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The impact of an introduced population of elk upon the biota of Banff National Park Mair, William Winston 1952

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THE IMPACT OF AN INTRODUCED POPULATION OF ELK UPON THE BIOTA OF BANFF NATIONAL PARK by WILLIAM WINSTON MAIR A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS i n the Department of ZOOLOGY We accept t h i s t h e s i s as conforming to the standard r e q u i r e d from candidates f o r the degree of MASTER OF ARTS. Members of the Department Zoology THE UNIVERSITY OF BRITISH COLUMBIA March, 1952 ABSTRACT In 1949, a study was c a r r i e d out i n B a n f f N a t i o n a l Park to determine the r e l a t i o n s h i p e x i s t i n g between Introduced e l k (Cervus canadensis n e l s o n l ) and the other b i o t a o f the r e g i o n ; p a r t i c u l a r emphsis was p l a c e d upon p o s s i b l e c o m p e t i t i o n w i t h the indigenous moose ( A l c e s americana americana). Subsequently, the h i s t o r i c r e l a t i o n s h i p s of the major f l o r a l and f a u n a l s p e c i e s were f u r t h e r s t u d i e d . E l k were introduced i n t o the Park In I 9 I 7 and 1 9 1 9 - 2 0 . They f l o u r i s h e d and became so numerous t h a t i n 1 9 ^ 3 t h e i r c o n t r o l by n o n - s e l e c t i v e mechanical removal became necessary. T h i s con-t r o l has been continued t o date but the c o n d i t i o n of the main ranges i s s t i l l u n s a t i s f a c t o r y . Most e l k w i n t e r ranges are h e a v i l y over-browsed and over-grazed and summer ranges are now b e g i n n i n g to s u f f e r . Unorganized p r e d a t o r c o n t r o l has p o s s i b l y worsened the s i t u a t i o n . Moose f i r s t entered the ar e a ( i n recent times, at l e a s t ) i n 1 9 1 6 and pro b a b l y i n c r e a s e d s t e a d i l y to the l a t e 1 9 3 0 ' s . They then a p p a r e n t l y d e c l i n e d somewhat to re a c h a s t a b l e maximum by about 1 9 ^ 3 . T h e i r r e p r o d u c t i v e r a t e appears to be e x c e l l e n t . They show no p r e s e n t apparent d e t r i m e n t a l e f f e c t a r i s i n g from the e l k p o p u l a t i o n , a l t h o u g h the l a t t e r e x e r t s a steady p r e s s u r e by encroachment upon the d r i e r p o r t i o n s o f the moose range 1; and by almost complete removal of aspen, w i l l o w and dwarf b i r c h r e p r o d u c t i o n i n the Bow V a l l e y and some adjacent a r e a s . They w i l l probably e v e n t u a l l y a f f e c t the beaver-moose complex t o the f i n a l detriment of these s p e c i e s . 2 TV Mule deer (Odocolleua hemlonus hemlnonus) a r e indigenous to the Park, summering throughout the a r e a but w i n t e r i n g , i n the main, outside i t s c o n f i n e s . D e s t r u c t i v e use, by e l k , of much of the main Park w i n t e r range, below the 7 mile Beaver Dam i n the Bow V a l l e y , has depressed the r e s i d e n t d e e r p o p u l a t i o n to a near s t a t i c low. Herd Increment appears t o be low compared to ot h e r mountain r e g i o n s such as i n Utah. Bi g h o r n sheep (Ovls canadensis canadensis) have been i n the Park area s i n c e e a r l i e s t r e c o r d , but have been dependent, f o r much of t h e i r range upon openings c r e a t e d by f i r e s . Thus, Park p o l i c y of f i r e c o n t r o l has brought about containment of s u i t a b l e b i g h o r n sheep areas; these areas have, more r e c e n t l y , been invaded by e l k . B i g h o r n have been numerous, and reached a p o s s i b l e maximum i n the e a r l y l930 ,s. They then r a p i d l y d e c l i n e d , p o s s i b l y due to some e p i z o o t i c . Recovery, i f any, has