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Some factors affecting the winter range of Jasper National Park Pfeiffer, Egbert Wheeler 1948

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& 3 a 7 SOME FACTORS AFFECTING THE WINTER RANGES OF JASPER NATIONAL PARK by Egbert Wheeler Pfeiffer, B. A., (Cornell University) A Thesis Submitted in Partial Fulfilment of the Requirements for the Degree of Master of Arts in the Department of Zoology. University of British Columbia 1948 SOME FACTORS AFFECTING THE WINTER RANGES OF JASPER NATIONAL PARK ABSTRACT OF THESIS SOME FACTORS AFFECTING THE WINTER GAME RANGES OF JASPER NATIONAL PARK by E. w. Pfeiffer Recent studies by Dr. I. MoT. Cowan in Jasper National Park have revealed that many of the winter game ranges of the Park are heavily overutilized by grazing and browsihg animals. In order to correct this situation, i t was f i r s t necessary to obtain accurate information on existing range conditions. Therefore, study of certain key areas of the Athabaska Valley was undertaken i n the summer of 1946 and 1947 by Dr, Ccwan and the author, for the purpose of describing existing plant growth, investigating factors affecting this plant growth, and recommending measures for improving range conditions. In carrying out the project;} standard range survey methods were used on nine representative areas. Point sampling revealed the composition of the ground vegetation while cover estimates and numerical counts gave the status of trees and shrubs. Analyses of s o i l samples indicated the f e r t i l i t y of the s o i l of the various ranges. The relative u t i l i z a t i o n of the ranges by herbivores was estimated by defecation counts. In order to determine the productivity of the ranges equal amounts of ground vegetation were clipped, air-dried, and weighed. Exclo8ure plots, established in 1942, served as a type of control as productivity within the plots was compared to that outside. The Jasper cemetery was used as a r e l i c t area. As a result of the investigations i t was concluded that the winter ranges of the Athabaska Valley vary greatly i n their capacity to produce vegetation, and that a l l the ranges are overutilized to a greater or lesser degree, principally by elk and horses. This over-u t i l i z a t i o n i s producing a retrogressive succession of the ground vegetation which is rendering the ranges increasingly unproductive. Destruction of browse species and s t r i c t f i r e protection are hastening development of climax spruce forests on certain range areas. In order to correct present trends i n the ecology of the Athabaska Valley i t was recommended that the numbers of elk be greatly reduced, that annual censuses of game populations be made, that a cover map of the area be prepared and new exclosure plots set up, that periodic range studies be made, that horse grazing be prohibited, and that conif-ers be controlled on certain game ranges. TABLE OF CONTENTS PAGE Acknowledgements I. Introduction 1 II. History of the Area 3 III. Methods used 8 IV. Description of the ranges 12 1. Range No. 1 13 2. Range No. 2 14 3 . Range No. 3 15 4 . Range No. 3A 16 5 . Range No. 4 17 6 . Range No. 5 18 7. Range No. 6 20 8. Range No. 7 2 0 9. Range No. 8 22 10. Cemetery 23 11. Comparative animal u t i l i z a t i o n of the Ranges 23 12. Productivity of the Ranges 24 V. Analysis of operative factors affecting the Ranges 1. Topographic 2 0 2. Edaphic 2 8 3 . Climatic 2 8 4 . Biotic 33 a. Faunal 36 b. Phytobioligical 44 TABLE OF CONTENTS (cont'd) PAGE VI. Conclusions and Recommendations 53 Plates Literature Cited Appendix A Appendix B ACKNOWLEDGEMENTS I wish to express my gratitude to Dr. I. McT. Cowan of the Department of Zoology, University of Brit i s h Columbia, for his suggestion of the subject of this thesis, for his aid i n the gathering of material, and for his invaluable advice i n i t s preparation. I am also greatly indebted to him for the photographs included i n the thesis. To Dr. W. A. Clemens, head of the Department of Zoology, University of British Columbia, I extend my thanks for his aid i n preparation of the work. I wish to thank Dr. V. C. Brink of the Faculty of Agriculture, University of British Columbia and Dr. T.M.C. Taylor of the Department of Biology and Botany, University of British Columbia, for much time spent by them on the identification of many of the plants l i s t e d and for their interest and help i n the work. To Mr. L. Farstad and Mr. J. T. Gillingham of the Faculty of Agriculture, University of British Columbia, I express my gratitude for aid i n the analysis and evaluation of the s o i l samples. 1. Introduction. Recent studies by Dr. I. McT. Cowan i n the Athabaska Valley of Jasper National Park have revealed that many of the winter game ranges of the Park are i n a deplorable condition from the standpoint of food productivity. The situation became so acute that mass slaughter of elk was begun i n 1942 i n order to attempt to relieve pressure on the winter ranges. It became apparent that a long term policy of range manage-ment was needed, but, i n order to i n i t i a t e such a policy, accurate information on the current situation was necessary. Therefore, a study of certain key areas of the Athabaska Valley was undertaken i n the summer of 1946 by Dr. Cowan and the author with the following ends i n vievf: A. Description of the existing plant growth i n order to have a standard by which to measure any changes i n the future. B. Investigation of the various factors affecting this plant growth with a view towards ascertaining which factors are i n any degree controiable. C. Recommendations as to the procedure to be adopted for improving the range conditions. The following material i s the result of studies made during the summer of 1946 supplemented with f i e l d work during two weeks of the winter of 1946-47 and three weeks i n August and September 1947. The areas chosen for study are samples of some of the principal winter ranges of the ungulates of Jasper Park. The study was confined - 2 -to winter ranges because they are the limiting factor as regards the carrying capacity of the area under consideration. This i s due to i t s climate and topography. On the higher ranges of the Park snow accumul-ates to great depths making food unavailable to the elk, sheep and deer. On the floor of the Athabaska Valley i n the Jasper area, however, snow seldom reaches very great depths (see Table I) and any food plants that may be present are readily accessible. The animals, therefore, descend i n the f a l l from the summer ranges to those areas where there i s limited snow and a good supply of food. In addition to these two conditions bighorn sheep ranges are generally characterized by steep slopes. Such terrain offers escape from predator attacks and is subject to maximum wind action, which prevents snow from burying the grasses. Since the summer ranges which are v i r t u a l l y limitless, are un-available to the ungulate herds during many months of the year, i t i s apparent that the number and condition of the herds i s dependent upon the area and productivity of the winter ranges. The carrying capacity of the winter ranges i s , therefore, the carrying capacity of the area. It i s hoped that the following study w i l l aid the appraisal of the carrying capacity of the Athabaska valley and indicate methods of i n -creasing i t . - 3 - • I I . HISTORY OF THE AREA A survey of the history of the Rocky Mountains of Canada indic-ates that more human activity of a commercial nature has taken place through the Athabaska Valley and Yellowhead Pass than anywhere else i n these mountains. This activity has had a considerable effect on the ecology of the area under examination and, therefore, i t i s pertinent to the present study to examine the extent and nature of the major his t -o r i c a l events. In 1811, Athabaska Pass was discovered by David Thompson and soon became a new route to the trading posts of the Columbia River. Follow-ing this discovery, twice a year for many years, brigades with goods and passengers moved through the pass between the Pacific and Hudson's Bay. S i r James Hector (16) writes that the Athabaska was ascended i n boats to Jasper House where pack horses were then used for the t r i p over the pass. The discovery of the Yellowhead Pass route i n 1826 resulted i n s t i l l greater activity i n the Athabaska valley. The Hudson's Bay Company played a leading role i n the development of this area, as the writings of H. J. Moberly (24), long an employee of the Company, indicate. In the early days Jasper House (see Map Figure l ) served as headquarters for operations i n this area and as a trading post. There were many good t r a i l s throughout the surrounding country as whole camps travelled them annually i n search of meat. The procedure was to remain i n a good loc a l i t y where feed for horses was plentiful, u n t i l the area was hunted - 4 -out. Despite a l l this hunting, there was never a shortage of meat while Moberly was at Jasper House. Moberly describes one such t r i p up the Smoky River on which he led a party of nearly one hundred people with some one hundred and f i f t y horses. He states that they k i l l e d over seventy moose, besides many bighorn, caribou, and mountain goat. In addition to hunting parties organized by white traders, Indians also operated extensively i n this area. According to Moberly, "These Indians a l l had bands of horses of their own." He estimates that i n 1855 some 350 mares were kept i n the Athabaska Valley as far as the confluence of the Athabaska and Miette Rivers. In the late f i f t i e s , a gold rush to the Cariboo placed Jasper on the t r a i l of the rush with hundreds of people and their animals travers-ing the valley. The construction of two railways, the Canadian Northern (National) and the Grand Trunk Pacific, i n the f i r s t and second decades of the 1900's brought further great .activity to the valley and practical extermination to i t s large game. The human activity was gradually stabilized, however, with the formation of a National Park i n the area in 1907, followed by protection of almost a l l wild l i f e . The last historical event of significance to this study occurred i n 1920 when 96 elk of both sexes were liberated at Jasper. In considering the above his t o r i c a l events, i t i s important to realise that great numhers of horses were involved i n a l l the successive human undertakings. As is shown below, they were supported on the open - 5 -ranges of the valley along with dwindling numbers of game. On the basis of these facts, the conclusion is reached that for a period of over one hundred years, the Athabaska Valley was subjected to an abnormal amount of grazing. From the accounts of early travelers through the Athabaska Valley i t i s apparent that this area, i n i t s natural state, was capable of supporting large numbers of ungulates. Describing his t r i p across the mountains to the Columbia i n 1811, David Thompson writes (33): "We are now entering the defiles of the Rocky Mountains by the Athabaska river, the woods of Pine are stunted, f u l l of branches to the ground, and the Aspin and Willow, etc. not much better ....We came to the last grass (about Buffalo Prairie) for the horses i n marshes and ponds where a herd of bison had lately been feeding, and here we l e f t the horses, poor and tired, and notwithstanding bitter cold, (they) lived through the winter, yet they have only a clothing of close hair, short and without any fur." David Douglas (13), in 1827, found bighorn sheep "in abundance" i n the Jasper area, and also writes of willows six to ten feet high on the river banks i n company with poplars and pines. The area appears to have remained highly productive i n the follow-ing years. W. Moberly (21), who spent the winter of 1872-3 i n the Atha-baska Valley, 27 miles east of Yellowhead Pass, says i n his report to Mr. L. Fleming, "The animals (horses) were turned out (worn out and nearly starving) about Jan. 20th to shift for themselves, as we had no fodder for them. Not a single one of them died, and they were a l l i n f a i r - 6 -condition when they resumed work the following March." The general productivity of the s o i l of the Athabaska Valley was indicated i n 1900 i n a geological survey report by James McEvoy (22). He cites Mr. L. Swift who demonstrated on a piece of land two and a half miles below the Maligne River's confluence with the Athabaska, that the country was capable of producing wheat, potatoes, and various other kinds of vegetables. McEvoy further mentions that poplar and cottonwood were generally distributed throughout the valley. Specific references to the game that has been supported i n the area are numerous. Successful hunting, as described by Moberly, has already been cited, and i t i s obvious that game was abundant during these early days of human exploitation. In the above report, McEvoy outlines the status of certain species as follows: ••Elk sparingly found i n the f o o t h i l l s of the mountains, moose and deer throughout rather scarce, mountain sheep scarce i n the f i r s t ranges of the mountains." He thus gives us a summary of the status of game after seventy-five years of human exploitation. E. A. Preble (26), reporting on game conditions i n the area i n 1896 suggested that a few elk might s t i l l exist near the head of the Brazeau River. Deer were present but scarce i n