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Interactions between moose, elk and buffalo in Elk Island National Park, Alberta Holsworth, William Norton 1960

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INTERACTIONS BETWEEN MOOSE, ELK AND BUFFALO IN ELK ISLAND NATIONAL PARK, ALBERTA by WILLIAM NORTON HOLSWORTH B.Sc, University of British Columbia, 1958 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of ZOOLOGY We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA April, I960 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r an advanced degree a t the U n i v e r s i t y o f B r i t i s h C o l u m b i a , I agree t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r agree t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y purposes may be g r a n t e d by t h e Head o f my Department o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l not be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . Department o f The U n i v e r s i t y o f B r i t i s h Columbia, Vancouver B, Canada. Date VVy ^ 3tJ  ABSTRACT A study of the interactions between moose (Alces alces), elk (Cervus canadensis) and buffalo (Bison bison) was carried out during the summer of 1959 in Elk Island National Park, Alberta. The interactions studied were the possibility of competition for food and habitat, transmission of disease and dominance relations. A description of the Park is given. The distribution, habitat selection and food habits of each of the three species are discussed. The greatest interaction between these animals occurs through their effects on the range* Data, collected over the past years, on the incidence of brucellosis and i t s effect on the reproductive rate and health of the animals are presented. The high incidence of brucellosis in buffalo indicates that they are the source of the organism infecting moose and elk. Buffalo were found to be dominant over moose and elk. The effects of the existing dominance relations could not be evaluated. - i i -TABLE OF CONTENTS Page INTRODUCTION ... 1 A. Historical Background 4 DESCRIPTION OF THE PARK 7 A. Physical Features 8 B. Vegetation '. 9 C. Classification of Ruminants in the Park 23 METHODS AND DATA 24 A. Distribution of the Animals 25 B. Habitat Selection by the Animals 30 C. Food Habits of the Animals 35 D. Disease 45 E. Mortality Due to Other Causes 48 F. Behavior 48 DISCUSSION... 52 A. Effects of the Animals on the Range 53 B. Effects of Range Conditions on Food Habits of the Animals 58 C. Effects of the Animals on Each Other 59 D. Influence of Environment on Habitat Selection and Distribution of the Animals and Carrying Capacity of the Range. 61 E. Anatomical Reasons for Habitat Selection 62 SUMMARY 64 LITERATURE CITED. 67 APPENDICES 70 Appendix A. List of Plants of Elk Island National Park.... 70 Appendix B. Summary of Range Analysis 77 Appendix C, Photographs of Range and Habitat Types 87 - i i i -LIST OF TABLES TABLE PAGE I Long-term Temperature and Precipitation Bata for Edmonton.. 9 II Relation Between Density and Age of Aspen Trees . 14 III Incidence of Brucellosis in Moose, Elk and Buffalo. 46 IV Calving and Conception Rates in the Elk Island National Park Buffalo Herd, 1951 to 1956 47 V Summary of Data from Transect 1. 78 VI Summary of Data from Transect 2 79 VII Summary of Data from Transect 3. 80 VIII Summary of Data from Transect 4., 81 IX Summary of Data from Transect 5 . • 82 X Summary of Data from Transect 7 83 XI Summary of Data from Transect 9...... 84 - iv -LIST OF FIGURES FIGURE PAGE 1 Percentage Occurrence in the park of Habitat Types........ 19 2 Distribution of Habitat Types (Map) 20 3 Layout of Range Transect Lines. 21 4 Summer Concentration Areas of Moose, Elk and Buffalo (Map) . 27 5 Winter Concentration Areas of Moose and Elk (Map)......... 28 6 Winter Concentration Areas of Elk (Map)o . 29 7 a) Summer Habitat Selection by Moose....... ^ . 32 b) Winter Habitat Selection by Moose 32a 8 a) Summer Habitat Selection by Elk.... 34 b) Winter Habitat Selection by .Elk.. 34 a 9 a) Summer Habitat Selection by Buffalo.... 34b b) Winter Habitat Selection by Buffalo....... 34c 10 Summer Diet of Moose.. 39 11 Summer Diet of Moose, Elk and Buffalo..... 40 12 Winter Diet of Moose, Elk and Buffalo. 41 13 Location of Range Transects (Map)... 85 14 Sample of Form Used to Record Range Data., 86 - V -LIST OF PHOTOGRAPHS PHOTOGRAPH PAGE 1 Large slough near the north end of Elk Island National Park. .. 87 2 Prairie meadow near "soap-holes",. 87 3 Grass-forb meadow north end of Tawayik Lake. 88 4 Grass meadow formed by cultivation. ... 88 5 Aspen-balm shrub meadow 89 6 Aspen-balm shrub meadow 89 7 Semi-open aspen habitat 90 8 Aspen forest in isolation area... 90 9 Open aspen stand with understory of Calamagrostis grass. 91 10 Open aspen stand after burn in 1946........... 91 11 Open aspen stand after burn in 1946 V 92 _ yi. _ ACKNOWLEDGEMENTS I wish to thank Dr. I. McT. Cowan who served.as my principal advisor and Dr. J.R. Adams, Dr. J.F. Bendell and Dr. W.D. Kitts for providing help and ideas during the preparation of the thesis manuscript,.. I am indebted to Mr, Donald R, Flook,.Canadian Wildlife Service, who suggested this problem, provided advice and assistance in planning the fi e l d work, gave encouragement and help during the.summer, and provided me with the aerial survey data. For their help and interest in this study I wish to thank Mr. H.R, Webster, Superintendent, Elk Island National Park, Chief Warden Armstrong, and other employees of the Park. The f i e l d work was financed by the Canadian Wildlife Service who employed me during the summer, I was supported by a Canadian Industries Limited "Wildlife Fellowship" while at the University of British Columbia during the winters of 1958-59 and 1959-60, INTRODUCTION - 2 -"This summer I continued my investigations as to which plants are consumed by cattle, which are ignored, and which are avoided; this work in my opinion, is of fundamental importance both for private owners of livestock and for animal husbandry as a whole," Carolus Linnaeus, 1748 This study i s concerned with the interactions of moose (Alces alces), elk (Cervus canadensis), and buffalo (Bison bison), on a woodland range i n central Alberta. These three species, and deer, are pot e n t i a l l y capable of using the same foods and same habitat* Interaction between them i s , therefore, possibles Although there have been many previous studies concerned with various problems of in t e r s p e c i f i c interactions between different species of wild ruminants, and between wild ruminants and domestic animals, few studies have considered more than one type of interaction* The present study combines the effects of several environmental factors. Interactions considering competition for food are numerous (Cowan, 1947} Julander, 1958; Ma.ir, 1952; McMillan, 1953; Morris and Schwartz, 1957; Murie, 1951; Peterson, 1955; and many others). McHugh (1958) and Altmann (1951, 1956) studied i n t e r s p e c i f i c behavioral interactions of buffalo and elk respectively. Crrzimek and Grzimek (i960) censused the plains animals of the Serengeti National Park, Tanganyika, and found each of the ten common herbivores concentrated i n certain places and missing i n other regions. Studies of food habits, habitat selection and behavior have been conducted on a variety of birds and mammals. Lack (1944) c i t e s several instances of potential ecological overlap i n similar sympatric species, but i n each case where they are occupying similar habitats, close observation reveals that they are r e a l l y using separate portions of the environment. Lack suggests that overlap i n food habits indicates a superabundance of food, and competition between the species i s seldom involved. In North America, prior to the extermination of the large plains animals by white man, seven native ruminants and the f e r a l horse roamed the great plains. Elimination of elk and buffalo over much of their former range has precluded such studies of interaction as might have been carried out 100 years ago. Only i n Yellowstone National Park, Wyoming, and Elk Island National Park, Alberta, do large herds of the former dominant plains species co-exist i n areas which they formerly occupied and under conditions similar to the primitive state. Even i n these parks, the situation i s somewhat a r t i f i c i a l owing to selective stocking, predator control, selective harvesting and r e s t r i c t i o n of movements of the animals. During the summer of 1959 (May to September) and during a b r i e f period i n December, 1959, the interactions of moose, elk and buffalo i n E l k Island National Park were studied. A short v i s i t was made i n September to the National Bison Range, Montana, and to Yellowstone National Park, Wyoming. To understand the implications of the data presented i n t h i s study, i t i s necessary to be familiar with the habitat i n which the animals l i v e . Following a description of the habitat, the results of determining d i s t r i b u t i o n , habitat selection, food habits, diseases, predation and behavior of the animals are presented* Only after correlation of these factors are some suggestions warranted on how moose, elk and buffalo interact. In the following pages, each of the factors w i l l be considered i n turn. The i n t e r s p e c i f i c effects of each w i l l be discussed and f i n a l l y , the effects of a l l factors w i l l be evaluated. A. H i s t o r i c a l Background For ten thousand years, or more, buffalo, elk, pronghorn antelope, moose, and deer have l i v e d together i n the aspen parkland and sub-climax aspen forest of the coniferous biome of Alberta, Saskatchewan and Manitoba. U n t i l the coming of white man to the P r a i r i e Provinces, buffalo, elk and moose were numerous. Anthony Henday, the f i r s t white man to enter Alberta, records that his Indians k i l l e d eight elk, three buffalo and four moose on h i s f i r s t two days i n "Alberta". Paul Kane, the a r t i s t , came to Edmonton 89 years l a t e r , i n 1845. He records the nurabers.of buffalo as follows "...three days i t took us to reach Edmonton House, we saw nothing else but these animals (buffalo) covering the plains as f a r as the eye could reach, and so numerous were they, that at times they impeded our progress e.»" By 1900 a l l buffalo were gone from the plains and parklands of Alberta, Settlers, which had been moving into the area since the early 1880's had also extirpated most of the elk and moosee In March, 1906, f i v e pioneers east of Edmonton, cognizant of the pending loss of these native ungulates approached the Minister of the Interior to have 16 square miles set aside as a deer park and game preserve. On August 4, 1909, Elk Island Game Preserve became Elk Island National Park, This Park did not "just happen" to be located i n an area that s t i l l had a few moose and elk* In contrast to the surrounding parkland where trees are limited to stream valleys and depressions, the Beaverhills have a forest of aspen, balm and birch, and a climax forest of white spruce. The heavy forest discouraged the early s e t t l e r s from farming t h i s r o l l i n g ground, and gave protection to the few elk and moose s t i l l presents The Pablo—Allard herd of buffalo i n Montana came up for sale i n 1906, and the Dominion of Canada bought them to stock a new Park which was being created at Wainwright, Alberta, Because the Wainwright park was not completely fenced, 250 buffalo were released i n the E l k Island Game preserve for temporary holding. In 1909, most of these buffalo were removed and transported to Wainwrighte About 50 buffalo, mostly old b u l l s too wild, to catch, remained i n the park. Thus the present herd had i t s beginning from the enclosed moose and elk and the remaining 50 buffalo, A purchase of 34 square miles immediately south of the i n i t i a l 16 square mile block was made i n 1922, The old south boundary fence was removed, providing the herds with nearly 50 square miles of range. In 1947, a 24 square mile area was bought from the Blackfoot Grazing Association. That area was fenced and crossfenced dividing i t into three 8 square mile ranges. Most of the moose and elk were removed from the new area. In the spring of 1951, 67 young buffalo cows and 10 young b u l l s were introduced into the two western ranges. In 1959, the eastern t h i r d was opened to the buffalo. The 24 square mile area was o r i g i n a l l y designed as an "Isolation Area" to protect the buffalo herd from extermination should an epizootic occur (e.g. foot and mouth disease). The part of the Park north of Highway 16, w i l l be referred to as the "park" without a c a p i t a l "p"o For many years the stocking l e v e l of the range has been very high. As many as 1,500 buffalo, possibly 1,200 to 1,500 elk and 600 or more moose have occupied the park (considered as 45 square miles of useful range) at one time 0 The effect these large herds on the range has been to change the composition and physiognomy of the plants. I t i s through these changes i n f l o r a l structure that i n t e r s p e c i f i c interactions are most e f f e c t i v e . The number of animals i n the park during the summer of 1959 was estimated as 650 buffalo, 600 elk, and 300 moose. The population of the i s o l a t i o n area was i considered to be approximately 450 buffalo, 40 elk, 100 moose and 150 deer. DESCRIPTION OF THE PARK - 8 -A. Physical Features 1, Geology During the Keewatin glac i a t i o n the ice sheet moved i n a generally south-westerly direction, and i n passing over the area, materials i t had carried f o r a considerable distance were mixed with the materials from the underlying bedrock. The g l a c i a l d r i f t was deposited as a mantle over the Park. The Beaverhills represent a terminal moraine located on a bedrock high. Material i n t h i s moraine i s mainly of Edmonton formation o r i g i n (Allen). 2* Topography Most of the park i s h i l l y * The i s o l a t i o n area i s gently r o l l i n g . Extremes in elevation are from 2,375 to 2,475 feet i n the park and from 2,350 to 2,425 feet i n the i s o l a t i o n area. The northern half of the park tends to be very h i l l y with numerous potholes and sloughs. The south-east quarter i s leve l to gently r o l l i n g , and the south-west quarter i s characterized by several large h i l l s , and numerous morainic ridges running approximately i n an east-west direction. The h i l l s i n the i s o l a t i o n area are confined to the south-west portion where again east-west morains occur. Although there i s l i t t l e a l t i t u d i n a l change, because of the latitude, the actual i n c l i n a t i o n of the ground has a great effect on the amount of heat and l i g h t received from the sun, and a difference i n vegetation i s noticeable on north and south slopes, 3. Soils The s o i l over most of the area of Elk Island National Park i s classed as Cooking Lake Loam which i s an orthic grey wooded s o i l . The loam i s r e l a t i v e l y low i n natural f e r t i l i t y . Most of the soluble plant food has been leached from the upper horizons and deposited i n the B horizon. The s o i l has a high lime content,and i s - 9 -not low in sulphur and magnesium as are many of the grey wooded soils. Salt content of the C horizon is relatively low. Detailed analysis of a very similar s o i l , the Breton Loam, indicates that there is no one essential element distinctly deficient, although phosphorous is in lower supply than are the others (Bowser and Erdman, 1947). Areas of deep organic s o i l , many of them caused by gradual f i l l i n g in of beaver ponds with vegetation, occur throughout the Park (about 15% of the total area). 