been slow, pr o b a b l y due to the encroachment of the e l k i n r e c e n t y e a r s . Thus the sheep, s u f f e r i n g from d e b i l i t a t i n g p a r a s i t i s m and range impoverishment, are f a i l i n g i n the c o m p e t i t i o n a g a i n s t the more a g g r e s s i v e and v e r s a t i l e e l k , t h a t a p p a r e n t l y s u f f e r s l e s s from p a r a s i t i s m and o t h e r b i o l o g i c a l l i m i t i n g f a c t o r s . I t i s suggested t h a t few of the components o f the Park b i o t a w i l l e v e n t u a l l y escape the i n f l u e n c e of the e l k . Man, as a member o f the b i o t i c community, by h i s , a c t i o n s and h i s very presence, i n f l u e n c e s t h a t community o f t e n beyond the co n f i n e s of h i s present p e r c e p t i o n . ACKNOWLEDGMENTS I wish to express my thanks and a p p r e c i a t i o n to those who have a i d e d me i n t h i s work. My thanks; To Dr, W. A, Clemens and Dr„ I . McT. Cowan of the Department of Zoology, U n i v e r s i t y o f B r i t i s h Columbia, f o r t h e i r encouragement and a s s i s t a n c e . To the Canadian W i l d l i f e S e r v i c e which made t h i s study p o s s i b l e . To Mr. J . A. Hutchison, who, while Superintendent of Banff N a t i o n a l Park, extended to me every p o s s i b l e a s s i s t -ance and courtesy d u r i n g t h i s s t u d y — a n d a l s o t o : The Park Wardens of Banff N a t i o n a l Park. I t i s not p o s s i b l e to mention by name i n d i v i d u a l s who a s s i s t e d me. A l l were most h e l p f u l and c o n s i d e r a t e . To Mr. H. U. Green, S p e c i a l Warden i n Banff N a t i o n a l Park, who deserves p a r t i c u l a r mention f o r h i s whole-hearted i n t e r e s t and c o - o p e r a t i o n . To my w i f e , who a s s i s t e d me both i n the f i e l d and i n c o m p i l a t i o n and p r e p a r a t i o n of a l l d a t a . TABLE OF CONTENTS INTRODUCTION 1 SCOPE AND METHODS 3 THE PROBLEM AREA H i s t o r y 6 Geology and Geography .» 6 Climate , 10 F l o r a 11 Fauna 13 ELK AND THEIR INTER-RELATIONSHIP WITH PARK BIOTA Movements and Food H a b i t s of E l k 25 V i t a l S t a t i s t i c s of E l k 30 E l k Loss F a c t o r s 33 E l k Ranges 3D Movements and Food H a b i t s of Moose .... 4-5 V i t a l S t a t i s t i c s of Moose....... 53 Moose Loss F a c t o r s 5° Moose Ranges 57 Moose i n R e l a t i o n to E l k o l Movements and Food H a b i t s of Mule Deer 63 V i t a l S t a t i s t i c s of Mule Deer 65 Mule Deer Loss F a c t o r s 6? Mule Deer Ranges 68 Mule Deer In R e l a t i o n to E l k 69 Movements and Food H a b i t s of Bighorn Sheep .... 71 V i t a l S t a t i s t i c s of Bighorn Sheep ..... 73 Bighorn Sheep Loss F a c t o r s 78 Bighorn Sheep Ranges 82 Bighorn Sheep i n R e l a t i o n to E l k 84-SUMMARY DISCUSSION OF DYNAMIC INTER-RELATIONSHIPS OF ELK WITH PARK BIOTA 8b BIBLIOGRAPHY 95 i i LIST OF TABLES Table No. Page I . Sight r e c o r d s of e l k cows and c a l v e s In the Bow V a l l e y , Banff N a t i o n a l Park, i n 1949. .... 3 1 I I . E l k c a l f and y e a r l i n g counts (percentage of t o t a l herd) f o r Banff N a t i o n a l Park. 3 2 I I I . E l k removed by sl a u g h t e r from Banff N a t i o n a l Park. 3 3 IV. Data on square metre quadrats c l i p p e d i n H i l l s d a l e area, Banff N a t i o n a l Park, i n 1 9 ^ 9 . 3 9 V. Pond p l a n t s c o l l e c t e d from some moose summer ranges i n Banff N a t i o n a l Park i n 1 9 4 9 . 4 9 VI. C l a s s i f i c a t i o n , by age and sex, of moose known to be i n - t h e V e r m i l i o n Lakes - 1 7 M i l e Meadow area, Banff N a t i o n a l Park, from May to J u l y , 1 9 4 9 . 