the Henry House and Jasper House areas, and mountain sheep were common as far west as Henry House. It i s quite apparent that by the turn of the century changes i n the relationships of game and range conditions had altered the ecological picture. A Department of Interior publication (14) states: "During construction of the railways, a great portion of the big game was either destroyed or driven away from the main valley of the Athabaska, where the open flats and grassy meadows had afforded excellent ranges for deer." This publication also gives a brief picture of the tree growth of the valley i n 1917. It mentions the jack pine (Pinus contorta) as the commonest tree, attributing this to forest f i r e s . Poplar, usually of small size, cottonwood, willow and alder were also prevalent, with much spruce i n the swamps. In discussing elk, the publication states that no elk were present in the Park i n 1917* However, i t asserts that the Athabaska Valley was said to have been noted i n the early days for excellent elk hunting. In 1914 the Park superintendent found remains of over one hundred elk near Mount Kerkeslin. However, no other records vindicate the assump-tion that the Athabaska Valley was ever well stocked with elk, and i t i s presumed that the above-mentioned remains were improperly identified. From the foregoing information, i t i s evident that the area under consideration was capable of supporting, i n i t s original state, large numbers of ungulates; with the coming of the white man, with large numb-ers of horses, much of the game was driven off or k i l l e d . - 8 -IU. METHODS USED 1. Point Samples; The point sample, as developed by Clarke (6), was the method by which the composition and density of the ground vegetation on each range was determined. Briefly, this method may be described as follows: points are taken at intervals on transects of the range, l a i d out to cross the maximum variety of ground vegetation. The points are taken with a point sampler, a frame holding a l i n e of ten long pins arranged so as to move up and down i n conformity with the surface of the ground. Each time the frame i s set down hits are counted. A plant i s hit when the point of a pin touches i t s base. Hits for each species are recorded and are expressed i n terms of percentages of t o t a l number of points for each area, With one exception, the tot a l number i n this study is 1000, This i s adequate, according to Clarke (op. c i t . ) , to obtain a truly representative picture of the vegetative composition and density. Due to the small area, only 300 points were counted i n the cemetery. 2. Aldous Browse Survey: The purpose of this survey was to obtain by a standard method an approximate indication of the amount of available browse on each range that has a sufficient tree and shrub growth to be significant. The method consists of surveying milacre plots selected at regular intervals on transects of the range as described by Aldous ( l ) . In the survey of each plot, the species observed are recorded as being Dominant, Moderate, or Sparse. I f the species covers 50-100 per cent of the plot, i t i s dominant, i f 10-50 per cent, moderate, and i f 0-10 per cent, sparse. The average values are calculated as 70 per cent for dominant, 30 per cent for moderate, and 5 per cent for sparse. The average densities for each class are then added and- the sums divided by the number of plots surveyed to give an average density for each species on the various ranges. To facilitate comparison of the relative abundance of the species of trees and shrubs on different areas, the number of times each species occurred on each area is added and the sum divided by the total number of plots surveyed. As the number of plots surveyed was in each instance 1 0 0 , the resulting figure, expresses frequence of occurrence as a percentage. 3 . Numerical Survey: Each milacre plot surveyed by the Aldous method was also sub-jected to a direct count of individual trees and shrubs. Species were classified and counted, according to size, so that a picture of the tree and shrub reproduction could be obtained. 4 . C l i p p i n g ; In order to obtain an estimate of the productivity of the ranges, standard areas of ground vegetation were clipped, air-dried, and weighed during the summer of 1946. The unit of measure for clipping was one square yard, and in most cases a total of ten square yards was clipped from each range area. The locations of the clip plots were picked more or less at random, although the poorest areas on each range were avoided. Therefore, i t is reasonably accurate to state that the follow-average ing productivity figures represent an above/production of the ranges at the present time. 5. Defecation Countst In order to establish a basis for relative u t i l i z a t i o n of the ranges by different species, counts were made of the various ungulate or equine defecations encountered on a 1000 pace diagonal transect of each range. It must be emphasized that these are not individual pellet counts, but numbers of defecations. No attempt has been made to use these figures to show absolute numbers of animals as the adjustment involves too many variables and unknowns to be of value. They are i n -cluded merely to demonstrate the approximate relative usage of each range by the species involved. 6 . Exclosure plots Several exclosure plots, constructed for the most part i n 1942, i n various areas of the Park, were clipped i n the summer of 1946 and 1947 to demonstrate the difference i n production on grazed and ungrazed areas. A l l the exclosure plots are located i n positions representative of the prevailing range conditions and the difference i n productivity i n and outside the plots was i n no case due to the advantageous positions of the plots. They are, however, about 6 f t , by 6 f t . i n size, which is too small for optimum study. The Jasper town cemetery, about two miles east of Jasper, served as one exclosure plot, as i t has been surrounded by a game-proof fence for at least twenty years. In 1947 the amount clipped from each square yard was weighed and the mean determined from the sum of the clippings. In 1946 the t o t a l amount from each area was weighed and divided by the number of square yards clipped. In order to obtain an indication of the effect of grazing upon grass reproduction, counts of seed heads of the Graraineae were made within the exclosure plots and i n equal areas immediately adjacent to the exclosures. As this count was made i n late August 1947 a l l grasses had matured. 7. Relict Area In addition to serving as an exclosure plot the cemetery i s large enough ahd has been fenced i n for a sufficiently long time to serve as a r e l i c t area. It was point sampled to obtain an indication of the ground vegetation that i s developed without the grazing factor. 8. S o i l Analyses During August of 1947 samples were collected from typical locafc— tions on the nine areas. One profile per range was exposed to a depth of from 2 | to 3 f t . and samples taken from the various horizons thus exposed. The horizon designated C or D was generally at least 2 f t . beneath the surface. These samples were then analyzed as follows: 1 . Texture was determined by manual means by an expert agron-omist. 2 . pH was determined by the electrometer test. 3 . Available potash, phosphorus and nitrates were determined by the Spurway method ( 3 0 ) , as these are the elements of major importance from the agricultural point of view. - 12 -4. Percentage of organic matter wqs determined by the wet combust-ion method as described by Schollenberger (28). IV. Description of the Ranges The Athabasfci valley from the town of Jasper east to the eastern bound-ary i s the general area under consideration. According to Cowan's report for 1943 (8)> this area provides suitable winter ranges for deer, sheep, and elk (also domestic horses) and may be considered typical. The valley l i e s between highly folded mountains which consist largely of rocks that were l a i d down originally aa sand, mud and calcar-eous ooze. They are, therefore, sedimentary and date back to the Paleo-zooic and Masozooic eras (16). The soils of the valley are strongly alkaline i n reaction and medium to light i n texture. They are transported soils derived from aeolian, a l l u v i a l , or glacial deposition, of high lime content and with varying amounts of organic matter. As they are young the nature of the parent material has a pronounced effect on the s o i l s , which, because of the dominance of carbonates, show many of the character-i s t i c s of the Rendzina s o i l group. Table II gives certain qualities as exhibited by the s o i l samples from the individual ranges. These qualities are discussed i n detail i n the section dealing with the analysis of the edaphic factors• Despite the northern latitude and the considerable altitude (approximately 3300 f t . ) this area enjoys a climate that is so tempered by the "chinook winds" that i t compares favorably with more easterly regions lying much fafcther to the south (26). On the basis of annual precipitation the climate may be classified as semi-arid (20jl Table I gives pertinent data on weather conditions in Jasper Park, For the intensive study nine local range areas were chosen on a basis of winter utilization as observed during earlier field work. The nine areas selected are as heavily utilized as any in the Park, and are, therefore, excellent indicators of the general status of the winter ranges in the Park. They have been assigned numbers beginning with number 1. at the eastern boundary of the Park and progressing westward td fiumber .8 about six miles east of Jasper. (Fig. 1.) 1. Range No. 1. Miette Slopes This area comprises the open,grassy, southern slopes of the steep hills lying in the valley of Moosehorn Creek at its confluence with the Athabaska river. It is bounded to the south by the flood plain constituting Range No. 2, to the east by Moosehorn Creek, to the north by the heavy tree and shrub growth that occurs on the crest of the ridge and extends down each northern exposure, and to the west by Roche Ronde Mountain range. The hills rise to elevations of 600 feet above the valley and are characterized by a gently undulating surface (Fig. 3)« The soil is the most fertile of any area studied (Table II), On this range area very little tree and shrub growth has occurred, except in scattered locations where the local undulations have favored limited growth. For the purpose of this study, the tree and shrub growth was of too little significance to be surveyed. TABLE I Weather Data (average for 22 years) for Jasper Park daily Mean maximum daily Mean minimum Precipitation Precipitation Total Rain Snow Precipitat-Temperature Temperature ion. Jan. 20 4 0.05 inches 10.5 inches 1.10 Feb. 29 8 0.08 6.7 0.75 Mar, 37 17 0.07 5.3 0.60 Apiil 50 26 0.48 2.1 0.69 May 61 34 0.86 1.9 1.05 June 69 41 1.45 trace 1.45 July 74 45 1.73 0.0 1.73 Aug, 71 44 1.69 trace 1,69 Sept. 59 37 1.32 6.3' 1.35 Oct. 48 30 0.88 1.8 1.06 Nov. 30 19 0.2© 4.8 0.71 Dec. 16 6 0.19 6.9 0.88 Total annual 9.03 40.3 33.06 Mean Precipitatidfi per Week May-September 1946 .10 inches .44 inches Jasper Townsite East Gate May-August 1947 Jasper Townsite East Gate .66 inches 1.20 inches Temperatures at Jasper, winter of 1942-43 Mean Maximum Mean Minimum Nov. 28.4 degrees F. 11.0 degrees F Dec. 23.2 degrees F 5.0 degrees F TABLE I cont'd. Jan. 11.2 degrees F. 7 .7 degrees F Feb. 3 6 . 4 degrees F. 15 .1 degrees F Mar. 3 3 . 1 degrees F. 10 .6 degrees F Apr. 5 5 . 7 degrees F. 27 .9 degrees F TABLE I I SOIL QUALITIES Range Horizon *pH Effervescence Nitrates Ph K Organic matter Texture Parts per million 1 A 8.1 violent 10-15 .5 low 7.52f* very fine sandy loam to s i l t B 8.2 violent 10 0 very low s i l t loam loam C 8.4 violent heavy s i l t loam 2 A 8.3 violent 5 0 very low 2.65 s i l t loam B 8.6 violent 10 0 very low fine sandy loam D 8.6 violent sandy loam (gravelly) 3 A 8.2 violent 2 0 very low 4.28 fine sandy loam B 8.2 violent 0 0 very low loamy sand D 8.4 violent s i l t loam A 8.2 violent 5 0 very low 0.80 very fine sandy loam to s i l t loam 4 A 8o4 violent 5-10 1.0 medium-5 5.01 sandy loam B 8.9 violent 2 .5 very low clay loam C 9.1 violent clay loam mixed with gravel 5 A 8.0 . violent 10 0 low 9.40 loam B 8.2< violent 25 0 medium-5 clay C 8.0 violent heavy s i l t loam to s i l t y clay loam S A 8.3 violent 5 0 very low 1.91 fine sandy loam B 8.2 violent 25 0 very low fine sandy loam C 8.1 violent fine sandy loam 7 A 8.2 violent 2 0 very low 6.35 Coar.se sandy loam B 8.6 violent 0 0 very low loamy sand (some gravel) C 8.9 violent sand 8 A 8.2 violent 10 .5 low 6.40 loam B 8.4 violent 5 .5 very low loam D 8,7 violent gravel * The soils were air-dried, which reduces the pH i n alkaline s o i l s . ** This s o i l exhibited only one horizon to a depth of at least 3 f t . - 14 -However, as demonstrated by the surcey on June 1 0 , 1946, the results of which are given in Table III, the ground vegetation exhibits a luxuriant growth. Wheatgrasses comprise a large percentage of the growth (10 .1 #), and the presence of a variety of species of succulent forbs is further evidence of a healthy plant community. Over-use in-dicators, such as pasture sage, show a very low percentage « f pecurrenfcd. 