4. Climate The Park is in a region of typical continental climate of northern latrhdes. It is characterized by a long, cold, dry winter and a growing season of about three months, during which the days are long, bright and moderatley warm. The mean annual precipitation is about 18 inches, one-third falling in June. Owing to the prevailing low humidity during the day, the rate of loss of water from the soil and vegetation is high. However, very heavy dews f a l l every night during the summer. Weather dataare presented in Table I, as recorded at Edmonton, 30 miles west of the Park (Canada Year Book, 1957-58). Table I, Long-term Temperature and Precipitation Data for Edmonton. Mean Temperature°F Mean Frost-free Period Precipitation . Ann. Jan. July Total Ann. Ann. Snow Jan. Apr. July Oct. 36.8 7.7 62.9 May 29-Sept,6 17.6 52.9 0.9 1.1 3,1 0.84 B. Vegetation With the exception of the organic soil areas, a l l soils in the Park would naturally support either a forest of aspen and balm, or white spruce. (Martin and 10 -vernacular names of a l l plants mentioned i n text are l i s t e d i n Appendix.A). The theoretical climatic climax vegetation of the grey-wooded s o i l i s white spruce. However, prior to settlement of western Canada, grass and forest f i r e s were a normal occurrence and seldom was a spruce forest maintained over a large area for a long time. Consequently, the area i s a checkerboard of serai stages; each habitat type r e f l e c t i n g i t s past history. Demonstrating the regular occurrence of f i r e s , a p r o f i l e of a sphagnum bog i n the north end of the park showed fourteen layers of charcoal i n the f i r s t three feet of moss. Below t h i s depth, the charcoal layers were indeterminable. 1. Description of habitat types. Seven habitat types occur within the Park. These are: a. Lakes, b. Sloughs, c. Wet meadows, d. Bogs, e. Grass meadows, f . Grass-shrub meadows, g. Aspen-balm for e s t s . A detailed description of each habitat type follows. a. Lakes The lakes present i n the Park appear to be due to hollows i n the s u p e r f i c i a l g l a c i a l deposits and not to excavation by g l a c i a l action. The largest and deepest lake, Astotin, has several small rocky islands devoid of g l a c i a l debris. Tawayik, L i t t l e Tawayik, and Flyingshot Lakes are very shallow and have gently sloping shores. The margins of these lakes have extensive sedge meadows i n d r i e r years. Oster and Adamson lakes are located i n an area of morainic ridges and consequently have more sharply defined shorelines. A l l lakes have an abundance of submerged vegetation; however, much of i t i s i n water too deep to be of use to moose. The other named lakes i n the Park must be classed as temporary water bodies or sloughs. b. Sloughs There are s i x large, temporary water bodies i n the i s o l a t i o n area and many small ones i n the park. During the average summer there are over 200 small potholes i n the park, many of them less than 50 feet i n diameter. T 11 -Floating vegetation was not abundant i n 1959. Emergent vegetation was lacking from many of the sloughs and confined to the periphery of others. Submergent aquatic plants common to most of the sloughs were Ranunculus aquatillus, Ceratophyllum, Myriophyllum, and some Potamogetons. Typha. Scirpus. Spargnum and Eleocharis were present on only a few ponds, mostly those i n the north end of the park. Peripheral vegetation was primarily Carex, Glyceria and Calamagrostis. Forbs included Galium.  Petasites and Caltha. The commonest shrubs on the margins of sloughs were willow (several species) and alder. Bog birch, paper birch and balm were present i n a few areas. Alder was most common i n the north end of the park and almost absent i n the i s o l a t i o n area where willows were dominant. c. Wet meadows About one-quarter of the sloughs are the result of previous beaver a c t i v i t y . Several Carex-Calamagrostis-Glyceria meadows, one to f i v e acres i n size occur i n chains, each separated from the adjacent one by the remnants of a beaver dam. Often the only open water i n the meadow i s a small pond a few yards across on the "upstream side" of the dam. The old beaver dams are ea s i l y recognized by their position and by the vegetation, either balm, willow or alder, growing on them. Other wet meadows, having similar f l o r i s t i c composition to the f i l l e d - i n beaver ponds, have been produced by hydrosere development. The margins of some of the wet grass meadows have shore li n e s which show considerable erosion. Banks four feet high, or more, occur on the edge of some meadows which now have no open water throughout the year. There are few meadows i n the park which nave extensive willow cover. The most common shrubs are gooseberry, bog birch and raspberry. Triglochin,  P o t e n t i l l a p a l u s t r i s , Parnassia, Petasites and other forbs are common on these meadows. d. Bogs About twenty black spruce-sphagnum bogs, more or less similar to the bog - 12 -forests of the Cordillaran area, occur i n the Park* The bog forest i s characterized by an association of black spruce 15 to 30 feet high, about seven feet apart, and growing on a wet bed of Sphagnum, Ledum and other bog plants, Carex, and often willow, growing i n water, encircle each bog* Within this wet ring, Cladonia. Ledum and Vaccinium grow on mounds of Sphagnum* Other plants of the bogs are Ribes chamaemorus. Vaccinium oxycoccus, Eriophilum, Carex spp* and many others. Although black spruce i s the most common tree of the bogs, tamarack grows i n association with i t i n the north end of the park. Both paper birch and scrub birch are present as shrubs i n many of the bogs* No l i v i n g paper birch trees are present, although trunks of dead trees s t i l l stand. Heavy browsing of t h i s birch has prevented i t s maturation. e. Grass meadows Most of the grass meadows are the result of man's a c t i v i t y . In 1944, a series of small meadows along the road were created and sowed to tame grasses (probably Bromus and Poa)„ The exact extent of t h i s "farming operation" could not be determined. Other meadows, created by early s e t t l e r s are recognizable. Heavy browsing on these meadows has prevented regeneration of aspen trees. True p r a i r i e grasses are absent except on a small patch of " p r a i r i e ? " i n the south-east corner of the park where Koeleria can be found. That area i s the only part of the park which might be considered to be a true grassland association. Even i t , however, i f protected from the effects of browsing would support a forest of aspen or spruce. f . Grass-shrub meadow There are two types of grass-shrub meadows. On the sandy s o i l , a hazel-rose shrub meadow occurs (see Appendix B, Transect 4). On the heavier s o i l s , the shrub i s mostly aspen and balm (see Appendix B, Transect 5). A l l meadows were covered with an aspen forest before the l a s t f i r e . Their present condition i s the re s u l t of browsing and grazing. - 13 -i . Hazel-rose shrub meadow.— On the sandy soils on the east side of the park, and in the regions west of Tawayik Lake, areas up to 300 acres are covered with hazel and rose, interrupted by grass meadows and aspen groves. These meadows provide a large part of the forage for moose and elk. A l l shrubs are kept browsed to a height of not more than four feet. Aspen, balm, willow and saskatoon have been largely eliminated from these areas by browsing and possibly by competition with hazel, i i . Aspen-balm shrub meadow.— Associated with aspen, balm, birch, willow and snowberry shrubs, i s a carpet of Poa pratensis, and Trifolium repens. The tree species are heavily browsed so that forest regeneration has been completely prevented in the past 50 years. Browsing, heavy enough to k i l l aspen and balm trees, has permitted some of the grass meadows, particularly those in the grass meadow-aspen grove association to persist. The introduced meadow plants (Poa and Trifolium) are more resistant to heavy grazing than the native Calamagrostis. Agropyron and Bromus. Consequently, as an area became denuded of its native vegetation, the introduced species were able to become established, g. Aspen-balm forests Fires started by lightning, by Indians and by white man have altered the fl o r i s t i c composition to some extent almost everywhere in the Prairie Provinces. The great expanse of aspen forest in Canada owes i t s presence to the effect of forest fires in preventing the spread of spruce. After a f i r e , aspen and balm are able to regenerate quickly from the underground parts of the burnt trees. This results in aspen forests being an even age stand. Spruce, on the other hand, is kill e d completely by severe f i r e and requires dispersion of seed to increase its range. Spread of spruce i s , consequently, very slow. Aspen is found in the Park in two situations: as groves in the grass meadow - aspen grove association, and as an aspen forest. Most of the aspen forest - 14 -i n the Park originated as the primary succession following a f i r e . Some areas, possibly parts of the meadow - grove association, developed from a shrub meadow which had been released from the effects of heavy browsing. Measurements of trees show that aspen i n the park grow i n 70 years to a height of 60 to 75 feet and attain a diameter of not more than 18 inches. The great majority of the trees never reach t h i s s i z e . The trees are shade intolerant and thus require considerable thinning before new growth can take place. Table II shows the rate of thinning of aspen stands as determined by the approximate numbers of trees i n stands of different ages (Moss, 1932). Table I I . Relation Between Density and Age of Aspen Trees. Number of Trees i n Years Trees per 100 Square Meters 5 500 10 250 15 175 20 80 30 45 50 25 70 10 110 3 120 0 In addition to the competition between the trees, thinning of the stand i s hastened by the polypore, Pomes igniarius which attacks nearly a l l aspen over 50 years of age. The beetle Saperda sp. infects most trees about the same time and compliments the effects of Fomes. The second most abundant tree, balm, i s present i n association with aspen nearly everywhere i n the Park. Balm grows s l i g h t l y faster than aspen on wetter s i t e s , l i v e s a few years longer, and i s less subject to heart rot by Fomes (Moss, 1932). - 15 -Like aspen, i t i s shade intolerant. The number of balm present i n r e l a t i o n to the number of aspen varies from p r a c t i c a l l y none to 50$ or more i n some of the wetter areas* Paper birch has greater tolerance to wet and acid conditions than either aspen or balm and consequently i s found i n wetter areas and occasionally on spruce bogs* Birch i s more tolerant of shade than either balm or aspen. Outside the Park, birch trees are found scattered under a canopy of aspen and balm. In the park, only a few clumps of old trees occur* Although much birch i s present i n some areas of the park, i t usually occurs only as a very heavily hedged shrub* The oldest trees i n the park occur i n the north end around the area which was swept by f i r e i n 1946* The trees i n t h i s area are about 70 to 85 years old, indicating that they followed a f i r e about 1880* Aspen and balm, about 50 to 60 years old, and most of the spruce are i n the north-east side of the park* A f i r e occurred i n 1900, or thereabouts, which cleared most of the trees from the remainder of the Park, Regeneration was somehow retarded i n areas south of the o r i g i n a l game preserve boundary* The presence of the younger trees i n the southern portion of the park and most of the i s o l a t i o n area may be the result of more recent burning by se t t l e r s * The age of the trees of most of the southern portion of the park and the i s o l a t i o n area i s 25 to 40 years* A few spots exist where the trees are 50 to 60 years old, but most of these trees show some f i r e scars* There are many areas where balm and aspen, three to f i v e feet high, are over 15 years old. This indicates prolonged heavy browsing* Within the aspen forest, small clearings exist* Vegetation of the forest f l o o r varies considerably from one s i t e to another* The composition of the under-story i s correlated with the age and density of the dominant trees. There appears to be no correlation between form and composition of the aspen forest and s o i l typesj therefore, i t would appear that the different aspects of the forests are a r e f l e c t i o n - 16 -of t h e i r individual h i s t o r i e s . A similar conclusion was reached by E l l i s o n and Houston (1958), "There i s no indication that inherent s o i l differences are great enough or consistent enough to account for the marked differences that generally exist i n the ground vegetation." and "... the tendency suggests that the cause of the type boundaries are h i s t o r i c a l rather than environmental...." The aspen forest can be divided into f i v e types. i . Grass meadow - aspen grove a s s o c i a t i o n . — Heavy browsing by moose and elk, 15 to 25 years ago, k i l l e d most of the aspen and balm i n the area west of the Tawayik lakes. On the southern slopes, a cover of grass or hazel shrub i s established. On northern slopes and i n wetter areas, aspen groves occur. In a few places, aspen i s spreading from these clumps of trees onto the grassland meadow. i i . Open a s p e n , — The habitats, with well spaced, old trees, have a lower stratum composed of varying amounts of shrubs. Some aspen, balm and birch are present, but more common are raspberry, rose, hazel and snowberry. The dominant understory vegetation i s Calamagrostis grass (see Appendix C, Photograph 9). i i i . Semi-open aspen.— Under the "semi-open" aspen forest (trees 25 to 50 years old) the understory i s composed of either s a r s a p a r i l l a (Aralia) and snowberry or a mixture of hazel, rose, gooseberry and raspberry. Grass i s absent under the more dense canopies, but i n the stands approaching the "open" aspen type Calamagrostis becomes prevalent. Lacking i n the understory of a l l aspen forests i n the park i s red-osier dogwood, l i s t e d by Bir d (1930) as one of the two most outstanding subdominants of the shrub substratum i n aspen forests. Well-browsed dogwood i s present i n the i s o l a t i o n area. Saskatoon, chokecherry and Viburnum are likewise f a i r l y common i n the i s o l a t i o n area but largely absent i n the park. The reason for the difference i n plant composition between the park and i s o l a t i o n area, i s the high preference moose, elk and deer have for these plants and the low tolerance of the plants to heavy browsing. Hazel i s maintained at a height of less than three feet - 17 -throughout the park. In the i s o l a t i o n area, hazel over f i v e feet high i s not uncommon. Other shrubs, less palatable, are not browsed to such a constant l e v e l . The height of rose, gooseberry and buffalo-berry in the park varies between two and s i x fee t . With the exception of rose, these plants are eaten very l i t t l e * Photograph 7 i n Appendix C represents t h i s habitat type. i v . Dense aspen.