55 V I I . Number of counts and aggregate t o t a l s of bi g h o r n sheep seen a t V e r m i l i o n l i c k , g a n f f N a t i o n a l Park, f o r years 1 9 4 4 to 1 9 4 § i n c l u -s i v e . ...... 76 LIST OF MAPS Map 1 . Banff N a t i o n a l P a r k — s h o w i n g salient;-. . g e o g r a p h i c a l d e t a i l s • 5a Map 2. Banff N a t i o n a l P a r k — s h o w i n g main b i g h o r n sheep ranges a c c o r d i n g to Green ( 1 9 4 9 ) . . . . . 2>la THE IMPACT OF AN INTRODUCED POPULATION OF ELK UPON THE BIOTA OF BANFF NATIONAL PARK INTRODUCTION In an annual r e p o r t on the Canadian N a t i o n a l Parks (Canada, 1 9 4 4 : 7 9 ) i t i s s t a t e d t h a t "The Parks are hereby d e d i c a t e d to the people of Canada f o r t h e i r b e n e f i t , education and enjoy-ment, sub j e c t to the P r o v i s i o n s of t h i s Act and R e g u l a t i o n s , and such Parks s h a l l be maintained and made use o f so as*to l e a v e them unimpaired " f o r the enjoyment -of f u t u r e g e n e r a t i o n s " . Thus Banff N a t i o n a l Park, i n common w i t h most of our N a t i o n a l Parks, has been d e d i c a t e d to the maintenance of n a t u r a l con-d i t i o n s , where indigenous f l o r a and fauna may pursue t h e i r e v o l u t i o n a r y course, and n a t u r a l phenomena may be observed i n the f u l l p l a y of s e r a i p r o g r e s s i o n and r e t r o g r e s s i o n ; i n the ebb and flow of animal p o p u l a t i o n s . But man, i n h i s v i s i o n , f o r g o t perhaps that he h i m s e l f Is an i n t e g r a l p a r t of any b l o t i c community, and h i s p r e s e n c e — e v e n as an humble 2 student—must n e c e s s a r i l y b r i n g i n t o p l a y c e r t a i n f o r c e s too f r e q u e n t l y n e g l e c t e d i n the common concept o f n a t u r a l c o n d i -t i o n s . I f h i s numbers be few, and h i s a c t i v i t i e s be attuned to the n a t u r a l complex, then he may, through i n t e r p r e t a t i o n of the past and study o f the pre s e n t , delve w i t h some c o n f i -dence i n t o the f u t u r e . But i f h i s numbers be many, and h i s a c t i v i t i e s be merely those of a c a s u a l bystander, to be t h r i l l e d and amused, then he may w e l l warp the e v o l u t i o n a r y t r e n d beyond the l i m i t s o f present v i s i o n . C i v i l i z a t i o n has a l r e a d y taken i t s awful t o l l upon our r i c h f l o r a and fauna. I t s advent i n t o our N a t i o n a l Parks, with the consequent d e v e l -opment of highways, g o l f courses, s k i l i f t s , h o t e l s and motels can s c a r c e l y f a i l to leave i t s mark. F i r e , t h a t g r e a t f o r c e o f e v o l u t i o n , i s contained and stamped out. F i n a l l y , indigenous l i f e becomes inadequate, and e x o t i c s p e c i e s are p l a n t e d to f u r t h e r charm the Jaded eye. Thus i t i s i n Banff Park today, where hu n t i n g remains one of the few human a c t i v i -t i e s s t i l l f o r b i d d e n , and where e l k p l a n t i n g s have r e l e a s e d f o r c e s t h a t cannot f a i l to I n f l u e n c e the e v o l u t i o n a r y p a t t e r n of p l a n t s and animals a l i k e . I t I s , perhaps, time to take stock o f the s i t u a t i o n ; to ask where we are go i n g i n our con-cept o f N a t i o n a l Parks. T h i s study, having I t s beginning i n a c o n s i d e r a t i o n o f the moose-elk r e l a t i o n s h i p i n Banff Park, attempts to take us one step forward i n our search f o r the t r u t h . 