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

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& 3  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 i n the Department of Zoology.  University of British Columbia  1948  a7  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. P f e i f f e r Recent studies by Dr. I . MoT.  Cowan i n Jasper National Park  have revealed that many of the winter game ranges of the Park are heavily o v e r u t i l i z e d by grazing and browsihg animals.  In order to  correct this s i t u a t i o n , 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 V a l l e y was undertaken i n the summer of and 1947  1946  by Dr, Ccwan and the author, f o r the purpose of describing  existing plant growth, investigating factors a f f e c t i n g t h i s plant growth, and recommending measures f o r 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. u t i l i z a t i o n of the ranges by herbivores was counts.  The r e l a t i v e  estimated by defecation  In order to determine the productivity of the ranges  equal  amounts of ground vegetation were clipped, a i r - d r i e d , and weighed. Exclo8ure p l o t s , established i n 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 r e s u l t o f the investigations i t was concluded that the winter ranges of the Athabaska V a l l e y vary greatly i n t h e i r capacity to produce vegetation, and that a l l the ranges are o v e r u t i l i z e d t o a greater or lesser degree, p r i n c i p a l l y by elk and horses. utilization  This over-  i s producing a retrogressive succession of the ground  vegetation which i s rendering the ranges i n c r e a s i n g l y 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 c e r t a i n range areas.  In order to correct present trends i n the ecology o f the Athabaska V a l l e y 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 c o n i f ers be c o n t r o l l e d on c e r t a i n game ranges.  TABLE OF CONTENTS PAGE Acknowledgements I. Introduction  1  I I . History of the Area  3  I I I . 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 9. Range No. 8  2  0  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 o f operative factors a f f e c t i n g the Ranges 1. Topographic  2  0  2. Edaphic  2  8  2  8  3 . Climatic 4. Biotic a.  33 Faunal  b. P h y t o b i o l i g i c a l  36 44  TABLE OF CONTENTS (cont'd)  PAGE VI. Conclusions and Recommendations Plates L i t e r a t u r e Cited Appendix A Appendix B  53  ACKNOWLEDGEMENTS I wish to express my gratitude to Dr. I. McT. Cowan of the Department of Zoology, University of B r i t i s h Columbia, f o r his suggestion of the subject of this t h e s i s , for his a i d i n the gathering of material, and for h i s 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 B r i t i s h Columbia, I extend my thanks f o r h i s a i d i n preparation of the work. I wish to thank Dr. V. C. Brink o f the Faculty of Agriculture, University of B r i t i s h Columbia and Dr. T.M.C. Taylor of the Department of Biology and Botany, University of B r i t i s h Columbia, f o r much time spent by them on the i d e n t i f i c a t i o n of many of the plants l i s t e d and f o r their i n t e r e s t and help i n the work. To Mr. L. Farstad and Mr. J . T. Gillingham of the Faculty o f Agriculture, University of B r i t i s h Columbia, I express my gratitude for a i d 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 V a l l e y 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 s i t u a t i o n became so acute that mass slaughter of e l k  was begun i n 1942 i n order to attempt to r e l i e v e pressure on the winter ranges.  I t became  apparent that a long term p o l i c y of range manage-  ment was needed, but, i n order t o i n i t i a t e such a p o l i c y , accurate information on the current s i t u a t i o n was necessary.  Therefore, a study  of certain key areas of the Athabaska V a l l e y 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 e x i s t i n g plant growth i n order t o have a standard by which to measure any changes i n the future. B. Investigation of the various factors a f f e c t i n g t h i s plant growth with a view towards ascertaining which factors are i n any degree controiable. C. Recommendations as to the procedure to be adopted f o r improving the range conditions. The following material i s the r e s u l t o f studies made during the summer o f 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 f o r study are samples of some of the p r i n c i p a l winter ranges of the ungulates of Jasper Park.  The study was confined  - 2 to winter ranges because they are the l i m i t i n g 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 f l o o r of the Athabaska V a l l e y i n the Jasper area, however, snow seldom reaches very great depths (see Table I) and any food plants that may be present are r e a d i l y accessible.  The animals, therefore, descend  i n the f a l l from the summer ranges to those areas where there i s l i m i t e d snow and a good supply of food. bighorn  In addition to these two conditions  sheep ranges are generally characterized by steep slopes.  Such  t e r r a i n offers escape from predator attacks and i s 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 l i m i t l e s s , are unavailable 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. I t i s hoped that the f o l l o w i n g study w i l l a i d the a p p r a i s a l of the carrying capacity of the Athabaska v a l l e y and indicate methods of i n creasing i t .  - 3 - • II.  HISTORY OF THE AREA A survey of the history of the Rocky Mountains of Canada i n d i c -  ates that more human a c t i v i t y of a commercial nature has taken place through the Athabaska V a l l e y and Yellowhead Pass than anywhere else i n these mountains.  This a c t i v i t y 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 h i s 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 f o r many years, brigades with goods and passengers moved through the pass between the P a c i f i c 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 f o r the t r i p over  the pass. The discovery of the Yellowhead Pass route i n 1826 r e s u l t e d i n s t i l l greater a c t i v i t y i n the Athabaska v a l l e y .  The Hudson's Bay Company  played a leading r o l e i n the development of t h i s area, as the writings of H. J . Moberly  (24), long an employee of the Company, i n d i c a t e .  In  the early days Jasper House (see Map Figure l ) served as headquarters f o r 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 t r a v e l l e d them annually i n search of meat.  The procedure was to remain i n a good  l o c a l i t y where feed f o r horses was p l e n t i f u l , 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 t h i s area.  According to Moberly, "These Indians a l l had bands of horses of  t h e i r own."  He estimates that i n 1855  some 350  mares were kept i n the  Athabaska V a l l e y as f a r as the confluence of the Athabaska and Miette Rivers. In the l a t e 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 t h e i r animals t r a v e r s ing the v a l l e y .  The construction of two railways, the Canadian Northern  (National) and the Grand Trunk P a c i f i c , i n the f i r s t and second decades of the  1900's brought  further great .activity to the v a l l e y and p r a c t i c a l  extermination to i t s large game. The human a c t i v i t y was gradually s t a b i l i z e d , however, with the formation of a National Park i n the area i n 1907,  followed by p r o t e c t i o n  of almost a l l w i l d l i f e . The l a s t h i s t o r i c a l event of significance to t h i s study occurred i n 1920  when 96 elk of both sexes were l i b e r a t e d at Jasper. In considering the above h i s t o r i c a l events, i t i s important to  r e a l i s e that great numhers of horses were involved i n a l l the successive human undertakings.  As i s shown below, they were supported on the open  - 5 ranges of the v a l l e y along with dwindling numbers of game.  On the basis  of these f a c t s , the conclusion i s reached that f o r a period of over one hundred years, the Athabaska V a l l e y was subjected to an abnormal amount of grazing. From the accounts of early travelers through the Athabaska Valley i t i s apparent that t h i s area, i n i t s natural state, was capable of supporting large numbers of ungulates.  