— The dense aspen forest has an understory composed of almost 100$ s a r s a p a r i l l a . Other lesser vegetation i n this habitat, as i n the semi-open habitat, consists of snowberry, strawberry, wintergreen, bunchberry, v i o l e t s and many other forbs. The trees are 15 to 35 years old and are closely spaced. v« Aspen-balm forest outside the p a r k . — Outside the park, and i n the i s o l a t i o n area, the shrub strata i s well developed, and the forb understory i s greatly reduced. Sarsaparilla was found to be nowhere as abundant outside the park as i t was i n the dense aspen forest i n the south-east quarter of the park. Birch, willow, saskatoon, hazel, red-osier dogwood and other shrubs are common i n the areas free from heavy browsing. 2. Percentage occurrence of habitat types. The percentage each habitat type represented i n the park was determined from a i r photographs. The proportions were determined by examining, under a stereoscope, a series of east-west transect l i n e s spaced at one-half mile intervals over the whole park. Permanent water bodies and larger sloughs were excluded from the transects. The percentage habitat occurrence as shown i n Figure 1, i l l u s t r a t e s the dominance of the aspen forest habitats (nearly 70$). Wet meadows and grass meadows, which are interspersed i n the aspen forests account for 14.5$ and 9$ of the area of the park. The i s o l a t i o n area was not included because i t i s nearly 100$ aspen f o r e s t . - 18 -3. Distribution of habitat types. The d i s t r i b u t i o n of the habitat types i s shown on the map (Figure 2). The l i m i t s of these regions of vegetation types were determined by examination of a i r photographs taken i n 1948. Corrections for recent changes i n vegetation were made where possible. 4. Analysis of Range. Seven permanent range plots were established so that a quantitative description of vegetation i n key areas could be made. These plots also serve as the beginning of a long term range trend study. Plots were located i n the following habitat types: 1. and 2. Grass meadow; 3. 4. and 5. Grass - shrub meadow; 7. Burn i n mature aspen forest; 9. Semi-open aspen association. Range transect 8 was located i n an aspen forest i n the i s o l a t i o n area, but was not established as a permanent transect. A map showing location of the transects and a summary of the transect data are presented i n Appendix B. The raw data and a l l photographs of range are on f i l e i n the of f i c e s of the Canadian W i l d l i f e Service, Edmonton, a. Location of transects Transect locations were selected on the basis of being representative of the major habitat types existing i n the park, or i n places considered to be key areas. Four permanently located 100 foot transect l i n e s were arranged at each range p l o t . Location of the transect l i n e s i n a cluster as shown i n Figure 3, provides a measure of variation with a sampling s i t e , more information per man-hour, and makes for easier relocation (Parker, 1954). The arrangement of the transects within the cluster was designed so as to sample as large an area as possible while remaining i n a uniform habitat type. The distance between the center of the transect cluster and the "near" end of the transect l i n e was designed to minimize the - I 9 -6 0 -IS 50 -fe-ci w • a ! E H H o 30 20 10 O ' o o p 'd c o cd cd cd O 0) • a CD 2 2 S* PU cd a a a to CO CD cd 0 CO E CO CO CO C - P cd cd P. CD CD Sh 6 tj CO CD (X ? c • H CD a p. CD CO CO cd C CD P« CO cd CD CO & <D o p. CO o cd M r H O CO co HABITAT TYPES Figure 1. Percentage occurrence i n the park of habitat types as determined, by 91 miles of li n e transects o n a i r photographs. Permanent water bodies and i s o l a t i o n area are not included. - 20 -ELK ISLAND NATIONAL PARK DISTRIBUTION OF HABITAT TYPES Scale: LEDCrEND 1.5cm. to 1 mile Habitat types Lakes and sloughs o o 0 • £ a al Cx. Wet meadow G.M. Grass meadow Sh.Jw-A.Gr. Shrub meadow-Aspen grove O.A. Open aspen S.O.A. Semi-open aspen D.A. Dense,  aspen C.L. Cultivated Land Highway Firure 2. Di s t r i b u t i o n of Habitat Types - 21 -N 1 P i t l«" . ««»'»••• i n c a n t « r < yr —1 - Transect \ ; „ t Figure 2. Layout of transect lines within range p l o t s . 22 importance of the location of the hub stake but not oversampling the center of the p l o t , b. Analysis of the grassland vegetation A transect l i n e was run between the pairs of stakes, and at each foot in t e r v a l , the vegetation (root crown only) occurring within a three-quarter inch c i r c l e d i r e c t l y below the foot mark, was recorded. The three-quarter inch c i r c l e has a l l the advantages of both the "point" and the "area" method of determining the sub-sampleo The "point" has the disadvantages of being a point, theoretically determinable, but i n practice i t must be an area. The size of the area of the "point" makes considerable difference i n the results obtained (Greig-Smith, 1957; p. 6), The "area" sub-sample has the disadvantages of requiring subjective interpretation as to percent of area occupied by each plant species. The 3/4 inch "point" seldom contains more than one species, and when i t does, the per icent can be estimated very quickly. In practice, i f two species are present, each i s credited with 1/2 the area. Using the 3/4 inch c i r c l e has been found to be faster and less subjective than the "area" sub-unit, more accurate, more informative, and faster than the "point" sub-unit (Parker, 1954), Permanent location of transect lines permits absolute comparisons of range conditions from one year to the next, c. Analysis of browse vegetation The method outlined above i s not suitable for sampling heavy woodland, or browse habitat. Browse was sampled by counting the browse plants i n 3 by 100 foot plots adjacent to transect l i n e s , and segregating the plants into species and height classes. Notes were made on the vigor and extent of browsing on the plants. Species present i n the s i t e , but not included i n the transect were noted, and their significance, i f any, was recorded. - 23 -C, C l a s s i f i c a t i o n of Ruminants i n the Park The common North American vernacular for the ruminants discussed i n th i s study w i l l be used. These names are: elk (Gervus canadensis manitobensis M i l l i a s . C. c_. nelsoni Bailey); mule deer (Odocoileus hemionus hemionus Rafinesque); white-t a i l e d deer (0. virginianus dacotensis Goldman and Kellogg); moose (Alces alces  andersoni Peterson); and buffalo (Bison bison bison (L.))„ - 24 -METHODS AND DATA - 25 A. Distribution of the Animals Three methods were used to determine the di s t r i b u t i o n of the animals i n the park and their habitat selection throughout the year. 1. Winter d i s t r i b u t i o n (Figure 5) and population estimates were made by ae r i a l census i n February, I960, by Mr. D.R. Flook, 2. Summer di s t r i b u t i o n (Figure 4) was determined from direct observation during the summer of 1959 (May to September), 3. Late f a l l and winter d i s t r i b u t i o n was determined by a defecation-site census conducted during the f i r s t two weeks of May, 1959, The "defecation-site" census was used to determine d i s t r i b u t i o n and habitat selection during the winter. An explanation of the method and discussion of the limitations of the information i s necessary. The new vegetation appears about the end of May, Before that time, the ground i s quite bare and the winter's accumulation of dung i s v i s i b l e . Transects, 600 feet long and 12 feet wide were placed approximately at the center of every other quarter sections Location of the center of the transect was made from a i r photographs. Only dung, lying on top of the leaves and l a s t summer's grass, was counted. The c r i t e r i o n f o r a p e l l e t group was at least 10 pel l e t s i n a square foot, or for buffalo, a winter "chip" more than s i x inches i n diameter. Where i t was obvious that several chips belonged to one defecation, they were counted as one. Proportions of t o t a l use made by moose, elk and buffalo on a single habitat type during the winter could not be determined exactly by defecation-site transects because several factors, f o r which v a l i d correction cannot be made, alt e r the number of defecation sites of each species i n a habitat. These factors are the defecation behavior of the species (e.g. do the animals defecate by their bed or on the i r feeding ground?) and frequency of defecation on a winter d i e t . Thus i t i s possible to make only a rough correlation between di f f e r e n t species and habitat use. - 26 -The elk counts tend to be too low. Much of the time the elk axe running when they defecate. The buffalo counts tend to be high because the period of accumulation i s longer than for moose or elk. The grass on which the buffalo defecates reaches maturity by mid August and the buffalo dung changes from the loose summer dung to the more compact winter dung shortly after the grass cures. The time when the moose and elk pe l l e t s become hard coincides closely with the loss of green herbage, which i s the same time the leaves f a l l . Moose and elk pe l l e t s look l i k e "winter p e l l e t s " from then u n t i l green leaves appear about the middle of May. The time f o r which the defecation s i t e d i s t r i b u t i o n data for moose and elk i s v a l i d was set as October 1, 1958, to May 15, 1959, or about seven and one-half months. The winter d i s t r i b u t i o n of moose as determined by p e l l e t group transect analysis i s very similar to that shown by the dis t r i b u t i o n of moose i n the a e r i a l survey flown i n February, 1960, (Figure 5). This correlation i s a r e f l e c t i o n of one sedentary habit of the moose. No areas of concentration of elk were indicated by the p e l l e t group transect analysis. The d i s t r i b u t i o n of elk shown by the a e r i a l survey (Figure 5) i s a r e f l e c t i o n of the position of elk only at the time of the survey. Herds of 20 to 150 elk are common during the winter. These herds move from one part of the park to another, apparently at random, u t i l i z i n g suitable habitat as i t occurs. Distribution of elk as indicated by p e l l e t group locations (Figure 6) gives a much better picture of the winters use of the range. There i s very l i t t l e seasonal change i n the di s t r i b u t i o n of elk. The apparent lack of use of range i n the portion of the park north and west of Tawayik Lake may not be si g n i f i c a n t , but only a product of the sampling techniques. - 27 -Figure 4 . Summer concentration areas of moose, elk and buffalo during the summer, determined by observations during the summer. Figure 5. Winter concentration areas of moose and elk determined by an a e r i a l survey during February, I960. Seventy percent of the t o t a l population occurs within the delimited areas. - 29 -Figure 6 . '..'inter concentration areas of elk as determined by p e l l e t group transects. - 30 -B. Habitat Selection by the Animals An attempt was made to cover a l l parts of the park evenly during the summer. The habitat type and location of each animal observed during the summer was recorded. Direct observation, and inferences gained by close examination of the habitat f o r signs of use by each species are summarized i n the following Figures (see Figures 7, 8 and 9). Habitat selection during the winter i s based on an analysis of the location of p e l l e t groups and buffalo chips. Although the method has certain biases, i t i s the best reference to habitat selection which could be made by indirect methods, A description of the habitat selection, and the reason for each species selecting given habitat types i s outlined i n the following paragraphs, 1, Moose a. Summer habitat selection Habitat selection by moose, based on observations of animals coincides with the d i s t r i b u t i o n of their preferred foods. This i s to be expected since the moose are most v i s i b l e when they are standing feeding. Presence of moose i n a given habitat i s a di r e c t response to available food. Moose on the wet meadows were eating willows, those on the shrub meadows and most other places were eating primarily aspen and balm. Lack of moose i n open aspen habitat i s to be expected since there i s very l i t t l e browse available. The few preferred browse plants which do.occur have been browsed heavily by elk. During the summer, 35$ of the moose groups (one or more animals) seen were on, or near, wet meadows. Aspen-balm and hazel shrub meadows were the second most common habitats selected by moose and nearly 20$ of a l l groups seen were i n th i s habitat. Selection of other habitat types was about equal. With the exception of the open aspen areas, i n which moose were never seen, the remaining 45$ of the direct observations indicated that moose were preferring equally the six other habitat types. - 31 -A summary of habitat selection based on the summer's observations i s presented i n Figure 7a« b. Winter habitat selection Rumen analysis indicate that moose feed on hazel during most of the winter and t h i s means that they are i n the semi-^pen aspen habitat and on the hazel shrub meadows8 Observations made during December, 1959, agree with the information yielded by the rumen analysis» i : Habitat selection of moose as determined by defecation-site transects (Figure 7b) shows a heavy use of shrub meadows, semi-open aspen and the grass meadow-aspen grove habitats. The location of moose as determined by the-aerial census i n February, 1960, show a d i s t r i b u t i o n similar to that determined from the defectatien-site transect, and coincides with the suitable browse habitat. 2. E l k a. Summer habitat selection The elk show the least amount of s p e c i f i c i t y i n habitat preferences. They were seen equally frequent on a l l habitat types except the spruce bog (Figure 8a). The use made of each habitat type was not, however, the same. Generally, the elk bed down i n the wooded areas and during late spring and early summer they feed on the sedge meadows, and l a t e r , on the grass and shrub meadows. The amount of "use" elk (or other species) makes of a particular habitat may be calculated by dividing the per cent of the t o t a l area that a given habitat i s represented i n the park by the proportion of times elk were seen i n that habitat. Such calculations c l e a r l y indicate that elk were not randomly distributed. The "use" elk make of the shrub meadow habitat i s twice that of the sedge- meadow and aspen forest combined. Use of sedge meadows drops very sharply during June. During the calving period i n early June, cow elk and young calves are seldom seen on the open meadows but can be encountered frequently i n the forest. During late summer elk return to - 32 -60 I fO g 30 20 10 JZL O o O CD ce t> T3 c d) o cd cd cd O CD c P, © P . <D CO P . cd a a S M CO P- cd CO CO cd O a co co C 1 fl CD hO co CO «) • d CD CO " o +^  u cd cd P , CD Pu cd M O CD IU n P-, CD CO CD >-i o i—I CO e> O CO cd O m ^) CO HABITAT TYPES Figure 7a. Summer habitat selection by moose. Determined by observations during May, June, July and August, 1959. - 52a -60 -ko -8 30 o s PL, 20 -10 -3= 3 JL :> O o o © 13 £• c o cj d a! O <D 4) 45 4) (H S to P. a a ro 4) al a to to C 03 10 V a -P c 01 td P. © (D Ja tn P . CO e> o © P, ? © © CO CO CJ C o> p, 0) <D CO C a CD o p. in o 05 bO o ro , o cn g C O HABITAT TYPES Figure 7b• Winter habitat selection by moose. Determined by defecation site transects. - 33 -the grass and sedge meadows* b. Winter habitat selection P e l l e t group locations, summarized i n Figure 8b, indicate that during the winter elk have a strong preference for the grass meadow habitats and a lesser preference for the shrub meadows. It i s s i g n i f i c a n t that only 15$ of the p e l l e t groups were found i n the semi-open aspen habitat which covers nearly 60$ of the park, whereas during the summer, semi-open aspen was the preferred habitat. The low use of the aspen forest during the winter corresponds to the high proportion of meadow grasses eaten during the winter as revealed by rumen analysis. The winter "use" factor for grass meadow, shrub meadow and aspen grove habitat types was 0.225 as compared to 0.08 for sedge meadow and 0.04 for aspen forests, 3. Buffalo a. Summer habitat selection The buffalo show a very decided preference for grassland meadows. On a l l habitats the buffalo were using, only the grass areas were used for feeding, Buffalo use forested areas only i n t r a v e l l i n g from one grass meadow to another. They do not go to the forests for shade even on the hottest days. In the park, the only buffalo feeding on the sedge meadows are the small b u l l herds* In the i s o l a t i o n area, the only available buffalo food i s on the sedge meadows and the buffalo must, therefore feed on these meadows. Use of emergent aquatic plants, mostly sedges, by buffalo i n the i s o l a t i o n area was observed several times* Habitat selection by buffalo during the summer as shown by Figure 9a, indicates a s l i g h t l y biased emphasis on the grass meadows. The most extensive system of grass meadows are those along the road, and consequently, i f buffalo were on them they were seen every time the road was t r a v e l l e d . Probably a more accurate representation of habitat selection would show a higher use of the grassy areas of shrub meadows and the grass meadow-aspen grove association. 