3 SCOPE AND METHODS On the a u t h o r i t y of the then Dominion W i l d l i f e S e r v i c e , an i n v e s t i g a t i o n was c a r r i e d out 1ft 194-9, d u r i n g the months May to August i n c l u s i v e , to determine what, i f any, competi-t i o n might e x i s t between the moose and e l k of Banff N a t i o n a l Park; and to g a i n some i n s i g h t i n t o the i m p l i c a t i o n s of such co m p e t i t i o n should i t he found to e x i s t . The s t u d i e s were mainly c o n f i n e d to the Bow V a l l e y r e g i o n west o f B a n f f , but surveys were made of many other areas to g a i n a g e n e r a l know-ledge of the problems e x i s t i n g i n the Park, and to o b t a i n com-p a r a t i v e d a t a by which to measure c o n d i t i o n s found i n the main study a r e a . Such widespread o b s e r v a t i o n s c o u l d not f a l l to suggest the complex i n t e r - r e l a t i o n s h i p s e x i s t i n g between fo u r , at l e a s t , of the major b i g games s p e c i e s and t h e i r e n v i r o n -ments. Thus the dynamic aspects of the b i o t a became the major i n t e r e s t ( i f not the major task) throughout the study,, The problem was approached from two a s p e c t s : study of the ranges and range c o n d i t i o n s , and d i r e c t o b s e r v a t i o n s of the two major s p e c i e s c o n c e r n e d — t o g a i n knowledge of l o c a l food and r e p r o d u c t i v e and l i f e h a b i t s . In these l a t t e r s t u d i e s g r e a t e r emphasis was l a i d upon the moose since they were not o n l y more r e a d i l y observed i n summer but were con-s i d e r e d most l i k e l y to be a f f e c t e d by c o m p e t i t i o n , Judging 4 from p o p u l a t i o n d a t a . D a i l y r e c o r d s were kept of a l l w i l d -l i f e seen, and of v i s u a l estimates (supported by frequent mechanical checks of browse he i g h t and branch u t i l i z a t i o n ) of browse and graze c o n d i t i o n s on a l l ranges. Some q u a n t i -t a t i v e range d e t e r m i n a t i o n s were c a r r i e d out by means of square metre quadrats. P e l l e t counts and o b s e r v a t i o n s were used e x t e n s i v e l y to determine game sp e c i e s i n v o l v e d as the animals themselves were f r e q u e n t l y not p r e s e n t . P r e v i o u s f l o r a l s t u d i e s were c o n s u l t e d f o r purposes of t h i s i n v e s t i -g a t i o n , but over two hundred shrubs, grasses and f o r b s were c o l l e c t e d . Underwater p l a n t s p e c i e s were o b t a i n e d from moose summer range by means o f s e v e r a l hours of wading, knee to neck deep, i n l a k e s and marshes. V i s u a l s t u d i e s , w i t h and without a i d of b i n o c u l a r s , were made of p l a n t s u t i l i z e d by s i n g l e animals and by groups; by animals s t a t i o n e r y and mov-in g . These o b s e r v a t i o n s were made a t times v a r y i n g from day-break to a f t e r sundown, to ensure the widest p o s s i b l e coverage of behaviour p a t t e r n . As before s t a t e d , c h i e f emphasis was p l a c e d upon the Bow V a l l e y . But o b s e r v a t i o n s were made i n some d e t a i l i n the Cascade V a l l e y , Spray V a l l e y and Waterfowl Lake a r e a s . Recon-naissance surveys were made i n t o the C a r r o t Creek, Johnston Creek, Boom Lake, Helen Lake and Howse River-Saskatchewan C r o s s i n g ( i n c l u d i n g Owen Creek) r e g i o n s . Numerous other short t r i p s were taken, to round out knowledge of the Park and to cross-check game movements and range use comparisons. In a l l 9 these s t u d i e s i n v o l v e d over 3»500 m i l e s of t r a v e l by s t a t i o n wagon, 300-400 m i l e s by f o o t , and some t r a v e l by horse and by rubber boat. The e l k were subjected to s c r u t i n y f i r s t ; to a s c e r t a i n t h e i r food h a b i t s d u r i n g the s p r i n g when new gra s s was not r e a d i l y a