2 . Range No. 2 : Miette Flats This area includes the alluvial flats from which rise the slopes of Range No. 1 (Fig. 3 ) » To the south, the area is bounded by the Athabaska river, while to the east Moosehorn Creek forms the bound-ary. An extensive and heavy stand of spruce arising from swampy terrain limits the range area to the west. The subsoil is too coarse in texture for good moisture retention.(Table II.) The plant growth of this area is in the most healthy condition of any studied (Fig. 4 ) « It is a sub-climax association of widely scatter-ed young spruce and deciduous trees, interspersed with heavy stands of shrubby plants, mainly silverberry, buffaloberry, and willow. Table IV. gives the composition of the tree and shrub growth as sampled on June 1 2 , 1946. There are large numbers of young aspen and poplar, indicating that successful reproduction is taking place. The buffaloberry and silverberry of this area have attained normal heights (Figs. 5 and 6) and willow is in a healthy condition (Fig. 7 ) . In view of these facts, the browse species of this range are considered to be in a satisfactory condition, and the data on them a r e used as a standard of proper use in TABLE III Bare ground 73 .7$ Grasses Thickspike wheatgrass 10.1 Wild ryegrass 2.7 Brome grass No. 1 0 . 3 Purple reedgrass 0 .3 13 .4 Shrubs Bearberry 6 . 2 Field sage 1 .0 Pasture sage 0 .5 '•• Rose 0 . 2 7 .9 Forbs Daisy 0 . 8 Pussytoes 0 . 7 Borage 0 .5 Violet 0 .5 Northern Bedstraw 0 .5 Goldenrod No. 2 0 . 4 Flax • 0 . 4 Strawberry /•' 0*3 Sedge °»3 Blue-eyed grass 0*2 Black-eyed Susan 0 » 2 Vetch 0«1 TABLE I V SPECIES HEIGHT NUMBERS DENSITY PERCENTAGE Trees Aspen -4 f t . 700 .95 18 Aspen 8fft. — — 2 Poplar 8+ft. ~ — — White Spruce --8 f t . 190 2.75 8 White Spruce 8fft. 110 — 8 Shrubs Buffaloberry — 360 5.80 20 Silverberry 1+ft 1740 3.55 9 Dogwood — 480 .40 — Rose — 1800 1.95 23 Shrubby Cinquefoil — 20 .10 2 High Bush Cranberry — 20 . 05 Willow — 480 2.55 13 OMISSION-TABLE IV Species Height Numbers Density Percentage Poplar -4 f t . 40 . 1 5 3 - 15 -discussions of the other areas. However, the existing relationships are far from st a t i c . The table above shows a large number of small spruce i n this area. This indicates a situation which appears quite obvious on inspection of the range (Fig. 8). The spruce to the west i s rapidly invading the area under study and within a few years w i l l transform the entire area into a climax spruce forest, thus establish-ing a hiome that would not include any of the present ungulate forms. The ground vegetation of this area is the heaviest of any area studied. Table v" gives the composition of the ground vegetation on August 20, 1946;-The outstanding feature of the above survey is the prevalence of pussytoes (28.7$). In many small areas, i t carpets the ground to the exclusion of other plant species. This i s evident i n the small percentage of forbs. The tree and shrub growth, as well as the sandy type of s o i l , undoubtedly accounts i n part for this condition. However, of pressure/animals grazing on the grasses and forbs i s probably a factor i n the existing prevalence of pussytoes. This area has long been the favorite winter range of a group of horses. 3. Range No. 3t Devona Flats This area is similar topographically and edaphically to Range No. 2. It is an a l l u v i a l f l a t formed as a result of the activity of the Snake Indian river at i t s confluence with the Athabaska river. The area is shaped like a triangle with the Athabaska river on the south as i t s base. The northeast side is bounded by the Snake Indian river, while the northwest side i s defined by the abruptly r i s i n g f o o t h i l l s Bare ground Grasses Fescue June grass Wild Rye Thickspike wheatgrass Shrubs Pasture sage Forbs Pussytoes Butterweed Goldenrod No. 1 Bluebell Sedge 1 - 16 -of the Roche de Smet mountain range. Local areas of swamp support heavy stands of spruce in the northwest portion of the area. The general character of plant growth resembles that of Range No.-2. However, trees and shrubs are much less evident and occur only in widely scattered clumps and in limited numbers. Silverberry and buffalo-berry are the most prevalent, but do not attain as luxuriant•growth as is found on Range No. 2. Spruce is not a factor in the plant community of this area. In view of the small numbers and the sparse distribution of trees and shrubs on this range, no survey of these was made. As this range supports a considerable stand of ground vegetation, it is of primary value as a grazing area. A survey made on August 22, 1946, of the ground vegetation on this range gave results as shown in Table VI. The percentage of grasses (9.0) on this area is reduced as com-pared to the two previous areas. Pussytoes is limited in extent, probab-ly by very local ecological conditions, but its place as the dominant plant on the range is taken by pasture sage (9.4$)> a sub-climax species which is a well recognized indicator of over-grazing. 4. Range No. 3a. Windy Pointt This area consists of steep slopes rising to over 400-foot elevations directly from the north bank of the Athabaska river at Jasper lake. It exhibits characteristics common to a large part of the most heavily utilized sheep ranges in the Athabaska valley (Fig. 9). The contours aire steep but not precipitous, and the fine aeolian soil TABLE VI. Bare ground 73.1$ Grasses and sedges June grass 5.4 Sedge 2.0 Sandburg bluegrass 1.9 Thickspike wheatgrass 1.5 Bluegrass »2 11.0 Shrubs Pasture sage 9.4 Bearberry 10.0 Forbs Pussytoes 2 ,5 White daisy 1.7 Flax I * 1 Milk vetch 1»° Clinquefoil sJL 6.8 - 17 -supports considerable ground vegetation. Tree and shrub growth is defin>-itely limited to hollows and to the less exposed northeast slopes of the small ridges, as was described for Range No. 1. Spruce, in limited numbers, is practically the only species of tree or shrub, and therefore no survey of this category was made. Due to the exposed location of this area, it is subjected to a maximum amount of wind and sun. This has produced the most arid section of the Park, and dust storms are a common occurrence throughout the year. Terracing is very extensive on a l l steep slopes, as a result of the nature of the soil and the large numbers of hoofed animals that utilize this type of range. Such conditions are not conducive to luxuriant plant growth. This is evident in the results of the survey made on June 19, 1946 which is given in Table VII. There is a very large percentage of bare ground and a greatly reduced amount of grass(2.8$) and forbs. The extreme prevalence of „ pasture sage and Russian thistle< (11.7%) suggests that the low percentage of grass and forbs is not solely the result of factors des-cribed above, but is also the result of overuse by grazing animals. Retrogressive succession has proceeded further on this area than on ranges previously discussed. 5. Range No. 4: Greenock This area is similar In general topography to Range No. 3 a . It consists of the steep south-western slopes of the benches at the foot of Mount Greenock. These benches rise 200 to 400 feet from the more gently sloping valley and are composed of coarse, morainal gravel under-T A B L E VII Bare ground 80.9$ Grasses Junegrass 1.5 Purple reedgrass .6 Thickspike wheatgrass .4 Needlegrass .3 2.8 Shrubs Pasture sage 10,4 Juniper 1.5 Russian this t l e 1.3 Bearberry .5 Forbs Aster 1.3 Flax • ^  Unidentified »5 2,6 - 18 -lying a shallow, loamy, highly alkaline surface soil as distinguished from the sandy, aeolian soil of Range No. 3a. Tree and shrub growth is almost non-existent on these slopes, as there are few protected locations. The special edaphic and topographic factors on this range have produced a ground vegetation quite different from that encountered on any other range. The survey made on August 14, 1946, shows this to be the case. (Table VHX) There is a veryolow percentage of grasses (1.4%) and forbs, which is due in part to topographic and edaphic factors which.favor the growth of bearberry. This species carpets large areas to the exclusion of a l l other plants. However, bearberry grows in many rocky parts of this range that would support very little else, and would probably be a dominant plant under any normal conditions on this type of range. 6. Range No. 5: Moberly Flats This area is a plain sloping gently from the foot of abruptly rising mountains on the west, to the Athabaska river on the east. Heavy timber growth limits the range on the north and south. On the lower flats the soil is a heavy alluvial type, while higher up i t is mixed with a considerable amount of gravel and sand. The comparatively heavy vegetation testifies to the favorable quality (high moisture retention capacity|of the soil of this area. Its potential plant growth can be compared to Range No. 2, although its composition is of a somewhat different character. It is an open, aspen parkland association supporting a large population, of shrub species. The dominant shrub is silverberry, with TABLE VIII Bare ground 76.6$ Grasses and sedges Thickspike wheatgrass 1.1 Sedge 1.1 Junegrass .3 2.5 Shrubs Bearberry - 18.4 Juniper 1.2 Pasture sage .1 2Q.7 Forbs Northern bedstraw .7 Legume •2 Alpine aster .2 Daisy »I 1.2 - 19 -service-berry and buffaloberry common throughout. Willow occurs sparsely scattered, but numbers of dead individuals indicate its former abundance. Table IX gives the composition of the trees and shrubs on August 7, 1946i~ Although the above figures demonstrate a free and shrub flora, they also show that browse species are being depleted. A comparison of the data for silverberry and buffaloberry on this range with those for Range No. 2 indicates the extent of this depletion. The numerical counts on Range No. 5 exceed those for Range No. 2, but the density figures are in inverse relation. This results from the short height of the majority of plants in Range No. 5, and it is accompanied by a correspondingly decreased amount of browse. The limited stature of silverberry and buffaloberry on Range No. 5, is almost entirely due to the browsing down of mature plants. There are no young plants. > The lack of aspen reproduction is shown by a comparison of figures for this range and Range No. 2. On the latter area there is an average of 20 aspen trees per acre over 8 feet (mature trees) and 700 aspen trees per acre under 4 feet (young trees). However, on Range No. 5, there are 330 trees per acre over 8 feet and only 120 under 4 feet. Prac-tically a l l these are yearlings and will be eliminated during the succeed-ing months. The ground vegetation of this area is comparatively dense and varied. The survey on August 6, 1946, gave the results, shown in Table X. As junegrass is the dominant species (17.3$), and as there is a large percentage of pussytoes (3.5%) and a scarcity of wheatgrasses (0.6%), i t is evident that overgrazing is an important factor on this area. TABLE IX SPECIES HEIGHT NUMBERS DENSITY PERCENTAGE OCCURRENCE Trees Spruce - 8 f t . 10 .05 1 Spruce 8+ft. 50 — 5 Aspen -4 f t . 120 .45 9 Shrubs Rose — 1910 2.55 46 Serviceberry — 1460 .60 13 Silverberry +1 f t . 1760 1.35 . 27 Silverberry 1+ft 320 1.25 12 Willow ' — 50 .20 4 Buffaloberry 1+ft. 110 2.65 11 Shrubby Cinquefoil — 120 .30 6 Common Juniper 10 .30 1 OMISSION-TABLE IX Species Height Numbers Density Percentage Aspen &> f t . 330 - 22 Buffaloberry -1 f t . 320 1.45 19 Bare ground Grasses and sedges Junegrass Sedge Thickspike wheatgrass Bearded wheatgrass Shrubs Bearberry Rose Forbs Pussytoes Purple Daisy Ginquefoil Puccoon Flax Bastard toad-flax Goldenrod No. 1 - 20 ~ *7. Range No. 6: Talbot Lake This area is almost identical to Range No. 3a, but l i e s on the south bank of the Athabaska river between Talbot and Edna lakes. It is a steeply sloping h i l l rising from the Athabaska river on the north, and extending to the precipitous slopes of Mount Ginquefoil on the South. Its exposed position has resulted i n a high degree of dessication from wind and sun. Consequently, the tree and shrub growth i s very limited and i s of l i t t l e significance. In certain areas on the crest of the h i l l , almost a l l plant l i f e has gone, and the fine-textured aeolian s o i l has been deeply eroded by wind. (Fig. 10). Such conditions have reduced the percentage of ground vegetation as shovm in the survey made on August 12, 1946. (Table XI) As on Range Area No. 3a, pasture sage i s the dominant plant, (5.0%), and grasses (8.9%) are limited i n occurrence. The general con-dition of this range indicates that i t i s i n a similar stage succession-a l l y to Range Area No. 3a, although i t i s associated with a completely different mountain system. 