penticton, after being captured on South Slope during- the early winter of the same year. 150 8. SUMMARY 8.1 Habitat utilization by mule deer in relation to California big-horn sheep and cattle was studied on the winter-spring ranges of • Flatiron Mountain from January, 1968, through November, 1969. Techniques used to evaluate habitat utilization and possible range competition included: (1) food habits of the three species, (2) measurement of forage production and utilization, (3) the effect of spring utilization on subsequent forage production and (4) temporal and spatial distribution of the ungulates on the range.< Only data from the f i r s t three factors is presented in the dissertation. 8.2 Cattle utilization of the Crater and Flatiron Mountain ranges was unexpectedly terminated following the 1968 grazing season. Thus it'was impossible to obtain information on forage utilization by cattle comparable to that for deer and sheep.' 8.3 Measurement of forage production, utilization and the effect of utilization was conducted in the spring and summer on the grasslands rather than the forested range since the bighorn prefer and rely upon the grasslands, mule deer utilize the ranges in the spring (to June 18 in 1968 and June 7 in 1969) and cattle utilize them during the spring (May 8 to'July 8, 1968) and f a l l (September 18 to October 2, 1968). 8.4 Total forage losses in the Poa pratensis community on Juniper Slope, including late summer weathering losses (33.1%; Harper, 1969) and use'by cattle and wildlife from spring until September 18, 1968, resulted in a 71.3% reduction in the standing forage crop. Further 151 losses by f a l l cattle grazing, over winter sheep grazing and winter weathering losses contributed to the 89.3% removal of forage produced in 1968. 8.5 In 1969, spring use, primarily by mule deer,.oftrePoa pratensis community, Juniper Slope, resulted in a 29.3% reduction of the standing forage crop in the f a l l . ' '' 8.6 Forage carryover in the Poa pratensis community, Juniper Slope, was 28.7%.in the f a l l of 1968 and 10.7% in the spring of 1969 as a result of combined cattle and wildlife grazing.; Forage carryover was increased to 70.0% in the f a l l of 1969 and 32.3% total forage carryover in the spring of 1970 as a result of only wildlife use, primarily deer spring grazing and sheep f a l l grazing, during the 1969 production year. 8.7 Wildlife grazing alone did not make efficient use of the forage produced in 1969, however, the return of cattle grazing, even at a reduced intensity, under the grazing system previously allowed would likely retard significant "improvements" in species composition of the bighorn winter range. 8.8 Spring forage utilization by deer on the Poa pratensis and Stipa-Arternesia communities, measured at the time deer departed from the range, was less than the increase in forage production due to the stimulatory effects of light spring grazing. At the time of maximum forage production no significant reduction in the standing forage crop was measurable at the Stipa-Arternes i a or Poa pratensis communities as a result of spring grazing by deer. 152 8.9 Spring forage utilization by sheep alone or in combination with deer on the Agropyron spicatum community, measured at the time sheep departed from the range, was intensive enough to exceed the rate of forage production and hence reduce the standing forage crop. Signifi-cant reductions were also measured at the time of maximum forage production on the same areas. Thus early spring grazing by sheep or sheep and deer in combination significantly reduce the amount of forage remaining in the Agropyron spicatum community for use by sheep wintering on South Slope. 8.10 Deer.and sheep utilization of the alpine summer range in the Joe Lake area had not significantly reduced the. standing forage crop in late August, 1969. Forage production of the Carex and the Danthonia- Carex communities was found to exceed that of the lower elevation winter-spring range communities with the exception of that from one Agropyron  spicatum study site. The percent crude protein levels (7.8 and 8.7% respectively) exceed those observed by Harper (1969) from three winter-spring range communities on Flatiron Mountain on August 30, 1967. 8.11 The spring diet of cattle on Juniper Slope consisted of 97.5% grass, 1.7% forb and 0.8% browse as indicated from rumen sample analysis. The general absence of shrubs on the Flatiron Mountain ranges restricts cattle to a diet of grass and forb. Chamaephytes, such as pasture sage, were not utilized by cattle during the spring. 8.12 The spring diet of bighorn on South Slope consisted of 91.8% grass, 7.0% forb and 2.1% browse as indicated from rumen sample analysis. 153 Buckwheat (Eriogonum heracleoides) constituted the observed browse utilization. Grazing observations conducted in early July 1969, in the absence of cattle and a l l observed deer grazing, indicated the frequency of use on grass (bluebunch wheatgrass, junegrass and blue-grasses primarily) was 68.6%, forb was 23.8% and browse (primarily pasture sage) was 7.7%. 8.13 The spring diet of mule deer consisted of 38.0% grass, 7.7% forb, and 53.8% browse as indicated from rumen sample analysis. There was a significant increase in use of forb in the latter part of the spring period. Douglas f i r was the most important browse species, (43.2%) of the spring diet. Pasture sage and buckwheat formed only 2.1% and 1.6% of the spring deer diet. Grazing observations conducted in mid June, 1969, on Juniper Slope indicated that the frequency of use on grass (primarily Poa pratensis and P. secunda) was 66.7%, forb 25% and browse 8..5% of the spring diet. Spring browse utiliza-tion was observed primarily on pasture sage in the Artemesia community on Juniper Slope; however, only 16%-of the pasture sage plants observed had been browsed. 