v a i l a b l e . At the same time, a g e n e r a l survey of range c o n d i t i o n s was made. Then, as the e l k groups commenced to break up p r e p a r a t o r y to c a l v i n g , and to the f i n a l movement toward summer range, a t t e n t i o n was turned more to the study of moose and f u r t h e r , more d e t a i l e d , study o f the ranges. As the summer progressed, s t u d i e s were d i v i d e d between moose and e l k , w i t h o b s e r v a t i o n s on sheep, deer and goat c a r r i e d on con-c u r r e n t l y . F i n a l l y , the v a r i o u s Wardens i n the Park were c o n t a c t e d and t h e i r o b s e r v a t i o n s , o p i n i o n s and suggestions noted. The old-time wardens were questioned to o b t a i n h i s t o r y of the r e g i o n . Many of the a v a i l a b l e d i a r i e s were scanned and moose and e l k occurrences noted. In p a r t i c u l a r , c o n s o l i d a t e d r e p o r t s of a l l b i g game and p r e d a t o r s , prepared by S p e c i a l Warden H. U. Green, were s t u d i e d to p r o v i d e b a s i s f o r comparison, and more p a r t i c u l a r l y to g a i n an i n s i g h t i n t o the e c o l o g i c a l com-p l e x i t i e s of the b i o t a of the Park. Reference has been to Mr. Green's r e p o r t s s i n c e t h i s survey, to b r i n g some of the data up-to-date. 6 THE PROBLEM AREA HISTORY The Rocky Mountains, f o r which the Rocky Mountains Park was named, o r i g i n a l l y were named A s s i n - w a t i by the Crees; t h i s was t r a n s l a t e d i n 1752 to the French Montagnes des Roches ( W i l l i a m s , 1929:7). But i t was not u n t i l n e a r l y a h a l f a century l a t e r that e x p l o r e r s f i r s t e n tered the r e g i o n now known as Banff N a t i o n a l Park. In November, 1800, David Thompson and Duncan M c G i l l l v r a y ascended the Bow R i v e r to "The Gap", w i t h i n about f i f t e e n m i l e s of the p r e s e n t B a n f f townsite, but d i d not proceed f a r t h e r west. In iBkO, Reverend R. T. Rundle, while working w i t h the Stoney Indians, spent a p a r t of June and J u l y camped i n f r o n t of what i s now Mount Rundle, and i n lglj-l S i r George Simpson passed through the Banff area and proceeded to the Columbia V a l l e y by way of Healy Creek and Simpson Pass. Next r e c o r d i s f o r 1&H-5, when Father P. J . DeSmet cros s e d White Man Pass from the south, camped at Canmore, and then f o l l o w e d the mountain t r a i l to Rocky Mountain House. Exact d e t a i l s of h i s r o u t e are not p r e s e n t l y a v a i l a b l e . (Warren, 1927:1-2.) In lS^S" C a p t a i n John P a l l i s e r headed an e x p e d i t i o n i n t o the Rockies, adjacent to and p o s s i b l y i n t o what i s now the Park, while In search of an a l l - B r i t i s h wagon road to the P a c i f i c c o a s t . He s p l i t h i s p a r t y i n three and one 7 member, S i r James Hector, a g e o l o g i s t , ascended the Bow Ri v e r t o i t s j u n c t i o n w i t h the V e r m i l i o n ( A l t r u d e Creek?), ascended to the D i v i d e and turned n o r t h to the K i c k i n g Horse Pass. ( W i l l i a m s , 1929:11-12.) The f o l l o w i n g year the E a r l o f Southesk went i n t o the a r e a (probably t o hunt, so i t i s recorded) and t r a v e l l e d over the Pipestone Pass (from the Clearwater or S i f f l e u r ? ) t o descend the Bow R i v e r . In 1880 the f i r s t surveyors entered the now Banff Park area. (Warren, 1927:1-2.) White men f i r s t saw and recor d e d the now famous hot sp r i n g s o f Banff i n 18&3. S o important was t h i s f i n d con-s i d e r e d t h at i n 1885, upon completion of the Canadian P a c i f i c Railway, the Government set a s i d e t