8. Range Area No. 7: Henry House Flats This i s one of the largest level areas i n Jasper Park (Figs. 11 and 12), a portion of i t being used as an a i r - f i e l d . It i s , generally, several feet above the river and is therefore well drained. The s o i l i s a very coarse, sandy, l i g h t , a l l u v i a l type, several feet deep through-out the area. The Athabaska river forms the eastern boundary, while the western and southern limit i s the rocky slope of the Palisade ridge. To the north, the area is bounded by a shallow arm of the Athabaska river, TABLE XI Bare ground 82.9$ Grasses and sedges Thickspike wheatgrass 3.5 Junegrass 3*1 Nfe'edlegrass 2.3 Sedge 1*1 10.0 Shrubs Pasture sage 5*0 Forbs Daisy .7 Butterweed «6 Flax .1 Unidentified *5 1.9 - 21 -and an extensive black spruce swamp and marsh. Henry House f l a t s , as the area i s loc a l l y known, i s typical of much of the Athabaska valley from Henry House to Jasper. It i s a lodgepole pine sub-climax assoc-iation with large areas, such as the landing f i e l d , without any tree or shrub growth. Beneath the pine, which grows i n pure stands, there i s a large population of browse shrubs. A survey of the trees and shrubs on June 17, 1946, gave the results shown in Table XII. Buffaloberry i s the principal browse shrub despite the high figure for serviceberry, which i s generally too small a plant to be significant. The individual buffaloberry plants are stunted and show much dead material (Fig. 13). The c r i t i c a l condition of this species i n this area is apparent i f the data for buffaloberry on this range is compared with the data for buffaloberry on Range No. 2. There are 700 buffaloberry plants per acre on Range No. 7 to 360 on Range No. 2. • However, the density of the cover grown by these plants amounts to only 2.80 on Range No. 7 as against 5.80 for Range No. 2. As shown i n the numerical counts for Range No. 7, the number of buffaloberry plants under 1 foot far exceeds that of those over 1 foot. The short plants are not young ones, but mature plants browsed down to this height. The ground vegetation i s comparatively heavy, as this area i s , potentially, moderately productive. However, the large percentage of sedge, junegrass, and overgrazing indicators such as butterweed, and pasture sage indicate over-use by grazing animals. The survey on June 17, 1946 gave the composition of the ground vegetation as shown in Table XIII. TABLE XII SPECIES HEIGHT NO. per ACRE . DENSITY ' PERCENTAGE OCCURRENCE Trees Aspen -1 f t . 10 .05 1 Lodgepole pine - 8 f t . 190 1.05 13 Lodgepole pine 8+ft. 870 — 36 Spruce -8 f t . 30 .15 3 :ubs Juniper — 60 .80 5 Buffaloberry -1 f t . 460 1.80 30 Buffaloberry 1+ft. 240 1.00 13 Rose — . 670 .60 12 Serviceberry — 1660 .70 14 Silverberry. — 30 .05 1 TABLE XIII Bare ground 75*2% Grasses and sedges Sedge 6.5 Junegrass 4 . 7 Pinegrass 2 .2 13 .4 Shrubs Pasture sage 3 . 0 Forbs Butterweed 3 . 8 Pussytoes 3 . 1 Flax .4 Wind flower .1 Canada pussytoes . 1 Cinquefoil .1 7.6 - 22 -9. Range No, 8; Maligne This area is the former delta of the large stream that runs off Mount Roche Benhomme, through the range to the Athabaska river. The area is a f l a t , triangular outwash plain sloping gently toward the river, which forms the west boundary. The s o i l i s a shallow loam, except on the upper reaches, where rock and gravel are i n evidence, A dense stand of spruce occupies the western portion of the area. This range i s a characteristic aspen-poplar parkland association, very prevalent in the Athabaska valley. These species grow i n almost pure stands, although the conifers, spruce and Douglas f i r are beginning to establish themselves. There i s a heavy growth of shrub species, main-l y buffaloberry and silverberry. The survey of tree and shrub growth on June 13, 1946, gave the results: shown in Table XIV. The counts i n Table XIV reveal that aspen reproduction has ceased. There are no trees between the heights of four and eight feet and the majority of those i n the 4-foot class, or less, are seedlings of only a year's growth and due to be browsed off during the succeeding winter. The figures on adult plants indicate that there are ample numbers of parent trees for reproduction. The same situation exists with poplar, although to a lesser degree. With regard to the shrubs, a l l silverberry are 1 foot, or less, in height,-the majority of the plants being mature plants browsed down to that height. The situation i s the same for buffaloberry. The great loss i n foliage that this reflects i s demonstrated by comparison of the density of buffaloberry on this range with that on Range No. 2, On the TABLE XIV SPECIES HEIGHT NUMBERS DENSITY PERCENTAGE OCCURRENCE Trees Aspen -4 f t . 240 .65 13 Aspen 8ff t . 500 35 Douglas f i r — 60 1.40 2 Poplar -4 f t . 490 .35 7 Poplar 8+ft. 70 ~ 2 Spruce -8 f t . 70 2.75 -* Spruce 8tft. 20 — — Shrubs Buffaloberry -1 f t . 500 1.45 29 Willow 40 .55 4 Silverberry -1 f t . 250 . 5 0 10 Rose — 1280 1.40 29 Juniper — 60 .40 3 Gooseberry — 80 .70 6 Serviceberry — 1080 1.00 12 TABLE XV Bare ground 88.0$ Grasses and sedges Thickspike wheatgrass 2,4 Bearded wheatgrass 1.9 Junegrass 1.2 Bluegrass 1.1 Richardson's need!egrass 1.0 Sedge 1.0 Brome No. 2 .8 9.4 Shrubs Bearberry 1.8 Pasture sage »6 2.4 Forbs Strawberry »2 - 2 3 -latter range there were 360 plants per acre with an average density of 5.80, while on the range area under consideration there were 500 shrubs per acre with an average density of only 1 . 4 5 . The present status of willow on this range is characteristic of this species on a l l other ranges, except Range No. 2 . As a highly preferred browse species i t has been practically exterminated. This area exhibits the lowest percentage of ground vegetation (according to point samples) of a l l the ranges studied. (Table XV.) The low percentage of ground vegetation (12%) i s due i n part to the density of tree growth i n various sections. Such grasses as Richard-son's needlegrass are characteristic of shaded areas. The rocky and gravelled areas are unfavorable to most plant growth except bearberry, which i s prevalent under these conditions. However, much of the area i s capable of supporting a heavier stand of grasses and forbs than i s evident at present. 10. Cemetery This i s a very small area (less than an acre) which has been protected from grazing for at least twenty years. Its topography and s o i l are similar to that of Range No. 7 . The survey on August 28,1947 revealed heavy ground vegetation with a wide variety of midgrasses. ( Table XVI.) 1 1 . Comparative Animal Uti l i z a t i o n of the Ranges as demonstrated by Defecation Counts. An indication of the relative numbers of animals of various species u t i l i z i n g these ranges was obtained by the defecation counts, which are given i n Table XVII. Therefore, a range was dominated by the TABLE XVI Bare ground 65.3 Grasses and sedges Junegrass 7.4 Thickspike wheatgrass 5.3 Bearded wheatgrass 2.6 Purple needgrass 1.0 Bromegrass No. 2 1.0 Needlegrass 0.3 17.6 Shrubs Pasture sage 1.6 Ground cedar 0.3 1.9 Forbs Pussytoes 2.3 Purple vetch 2.0 Cinquefoil 1.6 Aven 1.3 Goldenrod No. 2 0.7 Elack-eyed Susan 0.3 8.2 Moss 7.0 TABLE XVII Range Numbers Elk Mule Deer Horse Bighorn sheep TOTALS Elk 1196 Comparative Animal Utilization of the Ranges as demonstrated by Defecation Counts, 1 2 3 3a 4 5 6 7 8 208 104 156 102 24 199 38 213 161 7 22 — — — 10 — — — 1 60 21 25 — 21 34 23 42 6 i — — 140 79 — 155 — 4 Mule Deer Horse Bighorn Sheep 39 207 439 - 24 -species having the highest defecation count for that area. The figures indicate a considerable competition for winter range between sheep and elk with the horse a competitor principally on the elk ranges, as the sheep ranges are often quite precipitous. In order to f a c i l i t a t e more accurate interpretation of the above figures, i t i s pointed out that sheep, being the smallest animal under consideration, defecate more frequently, with frequency of defecation by the elk and the horse following i n that order. It is important to note, also that, despite t h i s , the horse shows up as a most important element in the grazing to which the Athabaska Valley winter ranges are subjected. 12. Productivity of the Ranges From the foregoing descriptions of the ranges studied i t i s apparent that,several factors influence production as revealed i n Table XVIII. A. The increased production within the exclosure plots demonstrates that grazing is a very important factor determining product-i v i t y . B. The differences i n production within exclosure plots, e. g. 82 grams per sq. yd. within the cemetery and 134 grams per sq. yd. within the exclosure plot on Range 1, show that factors other than grazing influence productivity. These other factors are indicated as follows: 1. The difference i n productivity within exclosure plots on Range No. 1 and No. 2 (134 and 105 grams produced per sq. yd. respectively) appears to be dependent on a very local factor such as s o i l . TABLE XVIII Productivity of the Ranges Area clipped Total wt. Mean productivity to (sq. yds) i n grams nearest gr.per sq.yd 1946 1947 1946 1947 1946 1947 A. Range No. 1 10 725 73 A. Range No. 1 exclosure plot 4 3 535 571 134 190 B. Range No. 2 10 530 93 Range No. 2 exclosure plot 2 3 210 146 105 49 C. Range No. 3 7 240 34 Range No. 3 upper exclosure plot 3 4 250 255 83 64 Range No. 3 Lower exclosure Plot 4 4 380 157 95 39 D. Range No. 3A 10 360 36 E. Range No. 4 10 100 10 F. Range No. 5 10 330 33 G. Range No. 6 10 290 29 H. Range No. 7 10 160 16 I. Range No. 8 10 300 30 J. Cemetery-outside 5 H5 23 Cemetery-inside 5 5 .410 301 82 60.2 - 25 -2. The difference i n productivity within the exclosure on Range No. 1 and the cemetery (134 and 82 grams per sq. yd. respect-ively) may be due to very local factors, as i n lrabove, or to the higher r a i n f a l l i n the more easterly area, or to a combination of these factors. 3. A comparison of the difference i n productivity (1946) inside exclosure plots to that outside reveals that the differences become smaller according to the proximity of the ranges to the Eastern boundary of the Park. Thus, productivity figures for the cemetery show an approx-imate ratio of one outside to four inside, while progressing eastward to Range No. 3 there i s found a ratio of somewhat less than three inside to one outside, and on Range No. 1 and No. 2 the ratio i s about two inside to one outside. It w i l l be shown i n the discussion of biotic factors that elk have only recently arrived on the eastern ranges. Correlating this fact with the different ratios of productivity inside and outside the exclosure plots, i t becomes apparent t hat productivity of the ranges i s influenced by the length of time that grazing by elk has been carried on. 4. The differences in productivity within exclosure plots in 1946 and 1947 demonstrate the va r i a b i l i t y i n amount of ground veg-etation from year to year. This i s probably due to variations i n the amount of moisture available to plant growth. Correlating productivity with the various factors which have been measured on the range nearest Jasper (Range No. 7) and on the range far-thest from Jasper (Range No. l ) , i t i s clearly demonstrated that a v a i l -able moisture i s the basic factor determining productivity (see Fig. 15). - 2 6 -The figures i n Table XVIII should be considered i n the light of the vegetative compositions of these areas as demonstrated by the point samples. I t w i l l then be apparent that the high productivity of certain areas, as for instance Range No.3A with 36 grams per sq. yd., is prin-cipally due to large percentage of poor fodder species (10.4% pasture sage) while on other areas, as for instance Range No. 7 with 16 grams per sq. yd., with a large percentage of grass i n the vegetative composition (6.0$) and a small amount of pasture sage (3.0$), the productivity figures appear low. However, they represent a large amount of good fodder species. IV. ANALYSIS OF OPERATIVE FACTORS AFFECTING THE RANGES 1. Topographic As these ranges are very similar i n altitude, topography affects them through the influence of slope and exposure. Fig. 1 ind-icates that the ranges studied are either f l a t s or steeply rising h i l l s . The foregoing descriptions of the areas show that a l l the flats support trees or shrub growth, while the slopes are limited to ground vegetation. This major difference i n vegetative cover can be attributed to the i n -fluence of slope and exposure. The angle of slope largely determines the amount and type of s o i l accumulated. It is only on nearly level ground or gentle slopes that considerable depths of s o i l accumulate, and undergo characteristic dev-elopment of a mature s o i l . The h i l l s of Ranges No. 3A and No. 6 are immature, a l l u v i a l deposits that w i l l probably never become stabilized due to constant drif t i n g of dust from the Athabaska plain and blowouts. Range No. 4 is too steep to permit the development of a mature s o i l under any conditions, as the continuous removal of surface s o i l by - 27 -erosion keeps exposing the lower horizons and so modifies the profile that would otherwise be present. This is one of the causes of the low productivity of this area. The above areas are subject to excessive runoff during and after precipitation as runoff increases directly with slope. In semi-arid areas such as that under study this means a considerable loss of soil moisture. Furthermore, the slopes of Ranges No. 3A, No. 4, and No. 6 are so situated that most of the snow with its moisture bearing poss-* ibilities is blown away. The exposure of slopes to the sun is a factor of extreme import-ance in regulating soil moisture and. temperature, and, thereby, plant growth. Bates (2) made a study of the microclimate in a valley running east-west through a small section of the Rocky Mountains. He found that marked difference in vegetation of opposing slopes was due to differences in isolation. The south slope exhibited much higher temperatures, evap-oration, and much lower soil moisture than the northern exposure. These factors produced moisture values in the soil of the south slope below the wilting coefficient of the soil. The temperature was found to be high enough to k i l l certain tree seedlings. Much the same situation exists through the Athabaska valley, and a l l the hillside ranges (No.'s 1, 3A, 4, 5, & 6) studied exhibit almost completely treeless growth on the southwestern exposures, which in general constitute the ranges. The opposite slopes are usually heavily populated with spruce which is similar to European conditions as des-cribed by Braun-Blanquet (4). Range No. 1 is thought to be influenced to a small degree by seepage of water from higher up the Moosehorn - 28 -valley. This situation would account for fluctuations i n productive capacity that were independent of other nearby areas. 