8.14 • The f a l l diet of mule deer consisted of 17% grass, 23.4% forb and 59.6% browse, from rumen sample analysis. Lodgepole pine, big sage and bearberry were the major browse species used by deer in a lodgepole pine area from which the f a l l samples were collected. '8.15 The winter diet of mule deer consisted of 12.9% grass, 19.7% forb and 66.8% browse. Deer spent the majority of the winter in the 154 Douglas f i r community, especially i f the grasslands were snow covered. The major constituent of their diet at this time was needles and bough tips of Douglas f i r (52.9%). 8.16 Animal equivalence ratios were calculated for the three ungulate species using the Flatiron Mountain ranges from metabolic equivalence and food habit data. In the spring, using the technique outlined, 12.9 deer or 5.4 sheep are equivalent to one A.U.M. ,• and 2.4 deer are equivalent to one sheep unit month (S.U.M.). In the winter, 6.7 deer are equiva-lent to one S.U.M. 8.17 The decadent condition of f i r regeneration, especially in the area north of Starvation Slope, indicates o. heavy - use of this species. This may result in intraspecific deer competition on the Flatiron Mountain winter .ranges. 8.18 Spring and summer utilization of Agropyron spicatum on South Slope, especially by the herd of bighorn summering on that slope may contribute to interspecific sheep competition during the winter on the winter'.ranges,. 8.19 Cattle utilization of the winter-spring ranges exceeded the "proper" level. As such, i t contributed to interspecific cattle-deer competition on the grasslands in the spring; and cattle sheep compe- . tition on the grasslands during the winter. 8.20 In the absence of cattle grazing, observations indicate that interspecific deer-sheep competition during the spring is very light. 155 In the spring the two ungulate species prefer and utilize different forage species and different communities on the open grasslands. 8.21 The possibility of interspecific deer-sheep competition arising during the winter is slight, as a result of differences in food habits and habitats occupied. However, deer w i l l make greater use of the grasslands during winter i f they are relatively free of snow as in 1968, than during a year of heavier snowpack as was observed in 1969. 156 9. LITERATURE CITED Beament, J.W.L. 1961. The role of physiology in adaptation and competition between animals. In,'Mechanisms in Biological Competition'! Symposia of the Society for Experimental Biology. XV: 40-61. Bergerud, A.T. and L. Russell. 1964. Evaluation of rumen food analysis for Newfoundland caribou. J. Wildl. Mgmt. 28: 809-814. Blood, D.A. 1961. An ecological study of California Bighorn Sheep. M.Sc. Thesis, Univ. of British Columbia. Blood, D.A. 1967. Food habits of the Ashnola Bighorn sheep herd. The Canadian field-naturalist, 81: (1) 23-29. Boeker, E.L., V.E. Scott, H.G. Reynolds, B.A. Donaldson. 1972.-Seasonal food habits of mule deer in Southeastern New Mexico. J. of Wildl. Mgmt.. 36 (1): 56-63. Brian, M.V. 1956. Segregation of species of the ant genus Myrmica. J. of Animal Ecol. 25:. 319-337. Brink, V.C. and L. Farstad. 1949. The physiography of the agricultural areas of British Columbia. Sci. Agric. 29: 273-301. Brink, V.C, A. Luckhurst and D. Morrison. 1972. Productivity esti-mates from Alpine Tundra. Can. J. Plant Sci. 52: 321-323. Brown,. D. 1954.. Methods of Surveying and Measuring Vegetation. Bull. 42. Commonwealth Bureau of Pastures and Field Crops. Franham Royal, Bucks., England, 223 pp. Buechner, H.K. 1947. Range use of the Pronghorned antelope in western Texas. T.N.A.W.C. 12: 185-192. Buechner, H.K. 1960. The bighorn sheep in the United States; its past, present and future. The Wildl. Society, Wildl. Monog. No. 4. 174 p. Capp, J.C. 1968. Bighorn sheep, Elk, Mule Deer Range relationships -A review of literature. Rocky Mt. Nature Association and Colorado State University. 157 Chamrad, A.D. and T.W. Box. 1964. A point frame for sampling rumen contents. J. Wildl. Mgmt. 28: 473-477. Chamrad, A.D. and T.W. Box. 1968. Food habits of White-tailed deer in south Texas. J. Range Mgmt. 21 (3): 158-164. Chapman, H.D. and P.F. Pratt. 1961. Methods of analysis for soils, plants and waters. University of Calif., Division of Agric. Sci., Chapter 17. pp. 150-152. Clements, F.E. and V.E. Shelford. 1939. Bio-ecology. J. Wiley and Sons, N.Y. 425 p. Coblentz, B.E. 1970. Food habits of George Reserve deer. J. Wildl. Mgmt. 34 (3): 535. Cole, G.F. 1958. Big game-livestock competition on Montana's mountain rangelands. Mont. Wildl. Apr.. Cook, C.W. , L.E. Harris, and M.C. Young. 1967. Botanical and nutritive content of diets of cattle and sheep under single and common use on mountain range. J. Animal Sci. 26: 1169 Cook, C.W. and L.H. Stoddart. 1953. The quandry of utilization and preference. J. Range Mgmt. 6: 329-335. Cook, C.W., L.A. Stoddart, L.E. Harris. 1953. Effects of grazing intensity upon the nutritive value of range forages. J. Range Mgmt. 6: 51-54. Cooperrider, A.Y. 1967. Competition for food between mule deer and bighorn sheep on Rock Creek winter range, Montana. M.Sc.-Thesis. Univ. of Montana. 92 p. Cowan, I. McT. 1940. Distribution and variation in the native sheep of North America. American Midland Naturalist. 24 (3); 505-580. Cowan, I. McT. 1947. Range competition between mule deer, bighorn sheep and elk in Jasper Park, Alberta. T.N.A.W.C. 12: 223-227. Cowan, I. McT. 1956. What and where are the mule and black-tailed deer? Ih"The Deer of North America'.' W.p. Taylor. Ed. 668 p. Cowan, I. McT. and C.J. Guiguet. 