Describing h i s t r i p across the  mountains to the Columbia i n 1811, David Thompson writes (33):  "We  now entering the defiles of the Rocky Mountains by the Athabaska  are  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 l a s t  grass (about Buffalo P r a i r i e ) f o r the horses i n marshes and ponds where a herd of bison had l a t e l y been feeding, and here we l e f t the horses, poor and t i r e d , and notwithstanding b i t t e r cold, (they) l i v e d through the winter, yet they have only a clothing of close h a i r , short and without any f u r . " David Douglas (13), i n 1827, found bighorn sheep " i n abundance" i n the Jasper area, and also writes of willows s i x to ten feet high on the r i v e r banks i n company with poplars and pines. The area appears to have remained highly productive i n the following years. W. Moberly (21), who spent the winter of 1872-3 i n the Athabaska V a l l e y , 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 s h i f t f o r 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 V a l l e y 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 h a l f 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 d i s t r i b u t e d throughout the v a l l e y . S p e c i f i c references to the game that has been supported i n the area are numerous.  Successful hunting, as described by Moberly, has  already been c i t e d , and i t i s obvious that game was abundant during these early days of human e x p l o i t a t i o n .  In the above report, McEvoy  outlines the status of c e r t a i n 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 a f t e r seventy-five years of human e x p l o i t a t i o n . 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. I t i s quite apparent that by the turn of t h e century changes i n the relationships picture.  of game and range conditions had a l t e r e d the e c o l o g i c a l  A Department of I n t e r i o r publication (14) states: "During  construction of the railways, a great portion o f the b i g game was either destroyed  or driven away from the main v a l l e y of the Athabaska, where the  open f l a t s and grassy meadows had afforded excellent ranges f o r deer." This publication also gives a b r i e f picture o f the tree growth of the v a l l e y i n 1917.  I t mentions the jack pine (Pinus contorta) as the  commonest t r e e , a t t r i b u t i n g t h i s to f o r e s t f i r e s .  Poplar, usually of  small s i z e , cottonwood, willow and alder were also prevalent, with much spruce i n the swamps. In discussing elk, the p u b l i c a t i o n states that no elk were present i n the Park i n 1917* was  However, i t asserts that the Athabaska V a l l e y  s a i d t o 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 v i n d i c a t e the assump-  t i o n that the Athabaska V a l l e y was ever w e l l 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 o r i g i n a l s t a t e , large numbers of ungulates; with the coming of the white man, with large numbers o f horses, much of the game was driven o f f or k i l l e d .  - 8I U . METHODS USED 1.  Point Samples;  The point sample, as developed by Clarke (6), which the composition was determined.  was the method by  and density of the ground vegetation on each range  B r i e f l y , t h i s method may be described as follows:  points are taken at i n t e r v a l s on transects of the range, l a i d out t o cross the maximum v a r i e t y o f 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 h i t s are counted. the point of a p i n touches i t s base.  A plant i s h i t when  Hits for each species are recorded  and are expressed i n terms o f percentages o f t o t a l number o f points f o r each area, With one exception, the t o t a l number i n t h i s study i s  1000,  This i s adequate, according t o Clarke (op. c i t . ) , to obtain a t r u l y 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 i n d i c a t i o n o f the amount o f available browse on each range that has a s u f f i c i e n t tree and shrub growth t o be s i g n i f i c a n t .  The  method consists of surveying milacre plots selected at regular i n t e r v a l s on transects of the range as described by Aldous ( l ) .  I n the survey of  each p l o t , the species observed are recorded as being Dominant, Moderate, or Sparse.  I f the species covers  50-100 per cent  of the p l o t , 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 subjected 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.  Clipping;  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 followaverage ing productivity figures represent an above/production of the ranges at  the present 5.  time.  Defecation Countst  In order to establish a basis f o r r e l a t i v e u t i l i z a t i o n of the ranges by d i f f e r e n t species, counts were made of the various ungulate or equine defecations encountered on a 1000 pace diagonal transect of each range.  I t must be emphasized that these are not i n d i v i d u a l p e l l e t  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 r e l a t i v e usage of each range by the species involved. 6 . Exclosure plots 1942,  Several exclosure p l o t s , constructed f o r the most part i n i n various areas of the Park, were clipped i n the summer of 1946 1947  and  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 p r e v a i l i n g range conditions and the difference i n p r o d u c t i v i t y i n and outside the plots was of the p l o t s .  i n no case due to the advantageous positions  They are, however, about 6 f t , by 6 f t . i n s i z e , which  i s too small f o r optimum study.  The Jasper town cemetery, about two  miles east of Jasper, served as one exclosure p l o t , as i t has been surrounded by a game-proof fence f o r at l e a s t twenty years.  In  1947  the amount clipped from each square yard was weighed and the mean determined from the sum of the c l i p p i n g s .  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 i n d i c a t i o n o f the e f f e c t of grazing upon grass reproduction, counts o f seed heads of the Graraineae were made within the exclosure plots and i n equal areas immediately adjacent to the exclosures. As t h i s count was made i n l a t e August 1947 a l l grasses had matured. 7.  R e l i c t Area  In addition t o serving as an exclosure plot the cemetery i s large enough ahd has been fenced i n f o r a s u f f i c i e n t l y long time to serve as a r e l i c t area.  I t was point sampled to obtain an i n d i c a t i o n of the  ground vegetation that i s developed without the grazing f a c t o r . 8. S o i l Analyses During August o f 1947 samples were c o l l e c t e d from t y p i c a l locafc— tions on the nine areas.  One p r o f i l e per range was exposed to a depth  of from 2 | t o 3 f t . and samples taken from the various horizons thus exposed.  The horizon designated C or D was generally at l e a s t 2 f t .  beneath the surface. These samples were then analyzed as follows: 1 . Texture was determined by manual means by an expert agronomist. 2 . pH was determined by the electrometer t e s t . 3 . Available potash, phosphorus and n i t r a t e s were determined by the Spurway method ( 3 0 ) , as these are the elements of major importance from the a g r i c u l t u r a l point of view.  - 12 4. Percentage of organic matter wqs determined by the wet ion method as described by Schollenberger IV.  combust-  (28).  Description of the Ranges  The Athabasfci v a l l e y from the town of Jasper east t o the eastern boundary i s the general area under consideration. According to Cowan's report for 1943  (8)>  t h i s area provides s u i t a b l e winter ranges f o r deer, sheep,  and elk (also domestic horses) and may be considered t y p i c a l . The v a l l e y l i e s between highly f o l d e d mountains which consist l a r g e l y of rocks that were l a i d down o r i g i n a l l y aa sand, mud and c a l c a r eous ooze.  They are, therefore, sedimentary and date back to the Paleo-  zooic and Masozooic eras (16).  The s o i l s of the v a l l e y are strongly  a l k a l i n e i n reaction and medium to l i g h t i n texture.  They are transported  s o i l s derived from aeolian, a l l u v i a l , or g l a c i a l 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 characteri s t i c s of the Rendzina s o i l group.  Table I I gives c e r t a i n q u a l i t i e s as  exhibited by the s o i l samples from the i n d i v i d u a l ranges.  These q u a l i t i e s  are discussed i n d e t a i l i n the section dealing with the analysis of the edaphic factors• Despite the northern l a t i t u d e and the considerable a l t i t u d e (approximately  3300 f t . ) this area enjoys a climate that i s so tempered  by the "chinook winds" that i t compares favorably with more easterly regions l y i n g 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 i n the Park. They have been assigned numbers beginning with number 1. at the eastern boundary of the Park and progressing westward tdfiumber.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 l i t t l e 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 l i t t l e significance to be surveyed.  TABLE I  Weather Data (average for 22 years) for Jasper Park daily Mean maximum Temperature  daily Mean minimum Temperature  Precipitation Rain  Precipitation Total Snow Precipitation.  