2. Edaphic The effect of s o i l upon the ranges may be studied according to i t s chemical and physical properties, which w i l l be treated separately. Because of i t s importance to this study, organic natter w i l l also be discussed separately. a. Chemical The samples indicate that the lime content of these soils i s sufficiently high to have a profound effect upon the plant growth of the ranges. Certain species of plants cannot survive on highly alkaline s o i l s , and thus many species are restricted or prohibited from growing i n this area. The high alkalinity of Range No. 4 undoubtedly inhibits much plant growth. However, on the other ranges there i s such uniformity of pH that i t can be disregarded i n comparisons of the various ranges. As an ind-ication of optimum reaction conditions, the pH range of certain plants studied i s given i n Table XIX. Alkalinity suppresses the solubility of some elements re-quired by plants. The insolubility of phosphates i n a s o i l containing calcium carbonate i s an example of this condition. E l l i s i n discussing high lime soils of Manitoba states (15) "In the high lime s o i l s , phos-phorus is either low or in a not readily available form with the result that plants do not thrive as well as on the soils where phosphorus i s available." It i s , therefore, not surprising to find very l i t t l e a v a i l -able phosphorus i n these s o i l s . As this element has a great effect upon the a b i l i t y of a plant to withstand adverse conditions such as over-TABLE X I X OPTIMUM pH Range of V a r i o u s P l a n t s (29) Aspen 3.8-5.5 Bearberry 5 . 0 -8 .0 Bedstraw 5 .0 -6 .0 B l a c k - e y e d Susan 5.5-7.0 Douglas f i r 6.0-7 .0 F l a x 5.0-7.0 J u n i p e r 5 . 0 -6 .0 S e r v i c e b e r r y 5.0-7.0 P o p l a r 6 . 0 - 8 . 0 - 29 -grazing, its lack in this area is an important factor in the condition of the ranges. In general high alkalinity tends to limit the supply of many nutrients as negatively charged nutrient ions are absorbed with difficulty by plants when the soil reaction exceeds pH 7.6. Due to the nature of the parent material potassium is found only in very small amounts. This element is limited in limestone, and thus, one cannot expect to find large quantities in the soils derived from this rock. The potassium that is present in this area is largely unavailable because of the coarse texture of the soils. It is certain, therefore, that very l i t t l e of this essential nutrient is present in the area under consideration. Potassium plays an important part in many of the vital physiological processes of the plant, and a deficiency has decided effects, such as lowering disease resistance, photosynthetic functions, and re-productive capacity. Nitrates are more plentiful than the above two elements, but the extreme instability of the nitrate content in a soil at any given time makes the results of the test less conclusive than the results for phosphorus and potassium. In general, available nitrogen is produced in proportion to the activity of microorganisms and to the amount of organic matter in the soil. Light soils poor in organic matter, such as Ranges No. 3A and No. 6, contain much fewer nitrifying organisms than heavy soils rich in organic matter, such as Ranges No. 1 and No. 5* The optimum soil reaction for the functioning of beneficial s o i l organisms i s approximately neutral, while temperature and moisture should be high. The opposite conditions of moisture prevail on these ranges and reaction i s above the optimum. In view of the above facts and the results of the tests, which reveal comparatively small amounts of nitrates, i t can be assumed that the nitrogen content i n this area i s low, b. Physical A study of the texture of these soils reveals that i t plays as important a part as the chemical reaction i n conditioning the general character of plant growth, and that i t appears to be the decisive factor i n the type of growth on certain of the ranges. Texture i s important by reason of the differential water holding capacities of the various textural classes. The water retention capacity of a s o i l composed of large particles (such as sand) is much less than that of a small particled s o i l (such as clay). E l l i s demonstrates this i n the following figures (15). Inches of water per foot available to plants that the various s o i l classes w i l l retain Sands Sandy loams 1" Fine sandy loams l l " ! 3/4" Loams 2" Clay loams 3" Clays 3k" With the exception of Range No. 1 a l l the ranges exhibit a coarse texture i n at least one of the horizons of the p r o f i l e . - 31 -It i s primarily because of i t s high water retention capacity that Range No. 1 has the most luxuriant ground vegetation of the area studied. In discussing similar soils i n Manitoba E l l i s states.(15) that " i t i s obvious that the lighter textured soils are submarginal soils i n a region of light precipation." The chemical activity of a s o i l i s greatly affected by the size of the separates. Thus sand separates take an almost neg-l i g i b l e part i n the chemical and physical a c t i v i t i e s unless the particles are composed of CaCO^ or some other appreciably soluble compound. Clay particles on the other hand are synthesized i n the s o i l and are very active chemically as demonstrated below. As the soils studied are c*om-posed for the most part of large separates the soluble elements are not found i n great amounts, even though they may be present i n the s o i l part-i c l e s . Comparative Nutrient Content of S o i l Separates from 5 soils derived from limestones and shales (23). P2°5 CaO MgO K20 sand s i l t clay sand s i l t clay sand s i l t clay sand s i l t clay 0.19 0.17 0.49 7.55 6.82 6.67 0.44 0.52 1.84 1.49 1.95 2.67 C. Organic Matter Content The test for organic matter i n the soils studied reveals great variation i n the percentages of the various ranges, (Table II) . As the source of s o i l organic natter is almost entirely plant growth i t can be asserted that those soils with high percentages have supported more plant growth i n the past than those with low percentages, provided that conditions for the functioning of s o i l organisms (temperature, moist-- 32 -. ure, reaction) are similar, as is the case in the Athabaska valley. Ranges No.'s 1, 5, 7, and 8 have, therefore, produced in the past considerably more vegetation than the other areas, although production at the present time is very low on certain areas. Ranges No. 3A and No. 6 on the other hand have never produced much plant growth and the climatic conditions combined with the texture of these soils pre-cludes any great accumulation of organic matter. Organic matter serves many useful functions and its presence increases the general fertility of the soil, both by physical and chemical means. Organic matter increases the water retaining power of soils, decreases water run-off losses, improves aeration , and produces a better soil structure.. It is very important in controlling wind erosion as i t serves to bind the soil particles. In this connection it is sig-nificant that those ranges most severely affected by wind erosion (Ranges No. 3A and No. 6) are also the lowest in organic matter content. In the semi-arid climate of this area any agency which serves to retain moisture in the soil is of great value. Thus, Range No. 1 with its high organic matter content and fine textured soil separates is far superior in productivity to Range No. 2 which is much lower in organic matter and coarser in texture. The chemical effects of organic matter result from the decay activities of microorganisms. Nitrogen producing activity has al-ready been referred to, and the conditions which inhibit the nitrifying process in the soils sampled also prevent rapid decay of any of the organic materials. Therefore, decomposition of organic matter on these - 33 -ranges i s greatly retarded, which accounts for the high percentages on some areas despite the comparatively limited current plant production. Because the activity provoked by organic matter de-composition is more important than i t s presence i n the s o i l , the varying amounts of organic matter are not necessarily indicative of the nutritive value of the various soils sampled. As favorable s o i l conditions for any plant demand a suitable s o i l texture, the right temperature, and an adequate supply of nutrients and water, i t i s apparent from the above discussion that most of the soils sampled are unfavorable to heavy plant growth due primarily to a low moisture retention capacity and an inadequate supply of nutrients. Assuming that the differences i n nutrient supplies are inconsequential, i t must be concluded that the low productivity present, and low organic matter of Ranges No.'s 2, 3f 3A U, and 6 are due to the poor water re-tention capacities of these soils or to the topography of the ranges while the low productivity of Ranges No.'s 5, 7 and 8 indicates some recently introduced factor. From the edaphic viewpoint, Range No. 1 i s far superior in productivity to the other ranges because of i t s superior moisture holding qualities. 3. Climatic Climate is the most important physical feature det-ermining vegetation, through i t s action i n forming soils and through i t s direct effect upon the plants. The relative rates of activity of the different forces of weathering and s o i l formation are determined by climate, which also limits the extent to which s o i l development proceeds. In arid regions,for example, the lack of water prevents s o i l development - 34 -from ever progressing to a mature type. Under the semi-arid conditions of Jasper Park, soil development under the best of conditions is very slow. Among the factors which comprise climate, the most important single influence in range productivity is precipitation. This influence is largely determined by the amount and seasonal distribution of precipit-ation. According to the U. S. Department of Agriculture (35)> the number of acres in good condition required to support one cow for one year in areas receiving various annual precipitations have been estimated as follows: Precipitation (inches)- Acres per animal per year 5-10 200 or more 10-15 70-200 Over 30 3- 12 As the annual precipitation at Jasper Park falls in the 10-15 inches class (Table I) it is apparent that the grazing animals using these ranges need a much larger area per animal over which to graze than they need on other ranges of higher precipitation. An important feature of the precipitation of this area is the local variation in amount. Primarily because of the topography of the val-ley the most easterly ranges (Range No. 1 and No. 2) receive considerably more rain during the growing season than those ranges nearest Jasper townsite (see Table I) The effect of this increased precipitation upon vegetation is augmented by the favorable water retention capacity of the soil texture of Range No. 1. Almost as important as the amount of precipitation is its seasonal - 35 -distribution. This determines whether vegetation receives moisture during i t s growing season, or whether the moisture must be stored i n the s o i l for use at some later period. Fig. 16 compares precipitation per month with the minimum, monthly temperatures. Assuming that the growing season of plants corresponds with the period of highest temperatures, i t w i l l be noted that the period of highest precipitation occurs during the growing season which i s , of course, a favorable situation for plant growth. Precipitation affects directly the animal l i f e of the area due to the varying depths of anow. Ranges such as No. 3A and No. 6 are heavily u t i l i z e d by sheep because the snow i s blown off regularly during the winter, exposing the grasses, while elk and deer migrate to the Athabaska valley bottom during the winter because of the generally shallow depths of snow on these ranges (see Table I) as compared to the deep snow of ranges at higher elevations. Temperature is a very important climatic factor with respect to both plants and animals. As mentioned above the growing season of plants is determined by temperature, which, i n Jasper Park, limits i t to a very few months. Vegetation i s also restricted by the extremes of heat and cold which are encountered i n this area. There are many plants which cannot stand the excessive cold, particularly the sudden drops i n temp-erature which are characteristic of the Jasper climate (see Table I ) . As the a b i l i t y of a plant to endure low temperatures depends on physio-logical condition, i t may be assumed that many of the overbrowsed and overgrazed plants i n this area have succumbed or w i l l succumb to periods of very low temperatures. - 36 -Temperature extremes have a decided effect upon the animal l i f e of the ranges. Heat would appear to be an important factor i n driving the sheep to higher ranges i n the summer, and prolonged periods of winter cold may k i l l grazing animals. Wind i s a climatic factor of considerable importance on certain of the ranges studied. The importance of wind in removing snow from sheep ranges has. already been alluded to. Erosion by wind following disappear-ance of the plant cover due to overgrazing is another influence having drastic effects on certain areas.:('FiglO)Death of these plants combined with trampling of the sod has destroyed the root systems and annual leaf carry-over that formerly held the s o i l intact. As a result of frequent high winds and semi-arid climate much of the top s o i l has been removed from large areas. This valuable material is blown about the valley or into the Athabaska river. Ranges No. 3A and No. 6 are typical of these areas, and their low productivity may be attributed primarily to wind and i t s effect on the s o i l . S o i l retrogression, as a result of wind erosion i s , therefore, taking place and could eventually reach the point where vegetation w i l l remain static (in an unproductive state) for many years, due to altered edaphic factors, even though grazing may have ceased entirely. 