1965. The Mammals of British Columbia. B.C. Provincial Museum, Handbook No. 11. 414 p. 158 Couey, F.M. 1955. Montana bighorn sheep. Proc. West. Assoc. of State Game and Fish Com. 35: 62-56. Dasmann, W.P. 1949. Deer-livestock forage studies on the interstate winter deer range in California. J. Range Mgmt. 2: 206-212. Demarchi, R. 1965. An ecological study of the Ashnola bighorn winter ranges. M.Sc. Thesis. University of British Columbia. De Vos, A. 1969. Ecological conditions affecting the .production of wild herbivorous mammals on grasslands. Advances in Ecological Research 6: 137-183. Ed. J.B. Cragg. Dice, L.R. 1952. Natural Communities. .Ann. Arbor: Univer. of Michigan Press. 547 p. Dorn, R.D. 1970. Moose and cattle food habits, on southwest Montana. J. Wildl. Mgmt. 34: 559-564. Dorrance, M.J. 1966. • A literature review on behavior of mule deer. Special Report No. 7, State of Colorado. Drawe, D.L. and T.W. Box. 1968. Forage ratings for deer'and cattle on the Welder Wildlife Refuge. J. Range Mgmt..21 (4): 225-228. Edwards, CE. 1893. Camp-fires of a Naturalist. Sampson Low, Marston and Co., London. Einarsen, H.S. 1965. Life of the mule deer. In"The Deer of North America!' N.P. Taylor ed. The Stackpole Co. and the Wildl. Mgmt. Institute Pub. 668 p. Elton, CS. 1927. Animal Ecology. The Macmillan Co., New York. (3rd Ed. 1947). .Flook, D.R. 1964. Range relationships of some ungulates native to Banff and Jasper National Parks, Alberta. Reprint from Grazing in Terrestrial and Marine Environments. Blackwells . Scientific Publications. Freyman, S. and A.L. van Ryswyk. 1969. Effect of fertilizer on Pinegrass in southern B.C. J. Range Mgmt. 22 (6): 390-395. 159 Gait, H.D. , P.R. Ogden, J.H., Earenreich, W.H. Hale and S.C. Martin. 1968. Estimating botanical composition of forage samples from fistulated steers by a microscope point method. J. Range Mgmt. 21: 397-401. Gait, H.D. , B. Theurer, J.H. Ehrenreich, W.H. .Hale and S^ C„ Martin. 1969. Botanical composition of diet of steers grazing a desert grassland range. J. Range Mgmt. 22 (1): 14-19. Gross, J.E. 1970. Competition and carrying capacity measurements: a credibility gap? Paper presented at The Wildlife Confer-ence Meeting, Victoria. Harper, F.E. 1969. Effects of certain climatic factors on the pro-ductivity and availability of forages on the Ashnola bighorn winter ranges. M.Sc. Thesis. University of British Columbia. Harris, R.W. 1954. Fluctuations in forage utilization on Ponderosa Pine ranges in eastern Oregon. J. Range Mgmt. 7: 250-255. Heady, H.F. 1964. Palatability of herbage and animal preference. J. Range Mgmt.. 17: 76-82. H i l l , R.R. 1965. Forage, food habits and range management of mule deer. In "The Deer of North America". W.P. Taylor, ed. The Stackpole Co. and the Wildl. Mgmt. Institute Pub. 668 p. Hitchcock, A.S. 1950. The manual of the grasses of the United States. 2nd ed. U.S.D.A. Misc. Publ. No. 200. Hitchcock, C.L., A. Cronquist, M. Ownbey and J.W. Thompson: 1955, 1959, 1961, 1964, 1969. The vascular plants of the Pacific Northwest. In five parts. University of Washington Press, Seattle and London. Ho, H.K. , T. Sakai, M.B. Jones and W.M. Longhurst. 1967. Effects of various essential oils isolated from Douglas f i r needles upon sheep and deer microbial activity. Applied Micro-biology 15 (4): 777-784. Holland, S.S. 1964./ Landforms of British Columbia, a physiographic outline. Bull. 48. B.C. Dept. of Mines and Natural Resources. Joint Committee of the Amer. Soc. of Range Mgmt. and the Agric. Board. 1962. Basic problems and techniques in range research, p. .318. Nat. Academy of Sciences- Nat. Res. Council. Pub. No. 890. 341 p. 160 Jones, D.A., W.L. Robinette and 0. Julander. 1956. The influence of . summer range condition, on mule deer reproduction in Utah, Proc. West. Assn. Game and Fish Comm. 35: 177-181. Julander, 0. 1955. Deer and cattle range 'relations in Utah. For. Sc. 1: 130-139. Julander, 0. 1958. Techniques in studying competition between big game and livestock. J. Range Mgmt. 11: 18-21. Julander, 0. • 1962. Range management in relation to mule deer habitat and herd productivity in Utah. J. Range Mgmt. 15: 278-281. Julander, 0. and W.L. Robinette. 1950. Deer and cattle range relation-ships on Oak Creek range in Utah. J. Forestry 48: 410-415. Julander, 0., W.L. Robinette and D.A. Jones. 1961. Relations of summer range condition to mule deer herd productivity. J. Wildl. Mgmt. 25: 54^60. Kelsall, J.P. 1968. The•migratory barren ground caribou of Canada. Dept. of Indian Affairs and Northern Dev., Canadian Wildl. Service. Leckenby, D.A. 1967.' Ecological study of mule deer: deer ecology study. Job completion report, State of Oregon. Project No. W-53-R-8. Job No. 1. Lesperance, A.L. and P.T. Tueller. 1969. Competitive uses of Nevada's range forage by livestock and big game. RNR. Reports 1969 reviewed in J. Range Mgmt. 22 (3):' 195 and 215. Lesperance, A.L. , P.T. Tueller and'V.'R. Bohman. 1970. Symposium on pasture methods for maximum production in beef cattle: competitive use of the range forage resource. J. Animal Sci. 30 (1): 115-121. Lovass, A.L. 1958. Mule deer food habits and range use, Little Belt Mountains, Montana. J. W. Mgmt. 22: 275-283. McCullough, D.R. and R.E. Schneegas. 1966. Winter observations on the Sierra Nevada Bighorn sheep. Calif. Fish and Game. 52 (2): 68-84. Mackie, R.J. 1970. Range ecology and relations of mule deer, elk and cattle in the Missouri River breaks, Montana. Wildlife Mono-graphs No. 20. 161 McLean, A. 1967. Beef production on Lodgepole Pine- Pinegrass range in southern British Columbia. J. Range Mgmt. 20 (4): 214-216. McLean, A., S. Freyman, J.E. Miltimore and D.M. Bowden. 