Jan.  20  4  0.05 inches  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  9.03  40.3  Total annual  10.5 inches 1.10  Mean Precipitatidfi per Week May-September 1946 Jasper Townsite  .10 inches  East Gate  .44 inches  May-August 1947 Jasper Townsite East Gate Nov. Dec.  .66 inches  1.20 inches Temperatures at Jasper, winter of 1942-43 Mean Maximum Mean Minimum 28.4 degrees F. 11.0 degrees F 23.2 degrees F 5.0 degrees F  33.06  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 1  Horizon A B  C  2  3  4  A B D  S  8.4 8.3 8.6 8.6  violent violent violent  very f i n e sandy loam to s i l t s i l t loam loam heavy s i l t loam  violent violent violent  5 10  0 0  very low very low  2.65  s i l t loam f i n e sandy loam sandy loam ( g r a v e l l y )  2 0  0  very low very low  4.28  0  f i n e sandy loam loamy sand s i l t loam  very low  0.80  very f i n e sandy loam t o s i l t loam  medium-5 very low  5.01  sandy loam clay loam clay loam mixed with g r a v e l loam clay heavy s i l t loam to s i l t y  8.4  violent violent violent  A  8.2  violent  5  0  A B  8o4  5-10 2  1.0  9.1  violent violent violent  A B  violent violent violent  10 25  0 0  low medium-5  9.40  C  8.0 . 8.2< 8.0  A B  8.3 8.2  violent violent violent  5  0 0  very low very low  1.91  25  f i n e sandy loam fine sandy loam f i n e sandy loam  violent violent violent  2  0 0  very low very low  6.35  0 sandy loam loamy sand (some gravel) sand  low very low  6.40  loam loam gravel  A B  C  8  8.2  K Organic matter Ph Parts per m i l l i o n low 7.52f* 10-15 .5 10 very low 0  Texture  Nitrates  8.2  C  7  8.1  Effervescence  A B D  C  5  *pH  A B D  8.2  8.9  8.1 8.2 8.6 8.9 8.2 8.4 8,7  violent violent violent  10 5  .5  .5 .5  * The s o i l s were a i r - d r i e d , which reduces the pH i n a l k a l i n e  soils.  ** This s o i l exhibited only one horizon to a depth o f at l e a s t 3 f t .  clay loam  - 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 ( 1 0 . 1 #), and the presence of a variety of species of succulent forbs is further evidence of a healthy plant community. Over-use indicators, 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 boundary. 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 12,  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 73.7$  Bare ground Grasses  10.1  Thickspike wheatgrass Wild ryegrass  2.7  Brome grass No. 1  0.3  Purple reedgrass  0.3 13.4  Shrubs Bearberry  6.2  F i e l d sage  1.0  Pasture sage  0.5 0.2 7.9  '•• Rose 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 Vetch  0» 0«1  2  TABLE SPECIES  HEIGHT  NUMBERS  IV  DENSITY  PERCENTAGE  Trees 700  .95  8fft.  —  —  2  8+ft.  ~  —  —  Aspen  -4 f t .  Aspen Poplar  18  White Spruce  --8 f t .  190  2.75  8  White Spruce  8fft.  110  —  8  360  5.80  20  1740  3.55  9  Shrubs Buffaloberry Silverberry  — 1+ft  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 Poplar  Height -4 f t .  Numbers 40  Density .15  Percentage 3  - 15 discussions of the other areas.  However,  the existing relationships  are f a r from s t a t i c .  The table above shows a large number of small  spruce i n t h i s area.  This indicates a s i t u a t i o n which appears quite  obvious on inspection of the range ( F i g . 8 ) .  The spruce to the west  i s r a p i d l y invading the area under study and w i t h i n a few years w i l l transform the entire area into a climax spruce f o r e s t , thus e s t a b l i s h ing a hiome that would not include any of the present ungulate forms. The ground vegetation of t h i s area i s 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 i s 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 w e l l as the sandy  type of s o i l , undoubtedly accounts i n part f o r this condition. However, of pressure/animals grazing on the grasses and forbs i s probably a factor i n the e x i s t i n g prevalence of pussytoes.  This area has long been the  f a v o r i t e winter range of a group of horses. 3. Range No. 3t Devona F l a t s This area i s s i m i l a r topographically and edaphically Range No. 2.  to  I t i s an a l l u v i a l f l a t formed as a r e s u l t of the a c t i v i t y  of the Snake Indian r i v e r at i t s confluence with the Athabaska r i v e r . The area i s shaped l i k e a t r i a n g l e with the Athabaska r i v e r on the south as i t s base.  The northeast side i s bounded by the Snake Indian r i v e r ,  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 buffaloberry 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 V I . The percentage of grasses (9.0) on this area is reduced as compared 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 V I .  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 Bearberry  9.4  10.0  Forbs Pussytoes White daisy Flax Milk vetch Clinquefoil  2,  5  1.7 I*  1  1»° 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, i t 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$) and forbs. The extreme prevalence of „ pasture sage and Russian thistle< (11.7%) suggests that  the low  percentage of grass and forbs is not solely the result of factors described above, but is also the result of overuse by grazing animals. Retrogressive succession has proceeded further on this area than on ranges previously discussed. 5. Range No. 4: Greenock This area is similar In general topography to Range No. 3 a . It consists of the steep south-western slopes of the benches at the foot of Mount Greenock. These benches rise 200 to 400 feet from the more gently sloping valley and are composed of coarse, morainal gravel under-  TABLE  Bare ground  VII  80.9$  Grasses Junegrass  1.5  Purple reedgrass  .6  Thickspike wheatgrass  .4  Needlegrass  .3 2.8  Shrubs Pasture sage  10,4  Juniper  1.5  Russian t h i s t l e  1.3  Bearberry  .5  Forbs Aster Flax Unidentified  1.3 •^ »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 l i t t l e 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 .3  Junegrass  2.5 Shrubs Bearberry Juniper Pasture sage  -  18.4 1.2 .1 2Q.7  Forbs Northern bedstraw  .7  Legume  •2  Alpine aster Daisy  .2 »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 i t 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. Practically a l l these are yearlings and will be eliminated during the succeeding 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  -4 f t .  120  .45  9  Aspen Shrubs Rose  —  1910  2.55  46  Serviceberry  —  1460  .60  13  Silverberry  +1 f t .  Silverberry  1+ft  Willow  —  '  Buffaloberry Shrubby Cinquefoil Common Juniper  1+ft. —  1760  1.35  . 27  320  1.25  12  50  .20  4  110  2.65  11  120  .30  6  10  .30  1  OMISSION-TABLE IX  Species  Height  Numbers  Aspen  &> f t .  330  Buffaloberry  -1 f t .  320  Density 1.45  Percentage 22 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 i s almost i d e n t i c a l to Range No. 3a, but l i e s on the south bank of the Athabaska r i v e r between Talbot and Edna lakes. I t i s a steeply sloping h i l l r i s i n g from the Athabaska r i v e r on the north, and extending t o 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 l i m i t e d  and i s of l i t t l e s i g n i f i c a n c e .  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.  ( F i g . 10).  Such conditions have reduced the percentage of ground vegetation as shovm i n the survey made on August 12, 1946.  (Table XI)  As on Range Area No. 3a, pasture sage i s the dominant p l a n t ,  (5.0%), and grasses (8.9%) are l i m i t e d i n occurrence.  The general con-  d i t i o n of t h i s range indicates that i t i s i n a s i m i l a r stage successiona 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 F l a t s  This i s one of the largest l e v e l 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 .  I t i s , generally,  several feet above the r i v e r and i s 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 throughout the area.  The Athabaska r i v e r forms the eastern boundary, while the  western and southern l i m i t i s the rocky slope of the Palisade r i d g e . To the north, the area i s bounded by a shallow arm of the Athabaska r i v e r ,  TABLE X I 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 l o c a l l y known, i s t y p i c a l of much of the Athabaska v a l l e y from Henry House to Jasper.  I t i s a lodgepole pine sub-climax assoc-  i a t i o n 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  r e s u l t s shown i n Table XII. Buffaloberry i s the p r i n c i p a l browse shrub despite the high figure for serviceberry, which i s generally too small a plant to be significant.  The i n d i v i d u a l buffaloberry plants are stunted and show  much dead material ( F i g . 13).  The c r i t i c a l condition of t h i s species  i n this area i s apparent i f the data f o r buffaloberry on this range i s compared with the data f o r 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  the numerical counts f o r Range No. 7,  f o r Range No. 2.  