4. Biotic ^ a. Faunal The ungulates of the Athabaska valley are the most important faunal factor affecting the ranges. This is because of their large size, great numbers, and feeding habits. In general, large numbers of any ungulate have certain effects which become increasingly damaging to the ranges as the numbers of grazing animals increase. Braun-Blanquet summarizes the injuries resulting from pasturing as follows: (4) 1. Withdrawal of large quantities of plant matter. 2. Mechanical harm to plants by trampling, grazing, and brushing against them. 3 . Selective destruction of preferred species, 4 . Disappearance of dung avoiding species and intro-duction of strongly nitrophilous communities. 5. Direct effect upon s o i l formation by s t i r r i n g of earth and changing the micro-relief. The description of the ranges and the discussions of the above factors have indicated that many of the areas exhibit some of the above injuries. It w i l l now be determined to what extent the various ungulates found on the ranges influence the conditions as they have been described. As the elk i s the major influent on the Jasper Park ranges, i t has the greatest effect upon the ecology of the area. It has been shown in the historical section of this study that the present numbers of elk r are atvery recent phenomenon. The extent of the increase is demonstrated i n Cowan's report of 1943 (8). He states that i n 1931 only two elk were seen in Jasper Park, while i n 1943> three hundred and forty-seven were observed. In 1944 ( 1 9 ) , he estimated that 250-280 elk were ranging i n the Henry House area alone. A further indication of the extent of elk reproductive success are the figures supplied by Warden F. A. Bryant from his journal. They are summarized below:-YEAR AVERAGE SEEN PER DAY DURING MONTH OF JANUARY 1927 0 - 38 -1930 3 . 4 1931 3 . 3 1933 3 . 3 1937 18. 3 1938 11. 3 A total figure for the ranges under study i s imp»essible to est-imate accurately with existing information. However, some idea of the large numbers i n the Athabaska valley may be obtained by citing figures of surplus elk destroyed by Park authorities. Practically an the k i l l i n g took place i n the general area under study. YEAR NUMBER KILLED 1942-43 143 1943- 44 84 1944- 45 275 1945- 46 197 1946- 47 355 Despite the numbers k i l l e d during the preceding four years, i t was possible to report (10) i n 1946 that "Elk were present i n the Atha-baska valley i n much the same numbers as i n previous years". This report also cites a calf survival of 11 per cent for the year 1946. Considering such a low reproductive rate in the light of the more or less constant and at times increasing k i l l figures, i t may be concluded that the numb-ers of adult elk ranging the Athabaska valley must be at least ten times as great as the number k i l l e d , or the annual census figures would drop. - 39 -The largest numbers of elk have in the past been centered around the liberation area within a few miles of Jasper townsite. Only recently have they progressed eastward. In 1944, Cowan stated (9) that "The occupation of the Moosehorn river has lagged behind that of the Snake Indian river, but elk of both sexes are now present i n considerable numb-ers and i t can be expected that this valley w i l l shortly see a heavy increase in population with resultant heavier pressure on the already over-grazed Miette ranges". The recent arrival of elk on the eastern ranges has already been indicated by the more healthy condition of the plant cover of these areas. Elk are primarily grass feeders, as demonstrated by the following figures:-Jasper Park (9) Grass 97$ Browse 3% Due to the large size of elk the amount of forage consumed per individual i s great compared with that of bighorn sheep, mule deer, and mountain goat. Because of the large numbers of elk and limited supplies of available grass, these animals are now forced to compete with deer for browse, thus i n f l i c t i n g the serious damage to aspen, willow, buffalo-berx'y, and silverberry that has been described. The forage consumption equivalents of the animals under study and some livestock are given below: (31). Elk 1.00 Bighorn sheep 0.33 - 40 -Mule deer 0.32 Cattle 1.88 Horse 2.35 There are indications that the existing numbers of elk are be-coming less able to support themselves i n a healthy condition on the ranges. Cowan (10) attributes a part of the low calf survival already mentioned to poor physical condition of the mother resulting in abortion, resorption of the embryo, poor calf survival, etc. He also states that, i n early May, 1946 the average nutritional condition of elk was poor. Without the factor of human k i l l i n g of surplus animals, undoubt-edly the lack of forage would create, i n time, an ecological balance of range capacity and elk by the wasteful process of death by starvation and disease. K i l l i n g of surpluses by Park authorities i s delaying this s i t -uation and can prevent i t . Another factor of no l i t t l e importance i n checking the numbers of elk in the Athabaska valley i s the considerable number of timber wolves ranging this area. They are of great value i n k i l l i n g surplus elk i n areas where human control is impractical and costly. On the basis of studies made i n the winter of 1946-47, of wolves at Jasper and Banff Parks, i t may be stated that a large part of their winter prey is elk. A pack of four or five wolves was estimated to k i l l at a mini mum f i f t y elk per year, or from one to one and one-half elk per wolf per month i n the midwinter months. (11).There are known to be several such packs operating i n the area under study. The domestic horse also figures prominently i n the range ecology of the Athabaska valley. As indicated by Table XVII, this animal competes mainly with the elk for forage as range requirements are much the same. - 41 -According to Stoddart and Smith ( 3 l ), "Horses are the most selective of the domestic animals. They are primarily grass eaters and u t i l i z e r e l a t -ively small amounts of other forage, though forbs and browse may become important where grass i s scarce." The importance of the horse i n the Athabaska valley i s indicated by the numbers of head of domestic stock that have been o f f i c i a l l y grazed in the past few years, as given below:-Year No. of Stock 1934-35 183 1935-36 181 1936-37 221 1937-38 214 1938-39 186 1939-40 193 1940-41 196 1941-42 175 1942-43 176 1943-44 143 1944-45 ' 147 1945-46 140 1946-47 128 Supplementary to these figures, the Superintendent of the Park writes that twenty of the above annual numbers were cows u n t i l 1946-47, when a l l were horses. He further adds that the above figures should be i n -creased by 25 per cent to account for the wild horses running in the Park and those not declared. The eighty-five government horses are not included in the above figures, as they are fed approximately 100 tons of hay each year. However, most of them spend from October to May on wild range. Considering the large numbers of horses present i n the Park and the fact that the horse/consumes double the forage of the elk (page 39) , i t is apparent that those horses are maintaining themselves at the ex-pense of the game population. I t i s also apparent that they have been a decided factor i n the past years i n hastening range deterioration. Bighorn sheep are an important factor on many of the areas studied being second in numbers to the elk. In 1943, Cowan estimated the sheep population as follows ( 8 ) : -Devona 350 Jasper benches 40 Miette 450 Snaring to East Gate...... 200 Like the elk and horse, sheep are primarily grazing animals. Cowan (9) gives the food of bighorn sheep in "Jasper Park as follows:-Grasses and sedges 83 per cent Pasture sage 10 per cent Bearberry... 6 per cent Mountain Juniper 1 per cent From defecation counts, i t i s apparent that elk and sheep are competing heavily for the range forage on several areas. In any long run competition of this type, i t is the author's opinion that the elk would become dominant at the expense of the sheep, due to the latter - 43 -species' more specialized environmental requirements. Cowan (8) ind-icates that a scarcity of lambs i n 1943 nay have been due to nutritional deficiencies of ewes resulting from over-populated ranges and severe weather. The large numbers of sheep and elk on sparsely vegetated ranges such as Hanges No. 3A and No. 6, are having a direct effect upon s o i l development. By changing the micro-relief through terracing and t r a i l formation ( f i g . 9) the hoofs of these animals are removing a l l plant cover from large sections of the ranges, thus aiding the degenerative forces of erosion by wind and water, already noted. In the light of the above facts i t must be pointed out that big-horn sheep are one of Jasper Park's greatest natural attractions from an aesthetic point of view. It is a species which has decreased greatly throughout much of i t s range in North America, and i t i s of primary importance to conservation that bighorn sheep flocks be maintained i n good condition at a l l times where this is feasible. According to historical records, the Athabaska Valley was orig-inally well populated with sheep, and i t would seem to be a mistake to permit elk to drive out any of the sheep flocks. Mule deer have always been one of the chief ungulate forms of the Athabaska Valley. At present they are found i n considerable numbers, particularly i n winter and spring, upon some of the areas studied. In a 13-day examination of the Athabaska Valley i n 1943, Clarke (5) saw 168 deer, while Cowan i n 1943 (8) found concentrations of deer on early spring ranges i n the Snaring-Devona area of 35-60 deer per square mile. The destruction of browse by elk has greatly affected potential - 44 ~ deer forage, as this species is primarily a browser. The following figures show the food of the mule deer in Jasper Park as determined by Cowan (9). Grass and sedge, 15% Pussytoes 2% Pasture sage....... t r . Forbs 4% Bearberry 58% Buffaloberry 9% Other browse. .12% The d i f f i c u l t i e s that deer are encountering on the ranges under study are described by Cowan (9): "Over-browsing of the species (willow and aspen) has removed them from the browse available to deer and there are few places where even elk can s t i l l obtain them." ©. Phytobiological As the descriptions of the ranges indicate, the interrelation-ships of the above factors plus other environmental influences have pro-duced a varied plant growth in this area. It i s the purpose of this section to discuss the contemporary effects of the various factors on the vegetation, indicate certain trends, i n the development of plant l i f e , and show the effects, i f any, of the plants on the above factors. A. Ground Vegetation It was pointed out i n the discussion of faunal factors that grasses are of primary importance to the ungulates of the Park. They are also among the f i r s t species to decrease on an overgrazed range be-cause they are highly preferred food plants of sheep and elk. As an aid in evaluating the palatability of the grasses of these ranges as compared - 45 -to other species of ground vegetation the following palatability ratings are given, as they concern sheep and cattle, animals with feeding habits somewhat similar to mountain sheep and elk respectively. (27). In these cases palatability i s the percentage of plant growth that animals w i l l u t i l i z e on a properly grazed range. . . Thickspike wheatgrass Cattle 80$ Sheep 60$ Junegrass 60$ 50$ Needlegrass 60$ 40$ Pussytoes 0 0 Pasture sage 10$ 20$ Following heavy grazing less palatable species w i l l invade the areas denuded of the preferred forms. As retrogression proceeds, with a lessening of forage, the less palatable species w i l l also be consumed. Because animals under sufficient pressure w i l l consume almost any plant, prolonged overgrazing may result i n the complete removal of a l l plant growth. The disastrous effects on the s o i l of such a situation have already been considered. From the data available, i t i s apparent that the ranges farthest from Jasper, exhibit a generally healthy ground vegetation, while a l l those nearest Jasper show that grasses have declined to such an extent that the grazing animals are increasing pressure on the less preferred invaders such as pasture sage and also upon the shrub and tree growth, A comparison of the present status of the ground vegetation on Ranges No. 1 and No. 2 (farthest from Jasper) with that on Range No. 7 (nearest Jasper) indicates the v a l i d i t y of the above assumptions. - 46 -Range Number 1. 