1969. Evaluation of pinegrass as range forage. Can. J. Plant Sci. 49: 351-359. McLean, A. and L. Marchand. 1968. Grassland ranges in the southern interior of British Columbia. Can. Dept. of Agric., Pub. 1319, 28 p. McLean, A. and E.W. Tisdale. 1972. Recovery rate of depleted range. sites under protection from grazing. J. Range Mgmt. 25 (3): 178-184. McMahan, CA. 1964. Comparative food habits of deer and three classes of livestock. J. Wildl. Mgmt. 28: 798-808. Martin, A.C. , R.H. Gensch and CP. Brown. 1946. Alternative methods in upland game bird food analysis. J. Wildl. Mgmt. 10: 8-12. Martin, S.C 1970. Relating vegetation measurements to. forage consump-tion by animals, p. 93-100. Range and Wildlife Habitat Evaluation, a Research Symposium. Arizona, 1968. U.S. Dept. of Agric. Misc. Pub. No. 1147. 220 p. Medin, D.E. 1970. Stomach Content Analysis: collections from wild herbivores and birds. Range and Wildlife Habitat Evaluation: A Research Symposium. U.S.D.A. , Forest Ser. Misc. Pub. No. 1147: 133-145. Miller, R.S. 1967. Pattern and process in competition. Adv. Ecol. Res. 4: 1-74. Milne, A. 1961. Definition of competition among animals. In "Mechanisms in- Biological Competition'.' Symposia of the Society for Experi-mental Biology. XV: 40-61. Morgan, J.K. 1968. Rocky Mountain Bighorn Sheep Investigation. Job Completion Report, W-85-R-18, Job.No. 11. Idaho Fish and Game Dept., Boise, Idaho. Morris, M.S. and J.E. Schwarty. 1957. Mule deer and elk food habits, on the National Bison range. J. Wildl. Mgmt. 21: 181. 162 Nagy, J.F., H.W. Steinhoff and G.M. Ward. 1964. Effects of essential oils of sagebrush on deer rumen microbial function. J. Wildl. Mgmt. 28: 785-790. Nellis, C.H. and R.L. Ross. 1969. Changes in mule deer food habits associated with herd reduction. J. Wildl. Mgmt. 33 (1): 191-195. Norris, J.J. 1943. Botanical analysis of stomach contents as a method of determining forage consumption of range sheep. Ecology 24: 244-251. Odum, F.P. 1959. Fundamentals of Ecology. 2nd Ed. W.B. Saunders Co., London, 546 p. Phillipps-Wooley, C. 1888. A Sportman's Eden. Richard Bentley and • Son, London. Phillipps-Wooley, C. 1894. Big Game Shooting. Vol. 1. Longmans, Green and Co., London. Reynolds, H.G. 1966a. Use of Ponderosa Pine forest in Arizona by deer, elk and cattle. U.S. Forest Service, Rocky Mountain For. and Range Exp. Sta. Res. Note RM-63, 8 p. Fort Collins, Colorado. Reynolds, H.F. 1966b. Use of openings in Spruce-fir forests of Arizona by elk, deer and cattle. U.S. For. Ser., Rocky Mountain For. and Range Exp. Sta. Res. Note RM-66, 4 p. Fort Collins, Colorado. Richens, V.B. 1967. Characteristics of mule deer herds and their range in northeastern Utah. J. Wildl. Mgmt. 31 (4): 651-666. Robel, R.J. and P.G. Watt. 1970. Comparison of volumetric and point analysis procedures to describe deer food habits. J. Wildl. Mgmt. 34 (1): 210-21,3. Scheffler, E.G. •'. 1972 ;•• Appraisal of-'.ungulate habitats .'in. the Ashnola-Re source Management Unit. M.Sc.Thesis, University of British Columbia, Vancouver, B.C. Scotter, G.W. 1966. Sieve mesh size as related to volumetric and gravimetric analysis of cariboo rumen contents. Can. Field Nat. 80: 238-241. 163 Scotter, G.W. 1967. Thewinter diet of Barren-Ground cariboo in Northern Canada. Can. Field Nat. 81:. 33-39. Shepherd, D.H. 1960. The ecology of mule deer on the Sheep River region. M.Sc. Thesis. University of Alberta. 123 p. •Shepherd, W.O. 1962. Herbage sampling for yield: Natural pastures and range, p. 102-105. In "Pasture and Range Research Tech-' niques". Joint-Committee of the Amer. - Soc. of Agronomy, Amer. Dairy Sci. Association, Amer. Soc. of Animal Production and Amer. Soc. of Range Mgmt. Comstock Pub. Assoc. Ithica, New York. 242 p. Skovlin, J.M., P.J. Edgerton and R.W. Harris. 1968. The influence of cattle management on deer and elk. T.N.A.W.C. 33: 169-181. Skovlin, J.M. and R.W. Harris. 1970. Management of conifer woodland grazing resources for cattle, deer and elk. In "Proceedings of the XI International Grassland Congress. Smith-, A.D. and D.D. Doell. 1968. Guides to allocating forage between cattle and big game on big game winter range. Utah State Division of Fish and Game. Pub. No. 68-11. Smith, D.R. 1954. The bighorn sheep in Idaho, status, l i f e history and management. Idaho Game and Fish Dept. Wildl. Bull. No. 1. 154 p. Smith, D.R., P.O. Currie, J.V. Basile and N.C. Frischknecht. 1963. Methods for measuring forage utilization and differentiating use by different classes of animals, p. 93-102. Range Research Methods, a Symposium Denver, Colorado. May 1962. U.S. Dept. of Agric. Misc. Pub. No. 940, 172 p. Smith, J.G. 1952. Food habits of mule deer in Utah. J.W. Mgmt. 17: 101-112. ' Smith, T.G. and 0. Julander. 1953. Deer and sheep competition in Utah. J. Wildl. Mgmt. 17: 101-112. Spalding, D.J. 1968. The boundary deer herd. Wildl. Mgmt. Pub. No. 2. B.C. Fish and Wildl. Branch. Spalding, D.J. and H.B. Mitchell. 1970. Abundance and distribution of California bighorn sheep in North America. J.' Wildl. Mgmt. 34 (2): 473-475. 164 Snyder, R.O. 1966. Competition between big game and cattle in the Grid Creek area of Montana. Quart. Report- Montana Coopera-tive Wildl. Res. Unit. Vol. 17 (2) Spilsbury, R.H. and E.W. Tisdale. 1944. Soil-plant relationships and vertical zonation in the southern interior of British Columbia. Sci. Agric. 24: 395-436. Stoddart, L.A. and D.I. Rasmussen. 1945. Big game-range livestock competition on western ranges. Trans. North Amer. Wildl. Confer. 10: 251-256. Sugden, L.G. 1961. The California bighorn in British Columbia, with particular reference to the Chum Creek herd. Brit. Columb. Dept. Recre. and Conserv. 58 p. Taber, R.D. and R.S. Hoffmann. 