As shown i n  the number of buffaloberry plants  under 1 foot f a r 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 t h i s area i s , potentially,  moderately productive. However, the large percentage  of sedge, junegrass, and overgrazing indicators such as and pasture sage indicate over-use by grazing animals. June 17, 1946  butterweed, The survey on  gave the composition of the ground vegetation as  i n Table X I I I .  shown  TABLE XII SPECIES  HEIGHT  NO. per ACRE  . DENSITY  ' PERCENTAGE OCCURRENCE  Trees  10  .05  Lodgepole pine - 8 f t .  190  1.05  Lodgepole pine  8+ft.  870  -8 f t .  30  .15  3  60  .80  5  Aspen  Spruce  -1 f t .  —  1 13 36  :ubs Juniper  —  Buffaloberry  -1 f t .  460  1.80  30  Buffaloberry  1+ft.  240  1.00  13  670  .60  12  .70  14  .05  1  Rose  —  Serviceberry  —  1660  Silverberry.  —  30  .  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 i s the former delta of the large stream that runs o f f Mount Roche Benhomme, through the range to the Athabaska r i v e r .  The area  i s a f l a t , triangular outwash p l a i n sloping gently toward the r i v e r , 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 c h a r a c t e r i s t i c aspen-poplar parkland association, very prevalent i n the Athabaska v a l l e y .  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 o f shrub species, main-  l y buffaloberry and s i l v e r b e r r y .  The survey of tree and shrub growth on  June 13, 1946, gave the r e s u l t s : shown i n Table XIV. The counts i n Table XIV r e v e a l 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 c l a s s , or l e s s , 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 f o r reproduction.  The same s i t u a t i o n exists with poplar,  although to a lesser degree. With regard t o the shrubs, a l l s i l v e r b e r r y are 1 foot, or l e s s , i n height,-the majority o f the plants being mature plants browsed down to that height.  The s i t u a t i o n i s the same f o r buffaloberry. The great  loss i n foliage that t h i s r e f l e c t s i s demonstrated by comparison of the density of buffaloberry on t h i s range with that on Range No. 2,  On the  TABLE XIV  SPECIES  HEIGHT  NUMBERS  DENSITY  PERCENTAGE OCCURRENCE  .65  13  Trees Aspen  -4 f t .  240  Aspen  8fft.  500  Douglas f i r  —  35  60  1.40  2  .35  7  ~  2  Poplar  -4 f t .  490  Poplar  8+ft.  70  Spruce  -8 f t .  70  Spruce  8tft.  20  2.75 —  -* —  Shrubs Buffaloberry  -1 f t .  Willow Silverberry  -1 f t .  500  1.45  29  40  .55  4  250  .50  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  -  23-  l a t t e r 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 t h i s range i s c h a r a c t e r i s t i c of t h i s species on a l l other ranges, except Range No. 2 . practically  As a highly preferred browse species i t has been  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 t r e e growth i n various sections. Such grasses as Richardson's needlegrass are c h a r a c t e r i s t i c 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. 1 0 . Cemetery This i s a very small area (less than an acre) which has been protected from grazing f o r at least twenty years. s o i l are s i m i l a r to that of Range No. 7.  I t s topography  and  The survey on August 28,1947  revealed heavy ground vegetation with a wide v a r i e t y of midgrasses. ( Table XVI.) 11.  Comparative Animal U t i l i z a t i o n of the Ranges as demonstrated by Defecation Counts.  An i n d i c a t i o n of the r e l a t i v e 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 Purple vetch  2.3 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  Comparative Animal U t i l i z a t i o n of the Ranges as demonstrated by Defecation Counts,  Range Numbers  1  2  208  Elk  3  3a  104 156 102  4  5  6  7  8  24  199  38  213  161  —  10  —  —  —  23  42  —  4  Mule Deer  7  22  —  —  Horse  1  60  21  25  —  21  34  —  — 140  79  —  155  Bighorn sheep  6i  TOTALS Elk 1196  Mule Deer 39  Horse 207  Bighorn Sheep 439  - 24 species having the highest defecation count for that area. indicate a considerable  competition  The f i g u r e s  for winter range between sheep and  elk with the horse a competitor p r i n c i p a l l y 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 i n t e r p r e t a t i o n of the above f i g u r e s , i t i s pointed out that sheep, being the smallest animal under consideration, defecate more frequently, with frequency of defecation the elk and the horse following i n that  by  order.  I t i s important to note, also t h a t , despite t h i s , the horse shows up as a most important element i n the grazing to which the Athabaska V a l l e y 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 i s a very important factor determining productivity. B. The differences i n production within exclosure p l o t s , 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 f a c t o r s other than grazing influence productivity.  These other factors are indicated as follows:  1. The difference i n productivity within plots on Range No.  exclosure  1 and No. 2 (134 and 105 grams produced per sq. yd.  respectively) appears to be dependent on a very l o c a l factor such as s o i l .  TABLE XVIII Productivity of the Ranges Area clipped (sq. yds)  A. Range No. 1  1946  10  1947  Total wt. i n grams  1946 725  1947  Mean productivity to nearest gr.per sq.yd  1946  1947  134  190  73  A. Range No. 1 exclosure plot B. Range No. 2 Range No. 2 exclosure plot C. Range No. 3  4  3  10  535  571  530  93  2 7  3  210 240  146  105 34  49  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  5  H5  23  J. Cemetery-outside 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. respecti v e l y ) may be due t o very l o c a l f a c t o r s , as i n l above, r  or to the higher  r a i n f a l l i n the more easterly area, or to a combination of these f a c t o r s .  3. A comparison of the difference  i n productivity (1946)  inside exclosure plots to that outside reveals that the differences become smaller according t o the proximity of the ranges to the Eastern boundary of the Park.  Thus, productivity figures f o r the cemetery show an approx-  imate r a t i o of one outside to four i n s i d e , while progressing  eastward to  Range No. 3 there i s found a r a t i o of somewhat l e s s than three inside to one outside, and on Range No. 1 and No. 2 the r a t i o i s about two inside to one outside. It w i l l be shown i n the discussion of b i o t i c factors that elk have only recently arrived on the eastern ranges.  Correlating t h i s f a c t with  the d i f f e r e n t r a t i o s of productivity inside and outside the exclosure p l o t s , i t becomes apparent t hat productivity of the ranges i s influenced by the length of time that grazing by elk has been c a r r i e d on. 4. The differences i n productivity within exclosure plots i n 1946 and 1947 demonstrate etation from year to year.  the v a r i a b i l i t y i n amount of ground vegThis 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 f a r thest from Jasper (Range No. l ) , i t i s c l e a r l y demonstrated that a v a i l able moisture i s the basic factor determining productivity (see F i g . 15).  -  26  -  The figures i n Table XVIII should be considered i n the l i g h t 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., i s p r i n c i p a l l y 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. F i g . 1 indicates that the ranges studied are either f l a t s or steeply r i s i n g h i l l s . The foregoing descriptions of the areas show that a l l the f l a t s 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. I t i s only on nearly level ground or gentle slopes that considerable depths of s o i l accumulate, and undergo characteristic development 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 d r i f t i n g of dust from the Athabaska plain and blowouts. Range No. 4 i s 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 importance 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 described 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 s i t u a t i o n would account f o r f l u c t u a t i o n s i n productive  capacity that were independent of other nearby areas. 2. Edaphic The e f f e c t 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 t h i s study, organic natter w i l l also be discussed  separately. a. Chemical The samples indicate that the lime content of these s o i l s i s  s u f f i c i e n t l y high to have a profound e f f e c t upon the plant growth of the ranges.  Certain species o f plants cannot survive on highly a l k a l i n e s o i l s ,  and thus many species are r e s t r i c t e d or prohibited from growing i n t h i s area. growth.  The high a l k a l i n i t y of Range No. 4 undoubtedly i n h i b i t s much plant 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 i n d -  i c a t i o n of optimum reaction conditions, the pH range of c e r t a i n plants studied i s given i n Table XIX. A l k a l i n i t y suppresses the s o l u b i l i t y of some elements r e quired by plants.  