60 -50 ko t . 30 20 10 1. |> d o O CD c -a T3 > 'CD o cj a5 cd o CD * c P T) at CD CD U P . CD CO Cj H g n « CO- P . C! CD cd o e x> CO • co C 1 c CD to CO CD « • d ® O +> U ca CJ P . CD a p-CD X u L , CO p* CD CO CD cn c!> t3 « ; o CO c5 HABITAT TYPES CD O P. CO o cd t o r—< O cn xi CO g' r - l cn Figure 8a. Summer habitat selection by elk* Determined by observations during May, June, July and August« 1959« - 34a -60 3 50 ko ofa o o g 30 ^ 20 o OH 10 -© O O O © o t l -a > © o c< CD Cj o © p . T 3 © © « S-, P, © CO p . Cg S U t-Q co p , 01 CO s, cu o a ,o CO co C 1 © ^! CT CO © n • H © "o - p cC CJ P , © r i P-. Cj t o U >-! CO © co © I—1 o ? = C O ci> tt o C O cj m Q CO f o o l-t oo HABITAT TYPES-Pigure 8b. Winter habitat s e l e c t i o n by elk. defecation s i t e t r a n sects. Determined by - 3 4 b -XL UdL :^ CD o o O CD O X I 'd > c CD ? O cd ca' cd O CD a p. C ia o CD CD P. CD CO p Co S a CO P, ri w CD cd O a CO CO 1 .C CD co CO CD • H 0 CO -p cd CJ P. CD p.. rd i2 U CO PH £ CO CD r - l CO Ct) tt O CO cd O m CO o r H CO HABITAT TYPES Figure 9a. Summer habitat selection by buffalo. Determined by observations during May, June, July and August, 1959* Figure 9b. Winter habitat selection by buffalo. Determined by defecation site transects. - 35-b» Winter habitat selection Winter habitat preference of the cows, yearlings and most of the other age classes is not known since the herds are on the feedlot shortly after the f i r s t snow f a l l s . The habitat selection of the solitary bulls appears to be very similar both summer and winter. The only segregation of buffalo chips that could be made with any constancy was whether or not they were deposited on last summers grass or earlier. Therefore Figure 9b, representing the habitat selection of buffalo based on chip counts, should be referred to as late summer, f a l l and early spring distribution. On this basis, the habitat selection of buffalo based on defecation sites is not significantly different from the summer's observations. The high number of occurrences of chips on the meadow-grove habitat may be due to sampling error. The transect data probably represents a truer picture of summer distribution than that recorded by direct observation in the summer. Less than 10$ of a l l buffalo chips were found in aspen forest habitats. Sedge meadows accounted for 8$ of the defecation sites. Seven per cent of the defecations occurred on the shrub meadow, but a l l these were on the interdigitating grassy areas of the shrub meadow. It is evident that over 80$ of a l l were on upland meadows during the f a l l and winter. The small percentage on the sedge meadow probably represents the solitary-bull component of the herds. C. Food Habits of the Animals Observation of animals feeding during the summer were made through eight power binoculars and a 20 power telescope. The limitations of such observations in determining food preferences and diet are as follows: a. t a l l plants are seen more easily than lower vegetation; b. long-stemmed plants which are pulled up when eaten (e.g. vetch) are more easily seen; c. browse and large forbs are easier - 36 -to identify from a distance than grasses and small forbs; d 0 animals feeding in open areas are more easily observed than animals in the forest, although the latter are easier to approach. The place the animals feed and the way they feed make a great deal of difference to the accuracy and amount of observations made. Buffalo graze much closer to the ground than do the elk. Buffalo tend to eat a l l grasses and soft forbs in a patch; elk are selective in their diet and wander as they eat. Moose strip and nip willow, aspen and balm of their leaves and terminal buds during the summer; elk nip only the ends of the branches. Direct observations were made on the plants to determine utilization. The important limitations in this method are listed as follows: (a) browsing on shrubs by stripping leaves is easier to spot than nipping; (b) removal of twigs from heavily hedged shrubs is very d i f f i c u l t to evaluate; (c) grazing on immature stages of grass is more di f f i c u l t to evaluate than grazing on mature grass; (d) removal of only the main stem of grass without eating leaves cannot be detected with accuracy; (e) grazing on rhizomatous species such as clover and strawberry is almost impossible to detect by direct observation of vegetation. The time spent feeding on a particular plant or in a particular site does not necessarily bear a direct relation to the amount of food consumed (Dzubin, 1951; Smith and Hubbard, 1954). Feeding minute counts, although producing data that looks "quantitative" actually means very l i t t l e . Winter food habits were determined by examination of rumen contents collected from animals killed during the herd reduction slaughter in November and December, 1959. The food habits of moose, elk and buffalo cannot be determined accurately by any one method. Only by considering observations of animals, examination of - 37 -individual plants and the range, and rumen analysis, can a reasonable impression of the animal's diets be achieved. 1. Moose a. Simmer food habits During the summer months a l l age classes of moose fed on leaves of aspen, balm and willows * Paper birch was eaten, bat birch browse has been severely reduced due to heavy browsing in the past, Alder was abundant, especially in the north end of the park, but i t was not eaten. Other shrubby plants, such as rose, gooseberry, raspberry, buffaloberry and snowberry were used slightly. Hazel provides not more than 25$ of the summer food, but comprises about 65$ of the winter diet. Three plants, saskatoon, red-osier dogwood and chokecherry are highly preferred browse species but have been nearly eliminated from the park by heavy browsing. The approximate composition of the moose's diet during the year is shown in Figure 10, Comparative values for elk and buffalo are presented in Figure 11* Aquatic plants were used very l i t t l e during the summer. The number of fl i e s and mosquitoes was abnormally low. Possibly the moose were not driven by the insects to seek the protection of the water, Park wardens report that during the f l y season in some years, moose can be seen nearly every day feeding in the potholes and lakes, b. Winter food habits The results of analysis of samples from 15 moose rumens clearly indicates that during December, the bulk (65$) of their diet is hazel (Figure 10), Aspen and balm were represented in 60$ of the rumens analysed, but responsible for only 25$ of the total rumen contents analysed. It is significant that willow, which represent the larger part of the moose's diet in the spring and early summer, is eaten only very l i t t l e (less than 5$) in the winter. The diet during the whole winter is not known but i t can be assumed that as long as the hazel lasts moose will - 38 -continue to eat i t . The cause of the changes i n food habits and/or habitat selection during the year may be a function of the p a l a t a b i l i t y and a v a i l a b i l i t y of the browse species. Once the leaves are out, the moose turn away from the willow and begin feeding on aspen and balm. When the moose switch from hazel to willow i s not known. A l l trees and shrubs i n the park drop th e i r leaves about the same time i n the f a l l . Aspen and balm are by t h i s time severely browsed; willows are scarcely touched. Hazel, a soft browse, which has been growing without much browsing a l l summer, now receives the attention of the moose and becomes the major food item of the moose u n t i l i t i s browsed down to the point where i t i s no longer e a s i l y available. By late winter, willow i s the only browse which has not been f u l l y u t i l i z e d , whereas aspen, balm and hazel are severely hedged (see Appendix C, Photograph 6), The heavy hedging of the palatable browse causes them to produce a large number of new shoots every year. The forage production of the park i s thus very high, and t h i s i s reflected i n the high carrying capacity of the range, high reproductive rate and the f a t condition of the moose. In the i s o l a t i o n area the browse i s much more p l e n t i f u l than i n the park. However, the willows are more heavily browsed. This may be the res u l t of the lack of hazel i n the understory of the aspen forest and the non-availability of aspen and balm, 2, E l k a. Summer food habits During the f i r s t two weeks i n May food was scarce. The elk were feeding on whatever green food they could f i n d . Observations suggest that elk were spending much more time feeding during May because 85$ of a l l elk seen i n May were W I L L O W I O ° / o W I L L O W 6 0 ° / o A S P E N & B A L M 2 5 % H A Z E L 5 ° / o O T H E R S P . IO° /o SPRING A S P E N & B A L M 4 0 ° /o H A Z E L 2S°to O T H E R B R O W S E I O % F O R B S , G R A S S E S , & A Q U A T I C S P . I5 °>b SUMMER W I L L O W 5 % A S P E N & B A L M 2 5 ° / o H A Z E L 6 5 ° > b O T H E R S P . 5 ° / o WINTER Figure 10. Diet of moose in Elk Island National Park based on spring and summer observations and analysis of contents from rumens of animals killed in.-December. WILLOW io°/o ASPEN & BALM 40 % HAZEL 2 5 % OTHER BROWSE IO% FORBS, GRASSES & AQUATIC SP. 15% M O O S E ROSE 1 0 % OTHER BROWSE 15% PEAVINE & VETCH 2 5 °/o WHITE CLOVER IO% OTHER FORBS 15% FINE GRASSES (POA ETC.) 15% OTHER GRASSES & CAREX IO°/o E L K WHITE CLOVER 1 5 % OTHER FORBS lO% FINE GRASSES (POA ETCJ 65% OTHER GRASSES IO°/< BUFFALO H' igure 11. D i e t s o f m o o s e , e l k and b u f f a l o i n filk I s l a n d N a t i o n a l P a r k b a s e d o n o b s e r v a t i o n s o f a n i r o i l s a n d p l a n t s d u r i n g t h e summer . A S P E N & B A L M 2 5 % H A Z E L 6 5 % O T H E R BROWSE, F O R B S , E T C . I O % A S P E N & B A L M 2 5 % OTHER BROWSE 5 % A L L F O R B S 1 5 % F I N E G R A S S E S ( P O A , E T C ) 3 0 % C A R E X 5 % G R O U N D L I T T E R ( L E A V E S , E T C . ) 2 0 % BROWSE, F O R B S & L I T T E R IO % F I N E G R A S S E S ( P O A , E T C . ) 2S°fo C O A R S E G R A S S E S , ( C A R E X . E T C . ) 6 5 % MOOSE ELK BUFFALO Figure 12. Diet of moose, elk and buffalo during December as determined by analysis of rumen contents. - 42 -feeding. In August only 50$ of the elk herds seen were feeding. Most of the elk during May were i n sedge meadows feeding on Carex shoots. Four elk were even seen feeding on sedge i n two feet of water. As the grass on the periphery of the sloughs started to grow, the elk l e f t the sedge i n favor of the green grass. By the end of May, much green vegetation was available and elk were seldom seen on the Carex-Calamagrostis meadows. With the onset of calving, about the f i r s t week i n June, cow elk retreated to the aspen cover. Adult b u l l elk became very inconspicuous. Most of the bulls observed during June and July were yearlings and two year olds which remained with the cow herds. Food during most of the summer was varied. Vetch and peavine, when they became available about the end of May, were selected ahead of a l l other plants and represented at least 25$ of the elk's d i e t . Feeding on browse consisted mostly of i d l y nipping rose, snowberry and buffaloberry while searching f o r the legumes. Vetch and peavine must be considered the preferred food species and probably the most important food items during June. As the grass and upland meadow forbs became abundant during the l a t t e r part of June, an increasing number of elk was observed i n the grass-shrub meadow and grass meadow-aspen grove habitats. Summer food habits of elk are summarized i n Figure 11. Poa, the most common grass on the meadows, undoubtedly was the most important grass i n the elk's diet, representing about 15$ of the t o t a l food eaten. Agropyron, Bromus and Calamagrostis were used more by elk than by buffalo, but they were of minor importance i n the elk's d i e t . Although 10 to 15$ of the park has a cover of Calamagrostis under an open aspen forest, there was very l i t t l e use of the grass or the associated shrubs, gooseberry and raspberry. The preferred forbs, second to vetch and peavine, were white clover and strawberry. Sarsaparilla, although very abundant i n the aspen forest, was not eaten at a l l . Other forbs were taken i n small amounts. Forbs made up 40 to 60$ of the - 43 -diet during the summer. Prom the observations made during the summer, i t appeared that the common browse i n the diet of the elk was rose. Buffaloberry, hazel, aspen, balm, snowberry, paper birch, saskatoon, raspberry, willow and gooseberry were of secondary importance. Browse makes up 25$ of the elk's summer diet once the browse plants are i n f u l l leaf. One t h i r d of the browse di e t i s rose, b. Winter food habits Analysis of samples from 29 elk rumens showed the composition of winter food to be 30$ browse, 15$ forbs, 35$ grass and 20$ leaves of aspen and balm. The browse was predominantly aspen and balm with lesser amounts of willow, birch, saskatoon, raspberry, buffalo-berry and hazel. Identified forbs were t h i s t l e , dandelion, blueberry, strawberry, clover, goldenrod and moss.. Most of the forbs were represented only by leaf skeletons and a few stems. Mushrooms were found i n three rumens. A summary of winter food habits, based on rumen analysis, are presented i n Figure 12, 3. Buffalo a. Summer food habits Buffalo have the most r i g i d habitat requirements of the three species concerned i n this study. A l l feeding of buffalo, except that of some older b u l l s , occurs on the upland grass meadows. During the summer, almost a l l feeding buffalo were seen on grassland meadows, or closely cropped areas of wet meadows. The princip l e vegetation of the upland meadows i s introduced bluegrass and white clover, also occurring are a variety of native plants such as strawberry, yarrow, bunch-berry, anemone and miscellaneous other grasses and sedges. A l l grasses and smaller forbs appear to be u t i l i z e d . Yarrow, goldenrod, and similar t a l l , hardstemmed plants are not eaten. Snowberry, which i s the most common