1963. Behavioral adaptations of mammals to mountain environments. Proceedings, XVI International Congress of Zoology. 3: 54. Talbot, L.M. and M.H. Talbot. 1963a. The wildebeest in west Masailand, East Africa. Wildlife Monograph No. 12. Talbot, L.M. and M.H. Talbot. 1963b. The high biomass of wild ungulates of East Africa Savana. T.N.A.W.C. 28: 465 Taylor, E. 1957-1963. Monthly Reports to British Columbia Fish and Wildl. Branch. Victoria, B.C. Taylor, W.P. ed. 1965. The deer of North America. The white-tailed, mule and black-tailed deer, genus Odocoileus. Their history and management. Stackpole Co. and The Wildl. Mgmt.. Institute. Tisdale, E.W. 1947. The grasslands' of the Southern interior of British Columbia. Ecol. 28: 346-382. Tisdale, E.W. . and A. McLean. 1957. The Douglas f i r zone of southern British Columbia. Ecological Monographs. 27: 247-266. Trout, P.L. 1886. Prospector'.s Manual-Granite Creek and the Similkameen Country. Van Ryswyk, A.L. , A. McLean and L.S. Marchand. 1966. The climate, native vegetation, and soils of some grasslands at different elevations in British Columbia. Can. J. Plant Sci. 46: 35-50. 165 Vesey-Fitzgerald, L.D.E.F. 1965. The utilization of natural pastures by wild animals in the Rukwa Valley, Tanganyika. East African Wildl. J. J. .3: 38-148. Wagnon, K.A. 1963. Behavior of beef cows on a California range. Calif. Agric. Exp. Station Bull. 799. Wilkins, B.T. 1957. Range use, food habits and agricultural relation-ships of the mule deer, Bridger Mts. Montana. J.W. Mgmt. 21: 159-169. Williams, A.B. 1925. Game trails in British Columbia. Charles Scribner's Sons, N.Y.. Wishart, W.D., 1958. The bighorn sheep of the Sheep River Valley. M.Sc. Thesis. University of Alberta. 66p. Yeager, L.E. 1960. Factors affecting the quality and management of big game winter range in Colorado. State of Colorado. Dept. of G. £ F. Report No. 30. Appendix 4.2.1. Paired plot design for measurement of forage utilization by cattle in the f a l l of 1968, and sheep in the winter of 1968-69. I I II •39.5" quadrat I I II. I I 10 II 12 13 I I II Clipping treatment: I Before grazing. II After f a l l or winter grazing. Appendix 4.2.2. Field plot design for forage production and utilization measurements. AREA C AREA A AREA B TREATMENT NUMBER IN ONE METER SQUARE QUADRAT FENCE Appendix 4.2.3. Field plot design for measure-ment of alpine forage production and u t i l i zation, 1969. I , I 1 m -FENCE II II II. •39.5" quadrat II II II. II II 169 Appendix 5.1.1. Hereford cattle spring food habits, as indicated from rumen sample analysis, in gms. Sample 1968' -V. Number . 17 \ ... 18 J Species Totals Species " June 3 June 3 GRASS 1. Mature 2. Green 1.3 2.8 1.0 6.6 2.3 9.4 Subtotals 4.1 7.6 11.7 FORBS 1. Achillea millefolium 2. Unknown sp. 3. Zigadenus (venenosus) 0.1 Tr 0.1 Tr Tr 0.2 Tr Subtotals 0.1 0.1 0.2 BROWSE 1. Pinus contorta 2. Unknown sp. Tr ' 0.1 0.1 Tr Subtotals Tr 0.1 0.1 UNIDENTIFIABLE FIBER 14.0 13.1 27.1 Appendix 5.1.2. Bighorn sheep spring food habits as indicated by rumen sample analysis, in gms. -". Species c , 1968 1969-2* Sample Species Number 15 16 19 Total May 30 June 1 June 10 March"'" 7 GRASS 1. Mature 0.4 0.6 5. 7 6.7 -2. Green 2.0 4.3 0. 2 6.5 Total 2.4 4.9 5. 9 13.2 13.1 FORBS 1. Achillea millefolium _ 0.1 0.1 _ • 2. Lupinus sp. - 0.3 0.3 -3. Phacelia (hastata) - 0.2 0.2 -"4. Unknown sp. Tr 0.3 0. 1 0.4 -5. Viola sp. - Tr Tr , -Total Tr 0.9 0. 1 1.0 -BROWSE 1. Artemesia frigida — — — 0.5 2. Eriogonum heracleoides - 0.3 0.3 0.7 3. Pseudotsuga menziesii - - - 0.3 4. Unknown sp. Tr - Tr -Total Tr 0.3 0.3 1.5 Unidentified fiber 11.4 11.7 15. 3 38.4 *1969-2 Winter k i l l on Juniper slope Appendix 5.1.3. Forage species occurrence and utilization by Bighorn sheep on South Slope, Agropyron community, :as/'lnoicate July 7, 1969. Species. Average percent canopy cover Frequency of occurrence No. % Frequency of grazing No. \ Percent of a l l grazed plants GRASSES 1. Agropyron spicatum 12.4 138 46 40 29.0 26.0 2. Festuca idahoensis 3.0 42 14 15 35.7 9.7 3. Koeleria cristata 4.8 113 37.7 43 38.1 27.9 4. Poa ampla Tr 1 0.3 - - -5. P. secunda 0.3 7 2.3 - • - -Subtotals • 20.5 301 55.9 98 32.6 63.6 FORBS 1. Achillea millefolium 3.4 86 28.7 12 13.9 7.8 2. Allium cernuum 0.2 14 4.7 9 64. 3 5.8 3. Antennaria rosea 1.2 13 4.3 - - -4. Arabis sp. 0.2 11 3.7 - - -5. Artemesia michauxiana 0.5 5 1.7 1 20 0.7 6. Astragulus- miser 1.0 13 4.3 9 69.2 5.8 7. Cirsium sp. 0.3 4 1.3 - - -8. Heuchera cylindricata Tr 1 0.3 1 100 0.65 9. Lappula redowskii 0.2 15 5 - - -10. Lupinus sericeus 1.9 16 5.3 11 68.8 7.1 11. Microeris alpestris Tr 2 0.7 2 100 1.3 12. Phacelia linearis Tr 5 1.7 - - -13. Potentilla gracilis Tr 2 0.7 - - -cont. Appendix 5.1.3., cont. Species Average percent canopy cover Frequency of occurrence No. Frequency of grazing No. % Percent «of_- a l l grazed plants 14. Silene scouleri Tr 3 1 1 33.3 0.7 15. Taraxicum officinale Tr 1 0.3 1 100 0.7 16. Woodsia oregana 0.2 2 0.7 - - -Subtotals 9.4 193 35.9 47 24.4 30.5 BROWSE 1. Artemesia frigida 2.5 31 10.3 4 12.9 2.6 2. Eriogonum heracleoides 0.9 13 4.3 5 38.5 3.3 Subtotals 3.4 44 8.2 20.5 5.9 SUNDRIES 1. Rock 8.2 43 14..3 2. Soil 58.6 295 98.3 rO Appendix 5.1.4. Forage species occurrence and utilization by Bighorn sheep on South Slope, Agropyron-Artemesia community, as indicated from grazed plant loop-transect data. July 7, 1969. Species Average percent canopy cover Frequency of occurrence No. \ Frequency of grazing No. % Percent of a l l grazed plants GPASSES 1. Agropyron spicatum 9.7 116 38.7 50 43.1 34.0 2. Carex sp. Tr 3 1 1 33.3 0.7 3. Festuca idahoensis 0.9 15 5 6 40 4.1 4. Koeleria cristata 4.5 102 34 44 43.1 29.9 5. Poa ampla Tr 1 0.3 - - -6. P. secunda 0.4 10 3.3 7 70 4.8 Subtotals 15.6 247 65.3 . 