The i n s o l u b i l i t y of phosphates i n a s o i l containing  calcium carbonate i s an example of t h i s condition.  E l l i s i n discussing  high lime s o i l s of Manitoba states (15) "In the high lime s o i l s , phosphorus i s either low or i n a not r e a d i l y a v a i l a b l e form with the r e s u l t that plants do not t h r i v e as well as on the s o i l s where phosphorus i s available."  I t i s , therefore, not surprising to f i n d 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 e f f e c t upon  the a b i l i t y o f a plant to withstand adverse conditions such as over-  TABLE X I X OPTIMUM pH Range o f 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  Flax  5.0-7.0  Juniper  5.0-6.0  Service berry  5.0-7.0  Poplar  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 i s 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 i s 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 reproductive 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 b e n e f i c i a l s o i l organisms i s approximately neutral,  while temperature and moisture should be high.  The opposite  conditions  of moisture p r e v a i l on these ranges and r e a c t i o n i s above the optimum. In view of the above facts and the r e s u l t s of the t e s t s , which reveal comparatively  small amounts of n i t r a t e s , i t can be  assumed that the nitrogen content i n t h i s area i s low, b. Physical A study of the texture of these s o i l s 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 c e r t a i n of the ranges. Texture i s important by reason of the d i f f e r e n t i a l water holding capacities of the various t e x t u r a l classes.  The water  retention capacity of a s o i l composed of large p a r t i c l e s (such as sand) i s much l e s s than that of a small p a r t i c l e d s o i l (such as c l a y ) . demonstrates t h i s i n the following figures  Ellis  (15).  Inches of water per foot available to plants that the various s o i l classes w i l l r e t a i n Sands Sandy loams  1"  Fine sandy loams  ll"!  Loams  2"  Clay loams  3"  Clays  3k"  3/4"  With the exception of Range No. 1 a l l the ranges exhibit a coarse texture i n at l e a s t 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 o f the area studied.  In  states.(15) that " i t i s obvious  discussing similar s o i l s i n Manitoba E l l i s  that the l i g h t e r textured s o i l s are submarginal s o i l s i n a region of l i g h t precipation." The chemical a c t i v i t y of a s o i l i s greatly affected by the s i z e 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 p a r t i c l e s are composed of CaCO^ or some other appreciably soluble compound.  Clay  p a r t i c l e s on the other hand are synthesized i n the s o i l and are very active chemically as demonstrated below.  As the s o i l s studied are c*om-  posed f o r 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 p a r t icles. Comparative Nutrient Content of S o i l Separates from 5 s o i l s derived from limestones and shales (23).  2°5  CaO  P  MgO  K0 2  sand  silt  clay  sand  silt  clay  sand  silt  clay  sand  silt  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 t e s t f o r organic matter i n the s o i l s studied reveals great v a r i a t i o n i n the percentages o f the various ranges, (Table I I ) . As the source of s o i l organic natter i s almost e n t i r e l y plant growth i t can be asserted that those s o i l s with high  percentages have supported  more plant growth i n the past than those with low percentages, provided that conditions f o r 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 precludes 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 i t 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 a l -  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 f o r the high percentages on some areas despite the comparatively  l i m i t e d current plant  production.  Because the a c t i v i t y provoked by organic matter decomposition i s more important than i t s presence i n the s o i l , the varying amounts of organic matter are not necessarily i n d i c a t i v e of the n u t r i t i v e value of the various s o i l s sampled. As favorable s o i l conditions f o r 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 s o i l s sampled are unfavorable to heavy plant growth due p r i m a r i l y 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 matter of Ranges No.'s  2, 3  f  organic  3A U, and 6 are due to the poor water r e -  tention capacities of these s o i l s or to the topography of the ranges while the low productivity of Ranges No.'s  5, 7 and 8 indicates some  recently introduced f a c t o r . From the edaphic viewpoint, Range No. 1 i s far superior i n productivity to the other ranges because of i t s superior moisture holding q u a l i t i e s . 3. Climatic Climate i s the most important physical feature determining vegetation, through i t s action i n forming s o i l s and through i t s direct effect upon the plants.  The r e l a t i v e rates of a c t i v i t y of the  different forces of weathering and s o i l formation are determined by climate, which also l i m i t s the extent to which s o i l development proceeds. In a r i d 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 precipitation. 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 10-15 Over 30  200 or more 70-200 3- 12  As the annual precipitation at Jasper Park falls in the 10-15 inches class (Table I) i t 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 valley 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.  F i g . 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 o f f 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 i s a very important climatic factor with respect to both plants and animals. As mentioned above the growing season of plants i s determined by temperature, which, i n Jasper Park, l i m i t s i t to a very few months. Vegetation i s also restricted by the extremes of heat and cold which are encountered i n t h i s area. There are many plants which cannot stand the excessive cold, particularly the sudden drops i n temperature 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 physiol o g i c a l 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 e f f e c t 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 c e r t a i n of  the ranges studied.  The importance of wind i n removing snow from sheep  ranges has. already been alluded to.  Erosion by wind following disappear-  ance of the plant cover due to overgrazing i s another influence having drastic effects on c e r t a i n areas.:('FiglO)Death of these plants combined with trampling of the sod has destroyed the root systems and annual l e a f carry-over that formerly held the s o i l i n t a c t .  As a r e s u l t 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 i s blown about the v a l l e y or  into the Athabaska r i v e r .  Ranges No. 3A and No. 6 are t y p i c a l of these  areas, and their low productivity may be a t t r i b u t e d p r i m a r i l y to wind and i t s e f f e c t on the s o i l . S o i l retrogression, as a r e s u l t of wind erosion i s , therefore, taking place and could eventually reach the point where vegetation w i l l remain s t a t i c ( i n an unproductive  state) for many years, due to a l t e r e d  edaphic f a c t o r s , even though grazing may 4. ^  have ceased e n t i r e l y .  Biotic  a. Faunal The ungulates of the Athabaska v a l l e y are the most  important  faunal factor a f f e c t i n g the ranges.  large s i z e , great numbers, and feeding habits.  This i s because of t h e i r In general, large numbers  of any ungulate have c e r t a i n effects which become increasingly damaging  to the ranges as the numbers of grazing animals increase.  Braun-Blanquet  summarizes the i n j u r i e s r e s u l t i n g from pasturing as follows: 1.  (4)  Withdrawal of large quantities of plant matter.  2 . Mechanical harm to plants by trampling, grazing,  and  brushing against them. 3 . S e l e c t i v e 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 m i c r o - r e l i e f .  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.  I t 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 i n f l u e n t on the Jasper Park ranges, i t has the greatest effect upon the ecology of the area.  I t has been shown  i n the h i s t o r i c a l section of t h i s study that the present numbers of elk r  are atvery recent phenomenon. i n Cowan's report of 1943  (8).  The extent of the increase i s demonstrated He states that i n 1931  only two elk were  seen i n Jasper Park, while i n 1943> three hundred and forty-seven were observed.  In 1944  (19),  he estimated that 250-280 elk were ranging i n  the Henry House area alone.  