browse on the grassland - 44 -range i s eaten by elk but not by buffalo. Grasses growing i n the protection of snowberry, gooseberry and other shrubs are not eaten by buffalo. Figure 11 shows the summer food habits of buffalo and the corresponding data for elk and moose. The buffalo t y p i c a l l y feeds very close to the ground. As i t feeds, i t generally clears off an area one to two feet square before making a step. A l l plants are removed to within one to three inches of the s o i l . Buffalo feed very car e f u l l y around a l l small shrubs and t a l l forbs, never making any attempt to feed on them. The only times buffalo were observed u t i l i z i n g browse was when they were walking through the aspen-balm for e s t s . Occasionally hazel, or other browse, was nipped. The leaves were dropped from the mouth more often than they were swallowed. The more s o l i t a r y b u l l buffalo, although they feed most of the time on the meadows, are often found at the edges of meadows and sloughs. At times they even wade into the water and feed on emergent sedges. In the i s o l a t i o n area, a l l buffalo seen were either on the roads, or fence l i n e s , or on the sedge meadows of the larger sloughs. There are p r a c t i c a l l y no grass meadows i n the i s o l a t i o n area, so nearly 100$ of the food must come from the sedge meadows, b. Winter food habits Over 90$ of the winter d i e t of the free ranging buffalo i n the park and is o l a t i o n area i s composed of grass-like plants. The greater majority of the food i s sedge. Although not a l l small scraps of grass were examined i n the six rumens analysed, very l i t t l e Poa was found. A l l food that could be i d e n t i f i e d was Carex sp., but i t i s known from observation that the bulls do feed on the grass meadows to seme extent during the winter. From the rumens analysed, i t appears that less than 10$ of the winter diet of b u l l buffalo i s browse, forbs and leaves. The lack of leaves, forbs and Poa i n the buffalo rumens, and the presence of these foods i n the elk rumens indicate that the buffalo are feeding in different habitat types than the elk. The majority of the buffalo are on the feed lot during the whole winter. The buffalo are baited into the feed lot with feedlines as soon as the f i r s t snow f a l l s . During the winter, the buffalo have very l i t t l e effect on the sedge and grass meadows. The winter food habits of the buffalo are summarized in Figure 12, D. Disease Disease and parasites can be a significant factor in controlling the population of a herd. Conversely the population density may be one of the most important factors in determining the incidence of infection of a disease or parasite, A high incidence of a disease in one species may serve as a source of infection for the other species. This will be shown to be the case in Elk Island National Park. 1. Brucellosis Investigation of the prevalence of brucellosis in the Park started in 1946-47, but i t was not until 1957 that significant data had been collected. Corner and Connell (1957) found 42$ of the buffalo infected, and reported a "large number of abortions" in the buffalo feeding grounds. Elk had a 13$ incidence of brucellosis. The only reactors among moose were two sick animals; 124 healthy animals tested were negative. Data collected in 1958-9 have been added to that published by Corner and Connell and are presented in Table III. - 46 -Table I I I , Incidence of Brucellosis i n Moose, Elk and Buffalo K i l l e d During the Herd Reduction i n Elk Island National Park (December, 1956 to December, 1959). Number Tested Number Positive Number Suspect Number Negative % Reactors Buffalo 753 253 91 164 45.5 Elk 653 40 22 585 9.5 Moose 264 5 — 259 1.9 Effects of brucelloj sis on moose Brucella abortus i s apparently f a t a l to moose. Two of the f i v e reactors found i n E l k Island National Park were shot because they were obviously i l l . Symptoms of advanced brucellosis infection i n moose are described by Corner and Connell (1958), b» Effects of brucellosis on elk The impossibility of getting a s u f f i c i e n t l y accurate estimate of the b i r t h rate of elk to compare with the pregnancy rate found i n the cows during the herd reduction slaughter, precludes determining the effects of a 10$ incidence of brucellosis on the reproductive rate, c. Effects of brucellosis on buffalo The high incidence of Brucella abortus i n buffalo i s attributed to their grazing habit providing a greater p o s s i b i l i t y of the i r picking up the organism, and to the i r confinement to the feed l o t s during the winter. The f i r s t investigations of the rate of conception and calving was conducted by Corner during 1955-6. Pregnant and lactating buffalo were infected to the same degree as the barren cows. This situation i s similar to that found i n chronically infected herds of domestic ca t t l e (Corner and Connell, 1958). Table IV i s a summary of the data presented i n Table II of Corner and Connell. - 47 -Table IV, Calving and Conception Rates in the Elk Island National Park Buffalo Herd, 1951 to 1956, 1951 1955 1956 Number mature cows kil l e d 246 244 227 Percentage pregnant 85 67 89 Percentage mature cows raising calves (i.e, lactating cows) 77 32 13 By comparison, in the brucellosis-free herd at the National Bison Range, Montant, about 85$ of the mature cows are pregnant each year. Abortions must be few, because the percentage of cows having calves equals the pregnancy rate found during the herd reduction slaughter (Henry, pers. comm,), It is evident that the reproductive rate of buffalo in Elk Island National Park is seriously depressed by the high incidence of brucellosis in the herd, 2, Tuberculosis There has been no evidence in recent years of tuberculosis in the animals of Elk Island National Park,, 3* Parasites Assessment of the parasite load of the animals in the park is not complete enough to determine whether any parasites are having a significant effect on the health of the animals. During the past several years, only three or four bull buffalo have been shot because they were infected with lungworm (Dictyocaulus sp,). Holmes (in l i t t , , I960) examined 11 buffalo and 5 elk for parasites and did not find any animals having a seriously heavy infection. There has been no record of warble grubs (Hypoderma sp») in the animals in recent years. During the summer (1959) the animals were observed being bothered by f l i e s on only two occasions. Horse f l i e s (Tabanidae) were probably the cause of the annoyance. - 48 -E. Predators There are no effective predators i n the park. Only 25 coyote were counted during an a e r i a l survey i n February, 1960. During December and January the coyotes probably l i v e on the viscera of moose and elk shot i n the Park, but grouse, rabbits and mice are no doubt their main food source. A few calves and fawns may be taken i n the spring, but the loss cannot be very high. Black bear and wolves have been reported i n the v i c i n i t y of the Park, but the reports could not be s a t i s f a c t o r i l y confirmed. P. Mortality Due to Other Causes Very few recent skeletal remains can be found i n the Park so i t must be assumed that by f a r the highest mortality factor i s the herd reduction slaughter. During the past four years, 10 to 40$ of the population has been removed each year. Starvation i s thought to have accounted for the death of 60 or more buffalo on the feed lots during the winter of 1947-8. Similar losses occurred other years but records are not complete. Accidents are responsible for loss of some animals. Drowning, after breaking through the ice of the lakes, i s known to have caused the death of 15 buffalo and at least one elk i n the past 10 years. G« Behavior 1, Dominance relations Domination of one species by another, or of one age class by another, may affect the habitat selection and food habits of the animals. The frequency of such interactions depends on the frequency of contact between the animals. The t o t a l effect of these interactions i s very d i f f i c u l t to assess, but the presence of a "peck order" between species was evident. Three examples of observations made during the summer which indicate that there i s a r e a l social hierarchy - 49 -between buffalo, elk and moose are given below, a. Buffalo dominant over elk July 30, A herd of 180 buffalo had been at the "soap-hole" i n the south-east corner of the park for three hours. They had begun to move northward when a herd of 10 elk entered the meadow from the south. Two b u l l elk came to the soap-hole to drink and l i c k or eat the saline s o i l . About one minute after the elk arrived at the soaphole, two buffalo calves ran down to the soap-hole; the elk retreated to the edge of the forest, then slowly started back to the soap-hole. Almost as i f the elk and buffalo calves were playing a game, the calves would leave the soap-hole, the elk would approach and then retreat as the buffalo calves came towards them again. They maintained this play for nearly 10 minutes, advancing and retreating six times i n a l l . F i n a l l y the elk retreated and remained by the edge of a nearby slough u n t i l the buffalo l e f t the v i c i n i t y twenty minutes l a t e r . b. Buffalo dominant over moose Three b u l l buffalo were feeding on a grass meadow about 100 yards from an a r t i f i c i a l s a l t l i c k , A cow moose had been at the s a l t l i c k for 15 minutes when one of the buffalo walked over to the s a l t . The b u l l approached to within 40 feet of the moose, stopped and slowly swung his head from side to side (the usual buffalo threat display). The cow moose retreated. The buffalo walked to the s a l t blocks and began l i c k i n g s a l t . The moose remained about 50 feet away for f i v e minutes, then started to walk slowly up to the b u l l buffalo. The buffalo looked up, threatened, and the moose retreated. This was repeated twice more, ending i n f i n a l retreat of the moose. c. Elk dominant over moose Elk and moose were seen together 14 times during the summer. Generally neither paid much attention to the other. On two occasions the actions of elk showed i t s dominating effect on the moose. 50 -On June 18, a cow elk chased a cow moose from an a r t i f i c i a l s a l t l i c k . On July 29, a b u l l moose had been feeding on an aspen-balm shrub meadow for 30 minutes or more when three cow elk walked onto the meadow. The moose looked at them for a moment, then trotted o f f . The elk remained and fed f o r nearly one hour. No observations were made which were contrary to the above observations. The hierarchy of dominance appears to be buffalo dominant over moose and elk. Although the observations above concerning elk and moose suggest dominance. The r e l a t i o n between them i s nearly neutral. Prom the observations made, elk seem to be more tolerant of moose than moose were of elk. This dominance order i s i n agreement with that of Altmann (l952), McHugh (1958) and Brown and Simon (1947) (cited i n Peterson, 1955)). McHugh records instances of buffalo finding elk calves and chasing them or trampling them; buffalo herds chasing elk herds; buffalo chasing a moose c a l f and k i l l i n g i t ; and elk displaying dominance by charges, by swinging antlers or by st r i k i n g with the forelegs. Such actions indicate that dominance i s an active aggression. However, much of the social interaction between species i s of a passive nature where no treat or aggression takes place. Avoidance of the dominant i s the only indication of interaction, 2, Herding Behavior It i s well known that the buffalo, a plains animal i s strongly gregarious. The gregariousness of the elk depends on the type of habitat and the season of the year. The elk i s the only North American ruminant r e g u l a r i l y found i n dense forests i n herds. The moose i s a non-gregarious animal. During the summer the largest herd of moose observed was a group of three 2 year olds which are known to be t r i p l e t s . During the summer buffalo are i n mixed herds. Mature bulls remain i n small groups u n t i l the l a t t e r part of July when they begin joining the mixed groups. The decrease i n numbers of large b u l l groups corresponds to the increase i n numbers of bulls i n the mixed herds. The number of lone bulls and pairs changes very l i t t l e - 51 -during the summer. The result of the bulls joining the mixed herds is to decrease the territory occupied by buffalo. The greater use made of grass meadows by elk in late summer may be a reflection of the extra unoccupied territory, Similarily, the effective territory occupied per moose in the park is greater than the effective territory occupied per elk, or buffalo. The significance of this is not known. - 52 DISCUSSION - 53 -Summarizing the facts presented thus f a r , indicate that although there i s a very high population of moose, elk and buffalo i n the Park, there i s very l i t t l e overlap i n the d i s t r i b u t i o n , habitat selection and food habits of the animals. Moose are concentrated i n a belt across the center of the park. The elk, although they use some of the same foods as moose, seldom occur i n the region where moose congregate. Neither do the elk frequent the range used by buffalo. The overlap i n food habits of elk and buffalo are consequently minimized by a spacial separation of the herds. It appears that each species i s occupying a different niche i n the Park, and, therefore, d i r e c t "competition" i s of l i t t l e importance. Nevertheless, the interactions between them can be very real and can have si g n i f i c a n t , long-lasting effects. The greatest effect the animals have on each other i s through their effects on the range. A, Effects of the Animals on the Range 1. Browsing For the past 30 years, heavy browsing of aspen, balm and birch by moose during the summer has prevented regeneration of deciduous forests. By the end of each summer a l l aspen and balm saplings have been stripped of t h e i r available leaves, and the terminal bud removed preventing further upward growth. The areas i n the park protected from the effects of browsing show aspen, balm and willow growing at a normal rate and assuming their natural form. Elsewhere i n the park, large shrub meadows are present. The shrubs are predominantly hazel and rose on sandy s o i l s , and aspen, balm and willow on the clay s o i l s . Upon close examination of the shrub meadows, evidence of f i r e could be found. Charred aspen or balm, and roots of birch formed the major evidence. In 1946, a f i r e i n the north end of the park cleared 40 acres. Similar f i r e s have occurred at frequent intervalso "Old-timers" say that much of the area was burned over about 1900. - 54 -The exact date, cause and extent of f i r e s i n the Beaverhills could not be determined except by inference from the age of aspen and balm trees. The fact that there are s t i l l many areas of shrub meadow which have not been able to attain an aspen-balm forest, indicates that a high population of browsers must have been present for many years. Warden Roberts says that i n 1943, the condition of browse was much worse than now. Willows were eaten down and broken over i n many areas. Aspen and balm regeneration was i n a more c r i t i c a l condition than i n 1959. Warden Henderson remarked that i n 1947, trees and brush i n the south-west corner of the park were so tangled with dead branches that i t was d i f f i c u l t to ride a horse through the area. The area now supports a grass meadow-aspen grove association. Comparison of a i r photographs taken i n 1948 with the park i n 1959 indicate that some areas of shrub meadow are now t a l l saplings. Examination of trees i n such areas show the age of the trees above three feet to be about 15 to 18 years, The age of the trees near, ground level was indeterminable, but i n some cases as many as 30 rings could be counted. The k i l l i n g of aspen, balm and birch by browsing permits an increase i n available forage for buffalo and elk. Without the protection the grazing land receives by the browsing of the moose, the carrying capacity of the range for buffalo and to a lesser degree, elk, would be considerably reduced. Heavy u t i l i z a t i o n of hazel appears to have l i t t l e effect i n reducing i t s abundance i n the park. On the contrary, browsing causes an increase i n hazel. Because i t can withstand heavy browsing better than the other shrub plants i t i s more successful i n competition with them. If hazel i s absent i n an area, moose are not attracted to the area to the same degree. Consequently, the amount of browsing on aspen, balm and willow i s reduced. Heaviest browsing on aspen and balm occurs i n areas adjacent to large willow 55 -covered sloughs and spruce bogs i n the center of the park. Degree of browsing of willow during mid-summer was very l i g h t i n comparison to the extremely heavy use adjacent aspen and balm received. The moose were using the willow areas for bedding grounds and the shrub areas for feeding. 