108 43.7 73.5 FORBS 1. Achillea millefolium 0.7 11 3.7 3 27.3 2.0 2. Allium cernuum 0.4 8 2.7 8 100 5.4 3. Antennaria rosea 0.6 10 3.3 1 10 0.7 4. Arabis sp. 0.2 6 2.0 4 66.7 2.7 5. Astragulus miser 0.4 2 0.7 - - -6. Collinsia parviflora Tr 1 0.3 - - -7. Lappula redowskii 0.4 16 5.3 - - -8. Lapinus sericeus 0.6 7 2.3 6 85.7 4.1 9. Microseris alpestris 0.1 2 0.7 2 10.0 1.4 10. Phacelia hastata Tr 1 0.3 - - -11. P. linearis Tr 1 0.3 - - -12. -ZigadenusTYehehds^ us; . 0.2 1 0.3 1 100 0.7 Subtotals 3.6 66 17.5 25 37.9 17.0 cont. Appendix 5.1.4., continued. . . Frequency of Frequency of „ , c -, -, Average percent ^ . Percent of a l l 0 . to c occurrence grazing , , , Species • canopy cover ,T „ M O grazed plants i\o. a iNo. BROWSE 1. Artemesia frigida 5.9 59 19.7 12 20.3 8.2 2. Eriogonum heracleoides 0.3 6 2 2 33.3 1.4 Subtotals 6.2 65 17.2 14 21.5 9.5 SUNDRIES 1. Soil 65 296 98.7 2. Rock 5.1 27 9 3. Litter --F 175 Appendix 5.1.5. Mule deer f a l l food habits as indicated from rumen sample analysis, in gms.: Sample 1961 3 1969 Species _ . Number Specxes 21 22 5 Total Nov. 10 Nov."30 " Oct. 9 GRASS 1. Mature 0.3 _ _ _ 2. Green 0.1 - -Subtotals 0.4 o-.i • 2.4 2..:9 FORBS 1. Achillea millefolium Tr Tr 0.5 0.-5 2. Aster sp. - - 0.2 0.2 3. Equisetum arvense • - - 0.1 0.1 4. Heuchera cylindricata - - 0.1 0.1 5. Lupinus sp. 0.3 0.8 - 1.1 6. Silene sp. - . 0.1 - 0.1 7. Stellaria sp. - - Tr Tr 8. Taraxacum officinale - - 0.1 0.1 9. Trifolium (repens) - - 0.3 0.3 10. Unknown sp. 0.2 0.1 1.2 1.5 Subtotals 0.5 1.0 2.5 4.0 BROWSE 1. Arctostaphylos uva-ursi 1.0 1.4 - • 2.4 2. Artemesia tridentata 0.1 0.6 1.4 2.1 3. Eriogonum heracleoides 0.3 - 0.3 4. Pinus contorta 0.3 ' 3.6 - 3.9 5. Pseudotsuga menziesii 0.1 - - 0.1 6. Salix sp. - 0.1 0.2 0.3 7. Unknown sp. 0.6 - 0.2 0.8 8. Vaccinium scoparium - Tr 0.3 0.3 Subtotals 2.1 6.0 2.1 10.2 Unidentifiable fiber 29.9 33.8 28.8 92.5 Appendix 5.1.6. Mule deer winter food habits as indicated from rumen sample analysis, in gms. bampie 4' 25 Number 1 2 3 5 23 24 Species Jan. 13 "Jan. 13 Feb. 16 Feb. 16 Feb. 16 Dec. 19 Dec. 20 Dec. 20 GRASS 1. Mature - - - - 0.5 - - - -2. Green - - - - . 0.1 - - -Subtotals 2.5 0.4 0.3 0.5 0.6' 2.8 0.3 0.2 7.6 FORBS 1. Achillea millefolium - — - — — 0.1 - - 0.1 2. Antennaria sp. - - 1.6 2.5 0.5 - - - 4.6 3. Capsella bursa-pastoris - - - - 2.6 - - 2.6 4. Lupinus sp. Tr Tr - - - - Tr 0.9 0.9 5. Unknown sp. 0.1 1.2 Tr 0.1 0.1 0.1 1.5 0.3 3.4 Subtotals 0.1 1.2 1.6 2.6 0.6 2.8 1.5 1.2 11.6 BROWSE 1. Artemesia frigida - - 0.1 0.1 0.1 0.2 Tr - 0.5 2. Eriogonum heracleoides - - 0.8 2.5 1.2 0.4 - - 4.9 3. Juniperus scopulorum - - 0.1 - - - - - 0.1 4. Pinus contorta - - - - - - 1.4 0.9 2.3 5. Pseudotsuga menziesii 3.0 4.0 5.1 10.3 8.4 0.4 • - - 31.2 6. Sheperdia canadensis - 0.1 - - - - - 0.1 7. Unknown sp. - 0.2 - - - - - ... 0.3 8. Vaccinium scoparium - - - - - - Tr . Tr. Subtotals 3.0 4.3 6.1 12.9 9.7 1.0 1.4 1.0 39.4 LICHENS 0.1 0.2 - - - 0.1 - 0.4 Unidentifiable fiber 5.9 14.3 6.3 9.1 10.4 6.4 18.4 23.7 94.'5 Appendix 5.1.7. Mule deer spring food habits, as indicated from rumen sample analysis, in gms. 1969 1968 Sample : Number 1* 6 7_ 8 i . _ 1° 11 . 12 13 14 20 Species Total Species 1* 6 7 8 9 10 11 12 . 13 14 20 Mar. Mar. Mar. Mar. Apr. Apr. Apr. May May May June 14 15 16 16 16 16 17 12 15 16 10 GPASS 1. Mature - 0.7 1.0 0.1 1. 6 1.4 0.8 0.6 0.6 6.8+ 2. Green — 0.1 - Tr 1.1 1.1 0. 6 2.7 0.9 0.5 1.1 8.1+ Subtotals 0.1 0.8 1.1 0.2 2.1 1.2 2. 2 4.1 1.7 1.1 1.7 16.2 FORBS 1. Achillea millefolium _ _ _ _ _ _ — Tr Tr Tr Tr 2. Allium cernuum - - - - 0.1 - - - 0.1 3. Antennaria sp. - 0.1 0.1 0.1 0.1 - - Tr - 0.4 4. Arenaria capillaris - 0.1 - - - - - - 0.1 5. Erigeron compositus - - - - - - - Tr - Tr Tr 6. Geranium viscosissimum - - - - *- - Tr - - - Tr 7. Heuchera cylindricata - 0.1 - • Tr Tr - 0.1 - 0.2 8. Microserus alpestris - - - - - - Tr - - - Tr 9. Oxytropus campestris - - - - - - 0.3 0.1 - 0.1 0.5 10. Penstenon procerus - - - - - - 0.1 - - 0.1 0.2 11. Potentilla gracilis - - - - - - 0-...1 Tr Tr 0.2 0.4 12. Ranunculus sp. - _ 0.1 - Tr Tr Tr - 0.1 13. Sisymbrium altissimum - _ - - - • - - 0.1 0.1 14. Stellaria sp. - _ - - - - Tr Tr Tr 15. Taraxacum officinale - _ - - Tr - - - Tr 16. Unknown sp. 0.1 Tr 0.1 0.1 - • - 0. 1 0.2 0.1 0.2 0.3 1.2 17. Viola sp. - - - - - - - Tr - Tr Tr Subtotals 0.1 Tr 0.2 0.3 0.2 0.1 0. 1 0.8 0.3 0.4 0.8 3. 3 "Winter k i l l Lower Ashnola Appendix 5.1.7., continued. Q = , r ™ 1 _ 1 9 6 9 1 9 6 8 bample . __ Number 1* 6 7 8 9 10 II 12 13 14 20 Species. Species Mar. Mar. Mar. Mar. Apr. Apr. Apr. May May May June Total BROWSE 1. Arctostaphylos ura-ursi - - - - - 0.7 - 0.1 Tr 0.2 - 1.0 2. Artemesia frigida - - - - - 0.1 0.1 0.4 Tr - 0.3 0.9 3. Berberis repens _ _ _ _ . _ T r _ . _ _ _ _ i r 4. Chrysothamnus nauseosus 0.9 _ _ _ _ _ _ _ _ _ _ n.9 5. Eriogonum heracleoides - 0.1 0.2 - 0.1 Tr Tr 0.1 0.1 Tr 0.1 0.7 6. Juniperus scopulorum 0.2 _ _ _ _ _ _ _ _ _ _ n.2 7. Pinus contorta - - • - - - - - - 0 . 1 0.3 - 0.4 8. Pseudotsuga menziesii 5.4 2.2 2.3 1.8 4.1 0.2 0.2 0.9 0.2 1.1 Tr 18.4 9. Ribes sp. - - - - 0.2 0.2 - - - - - 0.4 Subtotals 6.5 2.3 2.5 1.8 4.4 1.1 0.3 1.5 0.4 1.6 0.4 22.9 LICHENS _ 0.1 0.1 - - - - - - - - 0.2 UNIDENTIFIABLE FIBER 10.2 32.4 12.2 29.2 34.8 22.5 28.6 23.7 16.2 15.8 15.4 24.0 '''Winter k i l l Lower Ashnola H. CO Appendix 5.1.8. Forage species occurrence and utilization by mule deer on Juniper Slope, Poa community, as indicated from grazed plant loop-transect data. June 13, 1969. Species Average percent canopy cover in 300 loops Frequency of occurrence No. S Frequency of grazing No. % Percent of a l l grazed plants GRASSES 1. Agropyron smithii 2. Bromus tectorum 3. Carex sp. 4. Festuca idahoensis 5. Koeleria cristata 6. Poa pratensis 7. Stipa sp. 0.6 Tr* Tr Tr 1.5 27.8 4.4 11 1 1 1 34 286 100 3.7 0.3 0.3 0. 