A further i n d i c a t i o n of the extent of elk  reproductive success are the figures supplied by Warden F. A. Bryant from his journal.  They are summarized below:YEAR 1927  AVERAGE SEEN PER DAY DURING MONTH OF JANUARY 0  - 38 -  1930  3. 4  1931  3. 3  1933  3. 3  1937  18. 3  1938  11. 3  A t o t a l figure f o r the ranges under study i s imp»essible to e s t imate accurately with existing information.  However, some idea of the  large numbers i n the Athabaska v a l l e y may be obtained by c i t i n g figures of surplus elk destroyed by Park a u t h o r i t i e s .  P r a c t i c a l l y a n 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 Athabaska v a l l e y i n much the same numbers as i n previous years".  This report  also cites a c a l f s u r v i v a l of 11 per cent f o r the year 1946.  Considering  such a low reproductive rate i n the l i g h t of the more or l e s s constant and at times increasing k i l l figures, i t may be concluded that the numbers of adult elk ranging the Athabaska v a l l e y must be at l e a s t 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 i n the past been centered around the l i b e r a t i o n area within a few miles of Jasper townsite. have they progressed eastward.  In 1944,  Only recently  Cowan stated (9) that "The  occupation of the Moosehorn r i v e r has lagged behind that of the Snake Indian r i v e r , but elk of both sexes are now present i n considerable numbers and i t can be expected that t h i s v a l l e y w i l l shortly see a heavy increase i n population with resultant heavier pressure on the already over-grazed Miette ranges". The recent a r r i v a l 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 s i z e 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 l i m i t e d 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, buffaloberx'y, and s i l v e r b e r r y that has been described. The forage consumption equivalents of the animals under study and some l i v e s t o c k 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 e x i s t i n g numbers of elk are becoming less able to support themselves i n a healthy condition on the ranges.  Cowan (10) attributes a part of the low c a l f s u r v i v a l already  mentioned to poor physical condition o f the mother r e s u l t i n g i n abortion, resorption of the embryo, poor c a l f s u r v i v a l , etc.  He also states that,  i n early May, 1946 the average n u t r i t i o n a l condition of elk was poor. Without t h e f a c t o r of human k i l l i n g of surplus animals, undoubtedly the lack o f 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 t h i s  uation and can prevent i t .  sit-  Another f a c t o r o f no l i t t l e importance i n  checking the numbers o f elk i n the Athabaska v a l l e y i s the considerable number of timber wolves ranging t h i s area.  They are of great value i n  k i l l i n g surplus elk i n areas where human control i s impractical and costly. On the basis of studies made i n the winter of 1946-47, o f wolves at Jasper and Banff Parks, i t may be stated that a large part of t h e i r winter prey is elk.  A pack o f four or f i v e 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 v a l l e y .  As indicated by Table XVII, t h i s animal competes  mainly with the elk for forage as range requirements  are much the same.  - 41 According t o Stoddart and Smith ( 3 l ) , "Horses are the most s e l e c t i v e of the domestic animals.  They are p r i m a r i l y grass eaters and u t i l i z e r e l a t -  i v e l y 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 v a l l e y 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 i n 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 f i g u r e s , the Superintendent of the Park writes that twenty of the above annual numbers were cows u n t i l 1 9 4 6 - 4 7 , 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 w i l d horses running i n the  Park and those not declared.  The eighty-five government horses are not  included i n the above f i g u r e s , 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 i s apparent that those horses are maintaining themselves at the expense of the game population.  I t i s a l s o 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 i n numbers to the e l k .  In 1943,  Cowan estimated  the sheep population as follows ( 8 ) : Devona Jasper benches  350 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 i n "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 f o r the range forage on several areas.  In any long  run competition of this type, i t i s the author's opinion that the elk would become dominant at the expense of the sheep, due to the l a t t e r  - 43 species' more specialized environmental requirements. icates that a s c a r c i t y of lambs i n 1943  Cowan (8) i n d -  nay have been due to n u t r i t i o n a l  deficiencies of ewes r e s u l t i n g 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 d i r e c t effect upon s o i l development.  By changing the m i c r o - r e l i e f 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 l i g h t of the above facts i t must be pointed out that b i g horn sheep are one of Jasper Park's greatest natural attractions from an aesthetic point of view.  I t i s a species which has decreased greatly  throughout much of i t s range i n 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 i s f e a s i b l e . According to h i s t o r i c a l records, the Athabaska Valley was  orig-  i n a l l y w e l l populated with sheep, and i t would seem to be a mistake to permit elk to drive out any of the sheep f l o c k s . Mule deer have always been one of the chief ungulate forms of the Athabaska Valley.  At present they are found i n considerable numbers,  p a r t i c u l a r l y i n winter and spring, upon some of the areas studied.  In a  13-day examination of the Athabaska V a l l e y i n 1943,  168  deer, while Cowan i n 1943  Clarke (5) saw  (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 p o t e n t i a l  - 44 ~ deer forage, as t h i s species i s p r i m a r i l y a browser.  The following  figures show the food of the mule deer i n Jasper Park as determined by Cowan (9). Grass and sedge,  15%  Pussytoes  2%  Pasture sage.......  tr.  Forbs Bearberry Buffaloberry Other browse.  4% 58% 9% .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 a v a i l a b l e 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 i n d i c a t e , the i n t e r r e l a t i o n ships of the above factors plus other environmental influences have produced a varied plant growth i n t h i s area.  I t i s the purpose of t h i s  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 e f f e c t s , i f any, of the plants on the above f a c t o r s . 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 because they are highly preferred food plants of sheep and elk.  As an a i d  i n evaluating the p a l a t a b i l i t y of the grasses of these ranges as compared  - 45  -  to other species of ground vegetation the following p a l a t a b i l i t y ratings are given, as they concern sheep and c a t t l e , animals with feeding habits somewhat s i m i l a r to mountain sheep and elk r e s p e c t i v e l y . (27).  In these  cases p a l a t a b i l i t y 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.  .  . Cattle 80$  Sheep 60$  Junegrass  60$  50$  Needlegrass  60$  40$  Thickspike wheatgrass  Pussytoes  0  Pasture sage  10$  0 20$  Following heavy grazing l e s s palatable species w i l l invade areas denuded of the preferred forms.  the  As retrogression proceeds, with a  lessening of forage, the l e s s palatable species w i l l also be consumed. Because animals under s u f f i c i e n t 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 s i t u a t i o n have  already been considered. From the data a v a i l a b l e , 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 l e s s 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 1.  Range Number % of ground cover  2  7  31.0  45.0  25.0  8.0  65.0  25.0  32.0  22.0  fo of pussytoes and pasture sage of t o t a l ground cover  % of grasses of t o t a l ground cover 3 2 . 0  The r e l a t i v e amount o f 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 i n f e r i o r s o i l .  On Range No. 7 there  i s a g r e a t l y reduced amount o f t o t a l ground cover, as w e l l as reduced amounts o f both overgrazing indicators (pussytoes and pasture sage) and grasses.  Further evidence o f t h i s trend toward reduction o f t o t a l ground  vegetation i s given i n Table XX which compares point sample counts o f 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 ( 3 1 ) , 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 p r a c t i c a l l y c e r t a i n that t h i s species i s invading areas where grass has died.  