2 7 % of ground cover 3 1 . 0 4 5 . 0 2 5 . 0 fo of pussytoes and pasture sage of total ground cover 8 . 0 6 5 . 0 2 5 . 0 % of grasses of total ground cover 3 2 . 0 3 2 . 0 2 2 . 0 The relative amount of pussytoes and pasture sage i s very small on Range No. 1 where horse grazing i s very l i g h t . However, oh Range No.2 where horse grazing has been and s t i l l i s heavy, (table XvXl) pussytoes are dominant due to the heavier grazing and inferior s o i l . On Range No. 7 there i s a greatly reduced amount of total ground cover, as well as reduced amounts of both overgrazing indicators (pussytoes and pasture sage) and grasses. Further evidence of this trend toward reduction of total ground vegetation i s given i n Table XX which compares point sample counts of two of the ranges made i n different years. The figures i n Table XX show a high percentage of pasture sage i n comparison to other species. According to Stoddart and Smith (31), a high density of this species indicates misuse of the range. Although i t cannot be s t a t i s t i c a l l y proved as yet that pasture sage has increased at the expense of grass on these areas, i t i s practically certain that this species i s invading areas where grass has died. From the above figures, i t can be seen that due to the lack of grass, pasture sage i s also being grazed, despite i t s unpalatability, to the point where i t i s no longer maintaining i t s e l f . That the decrease of ground vegetation on ranges nearest Jasper i s due in large part to grazing i s demonstrated by the heavy vegetation of the cemetery, where percentage of ground vegetation even exceeds that TABLE XX Range Types of vegetation: Grasses and. sedges 3.6% Pasture sage...... 12.1 Other plants 6.2 Total plant cover 21.9 Total bare ground 78.1 Range 1944 Types of vegetation: Grasses and sedges 22.3% Pasture sage.and antennaria 7.2 Other plants 4.3 Total bare ground 66.2 - 47 -of Range No. 1. $he species of grasses and their occurrences on Range No. 1 and in the cemetery are very similar, indicating that before heavy grazing over an extended period the ranges nearest Jasper were, in general, similar to Range No. 1 where heavy grazing is more recent. The composition of ground vegetation of Range No. 7, is, however, very different, although in other conditions Range No. 7 resembles the cemet-ery area. Range No. 1 and the cemetery both produce high percentages of midgrasses, particularly Agropyrons, while on Range No. 7 the short junegrass is most prevalent, and no Agropyrons are grown. Sedges are also common on the latter range. According to Weaver and Clements (34) midgrasses disappear under frequent cutting while short grasses and sed-ges are much less affected. Clements states (7) that the decrease or disappearance of Stipa or Agropyron or of both marks the first stages of overgrazing. The death of the grasses can be attributed to several causes stemming from overgrazing. Any grazing has ah influence on the metabolic function of the plant. When photosynthetic tissue is reduced, there is a decrease in carbohydrate and nitrogen reserves as well as a lowered rate of root and forage production (31). Hanson and Stoddart (17) demon-strated this in the case of Agropyron inerme. a close relative of one of the chief forage grasses of Jasper Park. A count showing the reproduct-ive rates of this grass on grazed and ungrazed areas is given below with counts of seed heads made inside and outside exclosure plots at Jasper. These latter counts support the findings of the above investigators. No. of viable seeds per sq. m. 1 2 . 2 6 3 0 . 2 - 48 -Agropyron inerme Condition No. of heads per Filled Florets sq. meter per sq. meter Grazed 7 . 1 19.6 Protected 120.4 972.6 Number of seed heads of Gramineae Range Area counted Grazed Protected 1 4 sq. yds 0 11 2 4 sq. yds 29 63 3 4 sq. yds 154 5 6 l 5 4 sq. yds 12 76 With such drastic reductions in reproductive potential a result of overgrazing, it is easily understood why certain grasses are not main-taining themselves on many areas. Williams (35) found that Agropyron. when too closely grazed developed very few i f any fertile culms. Added to this lack of young plants are the factors of death from old age and from the physiological and mechanical effects (trampling) of overuse by ungulates. The discussion of the soils, climate and topography of these ranges reveals great differences in the capacity of the various areas to support vegetation. This variation must be reflected in the ability of ground vegetation to withstand damage. On range No. 1 where factors are the most favorable to plant growth, much more grazing is possible without serious injury than on Range No. 7 3 for example, where certain factors are less favorable. Ranges such as No. 3A and No. 6 are incapable of supporting much ground vegetation even without the faunal factor. There-- 49 -fore, these ranges are the least able to endure any over-use. B. Shrubs. The browse species of highest p a l a t i b i l i t y i n the area are willow, and silverberry while buffaloberry i s heavily u t i l i z e d when the former are not available. For the browsing ungulates, deer and moose, they should be the chief sources of food. For the elk, primarily a graz-ing animal, they are a secondary source of food u n t i l the lack of grasses forces them to turn to browse species in addition to ground vegetation. This situation obtains at Jasper, and the great pressure upon the browse shrubs i s apparent from the individual surveys already considered. Willow, being a highly preferred browse species, has suffered f i r s t . Although i t was noted in the ecological history that willows were very evident some years ago, i t i s now almost impossible to find a l i v e willow on certain areas. Dead skeletons of this species remain as proof of i t s former abundance. Many have only recently died, while others s t i l l produce a certain amount of sucker growth, which is quickly browsed down. Unless this trend i s stopped soon i t w i l l be many years before willow can re-establish i t s e l f from the peripheral areas. The status of buffaloberry and silverberry i s somewhat more favor-able. However, most shrubs are resistent to heavy overbrowsing, and one must look for secondary symptoms. Dead branches remaining as stubs, a hedged appearance, greatly increased branching, and the absence of f r u i t -ing structures, are signs of over-browsing. A l l these symptoms are exhibited, to a varying degree, by the two plants under consideration. Under favorable conditions, buffaloberry - 50 -attains average heights of from 3 to 6 feet (Fig. 5) while silverberry grows to 6-8 feet i n height (Fig. 6 ) . However, the individual range area N surveys show the average heights of these plants to be greatly heneath the normal. Under existing circumstances, the plants are undoubtedly dying prematurely, and the lack of reproduction precludes their replacement. 6. Trees. From the standpoint of the mammal population i n the Jasper area, the aspen i s the most important tree. It is a highly preferred food of elk and i s also one of the principal foods of the beaver population. Records indicate that aspen has always been a common tree i n the Athabaska valley. However, the present survey shows this species to be i n danger of complete removal from those areas, where reproduction has ceased. This s i t -uation is indicated by the age of the trees surveyed. They are either mature or the current year's seedlings, with no intervening range of ages that would indicate steady reproduction. The seedlings are certain to be con-sumed before they reach a year's growth. That reproduction ceased from 15 to 20 years ago on typical over-browsed areas, such as Range No, 5 , was proven by ring counts on an average-sized aspen. This tree was found to be about fifteen years old - a figure that indicates the point at which successful reproduction ceased. This corresponds with the a r r i v a l of elk in numbers upon these ranges. The present pressure upon aspen i s greatly accelerating the tempo of forest succession. Heavy aspen growth i s characteristic of sub-climax forests. According to Harlow and Harrar (18), conifers, such as Douglas f i r , lodgepole pine, and white f i r , replace aspen in the Rockies. Inaa-much as these species are of no forage value, i t i s important to keep the Jasper ranges in a sub-climax dondition as long as possible. As has been pointed out, Range No. 2 i s rapidly developing into a climax spruce forest, at the expense of a l l other browse plants. In a discussion of forest succession and i t s relation to the game animals of Jasper Park, i t i s necessary to draw attention to the effect of s t r i c t f i r e protection upon the future forage value of the ranges. According to Harlow and Harrar (18), burnt-over sites are the most favorable for production of aspen, but after twenty to thirty years, much of the aspen dies due to i t s intolerance of the invading conifers. There is abundant indication that similar circumstances apply i n the Atha-baska valley. Therefore, s t r i c t f i r e protection, or lack of other con-t r o l s , on the conifers, w i l l produce a climax conifer forest of almost no forage value. The poplar, although formerly common, according to the records, i s not frequently found. Under existing circumstances, i t cannot maintain i t s e l f . . * . « * * . « # . . The above analysis of factors affecting the ranges shows that there i s a complex interaction i n which the relative importance of the various factors varies from range to range. The ultimate effect of this interaction i s measured by the productivity of the ranges as shown in Table XVIII. The study indicates that over-use by ungulates i s the most ims--portant single factor determining the present ground vegetation of the ranges. This i s clearly indicated by the reduced yields on unprotected areas, and by a syndrome of symptoms well-known as indicators of range misuse. These symptoms axe: 1. Reduction i n reproduction of grasses. 2. Reduced occurrence of midgrasses with increased occurrence of short grasses. 3. Increase in pasture sage. 4. Decrease in total amount of ground vegetation. Insolation and the amount of s o i l moisture determine the tree and shrub growth of the area with the result that only the more level areas (Ranges No. 2, 3> k, 5, 7, and 8) support a tree and shrub growth. On these areas browsing by ungulates i s fast becoming the limiting factor as i t has curtailed reproduction of palatable species (aspen, willow, buffaloberry, and silverberry) on certain areas, and as i t i s k i l l i n g the adult plants of silverberry, buffaloberry,and willow. Thus, on Range No. 2, where elk are a recent influent, the s o i l is of poor texture compared to Range No. 5, but this iiange supports a much more luxuriant tree and shrub growth than Range No. 5> exhibiting a higher percentage of young aspen and poplar than any other area. As the degree of destruction of browse species increases accord-ing to the duration of occupation by elk, i t i s apparent that this animal i s responsible for the present destruction. The mule deer (which is primarily a browser) has been a constant factor i n this area, arid the gradient i n overbrowsing cannot be attributed to this animal. While the ground vegetation at the present time i s primarily affected by animal use, other factors are of importance, as the product-i v i t y table reveals differences i n production i n the protected areas. Increased precipitation causes greater general productivity on the ranges to the east than on the areas near Jasper. However, very local differences i n production and ground vegetation composition, such as occur between Range No. 1 and No. 2, must be attributed to the s o i l factor. As pH and chemical properties were found to be f a i r l y uniform (except on Range No. 4 where the pH undoubtedly inhibits plant growth to a high degree) on a l l ranges, moisture holding capacity, resulting from texture and organic matter, i s the limiting factor-as regards s o i l . Wind was found to be a factor lowering the percentage of organic mat.ter and thus the moisture holding capacity of ranges No. 3A and No.. 6. Because the areas to the east can produce more vegetation than those nearest Jasper, they are better able to tolerate over-use and to recover from this over-use, VI. CONCLUSIONS AND RECOMMENDATIONS The foregoing information shows that ungulates are so numerous i n the general area studied that many of the ranges are i n an acutely over-browsed and /or overgrazed condition. Ranges nearest Jasper have been the most severely damaged because they have been u t i l i z e d more intensely for a longer period of time than the easterly ranges, and because edaphic and climatic factors are not as favorable near Jasper/on certain ranges to the east. Thus, ranges,such as Range No, ljare capable of supporting many more elk at the present time than areas such as Range No, 7. The ungulate chiefly responsible for the misuse of the area studied i s the elk. This animal has increased greatly i n a very short time on ranges that had been previously overgrazed as horse range. It i s competing for winter food supplies with bighorn sheep on,certain - 54 -ranges(No. 3A and No. 6) that are capable of supporting only a very