11. 95. •33. 12 121 9 27.3 35.3 42.3 9.0 1.4 5.7 57.9 4.3 Subtotals 34.4 434 64.9 145 33.4 69.4 FORBS 1. Achillea millefolium 2. Antennaria rosea 3. Arabis sp. 4. Arnica sp. 5. Aster sp. 6. Fragaria virginiana 7. Lappula redowskii 8. Lithophragma parvifiora 9. Oxytropis campestris 10. Penstemon procerus 11. Phacelia linearis 12. Potentilla gracilis 13. Rumex sp. 14. Stellaria sp. 1.0 Tr 0.2 Tr Tr 0.1 0.1 0.3 Tr 0.2 6.2 Tr Tr Tr 22 1 10 1 1 2 9 6 1 22 79 1 4 1 7.3 0.3 3 ,3 ,3 ,7 3. 0. 0. 0. 30 2.0 0.3 7.3 26.3 0.3 1.3 0.3 10 5 1 1 1 4 1 27 1 45.5 50.0 100 50 11.1 66.6 4.5 34.2 100 4.8 2.4 0.5 0.5 0.5 1.9 0.5 12.9 0.5 ID cont. Appendix 5.1.8, continued. Frequency of Percent of a l l grazing grazed plants No. % 15; Taraxicum officinale 2.2 46 15.3 8 17.4 3.8 16. Verbascum thapsus 0.1 2 0.7 - - -17. Viola adunca 0.7 21 7.0 4 19 1.9 18. Viola nut t a l l i i 0.1 1 0.3 - - -19. Forb (unknown) Tr 1 0.3 - - -Subtotals 11.2 231 34.6 63 27.3 30.1 BROWSE 1. Artemesia frigida 0.2 1 0.3 1 100 0.5 2. Eriogonum heracleoides 0.1 2 0.7 - - -Subtotals 0.3 3 1.0 1 33.3 0.5 SUNDRIES 1. Rock -2. Soil 0.2 0.3 3. Litter 53.9 99.6 *Tr.= less than 0.1 Average percent' Frequency of „ . canopy cover occurrence 1 3 6 0 1 8 3 1 1 1 3 0 0 l o°P s N o. % H CO o Appendix 5.1.9. Forage species occurrence and utilization on Juniper Slope, Artemesia community as indicated from grazed plant loop-transect data. June 15, 1969. Species Average percent canopy cover in 300 loops Frequency of occurrence No. Frequency of grazing No. %. Percent of a l l grazed plants GRASSES 1. Agropyron smithii 0.7 16 5.3 2 12.5 1.2 2. Bromus tectorum 0.2 3 1 - - -3. Festuca idahoensis Tr* 1. 0.3 1 100 0.6 4. Koeleria cristata 3.5 51 17 22 43.1 13.7 5. Poa ampla Tr 1 0.3 1 100 0.6 6. P. pratensis 0.4 8 2.7 4 50 2.5 7. P. secunda 4.2 91 30 45 49.5 27.9 8. Stipa sp. 3.8 59 . 19.7 28 47.5 ' 17.4 Subtotals 12.7 230 47.7 103 44.8 63.9 FORBS 1. Achillea millefolium 0.7 3 1 2. Allium cernuum 0.3 9 3 8 88.8 5.0 3. Antennaria rosea 1.3 22 .7.3 - - -4. Arabis sp. 0.6 17 '• 5.7 11 64.7 6.8 5.- Arenaria capillaris Tr 1 " 0.3 1 100 0.6 6. Arnica sp. 0.9 3 •1 .1 33.3 0.6 7. Astragulus miser 0.7 2 0.7 - - -8. Lappula redowskii Tr 9 ' 3 - - -9. Lepidium densiflorum Tr 2 0.7 1 50 0.6 10. Lomatium geyeri 0.2 4 1.3 2 50 1.2 11. Microseris alpestris Tr i " 0.3 1 100 0.6 cont... Appendix 5.1.9., continued. Species Average percent canopy cover in 300 loops Frequency.of occurrence No. % Frequency of grazing No. % Percent of a l l grazed plants 12. Oxytropis campestris 0.2 13. Phacelia linearis Tr 14-. Taraxicum officinale 0.7 Subtotals 5.7 7 2.3 5 71.4 3.1 7 2.3 1 14.3 0.6 3 1 1 33.3 0.6 90 18.7 32 35.6 19.9 BROWSE 1. Artemesia frigida 2. Eriogonum heracleoides 14.8 0.7 160 2 53.3 0.7 26 16.3 16.0 Subtotals 15.5 162 33.6 26 16.0 16.4 SUNDRIES 1. Rock 0.9 3 1.0 2. Soil 68.3 300 100 3.1. Litter • - - -''Tr = less than 0.1 r—1 CO ro 183 Appendix 5.2.1. Scientific and popular names for plants collected on the winter-spring ranges, Flatiron Mountain, 1968-1969 GRASS and GRASS-LIKE PLANTS Agropyron caninum (L.) Beauy Agropyron dasystachyum (Hook.) Scribn. Agropyron repens (L.) Beauv. Agropyron smithii Rydb. Agropyron subsecundum (I_ink) Hitchc. Agrostis alba L. Agrostis palustris Huds. Agrostis scabra Willd, Bromus. carinatus Hook. Bromus tectorum L. Calamagrostis rubescens Buckl. Carex sp. Danthonia intermedia Vasey Festuca idahoensis Elmer Festuca ovina L. Hordeum jubatum L. Juncus balticus Willd. Juncus mucronation  Koeleria cristata (L.) Pers. Phleum pratense var. bakeri (Greene) C.L. Hitchc. Poa alpina L. Poa ampla Merr. Poa pratensis L. Poa secunda Hitch Stipa columbiana Macoun. Stipa.comata Trin. and Rupt. Stipa richardsoni Link Trisetum spicatum (L.) Richt. Bearded wheatgrass Thickspike wheatgrass Quackgrass Western wheatgrass Bearded wheatgrass Redtop Creeping bent Bentgrass California brome Downy brome Pinegrass Sedge Timber oatgrass Idaho fescue Sheep fescue Foxtail barley Rush Rush Junegrass Timothy Alpine bluegrass Big bluegrass Kentucky bluegrass Sandberg bluegrass Columbia needlegrass Needle-and-thread Richardson needlegrass Spike trisetum 184 Appendix 5.2.1., continued FORBS Achillea millefolium L. Allium cernuum Roth Antennaria rosea Greene Arabis hirsuta (L.) Scop. Arabis holboeilii Hornem. Arabis holboeilii var. pendulocarpa Hornem. Arctium lappa L. Arenaria capillaris Poir Artemisia michauxiana Bess Aster campestris Nutt. Aster occidentalis (Nutt.) T. & G. Astragulus miser Dougl. Capsella bursa-pastoris: (L.) Medic. Castelleja miniata Dougl. Chenopodium album L. Cirsium. sp. Collinsia parviflora Lindl. Collomia linearis Nutt. Yarrow Nodding Onion Rosy pussytoes Rockcress Rockcress Rockcress Great burdock Mountain Sandwort Sagebrush species Aster Aster Timber milk vetch Shepherd's purse Paintbrush Thistle Blue-eyed Mary Collomia Crepis atrabarba Heller Delphinium bicolor Nutt. Descurainia pinnata (Walt.) Britt. Descurainia pinnata (Walt.) Britt. var. filipes (Gray) Delt Descurainia sophia (L.) Webb' Dadecatheon pauciflorum (Durand) Greene Low larkspur Dodecatheon pauciflorum var. cusickii (Greene) Mason Draba nemorosa, L. Epllobium adenocaulon Hausskn. Erigeron compositus Pursh. Erigeron corymbosus Nutt. Erigeron divergens T. S G. Shooting Star Shooting Star Whitlow grass Willow herb Ewarf mountain fleabane Fleabane Fleabane 185 Appendix 5.2.1. , continued Erigeron f i l i f o l i u s Nutt.. Erigeron linearis (Hook.) Piper Erigeron pumilus Nutt. Erigeron subtrinervis Rydb. Fragaria virginiana Duchesne var. platypetala (Rydb.) Hall Fritillaria.pudica (Pursh) Spreng. Gentiana propinqua Richards Geranium viscosissimum F. g M. Gilia minutiflora