From the above f i g u r e s ,  i t can be seen that due to the lack o f grass, pasture sage i s also being grazed, despite i t s u n p a l a t a b i l i t y , 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 i n 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 Pasture sage...... Other plants  3.6% 12.1 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 cemetery 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 sedges 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) demonstrated this in the case of Agropyron inerme. a close relative of one of the chief forage grasses of Jasper Park. A count showing the reproductive 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.  - 48 -  Agropyron inerme Condition Grazed Protected  No. of heads per sq. meter  Filled Florets per sq. meter  No. of viable seeds per sq. m.  7.1  19.6  12.2  120.4  972.6  630.2  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  56l  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 maintaining 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 for example, where certain factors 3  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 i n 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 first.  Although i t was noted i n the ecological history that willows  were very evident some years ago, i t i s now almost impossible to f i n d 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 i s 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 favorable.  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 o f from 3 to 6 feet ( F i g . 5) while s i l v e r b e r r y grows to 6-8 feet i n height ( F i g . 6 ) . However, the i n d i v i d u a l range area  N  surveys show the average heights of these plants to be greatly heneath the normal. Under e x i s t i n g circumstances, the plants are undoubtedly prematurely, and the lack of reproduction precludes t h e i r 6.  dying  replacement.  Trees.  From the standpoint of the mammal population i n the Jasper area, the aspen i s the most important tree.  I t i s a highly preferred food of  elk and i s also one of the p r i n c i p a l 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 t h i s species to be i n danger o f  complete removal from those areas, where reproduction has ceased. uation i s indicated by the age o f the trees surveyed.  This s i t -  They are either mature  or the current year's seedlings, with no intervening range o f 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 t y p i c a l over-browsed areas, such as Range No, 5 , was proven by r i n g counts on an average-sized aspen.  This tree was found  to be about f i f t e e n years o l d - a f i g u r e that indicates the point at which successful reproduction ceased.  This corresponds with the a r r i v a l of elk  i n numbers upon these ranges. The present pressure upon aspen i s greatly accelerating the tempo of forest succession. forests.  Heavy aspen growth i s c h a r a c t e r i s t i c of sub-climax  According t o Harlow and Harrar ( 1 8 ) , conifers, such as Douglas  f i r , lodgepole pine, and white f i r , replace aspen i n the Rockies.  Inaa-  much as these species are o f no forage value, i t i s important to keep the Jasper ranges i n a sub-climax dondition as long as possible.  As has  been pointed out, Range No. 2 i s r a p i d l y developing into a climax spruce f o r e s t , at the expense o f a l l other browse plants. In a discussion of forest succession and i t s r e l a t i o n to the game animals of Jasper Park, i t i s necessary to draw attention to the effect o f s t r i c t f i r e protection upon the future forage value o f the ranges.  According to Harlow and Harrar (18), burnt-over s i t e s are the  most favorable f o r production of aspen, but a f t e r twenty t o t h i r t y years, much o f the aspen dies due to i t s intolerance of the invading conifers. There i s abundant i n d i c a t i o n that similar circumstances apply i n the Athabaska v a l l e y .  Therefore, s t r i c t f i r e protection, or lack o f other con-  t r o l s , on the conifers, w i l l produce a climax conifer forest o f almost no forage value.  The poplar, although formerly common, according to the  records, i s not frequently found. maintain i t s e l f .  Under e x i s t i n g circumstances, i t cannot  .*.«**.«#. .  The above analysis of factors a f f e c t i n g the ranges shows that there i s a complex i n t e r a c t i o n i n which the r e l a t i v e importance o f the various factors varies from range to range.  The ultimate effect o f this  i n t e r a c t i o n i s measured by the productivity of the ranges as shown i n Table XVIII. The study indicates that over-use by ungulates i s the most ims-portant s i n g l e factor determining the present ground vegetation of the ranges.  This i s c l e a r l y indicated by the reduced y i e l d s on unprotected  areas, and by a syndrome of symptoms well-known as indicators o f range misuse.  These symptoms axe: 1. Reduction i n reproduction of grasses. 2. Reduced occurrence o f midgrasses with increased occurrence of short grasses. 3. Increase i n pasture sage. 4. Decrease i n t o t a l 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 r e s u l t that only the more l e v e l 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 l i m i t i n g f a c t o r as i t has c u r t a i l e d reproduction of palatable species (aspen, willow, buffaloberry, and silverberry) on c e r t a i n areas, and as i t i s k i l l i n g the adult plants of s i l v e r b e r r y , buffaloberry,and willow. Thus, on Range No. 2, where elk are a recent i n f l u e n t , the s o i l i s of poor texture compared to Range No. 5, but t h i s 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 o f browse species increases according to the duration of occupation by elk, i t i s apparent that t h i s animal i s responsible f o r the present destruction. The mule deer (which i s primarily a browser) has been a constant factor i n t h i s area, arid the gradient i n overbrowsing  cannot be a t t r i b u t e d 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 producti v i t y table reveals differences i n production i n the protected areas. Increased p r e c i p i t a t i o n causes greater general productivity on  the ranges to the east than on the areas near Jasper.  However, very  l o c a l differences i n production and ground vegetation composition, such as occur between Range No. 1 and No. 2, must be a t t r i b u t e d 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 i n h i b i t s plant growth to a high degree) on a l l ranges, moisture holding capacity, r e s u l t i n g from texture and organic matter, i s the l i m i t i n g 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 t h i s 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 overbrowsed 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 c e r t a i n ranges to the east.  Thus, ranges,such as Range No, l j a r e capable of supporting  many more elk at the present time than areas such as Range No, The ungulate studied i s the elk.  7.  c h i e f l y responsible for the misuse of the area This animal has increased greatly i n a very short  time on ranges that had been previously overgrazed as horse range. i s competing f o r winter food supplies with bighorn sheep on,certain  It  - 54 ranges(No. 3A and No. 6) that are capable o f supporting only a very l i m i t e d amount of plant growth. Unless present trends i n the ecology of the area are reversed i n the very near future, the health/conditions^ of the ungulate J  herds  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 i n d i v i d u a l s on ranges with r a p i d l y decreasing amounts of forage. Under existing conditions, a winter of deep snow w i l l almost c e r t a i n l y r e s u l t 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 c e r t a i n areas a l l plant cover has died, and the s o i l i s  being dissipated by wind.  This s i t u a t i o n w i l l r a p i d l y become widespread  unless the ground vegetation i s permitted to recover i t s o r i g i n a l character.  Destruction of vegetation also tends to slow down'or prevent the  formation of a mature s o i l . Plant succession i s i n a state of retrogression on the area studied.  This i s p a r t i c u l a r l y true of the ground vegetation, where  production of palatable species has been g r e a t l y decreased.  Invading,  less palatable^forms are also showing signs of retrogression, r e s u l t i n g i n very large percentages of bare ground. The p r i n c i p a l browse shrubs, s i l v e r b e r r y and buffaloberry are suffering great p h y s i o l o g i c a l and morphological damage from over-use of twigs and f o l i a g e .  This damage i s d r a s t i c a l l y c u r t a i l i n g reproduc>-  t i o n , as also i s the consumption of seedling plants.  Willow has been  p r a c t i c a l l y exterminated on the areas nearest Jasper.  - 55 The tree associations of the Athabaska v a l l e y are changing, due to the t o t a l 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 f o r 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 s i t u a t i o n i n the •Athabaska V a l l e y , and to maintain a s a t i s f a c t o r y balance. (1)  Continued reduction o f the elk to a point where the ranges  w i l l show a substantial increase i n grasses, p a r t i c u l a r l y i n the t a l l e r midgrasses, with a corresponding decrease i n the over-grazing i n d i c a t o r s , and u n t i l aspen and willow are able to reproduce s u f f i c i e n t l y to maintain or increase present numbers.  This reduction should be concentrated near  Jasper and on the sheep ranges. (2) Annual census of e l k , sheep, and deer, to determine population trends rather than absolute numbers. by Hunter (19),  would be applicable.  Methods such as those used  In addition, the annual elk slaugh-  ter by Park authorities lends i t s e l f p a r t i c u l a r l y w e l l to the use of a technique described by Olsen (25)  i n which c a l f elk would be tagged and  population trends calculated on the basis of a r a t i o between elk k i l l e d and t o t a l population.  Also p e l l e t group quadrats, as described by  Bennet, English, and;, McCain (3)  f o r deer studies could be established.  (3) The making of a cover-map of the p r i n c i p a l winter ranges of the Park by a w i l d l i f e technician, as described by Dalke  (12).  (4) A study of r e l i c t areas to determine o r i g i n a l climax  - 56 vegetation. (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  establishment  o f a d d i t i o n a l ones to determine t r e n d s i n p r o t e c t e d and u n p r o t e c t e d (6) P e r i o d i c s t u d i e s , with a view to  similar  to these,  establishing carrying capacity  of  the range  conditions  figures.  (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 the Athabaska  of  valley.'  (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 w i n t e r range 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 such  (9) I n i t i a t i o n o f a k e y a r e a ,  key p l a n t  and t h e a c t u a l use f a c t o r  on t h e r a n g e s .  c e r t a i n areas on r e p r e s e n t a t i v e  the degree of u t i l i z a t i o n o f the  game  range.  s t u d y as a n i n d i r e c t  method 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 between t h e p r o p e r use  selecting  areas.  factor  T h i s method w o u l d c o n s i s t winter ranges,  essential forage  plants  of  and a p p r a i s i n g present.  PLATE I  > '  Fig.  ^  7  j  "To  :he Area studied showing i n d i v i d u a l 1. Map of the Range Areas  F i g . 2.  The Athabaska V a l l e y showing area studied.  a  Section of the  PLATE III.  Fig. 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 No. 2.* Photograph by Dr. I . McT. Cowan.  Range  PLATE VI  Fig.9  Range No. 3A showing terracing by grazing animals. Photograph by Dr. I. McT. Cowan.  PLATE V I I  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 N o . 7 showing much dead m a t e r i a l . Photograph by D r . I* McT. Cowan.  PLATE IX  Fig.  14  Silverberry on Range No. 7 showing much dead material. Photograph by Dr. I. McT. Cowan.  PIATE X  F i g . 15.  Correlation of measured Factors of Ranges  No. 1 and No. 7 with the Productivity of those Ranges. (The figures f o r animal u t i l i z a t i o n were obtained by d i v i d i n g the t o t a l defecation counts f o r each range by five).  Prec i p i t a t ion(inches) 1111111111111111 m ~ | T m  4 j 11111  PLATE XI  F i g . 16.  Average monthly P r e c i p i t a t i o n  and minimum Temperatures at Jasper Park (22 years).  LITERATURE CITED 1. Aldous, Shaler E. , 1944. A deer browse survey method. Journ. MammalogjrVol.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 populations by use of pellet group counts. Journ. Wildlife Manag. Vol. IV, No.4,pp. 398-403.  j  4. Braun-Blanquet, J., 1932. Plant Sociology. McGraw-Hill Book Co., Inc. New York and London, pp. x v i i i 439* illustr. 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 Mountain 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. 16.  The S o i l s of Manitoba. Manitoba Economic Survey Board, Winnipeg. 112 pp. i l l u s t r .  Geology o f the National Parks o f Canada, 1940. National Parks Bureau, Dept. of Mines and Resources, Ottawa, 20 pp. illustr.  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. McGrawH i l l Book Co., New York and London, pp. xv 542 illustr.  19. Hunter, G. N., 1945* Methods of determining trends i n b i g game numbers and range conditions. Trans. 10th N. Amer. W i l d l i f e 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. 22.  23.  Manitoba and the Great Northwest. World Publishing Co., Guelph. pp. x x i i 687, f r o n t , i l l u s t r . maps. 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.  M i l l a r , C. E., Turk, L. M., 1943. Fundamentals of S o i l Science. John Wiley & Sons, Inc. New York. pp. x i 462 illustr.  24. Moberly, H. J . , 1929. When Fur was King. J . M. Dent & Sons, London & Toronto, pp. x v i i 25.  Olson, H. F., 1943.  237.  Deer tagging and population studies i n Minnesota... 3rd. N. Amer. W i l d l i f e Conf. pp. 280-286.  26. Preble, E. A., 1908. A B i o l o g i c a l Investigation of the Athabaska and Mackenzie Region. U. S. Dept. A g r i c , Bur. Geol. Survey Gov't P r i n t i n g O f f i c e , 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. P r i n t i n g O f f i c e , Washington, D. C. 28. Schollenberger, C. J . , 1945. Determination of organic matter. S o i l Science. V o l . 59, No. 1 pp. 53-56. 29.  Spurway, C. H. 1941.  S o i l reaction (pH) preference of plants,  Special B u l l . 306 Mich. Agric. Exp't. Stn., East Lansing, Mich. 30. Spurway, C. H. 1944.vSoil Testing - a p r a c t i c a l system of s o i l f e r t i l i t y diagnosis. Tech. B u l l . 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 illustr. 32. Talbot, M. W,,  1937. Indicators of southwestern range conditions. Farmers B u l l . 1782, U. S. Dept. of Agric. U. S. Gov't P r i n t i n g O f f i c e , 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 o f 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. B u l l . 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  S c i e n t i f i c names of Mammals r e f e r r e d to i n the Text. Elk  Cervus canadensis  Moose  Alees americanus  Mule deer  Odocoileus hemionus  Bighorn sheep  Ovis canadensis  Timber wolf  Canis lupus  Mountain Goat  Oreamnus montanus  Beaver  Castor canadensis  APPENDIX B S c i e n t i f i c Names of Plants referred to i n the Text Common Name Aspen  S c i e n t i f i c Name Populus tremuloides Michx,  Aster  Aster sp.  -Alpine  Aster alpinus L.  Aven  -. Geum t r i f l a r u m Pursh (G)  Bastard toadflax  Comandra sp.  Bearberry  Arctostaphylus uva-ursi Spring  Bedstraw, northern  Galium boreale L.  Black-eyed Susan  G a i l l a r d i a , sp.  Bluebell  Companula r o t u n d i f o l i a 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  P o t e n t i l l a pennsylvanica var. s t r i g o s a Pursch. Dasiphora f r u t i c o s a L  -shrubby Cranberry, high bush  Viburnum opulus L. var. americanum  Daisy  Erigeron coespitosus Nutt  -purple  Erigeron l i n e a r i s (Hook) Piper  -white  Erigeron compositus  Fescue  Festuca sp.  F i r , Douglas  Pseudotsuga  Pursch.  t a x i f o l i a (Poir)  APPENDIX B  (cont'd)  Flax  Linum usitatissimum L.  Goldenrod No. 1  Solidago l e p i d a D. C.  Goldenrod No. 2  Solidago u l i g i n o s a Nutt.  Gooseberry  Ribes sp.  Grass -Sandburg blue  Boa secunda  -Blue  Poa sp.  -Brome No. 1  Bromus suksdorfii?  -Brome No. 2  Bromus pumpellianus Scribn.  -June  Koeleria c r i s t a t a (L.Pers.)  -Needle  Stipa comata T r i n . & Rupr.  -Richardson's  Needle  S t i p a richardsoni Link. .  -Pine  Poa subpurpurea Rydb.  -Purple reed  Calamogrostis  -Bearded wheat  Agropyron subsecundum (Link)  -Thickspike wheat  Agropyron dasystachyum (Hook & S c r i b . )  -Wild-rye  Elymus innovatus Beal  Juniper  purpurascens R. Br.  Juniperus scopolorum Sarg.  -Common  Juniperus communis L.  Legume  Leguminosae  Lily  L i l i u m montanum A. Nels.  Pine, lodgepole  Pinus contorta var. Murayana Engelm. Populus tacamahaca M i l l .  Poplar  APPENDIX B (cont'd) Puccoon  Lithosperum angustifolium Michx.  Pussytoes  Antennaria p a r v i f o l i a Nutt.  -Canada  Antennaria canadensis L.  Rose  Rosa Macounii Greene.  Sage, pasture  Artemisisa f r i g i d a W i l l d .  -Field  Artemisisa camporum Rydb.  Sedge  Car ex sp.  Serviceberry  Amelanchier sp.  ; ,Silverberry  Eleagnus argentea Pursh.  Spruce, Black  Picea marianna ( M i l l . )  Spruce, white  Picea glauca (Moench.) Voss  Strawberry  Fragaria glauca (S. Wats) Rydb.  T h i s t l e , Russian  Salsola k a l i var., t e n u i f o l i a Meyer  Vetch  Astragulus sp.  -Milk  Astragulus alpinus L.  -Purple  Astragulus sp.  Violet  V i o l a sp.  Willow  S a l i x sp.  Wind Flower  Anemone sp.  


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