limited amount of plant growth. Unless present trends i n the ecology of the area are reversed in the very near future, the health/conditions^ of the ungulate herds J of the area w i l l be seriously menaced, due to malnutrition. At present the situation i s one of increasing numbers of competing individuals on ranges with rapidly decreasing amounts of forage. Under existing conditions, a winter of deep snow w i l l almost certainly result i n heavy game losses. Predators, such as timber wolves, are performing valuable services i n checking elk increases. The effect of misuse upon the s o i l of the ranges i s increasingly profound. In certain areas a l l plant cover has died, and the s o i l i s being dissipated by wind. This situation w i l l rapidly become widespread unless the ground vegetation i s permitted to recover i t s original charac-ter. Destruction of vegetation also tends to slow down'or prevent the formation of a mature s o i l . Plant succession is i n a state of retrogression on the area studied. This i s particularly true of the ground vegetation, where production of palatable species has been greatly decreased. Invading, less palatable^forms are also showing signs of retrogression, resulting in very large percentages of bare ground. The principal browse shrubs, silverberry and buffaloberry are suffering great physiological and morphological damage from over-use of twigs and foliage. This damage i s drastically curtailing reproduc>-tio n , as also is the consumption of seedling plants. Willow has been practically exterminated on the areas nearest Jasper. - 55 -The tree associations of the Athabaska valley are changing, due to the total lack of reproduction i n many areas of the favored species, aspen and poplar. The decline of the Salicaceae. plus r i g i d f i r e protection, tends toward a rapid succession to a climax spruce forest, unsuitable for the support of ungulates. On the basis of the above conclusions, i t i s recommended that the following steps be taken to correct the existing situation i n the •Athabaska Valley, and to maintain a satisfactory balance. (1) Continued reduction of the elk to a point where the ranges w i l l show a substantial increase i n grasses, particularly in the t a l l e r midgrasses, with a corresponding decrease i n the over-grazing indicators, and un t i l aspen and willow are able to reproduce sufficiently to maintain or increase present numbers. This reduction should be concentrated near Jasper and on the sheep ranges. (2) Annual census of elk, sheep, and deer, to determine popul-ation trends rather than absolute numbers. Methods such as those used by Hunter (19), would be applicable. In addition, the annual elk slaugh-ter by Park authorities lends i t s e l f particularly well to the use of a technique described by Olsen (25) in which calf elk would be tagged and population trends calculated on the basis of a ratio between elk k i l l e d and total population. Also pellet group quadrats, as described by Bennet, English, and;, McCain (3) for deer studies could be established. (3) The making of a cover-map of the principal winter ranges of the Park by a wildlife technician, as described by Dalke (12). (4) A study of r e l i c t areas to determine original climax - 56 -v e g e t a t i o n . (5) M a i n t e n a n c e o f e x i s t i n g e x c l o s u r e p l o t s a n d e s t a b l i s h m e n t o f a d d i t i o n a l o n e s t o d e t e r m i n e t r e n d s i n p r o t e c t e d a n d u n p r o t e c t e d a r e a s . (6) P e r i o d i c s t u d i e s , s i m i l a r t o t h e s e , o f t h e r a n g e c o n d i t i o n s w i t h a v i e w t o e s t a b l i s h i n g c a r r y i n g c a p a c i t y f i g u r e s . (7) P r o h i b i t i o n o f h o r s e g r a z i n g o n t h e w i n t e r game r a n g e s o f t h e A t h a b a s k a v a l l e y . ' (8) C o n t r o l o f c o n i f e r s w h e r e t h e i r e n c r o a c h m e n t o n v i t a l game w i n t e r r a n g e i s o c c a s i o n i n g a r e d u c t i o n i n s u c h r a n g e . (9) I n i t i a t i o n o f a k e y a r e a , k e y p l a n t s t u d y as a n i n d i r e c t m e t h o d o f i n d i c a t i n g t h e r e l a t i o n s h i p b e t w e e n t h e p r o p e r u s e f a c t o r a n d t h e a c t u a l u s e f a c t o r o n t h e r a n g e s . T h i s m e t h o d w o u l d c o n s i s t o f s e l e c t i n g c e r t a i n a r e a s o n r e p r e s e n t a t i v e w i n t e r r a n g e s , a n d a p p r a i s i n g t h e d e g r e e o f u t i l i z a t i o n o f t h e e s s e n t i a l f o r a g e p l a n t s p r e s e n t . PLATE I '> ^ 7 j "To Fig. 1. Map of the Range Areas :he Area studied showing individual Fig. 2. The Athabaska Valley showing a Section of the area studied. PLATE III. F i g . k Range No. 2 showing proper u t i l i z a t i o n of browse species. Photograph by Dr. I. McT Cowan. PLATE IV Fig. 5 Normal Growth of Buffaloberry on Range No. 2 Photograph by Dr. I. McT. Cowan. Fig. 6 Normal Growth of Silverberry on Range No. 2 Photograph by Dr. I. McT. Cowan. PLATE V Fig.8 Young Spruce invading the western portion of Range No. 2.* Photograph by Dr. I. McT. Cowan. PLATE VI Fig.9 Range No. 3A showing terracing by grazing animals. Photograph by Dr. I. McT. Cowan. PLATE VII Fig.11 Range No. 7. Photograph by Dr. I. McT. Cowan. PLATE VIII Fig.13 B u f f a l o b e r r y on Range No. 7 showing much dead m a t e r i a l . Photograph by D r . I* McT. Cowan. PLATE IX F i g . 14 Silverberry on Range No. 7 showing much dead material. Photograph by Dr. I. McT. Cowan. PIATE X Fig. 15. Correlation of measured Factors of Ranges No. 1 and No. 7 with the Productivity of those Ranges. (The figures for animal u t i l i z a t i o n were obtained by dividing the to t a l defecation counts for each range by fiv e ) . Prec ipitat ion(inches) 1111111111111111 m ~ | T m 4 j 11111 PLATE XI Fig. 16. Average monthly Precipitation and minimum Temperatures at Jasper Park (22 years). LITERATURE CITED 1. Aldous, Shaler E. , 1944. A deer browse survey method. Journ. Mammal-ogjrVol.25 No.2. pp.130-136. 2. Bates, C. G., 1923. The transect of a mountain-valley. Ecology. Vol. 4 No. 1. pp. 54-62. 3. Bennett, L. J., P. F. English, R. McCain, 1940. A study of deer pop-ulations by use of pellet group counts. Journ. Wildlife Manag. Vol. IV, No.4,pp. 398-403. 4. Braun-Blanquet, J., 1932. Plant Sociology. McGraw-Hill Book Co., Inc. New York and London, pp. x v i i i 439* illustr. j 5. Clarke, C. H. D., 1942. Wildlife investigations in Banff and Jasper National Parks in 1941. Nats. Parks Bureau, Mimeog. pp. 1-21. 6. Clarke, S. E., J. A. Campbell, and J. B. Campbell, 1942. An ecological and grazing capacity study of the native grass pastures in southern Alberta, Saskatchewan, and Manitoba. Tech. Bull. 44, Dom. Can. Dept. Agric, pp. 1-31, 12 tables, 9 figs. 7. Clements, F. E. 1920. Plant Indicators. Carnegie Institution of Washington, Washington, D. C, pp. xvi 388,illustr. 8. Cowan, I. McT., 1943. Report on game conditions in Banff, Jasper, and Kootenay National Parks, mimeog. pp. 1-72. 9. Cowan, I. McT., 1944. Report on game conditions in the Rocky Mount-ain Parks, Mimeog. 10. Cowan, I. McT., 1946. General report upon wildlife studies in the Rocky Mountain Parks. Unpublished Manuscript. 11. Cowan, I. McT., 1947. The Timber Wolf in the Rocky Mountain National Parks of Canada. Canadian Journ.of Research. Vol. 2 5 , pp. 139-174, illustr. 12. Dalkie, Paul, 1941. Development of a statewide system of covermapping in Missouri. Journ. Wildlife Manag. Vol.IV No. 1, pp. 103-107. 13. Douglas, David, 1914. Journal kept by David Douglas during his travels in North America. 1823-1827. W. Wesley & Son, London, pp. 364. 14. Description of and guide to Jasper Park, 1917. Dept. of Interior Ottawa, 97 pp front, (map) illustr. 15. E l l i s , J. H. 1938. The Soils of Manitoba. Manitoba Economic Survey Board, Winnipeg. 112 pp. i l l u s t r . 16. Geology of the National Parks of Canada, 1940. National Parks Bureau, Dept. of Mines and Resources, Ottawa, 20 pp. i l l u s t r . 17. Hanson, W. R., L. A. Stoddart, 1940. Effects of grazing upon bunch wheatgrass. Journ. Am. Soc. Agronomy. 32 (4) pp. 278-289. 18. Harlow, W. M., E. S. Harrar, 1941. Textbook of Dendrology. McGraw-H i l l Book Co., New York and London, pp. xv 542 i l l u s t r . 19. Hunter, G. N., 1945* Methods of determining trends i n big game numbers and range conditions. Trans. 10th N. Amer. Wildlife Conference pp. 234-241. 20. Klages, K. H. W., 1942. Ecological Crop Survey. The Macmillan Co., New York, pp. x v i i i 615, 108 maps. 21. Macoun, John, 1882. Manitoba and the Great Northwest. World Publish-ing Co., Guelph. pp. x x i i 687, front, i l l u s t r . maps. 22. McEvoy, James, 1900. Report on the Geology and Natural Resources of the country traversed by the Yellowhead Pass Route. Geological Survey of Canada. S. E. Dawson, 44 pp. 1 plate, map. 23. Millar, C. E., Turk, L. M., 1943. Fundamentals of S o i l Science. John Wiley & Sons, Inc. New York. pp. x i 462 i l l u s t r . 24. Moberly, H. J. , 1929. When Fur was King. J. M. Dent & Sons, London & Toronto, pp. x v i i 237. 25. Olson, H. F., 1943. Deer tagging and population studies i n Minnesota... 3rd. N. Amer. Wildlife Conf. pp. 280-286. 26. Preble, E. A., 1908. A Biological Investigation of the Athabaska and Mackenzie Region. U. S. Dept. A g r i c , Bur. Geol. Survey Gov't Printing Office, Washington, D. C , pp. 574, i l l u s t r . XXV plates. 27. Range Plant Handbook, 1937. Forest Service, U. S. Dept. A g r i c , U. S. Gov't. Printing Office, Washington, D. C. 28. Schollenberger, C. J., 1945. Determination of organic matter. S o i l Science. Vol. 59, No. 1 pp. 53-56. 29. Spurway, C. H. 1941. S o i l reaction (pH) preference of plants, Special Bull. 306 Mich. Agric. Exp't. Stn., East Lansing, Mich. 30. Spurway, C. H. 1944.vSoil Testing - a practical system of s o i l f e r t i l i t y diagnosis. Tech. Bull. 132, Mich. State College, East Lansing, Mich. 31. Stoddart, L. A., A. D. Smith, 1943. Range Management. McGraw-Hill Book Co., New York and London pp. v i i 547 i l l u s t r . 32. Talbot, M. W,, 1937. Indicators of southwestern range conditions. Farmers Bull. 1782, U. S. Dept. of Agric. U. S. Gov't Printing Office, Washington, D. C. pp. i i 35 i l l u s t r . 33. T y r r e l l , J. B., 1916. David Thompsons Narrative of his explorations i n Western America 1784-1812. The Champlain Society, Toronto, pp. x c v i i i 582. 34. Weaver, J. E., Clements, 1938. Plant Ecology. McGraw-Hill Book Co., Inc., New York and London, pp. x x i i 601 i l l u s t r . 35. Williams, T. A., 1898. A report upon the grasses and forage plants and forage conditions of the eastern Rocky Mountains Region. U. S. Dept. Agric. Div. Agrost. Bull. 12. Washington, D. C. 36. Yearbook of Agriculture, 1941. Climate andMan. U. S. Dept. A g r i c , Washington, D. C. pp, x i i 1248, maps, i l l u s t r . APPENDIX A Scientif i c names of Mammals i n the Text. Elk Moose Mule deer Bighorn sheep Timber wolf Mountain Goat Beaver referred to Cervus canadensis Alees americanus Odocoileus hemionus Ovis canadensis Canis lupus Oreamnus montanus Castor canadensis APPENDIX B Scientific Names of Plants referred to i n the Text Common Name Scien t i f i c Name Aspen Populus tremuloides Michx, Aster Aster sp. -Alpine Aster alpinus L. Aven -. Geum triflarum Pursh (G) Bastard toadflax Comandra sp. Bearberry Arctostaphylus uva-ursi Spring Bedstraw, northern Galium boreale L. Black-eyed Susan Gaillardia, sp. Bluebell Companula rotundifolia L. Blue-eyed grass Sisyrinhhyum angustifolium M i l l , Borage Boraginaceae Buffaloberry Sheperdia canadensis Nutt. Biiitterweed Senecio hydrophilus Nutt Cedar, ground Lycopodium, sp.? Cinguefoil Potentilla pennsylvanica var. strigosa Pursch. -shrubby Dasiphora fruticosa L Cranberry, high bush Viburnum opulus L. var. americanum Daisy Erigeron coespitosus Nutt -purple Erigeron linearis (Hook) Piper -white Erigeron compositus Pursch. Fescue Festuca sp. F i r , Douglas Pseudotsuga t a x i f o l i a (Poir) Flax Goldenrod No. 1 Goldenrod No. 2 Gooseberry Grass -Sandburg blue -Blue -Brome No. 1 -Brome No. 2 -June -Needle -Richardson's Needle -Pine -Purple reed -Bearded wheat -Thickspike wheat -Wild-rye Juniper -Common Legume L i l y Pine, lodgepole Poplar APPENDIX B (cont'd) Linum usitatissimum L. Solidago lepida D. C. Solidago uliginosa Nutt. Ribes sp. Boa secunda Poa sp. Bromus suksdorfii? Bromus pumpellianus Scribn. Koeleria cristata (L.Pers.) Stipa comata Trin. & Rupr. Stipa richardsoni Link. . Poa subpurpurea Rydb. Calamogrostis purpurascens R. Br. Agropyron subsecundum (Link) Agropyron dasystachyum (Hook & Scrib.) Elymus innovatus Beal Juniperus scopolorum Sarg. Juniperus communis L. Leguminosae Lilium montanum A. Nels. Pinus contorta var. Murayana Engelm. Populus tacamahaca M i l l . APPENDIX B Puccoon Pussytoes -Canada Rose Sage, pasture -Field Sedge Serviceberry ; ,Silverberry Spruce, Black Spruce, white Strawberry Thistle, Russian Vetch -Milk -Purple Violet Willow Wind Flower (cont'd) Lithosperum angustifolium Michx. Antennaria parvifolia Nutt. Antennaria canadensis L. Rosa Macounii Greene. Artemisisa frigida Willd. Artemisisa camporum Rydb. Car ex sp. Amelanchier sp. Eleagnus argentea Pursh. Picea marianna (Mill.) Picea glauca (Moench.) Voss Fragaria glauca (S. Wats) Rydb. Salsola k a l i var., tenuifolia Meyer Astragulus sp. Astragulus alpinus L. Astragulus sp. Viola sp. Salix sp. Anemone sp. 

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