2. Grazing The grassland ranges of the park, many of them the result of cu l t i v a t i o n , have a plant composition largely of introduced f l o r a . They are probably as productive as any grassland range i n the region, natural or cultivated, Kentucky bluegrass and white clover are considered to be excellent forage and able to withstand extremely heavy use, Sampson (1952) says that Kentucky bluegrass "endures close grazing better than most species and does best when kept closely grazed." Very few unpalatable forbs are present. Yarrow, the commonest, may be considered to be an indicator of heavy grazing, but nowhere on the grass range i s i t very abundant. The damaging effect of the buffalo and elk on the s o i l i s very s l i g h t . Some compaction of s o i l occurs on heavily used areas and t r a i l s , but the grass and clover seem to be able to grow even on the main t r a i l s across the meadow. The greatest disturbing factor of the s o i l i s that of the b u l l buffalo i n creating wallowing areas. Each meadow i s pockmarked with one to a dozen or more areas of bare s o i l about eight to ten feet i n diameter. Generally, the wallows are not on h i l l s i d e s , so the p o s s i b i l i t y of them i n i t i a t i n g erosion i s s l i g h t . Only on sandy s o i l i n shrub meadow areas on the east side of the park i s there any evidence of recent erosion. In that region, a few erosion g u l l i e s can be found. Generally, throughout the park, h i l l s i d e s are well covered with vegetation and no terracing i s evident. The increased productivity of the range, produced by feces of the animals, p a r t i a l l y offsets any damaging effects caused by compaction, erosion or wallows. 3. Combined effects of browsing and grazing The degree of browsing on the range w i l l determine the amount of available - 56 -grass. Complete removal of the suppressive effects of moose and elk on the browse would rapidly decrease the grazing area to nearly n i l . Heavy browsing, to the point where a l l shrubs were k i l l e d , would increase the available grazing area by about 100$. Clearly, the carrying capacity and physiognomy of the range can be determined by manipulation of the numbers of animals i n the park. Continued f i r e suppression w i l l , however, result i n only on habitat type - the white spruce climax. The trend towards a climax white spruce forest w i l l p e r s i s t i n spite of any stocking l e v e l imposed on the range. 4. Effects of other herbivores on the range a. Deer White-tailed and mule deer are present i n the region around Elk Island National Park, but i n the park they are seldom seen. Although the deer are able to enter and leave the Park at w i l l , they are largely absent from the park. The a e r i a l survey flown i n February, 1960, indicated that there were 100 deer i n the i s o l a t i o n area and six on the overbrowsed range of the park. In contrast, 213 elk were counted i n the park and only eight i n the i s o l a t i o n area. Deer cannot compete with elk when food i s l i m i t i n g . Elk browse higher, paw more eff e c t i v e l y , move through deeper d r i f t s and range further than deer. In addition they are less specific i n their food habits and w i l l eat more dry grass than deer ( C l i f f , 1939). The presence of a heavily browsed range, and/or the presence of elk appear to be the major factors i n preventing white-tailed deer from becoming established i n the park. White-tailed deer were unknown i n the Park as late as 1951, A steady increase i n numbers has been observed since then. During the summer, only white-tailed deer were seen. Since 1900 there has been a steady northward movement of the white-t a i l e d deer, on the p r a i r i e s (Soper, 1951) with a corresponding decrease i n the numbers - 57 -of mule deer. Whether the change in distribution is a cause and effect relation, or a different reaction to a common cause (e.g. increase in amount of land cleared, or temperature change) has not been determined. b. Rodents The varying hare (Lepus americanus) was not present in the park except around the periphery and in areas from which the large herbivores had been excluded. More hares could be seen in the isolation area in one day than were seen in the park a l l summer. However, even in the isolation area they were not sufficiently numerous to have an appreciable effect on the vegetation. The numbers of hares may vary considerably, but there is no record of a dense population ever having been present. There are several species of mice present in the Park, but generally they have l i t t l e effect on the vegetation. During the winter, Microtus runways were present on the meadows around Tawayik Lake. Elsewhere grazing had been-so heavy that there was l i t t l e suitable habitat. Ground squirrels and gophers were present in the park, and mounds could be found in a few areas. Their effect on the vegetation was small. Beaver (Castor canadensis) were re-introduced rather recently. In the .-past they have had a great effect in creating sedge meadows, but now the park is in such condition that beaver can have very l i t t l e effect except around the margins of the lakes. Muskrats (Ondatra zibethica), porcupines (Erithizon dorsatum), and marmots (Marmota monax) are not present in sufficient numbers to have significant affect on the range, c. Other factors affecting the range Apart from the effects of Pomes and Saperda on aspen, the only other non-mammalian factor which must be considered are the Chrysomelid beetles which defoliated large areas of willow in the isolation area during July, 1959. Aspen and balm were infected only lightly. - 5 8 -B. Effects of Range Conditions on Food Habits of the Animals Heavy browsing maintains much of the aspen, balm and birch at a height where maximum forage i s available. Consequently, there are more of these foods i n the diet of elk and moose than would occur i f browsing was l i g h t enough to permit tree growth. The heavy grazing the grassland receives probably forces elk and buffalo to make more use of the sedge meadow than would occur i f grass were p l e n t i f u l . I t i s very d i f f i c u l t to correlate accurately the food preferences of animals i n E l k Island National Park with food preferences of similar animals else-where. F i r s t , the combination of dominant herbivores i s unique. Only i n Yellowstone Park i s the situation even closely comparable. Second, the variety of available food plants i s characteristic of only a similar habitat. No other study i s known to the author which discusses the food habits of moose or elk l i v i n g together i n an aspen forest with a s i g n i f i c a n t understory of hazel, rose, buffaloberry, gooseberry, raspberry and s a r s a p a r i l l a . Buffalo, on the other hand, have less variation i n food preferences, and everywhere they are found to be eaters of grasses or grasslike plants. F u l l e r , (1957) and Soper (1941), record the food of buffalo i n Wood Buffalo National Park as being predominantly sedge. Grass i n Wood Buffalo National Park i s very scarce and the buffalo have l i t t l e choice. In the National Bison Range, Montana, buffalo eat primarily grass with the addition of some forbs (Henry, pers. comm.). In Alaska, sedge sloughs furnish the bulk of the winter food. Summer food i s p r a c t i c a l l y a l l Calamagrostis canadensis and C_. purpurascens. Only wolf-willow i s browsed to any extent. (Hogben, i n l i t t . , 1959). The U.S. Fish and W i l d l i f e Service (1955) reports that on southern ranges. Buchloe and Bouteloua spp. are the most "acceptable" foods. On northern ranges, Agropyron tPoa, and Festuca spp. are eaten i n addition to Buchloe and Bouteloua. They say that the shorter grasses appear to be preferred to the t a l l e r and coarser species. ' - 59 -The food habits of elk at different seasons are given i n various studies as being nearly 100 per cent grass or sedge (Murie, 1951) to nearly a l l browse (Baldwin and Patton, 1938). In other studies, various proportions of browse, grasses and sedges are l i s t e d as the common or preferred foods of elk (Murie, 1951} deNio,1938; Gaffney, 1941; Cowan, 1947; Morris and Schwartz, 1957; Anderson, 1958). Clearly, they are very adaptable i n food habits and are able to l i v e i n a wide variety of habitats and eat many types of food. It may be profitable to study elk i n several areas and relate food habits with food available. Possibly a direct r e l a t i o n exists between the percentage of each palatable forage species present and the importance of the species i n the d i e t of elk. Coupled with the l a b i l e nature of the elk's food habits i s i t s non-specificity i n habitat selection. The elk's a b i l i t y to use a wide variety of foods and habitats permits i t to compete with many of the other ungulates. C. Effects of the Animals on Each Other 1. Food The principle interactions between the animals occur through their effects on the range. Any action which changes the amount of browse available w i l l d i r e c t l y affect the moose by changing the amount or nature of i t s food. S i m i l a r i l y any action which affects the grass and sedge meadows w i l l d i r e c t l y affect the buffalo. The f l e x i b l e nature of the elk i n food and habitat choice suggests that reduction i n either available browse or grass, would affect the elk's diet but probably would not affect i t s health or reproductive rate. The consequence of unrestricted increase i n numbers of moose, elk and buffalo i n Elk Island National Park would be that food would become l i m i t i n g . Then the elk would have the best chance of survival. If grazing ranges became depleted f i r s t , the buffalo would starve. The elk and moose could feed on browse.. When browse became short, the elk could l i v e on what was l e f t of the grassland, but moose would - 60 -starve. This sequence of events i s not u n r e a l i s t i c . In several instances elk have increased at the expense of deer, moose, pronghorn antelope, and bighorn sheep, e.g. Mayr, 1952, and Murie, 1951, p. 296-300. 2, Social Interactions The dominance exhibited by one species over another was not s u f f i c i e n t to be considered a s i g n i f i c a n t factor i n affecting the survival of either species. If food was i n short supply the dominant species would have the best chance of survival. In f a c t , i t has been noticed by the wardens that, i f elk are enclosed i n the winter feed l o t with the buffalo they soon become very t h i n . They w i l l not feed u n t i l after the buffalo have finished feeding and l e f t the feeding area. In the feed l o t , the buffalo display a social hierarchy, which, i f food i s short, can affect certain members of the herd. The b u l l s , cows and calves have l i t t l e trouble i n getting s u f f i c i e n t food. The long-yearlings, two and three year olds have l o s t the protection of their mothers which they enjoyed during t h e i r f i r s t winter, and are not yet large enough to assert themselves. Thus, i t i s t h i s group of sub-adult buffalo which comes through the winter i n the poorest condition i f food i s scarce. During winters when large losses of animals occurred because of starvation i t was the group of sub-adults which suffered the heaviest mortality (Warden Henderson, pers. comm.), 3. Brucellosis Nearly half the buffalo population i s infected with brucellosis, but only 10$ of the elk have contacted the disease. Obviously, the buffalo must come i n contact with the disease organism more often than elk or moose, or must be considerably more susceptible to infection, or must have a greater a b i l i t y to survive when infected. The l a s t two suggestions have not been shown to be so. Since the disease organism i s transmitted by ingestion, i n t e r s p e c i f i c transfer of the disease must occur by contact with infected milk, placenta, newborn or aborted c a l f , - or with contaminated food. The likelihood of elk or moose coming i n contact with any of the above sources - 61 -during the six months the buffalo are on the feed l o t i s very remote. However, uninfected buffalo have an excellent chance of encountering viable bacteria. Since most abortions occur while the buffalo are on the feed l o t , t h i s area i s l i k e l y to be the most heavily contaminated area of the Park. The only chance elk and moose have for contacting the brucellosis organism i s on the open range during the summer. Because of the high incidence of Brucellosis i n buffalo and the s i m i l a r i t y i n habitat selection of elk and buffalo, the chance of elk contacting brucellosis from buffalo i s greater than i t s chance of contacting i t from another elk or a moose. S i m i l a r i l y moose must achieve most of their infections by contact with vegetation contaminated by buffalo. Possibly moose and elk are more resistant to infection. However, i n view of the information available there i s no need to postulate a difference i n s u s c e p t i b i l i t y to explain the variation i n incidence between species. Differences i n the probability of: contacting the organism are s u f f i c i e n t to explain the differences i n incidence between the three species. D. Influence of Environment on Habitat Selection and Distribution of the  Animals and the Carrying Capacity of the Range 1. Climate Snow i s probably the only climatic factor which can d i r e c t l y l i m i t the d i s t r i b u t i o n of moose, elk and buffalo. Low winter temperatures do not affect the healthy, well fed animal. Other climatic factors produce their effects on the d i s t r i b u t i o n through changes i n vegetation and a v a i l a b i l i t y of water. The amount of precipitation w i l l severely affect the a v a i l a b i l i t y of food. Seldom i s r a i n f a l l i n s u f f i c i e n t to produce adequate forage on upland meadows, but after a series of wet years the large sedge meadows become flooded. The present high carrying capacity of the buffalo range i n the i s o l a t i o n area i s due to a series of r e l a t i v e l y dry years, A few wet years would have drastic results on the buffalo - 62 -population unless a r t i f i c i a l feeding was conducted on a year-round basis. 