Benth. Lappula redowskii (Hornem.) Greene Lepidium densiflorum Schrad. Lewisia rediviva. Pursh Lithophragma parviflora (Hook.) Nutt. Lomatium geyeri (Wats.) Coult. S Rose Lomatium macrocarpum (Nutt.) Coult. S Rose Lupinus sericeus Pursh. Microseris alpestris (Gray) Q. Jones Microseris troximoides Gray Opuntia fragilis (Nutt.) Haw. • Orobanche californica Cham. £ Schlecht Oxytropus campestris (L.) DC. Pacelia linearis (Pursh) Holy Penstemon humilis Nutt. Penstemon procerus Dougl. Phacelia hastata Dougl. Phacelia linearis (Pursh) Holz Phlox diffusa Benth Plantago major L. Potentilla arguta Pursh. Potentilla diversifolia Lehm. Fleabane Fleabane Fleabane Fleabane Yellow bell Four-parted gentian Sticky geranium Gilia Western stickseed Bitter-root Lomatium Bigseed lomatium Silky lupine False agosenis Cactus Locoweed Pacelia Penstemon. Tinybloom penstemon Whiteleaf phacelia Phacelia Spreading phlox Plantago Cinquefoil Mountain meadow cinquefoil 186 Appendix 5.2.1., continued Potentilla gracilis Dougl. Ranunculus alismaefolius Geyer ex. Benth. Ranunculus glaberrimus Hook. Sedum lanceolatum Torr. Silene douglasii Hook. Silene parryi, :(Wats.) Hitchc. S McQwre Sisymbrium altissimum L. Taraxacum officinale Weber Tragapogon pratensis L. Trifolium repens L. Verbascum thapsus L. Veronica americana.(Raf.) Schwein. Viola adunca Sm. Viola adunca var. cascadensis (Baker) Hitchc. Viola n u t t a l l i i Pursh. Woodsia oregana P.C. Eat. Zigadenus veneosus Wats. Beauty cinquefoil Buttercup Buttercup Stonecrop Douglas campion Parry's silene Tumbleweed Common dandelion White clover' Great Mullein American Brooklime Blue violet Blue violet Johnny-Jump-Up Woodsia Deathcamas 187 Appendix 5.2.1. , continued BROWSE Alnus sp. Arctostaphylos.uva-ursi (L.) Spreng. Arteme'sia frigida :Willd. Artemesia tridentata Nutt. Ceanothus velutinus Dougl. Chrysothamnus nauseosus (Pall.) Britt. Eriogonum heracleoides var. angustifolium (Nutt.) . T. g G. Eriogonum heracleoides var. heracleoides Nutt. Eriogonum niveum Dougl. Ledum glandulosum Nutt. Ribes sp. Ribes cereum Dougl. Rosa nutkana Presl. Rosa woodsii Lindl. Rubus idaeus L. Salix bebbiana Sarg.. Shepherdia canadensis (L.) Nutt. Alder Kinnikinnick Pasture sage Big sagebrush Ceanothus Rabbit bush Buckwheat Buckwheat Buckwheat Mountain labrador tea Currant Squaw currant Wild rose Wild rose Red raspberry Willow Soopolallie TREES Abies lasiocarpa (Hook.) Nutt. Picea glauca (Moench) Voss Pinus contorta Dougl. Populus tremuloides Michx. Pseudostuga menziesii var. glauca (Beissn) Franco Subalpine f i r White spruce Lodgepole pine Trembling aspen Douglas f i r 188 Appendix 5.2.2. Scientific and popular names for plant species collected on the summer ranges, Joe Lake area, 1958-1969 GRASS and GRASS-LIKE PLANTS Agropyron saundersii Agropyron griffithsi Calamagrostis purpurascens R. Br. Carex albonigra Mackenzie Carex phaeocephala Piper Carex praticola Rybd. Carex scirpoidea Michx. Danthonia intermedia Vasey. Festuca ovina L. Koeleria cristata (L.) Pers. Luzula spicata (L.) DC. Phleum alpinum L. Poa alpina L. Poa pratensis L. Poa rupicola Nash. Poa secunda Presl. Trisetum spicatum (L.) Richt. Wheatgrass Wheatgrass Purple reedgrass Sedge Sedge Sedge' Sedge Sedge Sheep fescue Junegrass Woodruck Alpine timothy Alpine bluegrass Kentucky bluegrass Timberline bluegrass Sandberg bluegrass Spike trisetum FORBS Achillea millefolium L. ssp. lanulosa Piper Agoseris glauca (Pursh.) Raf. Antennaria anaphaloides Rydb. Antennaria lanata (Hook.) Greene Antennaria rosea Greene Antennaria umbrinella Rydb. Arenaria capillaris Poir. Arenaria obtusiloba (Rydb.) Fern Arnica fulgens Pursh Yarrow Smooth mountain dandelion White pussytoes Pussytoes Rosy pussytoes Pussytoes Mountain sandwort Sandwort Arnica 189 Appendix 5.2.2., continued Arnica mollis Hook. Aster sp. Erigeron- peregrinus (Pursh) Greene Gentiana amarella L. Gentiana propinqua Richards Geum triflorum Pursh Oxytropis campestris (L.) DC. Polemonjum pulcherrimum Hook. Potentilla diversifolia var. diversifolia Lehm. Potentilla drumrrondii Lehm. Sedum lanceolatum Torr. Silene acaulis L. Silene scouleri Hook. Solidago multiradiata Ait. Stellaria longipes var. attocaulis Hulten Arnica Aster Peregrine fleabane Gentian Four-parted Gentian Long-plumed purple avens Locoweed Blue skunkleaf Mountain meadow cinquefoil Cinquefoil Stonecrop Moss silene Silene Goldenrod Chickweed BROWSE Arctostaphylos uva-ursi (L.) Spreng. Artemesia tridentata Nutt. Eriogonum umbellatum Torr. Ledum glandulosum Nutt. Potentilla fruticosa L. Vaccinium scoparium Leiberg Kimikinnick Sagebrush Sulphur eriogonum Mountain labrador tea Shrubby cinquefoil Grouseberry 

Cite

Citation Scheme:

        

Citations by CSL (citeproc-js)

Usage Statistics

Share

Embed

Customize your widget with the following options, then copy and paste the code below into the HTML of your page to embed this item in your website.
                        
                            <div id="ubcOpenCollectionsWidgetDisplay">
                            <script id="ubcOpenCollectionsWidget"
                            src="{[{embed.src}]}"
                            data-item="{[{embed.item}]}"
                            data-collection="{[{embed.collection}]}"
                            data-metadata="{[{embed.showMetadata}]}"
                            data-width="{[{embed.width}]}"
                            async >
                            </script>
                            </div>
                        
                    
IIIF logo Our image viewer uses the IIIF 2.0 standard. To load this item in other compatible viewers, use this url:
http://iiif.library.ubc.ca/presentation/dsp.831.1-0101708/manifest

Comment

Related Items