2, Water Because of the very heavy dew which occurs every night, the animals seldom f i n d i t necessary to go to watering holes. Although several thousands of animal-hours of observations were made, drinking animals were observed on only four occasions. No watering places were found which were used frequently by moose and elk. Buffalo frequent a few areas which look l i k e watering holes, but they were never observed drinking at these places. Wide d i s t r i b u t i o n of water and the heavy dews indicate that available water has l i t t l e a f fect i n r e s t r i c t i n g the d i s t r i b u t i o n of the animals. On the contrary, it-probably permits as even a use of the range areas as could be attained by any method,, 3. Pood Food habits and habitat selection of moose, elk and buffalo i n Elk Island National Park are i n accordance with the' hypothesis "Sympatric species or strains either occupy different habitats within the same t e r r i t o r y , or exploit the habitat i n different ways..." (Dobzhanzky and Pavan, 1950), Under present range conditions there i s very l i t t l e overlap i n food habits, either i n summer or winter, between moose, elk and buffalo. As can be seen from Figures 4, 5 and 6, d i s t r i b u t i o n of the three species overlaps very l i t t l e . Differences i n habitat preference indicate that even where di s t r i b u t i o n overlaps, the animals are not occupying the same niche. E, Anatomical Reasons for Habitat Selection Structural modifications permit the buffalo to u t i l i z e the grassland range better than the forest. The buffalo has short legs and carries i t s head low. Conversely, moose with t h e i r long legs and long neck are able to reach higher than - 63 -a buffalo but are unable to graze short grass ranges. The large nose on the moose is an advantage in browsing, but a hindrance to close grazing. Allee et a l . (1949, p. 241) noted similar anatomical differences between the black rhinoceros which feeds on browse and the grazing white-rhinoceros» Elk and deer are less specialized and are able to use a l l types of food. The distribution of moose and buffalo in North America prior to the presence of white-man was limited by the distribution of particular forms of food, e.g. browse and grass-like plants respectively. Elk use a wider variety of plant forms for food than do moose or buffalo and has a correspondingly wider geographical distribution. - 64 -SUMMARY - 65 -1. The soi l , climate and vegetation of Elk Island National Park indicate that with continued fire suppression the Park will eventually support a climax forest of white spruce. The rate of formation of a spruce forest is dependent only on the rate of dispersal of spruce and the complete absence of f i r e . It is not affected by heavy grazing or browsing, 2. A l l grasslands in the Park are the result of cultivation or heavy browsing preventing establishment of an aspen-balm forest. 3. The stocking level of the range is heavier than the range would support under pristine conditions: a, buffalo have to be fed during the winter; b, browsing is heavy enough to nearly eliminate three common shrubs, viz, saskatoon, dogwood, and chokecherry; c, regeneration of an aspen-balm forest is prevented by heavy browsing, 4 . The principle interactions between the animals are through their effects on the range, i e Moose have the greatest effect in determining the physiognomy of the range. During the summer, browsing on aspen, balm and birch prevents a deciduous forest from becoming established on the grass-shrub meadows. Prevention of reforestation provides more grassland range for elk and buffalo. i i . The habits of the elk permit them to utilize either forest or meadow habitats. During the summer, elk were concentrated in the aspen forest and areas of aspen grove. The effects of elk during the summer on browse plants is very slight. During the winter, as grass ranges become depleted, elk turn more to browsing. Browsing by elk compliments the effects of moose* Heavy use by elk of sedge meadows during early spring prepare these meadows for buffalo. The f l e x i b i l i t y of the elk's habits would permit i t to survive at the expense of deer, moose, and buffalo i f food were in c r i t i c a l l y short supply. i i i . Buffalo prefer to feed on short grass and sedge plants rather than on - 66 -t a l l coarse grasses and sedges. They a s s i s t the elk i n consuming a l l available forage on grass meadows each year. Use of sedge meadows by elk and buffalo i s a response to the shortage of suitable grass. 5. The condition of the range i n the park, produced by the combined effects of moose, elk and buffalo, makes i t unsuitable habitat f o r deer and hares. 6. Brucellosis has a depressant effect on the reproductive rate of buffalo. Buffalo serve as a reservoir host of Brucella and are probably responsible for the occurrance of brucellosis i n elk and moose. 7. Interspecific dominance was not shown to be a sig n i f i c a n t factor i n determining the habits and survival of the animals under existing conditions. - 67 -LITERATURE CITED Allan, J.A S Geology. Alberta Research Council Report No. 34. Alle e , W.CB, A.E. Emerson, 0. Park, T. Park and K.P. Schmidt. 1949. Principles of Animal Ecology. Philadelphia, W.B. Saunders Co. 837 pp. Altmann, Margaret. 1951. Social behavior of elk, Cervus canadensis nelsoni, i n the Jackson Hole area of Wyoming. Behavior, 3(4): 116-144. 1952. Social behavior of elk, Cervus canadensis nelsoni, i n the Jackson Hole area of Wyoming. Behavior, _4, 2. 1956. Patterns of social behavior i n big game (of the United States and Europe). Trans. North Amer. Wildl. Conf., 21: 538-545, Anderson, C.C. 1958 (1959). The Elk of Jackson Hole. A review of Jackson Hole elk Studies. Wyo, Game and Fish Comm., B u l l , no, 10, 1—184. Bird, R.D, 1930. Bi o t i c communities of the Aspen parkland of central Canada, Ecology. 11: 355-442, Bowser, W.E. and R.L. Erdman. 1947. S o i l survey of Peace H i l l s sheet. Report No, 14 of the Alberta S o i l Survey, Univ. of Alberta B u l l e t i n No, 48, Canada Tear Book, 1957-58. 1958. Dominion Bureau of S t a t i s t i c s . Ottawa, C l i f f , E,P. 1939, Relationship between elk and mule deer i n the Blue Mountains of Oregon, Trans. N. Amer. Wildl. Conf. 4: 560-569. Corner, A.H. and R. Connell. 1958. Brucellosis i n bison, elk and moose i n Elk Island National Park, Alberta, Canada. Can. Jour. Comp. Med, and Vet. S c i . 22: 9-20. Cowan, I. MoT, 1947. Range competition between mule deer, bighorn sheep, and elk i n Jasper National Park, Alberta, Trans. N, Amer. Wildl. Conf, 12: 223-227. DeNio, R.M. 1938, Elk and deer foods and feeding habits. Trans. N. Amer, Wildl. Conf. 3: 421-427. Dobzhansky, Th., and C. Pavan. 1950. Local and seasonal variation i n relative frequencies of species of Drosophila i n Brazil<• Jour, Arum. Ecol., 19: 1-14. Dzubin, A. 1951* P a l a t a b i l i t y studies on some foods of the Columbian Black-tailed deer, Odocoileus hemionus columbianus (Richardson), B.A, thesis, Dept. of Zoology, University of B r i t i s h Columbia, Vancouver, B.C. pp. 44. E l l i s o n , L, and W.R. Houston, 1958, Production of herbaceous vegetation i n openings and under canopies of Western Aspens, Ecology, 39(2): 337-345, F u l l e r , W.A, 1957. The biology and management of the Bison of Wood Buffalo National Park, Ph.D. thesis, Univ. of Wise, 130 p. - 68 -Gaffney, W.S. 1941. The effects of winter elk browsing, South Fork of the Flathead River, Montana, Jour. Wildl. Mgmt, 5: 427-453. Greig-Smith, P. 1957. Quantitative Plant Ecology. London, Butterworths Scientific Publications* 198 pp, Grzimek, M. and B. Grzimek. I960, Census of plains animals in the Serengeti National Park, Tanganyika, Jour, Wildl. Mgmt., 24(1): 27-37, Hogben, J, 1959, In letter to the author. July, 1959. Holmes, J.C, 1960. In letter to Mr. H. Webster, Superintendent, Elk Island National Park, March 26, I960. Julander, 0. 1958. Techniques in studying competition between big game and livestock. Jour. Range Mgmt,, l l ( l ) : 18-21. Lack, D. 1944, Ecological aspects of species-formation in passerine birds. Ibis, 86s 260-280, Mair, W,W, 1952, The impact of an introduced population of Elk upon the biota of Banff National Park. M.A. thesis, Univ. of B.C. 113 pp. McHugh, T.C 1958. Social behavior of the American Buffalo (Bison bison bison). Z o o l o g i c a l : 1—40. McMillan, J.F, 1953, Measures of associations between moose and elk on feeding grounds. Jour. Wildl. Mgmt., 17(2): 162-166. Morris, M.S. and J.E, Schwartz. 1957, Mule deer and elk food habits on the National Bison Range. Jour. Wildl. Mgmt. 21(2): 189-193. Moss, E.H, 1932. The vegetation of Alberta IV, The poplar association of central Alberta. Jour. Ecol.. 20: 380-415. Murie, 0,J, 1951. Elk of North America. Stackpole Co., Harrisburg, Penn, 376 pp, Parker, K.W, 1954. A method for measuring trend in range condition on National Forest Ranges with supplemental instructions for measurement and observation of vigor composition and browse. U.S. Dept, of Agric, Forest Service. Washington, D.C. Peterson, R.L. 1955. North American Moose. Toronto, Univ. of Toronto Press. 280 pp. Sampson, A.W. 1952. Range Management. New York, John Wiley and Sons, 570 pp. Smith, A..D, and R.L, Hubbard. 1954. Preference rating of winter deer food. Utah. Jour. Range Mgmt., 7: 262-266. Soper, J.D, 1941. History, range and home l i f e of the northern Bison. Ecol. Monographs 11: 347-412, - 69 -Soper, J.D, 1951. Mammals of Prince Albert National Park, Wildl. Mgmt, Bull. Series 1, No. 5. Canadian Wildlife Service, Ottawa, 48 pp. United States Dept. of Interior, 1955, Management of Buffalo Herds. Fish and Wildlife Service, Wildlife leaflet 212. March, 1955. 8 pp., mimeo. ' APPENDIX A LIST OF PLANTS OF ELK ISLAND NATIONAL PARK For proper interpretation of the data presented in the sections on Park vegetation, food habits of the animals and analysis of the range (Appendix B), a l i s t of nearly a l l vascular plants commonly found in the Park has been included. Thus, the total range of foods available to the animals can be learned and the reasons for the animals not eating many of the foods eaten elsewhere are soon evident. For example, the Park is far outside the distribution of f i r , the common winter food of deer, elk and moose in many areas. Buffalo grass and gamma grass, once the mainstay of buffalo on the prairies, are now uncommon everywhere. However, they probably never did occur in the Park, The l i s t of plants also provides a quick reference to the names of plants referred to only by their common names in the main body of this paper. Conversely, for one not familiar with scientific plant names, i t provides an index to the common names of plants which have been mentioned only by their latin names. - 71 -List of Plants of Elk Island National Park Pinaceae (Pine Family) Larix laricina Picea glauca Picea mariana Pinus sp. Typhaceae (Cattail Family) Typha l a t i f o l i a Sparganaiaceae (Bur-reed Family) Sparganium eurycarpum Najadaceae (Pondweed Family) Potamogeton pusillus Potamogeton richardsonii Juncaginaceae (Arrow-grass Family) Triglochin maritima Gramineae (Grass Family) Agropyron subsecundum Agropyron trachycaulum Agrostis scabra Beckmannia syzigachne Bromus ciliatus Bromus inermis Calamagrostis canadensis Danthonia spicata Elyraus canadensis Elymus glauca Glyceria grandis Hordeum jubatum Koeleria cristata Oryzopsis asperifolia Phalaris arundinacea Fhleum pratense Poa nervosa Poa palustris Poa pratensis Puccinellia nuttalliana Schizachne purpurascens Tamarack "White spruce Black spruce Pine (one specimen, dead) Common cattail Broad fruited bur-reed Richardsons pondweed Arrow grass Awned wheat grass Slender wheat grass Rough hair grass Slough grass Fringed Brome Awnless brome Marsh reed grass Poverty oat grass Nodding wild rye Smooth wild rye Tall manna grass Foxtail barley June grass Rice grass Reed canary grass Timothy Wheelers blue grass Fowl bluegrass Kentucky blue grass Nuttall's salt meadow grass False melic Cyperaceae (Sedge Family) Carex aenea Carex antheroides Carex aquatilis Carex bebbii Carex canaescens Carex crawfordii Carex interior * A l l specimens collected by the author and identified by Dr. E.H. Moss and staff at Department of Biology and Botany, University of British Columbia. - 72 -Carex rostrata Carex sp. Eleocharis palustris Eriophorum angustifolium Eriophorum sp. Scirpus microcarpus Lemnaceae (Duckweed Family) Lemna minor Juncaceae (Rush Family) Juncus balticus Liliaceae (Lily Family) Disporum trachycarpum Lilium philadelphicum var. andinum Maianthemum canadense Smilacina stellata Smilacina t r i f o l i a Orchidaceae (Orchid Family) Corallorhiza striata Habenaria hyperborea Salicaceae (Willow Family) Popuius balsamifera Popuius tremuloides Salix bebbiana Salix commutata Salix discolor Salix drummondiana Salix maccalliana Salix melanopsis Salix myrtillifolia Salix pedicellaris var. hypoglaucus Salix pyrifolia Betulaceae (Birch Family) Alnus tenuifolia Alnus crispa ? Betula papyifera var, subcordata Betula pumila var, glandulifera Corylaceae (Hazel Family) Corylus cornuta Urticaceae (Nettle Family) Urtica gracilis Polygonaceae (Buckwheat Family) Polygonum achoreum Polygonum amphibium var, stipulaceum Creeping spike-rush Tall cotton-grass Small-fruited bullrush Lesser duckweed Wire rush Fairybells Western Red L i l y Two-leaved solomon's seal Star-flowered solomon's seal Three-leaved solomon's seal Striped coral root Green-flowered bog orchid Balsam poplar Aspen poplar Beaked willow Paper birch Scrub birch Beaked hazelnut Common nettle Striated knotweed - 73 -Chenopodiaceae (G-oosefoot Family) Axyris amaranthoides Monolepis nuttalliana Caryophyllaceae (Pink Family) Arenaria lateriflora Stellaria longifolia Ceratophyllaceae (Hornwort Family) Ceratophyllum demersum Ranunculaceae (Crowfoot Family) Actaea rubra Anemone multifida Caltha palustris Ranunculus aquatilis var, capillaceus Ranunculus gmelini Ranunculus sceleratus Cruciferae (Mustard Family) Descurainia sophia Draba nemorosa Lepidium densiflorum Saxifragaceae (Saxifrage Family) Chrysosplenium iowense Parnassia palustris Ribes lacustre Ribes oxyacanthoides Ribes triste Rosaceae (Rose Family) Agrimonia striata Amelanchier alnifolia Fragaria virginiana var. glauca Geum aleppicum var, strictum Potentilla gracilis var, rigida Potentilla norvegica Potentilla palustris Potentilla pensylvanica Potentilla platensis Prunus pensylvanica Prunus virginiana var, me'lanocarpa Rosa acicularis Rubus acaulis Rubus ehamaemorus Rubus pubescens Rubus strigosus Leguminosae (Pea Family) Lathyrus ochroleucus Lathyrus venusosus Russian pigweed Spear-leaved goosefoot Blunt-leaved sandwort Long-leaved stitchwort Hornwort Red baneberry Cut-leaved anemone Marsh, merigold Small yellow water-crowfoot Cursed crowfoot Flixweed Wood whitlow grass Common peppergrass Golden saxifrage Northern grass of parnassus Bristly black current Wild gooseberry Wild red current Agrimony Saskatoon berry Smooth wild strawberry Yellow avens Graceful cinquefoil Rough cinquefoil Marsh cinquefoil Prairie cinquefoil Pin cherry Choke cherry Prickly rose Dwarf raspberry Cloud berry Running raspberry = Dewberry Wild Red Raspberry Cream-coloured vetchling Wild pea-vine - 74 -Trifolium pratense Trifolium repens Vicia americana Balsaminaceae (Touch-me-not Family) Impatiens capensis = biflora Violaceae (Violet Family) Viola glabella Viola rugulosa Elaeagnaceae (Oleaster Family) Elaeagnus commutata Shepherdia canadensis Onagraceae (Evening Primrose Family) Epilobium angustifolium Haloragidaceae (Water-milfoil Family) Myriophyllum exalbescens Araliaceae (Ginseng Family) Aralia nudicaulis Umbelliferae (Parsley Family) Heracleum lana.tum Sanicula marilandica Cornaceae (Dogwood Family) Cornus canadensis Cornus stolonifera Pyrolaceae (Wintergreen Family) Pyrola asarifolia Pyrola secunda Ericaceae (Heath Family) Ledum groenlandicum Vaccinium microcarpos ? Vaccinium myrtilloides = Vaccinium oxycoccus Vaccinium vitus-idaea Primulaceae (Primrose Family) Androsace septentrionalis var. puberu-lenta Lysimachia c i l i a t a Gentianaceae (Gentian Family) Menyanthes tr i f o l i a t a Boraginaceae (Borage Family) Mertensia paniculata Red clover White clover American, vetch Spotted touch-me-not Western Canadian violet Silver willow = wolf willow Canadian buffalo berry Fireweed Spiked water milfoil Wild sarsaparilla Cow parsnip Snake root Bunchberry Red Osier dogwood Common pink winter-green One-sided wintergreen Laborador tea Small bog cranberry V. canadense - Blueberry Swamp cranberry Bog cranberry Fairy candelabra Fringed loosestrife Buck-bean Tall lungwort - 75 -Labiatae (Mint Family) Agastache foeniculum Mentha arvensis var, villosa Scrophulariaceae (Figwort Family) Castilleja rhexifolia Orthocarpus luteus Plantaginaceae (Plantain Family) Plantago major Rubiaceae (Madden Family) Galium boreale Galium labrodoricum Galium trifidum Galium triflorum Caprifoliaceae (Honeysuckle Family) Linnaea borealis var, americana Lonicera dioica var. glaucescens Lonicera involucrata Symphoricaxpos albus var. pauciflorus Symphoricarpos occidentalis Viburnum edule Campanulaceae (Bluebell Family) Campanula rotundifolia Compositae (Composite Family) Achillea millefolium ssp, lanulosa Achillea sibirica Antennaria neglecta Antennaria parviflora Arnica cordifolia Artemi s ia f r ig ida Artemisia ludoviciana var. gnaphalode Aster ciliolatus Aster conspicuus Aster modestus Aster pansus Aster puniceus Bidens cemua Chrysanthemum leucanthemum Cirsium arvense Cirisum undulatum Crepis tectorum Erigero philadelphicus Hieracium umbellatum Matricaria matricarioides Petasites sagittatus Senecio congestus var, palustris Giant hyssop Wild mint Common Indian paint-brush Owl clover Common plantain Northern bedstraw Laborador bedstraw Small bedstraw Sweet-scented bedstraw Twin-flower Twining honeysuckle Involucrate honeysuckle Few-flowered snowberry Western snowberry Lowbush cranberry = Mooseberry Bluebell Woolly yarrow Many-flowered yarrow A. campestris - Pussytoes = Everlasting Small leaved everlasting Heart-leaved arnica Pasture sage s Prairie sage Showy aster Purple stemmed aster Nodding beggar-ticks Ox-eye daisy Canada thistle Wavy-leaved thistle Annual hawks beard Philadelphia fleabane Narrow-leaved, hawks weed Pineapple weed Arrow-leaved coltsfoot Marsh ragwort - 76 -Senecio pauperculus Solidago decumbens var. oreophila Solidago lepida var. elongata ? Solidago pruinosa Sonchus aspen Sonchus uliginosus Taraxacum officinale Balsam groundsel Mountain goldenrod Graceful goldenrod Canadian goldenrod Annual sow thistle Perennial sow thistle Dandelion - 77 -APPENDIX B SUMMARY OP RANGE ANALYSIS In the following summaries of transect analysis a general description of the habitat i s given. The per cent of " h i t s " on each class of plant (grass, forb or shrub), segregated into an a r b i t r a r i l y determined c l a s s i f i c a t i o n of p a l a t a b i l i t y , are l i s t e d i n a table. The percentage plant cover indicates "the number of points which h i t vegetation other than moss. The percentage composition of the plants l i s t e d i n the f i n a l paragraph of each transect summary i s based on the number of h i t s on each species i n rel a t i o n to t o t a l plant cover. For the purposes of t h i s summary, "palatable species" i s used to mean a plant which i s commonly eaten by one of the ruminants. Conversely, "unpalatable species" refers to a plant which the animals seldom eat or avoid eating. Palatable grasses include Poa, Agropyron, Bromus, Agrostis, Koeleria, and most Carex species. Unpalatable grasses are Calamagrostis, Phalaris, Hordeum, Glyceria and the t a l l rank Carex species. Of the common forbs, a l l are palatable except A r a l i a , Solidago, Artemisia, Antennaria and Sonchus» Palatable browse species are aspen, balm, birch, willow, cherry dogwood, saskatoon, hazel, rose and Vibrunum, The major unpalatable species are gooseberry, raspberry, buffaloberry, snowberry and alder. Figure 13 shows the location of the range transects and following i t (Figure 14) i s a sample of the form used to record the range analysis data. - 78 -SUMMARY OF RANGE ANALYSIS Transect 1« Tawayik Lake Meadow. This meadow is a natural grass-forb meadow at the north end of Tawayik Lake. The presence of charred logs on the meadow indicate that this meadow, like many of the others, is a stage in a fire succession. There are very few shrubs, and those present are very heavily browsed. See Photograph 3. Soil The meadow has a heavy hydromorphic s o i l . Nearby areas have sandy or gravelly soils. Vegetation Table V, Summary of Data From Transect 1, Palatable species Unpalatable species Grasses 45$ t r . Forbs 29$ 2$ Shrubs t r , t r . Total plant cover 77$ Of the total palatable forage, 40$ was Poa pratensis, and 29$ was Trifolium repens. Other species present in amounts greater than 1$ were: Agropyron  subsecundum, 7$; Agropyron trachycaulum, 4$j Agrostis, 1$; Taraxacum, 3$; Fragaria, 4$ and Achillea, 1$. Transect 2. Soap-hole Meadow, This meadow, located in the south east corner of the park, is possibly the only true parkland meadow in the Park, This is suggested by the absence of any indication of fire and. the presence of at least one characteristically prairie - 79 -grass (Koeleria sp,). The range is less productive than the a r t i f i c i a l grasslands located elsewhere in the park. See Photograph 2, Soil The soil has a five inch layer of black soil with a thick mat of grass roots* Below this, there is a very sticky, wet clay horizon with a lime layer at 21 inches, (The lime layer of grey-wooded soils is generally at 36 to 40 inches). Vegetation Table VI, Summary of Data Prom Transect 2, Palatable species Unpalatable species Grasses 47$ 2$ Forbs 4$ 16$ Shrubs tr. 0 Total plant cover 70$ Of the total forage, 18.5$ was Poa pratensis, 36$ was Koeleria and only 2$ was Trifolium. Other species present in lesser amounts were: Agrostis, 8$j Agropyron sp., l$j Poa nervosa, 2$; Potentilla, 5$; Achillea, 4$; Fragaria, 2$ and Antennaria, 1$. Transect 3. Grass-shrub meadow. This meadow represents one of the cultivated grass meadows along the main road through the park. Appearance and species composition appear similar to the other meadows along the road. Considerable Symphoricarpos has established in spite of the heavy browsing and grazing the meadow receives. Growth of aspen, balm and willow present on the meadow is completely controlled by browsing. See Photograph 4. - 8 0 -S o i l Black s o i l extended to a depth of three to f i v e inches; below t h i s , a s i l t y clay layer, Vegetation Table VII, Summary of Data Prom Transect 3, Palatable species Unpalatable species Grasses 60$ 0 Forbs 17$ 1$ Shrubs 1$ 0 Total plant cover 78$ Shrub intersect: 71 feet Symphoricarpos Of the t o t a l vegetation 57$ was Poa pratensis; Trifolium accounted for 13$. Other plants were: Agropyron, 6$; Taraxacum, 4$; Phleum, 2$ and Carex sp,, 2$. Transect 4, Grass-shrub meadow. Hazel and balm shrub are the dominant plants of th i s meadow, Grass occurs only i n small patches interspersed i n the thick shrub cover. This meadow on the east side of the park, and similar ones, one west of Tawayik Lake and one southwest of Astotin Lake, are a l l of the same general appearance. A l l are the result of a supressed f i r e succession. The age of surrounding trees indicates that the f i r e burned the area about 1900, So i l Black s o i l two inches; sandy clay below. - 81 -Vegetation Table VIII. Summary of Data Prom Transect 4. Palatable species Unpalatable species Grasses 18$ 0 Porbs . 20$ 6$ Shrubs t r . 4$ Total plant cover 48$ Shrub intersect: 210 feet The meadow was composed of 30$ Poa, 17$ Fragaria, and 15$ Trifolium, Several other species were present i n amount of 3$ or less . The dominant browse plants were predominantly hazel, but with significant amounts of willow, balm, aspen, snowberry and birch. A l l but a few shrubs were less than three feet high. Transect 5. Grass-shrub meadow. Transect 5 i s located about one mile east of the north end of Tawayik Lake, Aspen and balm are the dominant shrubs. Hazel, which was most abundant on Transect 4, i s represented by only one plant i n the transects on t h i s meadow. Age of aspen trees and shrubs varies from eight to twenty years. The height of a l l shrubs i s less than six fe e t . S o i l The meadow has a black s o i l two inches deep, a leached A horizon of ten inches and a B horizon of brownish clay with some sand. - 82 -Vegetation Table IX. Summary of Data Prom Transect 5. Palatable species Unpalatable species Grasses 29$ 1$ Forbs 25$ 2$ Shrubs 0 1$ Total plant cover 58$ The vegetation of the meadow was composed of 35$ Poa. 20$ Trifolium, 11$ Agropyron, and 9$ Fragaria. Many other plant species were present in lesser amounts. Solidago sp. was abundant on the meadow. In the browse transects (1,200 square feet) there were the following number of shrub plants: 96 aspen, 80 balm, 52 rose, 58 willow, 39 buffalo berry, 33 snowberry, 26 birch, 19 raspberry, and 22 hazel. Lesser numbers of gooseberry, saskatoon and chokecherry were present. Photograph 5 represents a typical portion of this meadow. Transect 7. 1946 Burn. This transect is located in a burned over stand of mature aspen. In 1946, fire killed a l l live trees on about 40 acres in the north end of the park. No regeneration of aspen or balm has taken place. Tree trunks which are s t i l l standing can be pushed over easily. The ground is covered with a tangle of rotting logs and a heavy cover of grass, (See Photograph 10). Soil The soil has considerable sand and gravel* - 83 -Vegetation Table X. Summary of Data Prom Transect 7. Palatable species Unpalatable species Grasses 7$ 11$ Forbs 13$ 3$ Shrubs 1$ - 2$ Total plant cover 37$ Phalaris and Calamagrostis accounted for 27$ of the vegetation. Other species were: Agropyron, 8$; Poa, 4$; Fragaria, 20$; Rubus, 10$; and several other species present i n amounts less than 4$. Eighteen per cent of the t o t a l transect points were rotting logs. Raspberry, rose, snowberry and hazel were the most abundant sirubs. A l l shrubs were less than four feet high. Transect 8. Isolation area - Aspen forest. This transect d i f f e r s from the others i n that no permanent transect li n e s were located. The shrub vegetation was analysed i n the usual way, i . e . by four rectangular plots 3 feet by 100 feet. A l l shrubs, trees and t a l l forbs occurring within the plots were divided into height classes and counted. The forest i s composed of aspen and balm, 35 to 40 years old, with an understory three to ten feet high. In the 1,200 square feet (approximately 1/30 acres) there was the following number of tree, shrub and forb plants: Solidago, 200; rose, 108; raspberry, 88; saskatoon, 84; willow, 75; snowberry, 54; hazel, 51} A r a l i a , 45} aspen, 33; balm, 18 and cherry, 13. The willows and saskatoon were the t a l l e s t shrubs. They averaged three to six feet high. The general form of the forest i s represented by Photograph 8. - 84 -Transect 9. Semi-open aspen forest. This transect i s located about two miles south of the north boundary of the park and 1.25 miles from the east boundary. The forest i s typi c a l of most of the area designated on Map 2 as serai-open aspen 0 Aspen and balm trees are 38 to 50 years old, 50 to 60 feet high and four to nine inches i n diameter. S o i l The s o i l i s a very sandy loam to a depth of at least eight feet. Vegetation Table XI. Summary of Data Prom Transect 9. Palatable species Unpalatable species Grasses 1$ 3?° Forbs 6$ Jfo Shrubs 2$ 3$ Total plant cover 17$ Most of the grass present i n the transects was Calamagrostis. Rose and hazel were present i n the shrub understory i n equal amount. Raspberry was the t h i r d most abundant browse species. Nearly a l l browse plants were less than four feet. Photograph 7 represents the semi-open aspen habitat. - 8 5 -Figure I J . Locations of range t r a n s e c t s e s t a b l i s h e d i n 195 - 86 -Record of Permanent L i n e T r a n s e c t P a r k C l u s t e r No. Date measured Observer S p e c i e s T r a n s e c t No. T r a n s e c t No. H i t s T o t a l H i t s T o t a l P l a n t d e n s i t y Index Bare S o i l E r o s i o n Pavement Rock L i t t e r Moss T o t a l 100.0 T o t a l 100.0 For a g e d e n s i t y i n d e x Ground c o v e r i n d e x . O v e r s t o r y . . . . U n d e r s t o r y . . . An n u a l s Figure 14. Sample of form used to record range transect data. P l a n t V i g o u r Measurements S p e c i e s T r a n s e c t No. T r a n s e c t 1 1 2 3 4 1 '" 6 . 7 8 9 10 T o t a l Av. Max. P e l l e t G roup Counts Width • Length t o S p e c i e s T r a n s e c t No. T r a n s e c t No. APPENDIX C PHOTOGRAPHS OP RANGE AND H4BITAT TYPES Photograph 1, Large slough by road near north end of Elk Island National Park, June 23, 1959. Typical slough, sedge meadow with willow, birch and goose-berry shrub on periphery. Photograph 2. Prairie meadow near "soap-holes", August 28, 1959. Possibly the only area of true prairie type vegetation. Grass is mostly Poa and Koeleria. See summary of Trar-soct 2. Photograph 3. Grass-forb meadow north end of Tawayik Lake, August 14, 1959. This i s the natural meadow type for the BeaverMlls. Note the large number of forbs, (mostly Yarrow), and the presence of Agropyro^. See summary of Transect 1, Photograph 4. Grass meadow formed by cult i v a t i o n i n 1044, Agust 14, 1959. This grass meadow consists largely of Poa and white clover. Aspen, balm, willow, saskatoon and other shrubs are present but cannot mature because of heavy browsing. See summary of Transect 3. - 89 -Photograph 5. Aspen-balm shrub meadow, August 25, 1959. This is a thick stand of aspen and balm on a grass meadow. A l l browse is maintained within reach of moose and elk. See summary of Transect 5. Photograph 6. Aspen-balm shrub meadow, May 5, 1959. This meadow is similar to the one in photograph 5. Note heavily browsed condition of aspen and extreme close cropping of grass. Also note the presence of grass stems where protected by the shrubs. Similar close grazing is found on a l l grass meadows in early May. Photograph 7. Semi-open aspen habitat, August 17, 1959. Note thick understory of hazel and sarsaparilla. Height of a l l browse, except rose, is less than three feet. Very l i t t l e grass is present. See summary of Transect 9. Photograph 8. Aspen forest in isolation area, August, 1959. Edge of clearing cut in 1958. Note the lack of leaves on the lower two-thirds of the aspen trees, and the presence of dead trees in the stand. Photograph 9. Open aspen stand with understory of Calamagrostis 'jrass. West of Tawayik Lake, May 6, 1959. Note heavy browsing of shrubs in fore-ground and lack of use of grass. Photograph 10. Open aspen stand after burn in 1946, August 29, 1959. Note complete lack of regenerating aspen and balm. Photograph 11. Open aspen stand after burn in 1946, August 29, 1959. Calamagrosti s, Bromus and Agropyron grasses present show very l i t t l e evidence of grazing. A l l browse species are supressed. See summery of Transect 7. 

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