en square m i l e s surround-i n g the s p r i n g s , to keep the area f i t f o r a f i r s t c l a s s r e -s o r t . Thus Banff Park had i t s g e n e s i s . In 1887, impressed by the beauty of the area, a group of i n f l u e n t i a l men and p o l i t i c i a n s were i n s t r u m e n t a l In having set a s i d e , by a c t of Parliament, the Rocky Mountains Park, a r e c r e a t i o n a l and sc e n i c a r e a of 260 square m i l e s ( i n c l u d i n g the o r i g i n a l ten square m i l e s ) . In 1902 t h i s was i n c r e a s e d to 5»000 square m i l e s , but was, i n 1911, reduced a g a i n t o 1,8>00 square m i l e s by passage of the F o r e s t Reserves and Parks Act of t h a t year. For game p r o t e c t i o n purposes i t was a g a i n i n c r e a s e d , some time p r i o r to 1921, to 2,751 square m i l e s . ( N a t i o n a l Parks of Canada, 1938:5 and W i l l i a m s , 1921:13-14.) In 1929, the name of the Park was changed to Banff N a t i o n a l Park. F i n a l l y , i n 193°> the Park was extended from i t s north-west terminus at Bow Summit, to Sunwapta Pass, southern terminus of J a s p e r N a t i o n a l P a r k — s o c r e a t i n g a short common boundary. Thus the Park today embraces 2,5$5 square m i l e s of t e r r i t o r y e n t i r e l y w i t h i n the Rocky Mountains and s i t u a t e immediately east of the C o n t i n e n t a l D i v i d e . I t s western boundary i s l a r g e l y shared w i t h the e a s t e r n boundaries of Yoho and Kootenay N a t i o n a l Parks* GEOLOGY AND GEOGRAPHY I t i s probable that t h i s s e c t i o n of the Rockies l a y under water d u r i n g p a r t of the Cambrian p e r i o d , and a g a i n d u r i n g the O r d o v i c i a n p e r i o d when an i n l a n d sea s t r e t c h e d from the r e g i o n of the S e l k i r k s to the L a u r e n t i a n s . During the ages s i l t was l a i d down, to form i n some p l a c e s a l a y e r 7 ? • ' ( over 50,000 f e e t i n t h i c k n e s s . In the C a r b o n i f e r o u s p e r i o d , p r e s s u r e over m i l l i o n s of years caused upheaval, and a g a i n toward the end of the Mesozoic e r a another t h r u s t from the west crumpled the area, f o l d s were r a i s e d up and f a u l t s o c c u r r e d . In some areas rocks from the west s i d e were pushed up over the e a s t e r n formations, l e a v i n g grey l i m e -stones s l o p i n g to the west and steep escarpments f a l l i n g away to the e a s t . Older formations were, i n p l a c e s , t h r u s t over new. Then f o l l o w e d the g l a c i a l p e r i o d , when thousands of f e e t o f i c e capped the area, g r i n d i n g and moving. F i n a l l y the ice. receded, l e a v i n g the contours p r e t t y much as they are today, though e r o s i o n and avalanche have made some marks. (W i l l i a m s , 1929:9-11.) Today there may be seen w i t h i n the Park some t r a c e of almost every g e o l o g i c a l system from the Prf-Cambrian t o the Cretaceous i n c l u s i v e , e x c e p t i n g o n l y the T r i a s s i c ( A l l a n , 1913:167); The c o a l beds a t the e a s t e r n boundary of the Park are of Cretaceous o r i g i n . The Park, as c o n s t i t u t e d today, i s t r a n s e c t e d by s e v e r a l mountain ranges which, i n the main, f o l l o w a northwest to southeast t r e n d and l i e p a r a l l e l to the C o n t i n e n t a l D i v i d e . The P a l l i s e r , Cascade and Sulphur ranges show w e s t w a r d - t i l t e d f a u l t b l o c k of P a l a e o z o i c and Mesozolc r o c k — w i t h east face Jagged escarpment and west sweeping sheer s t r a t a . The Sawback range shows Ca r b o n i f e r o u s and Devonian rock f o l d e d i n t o compressed a n t i c l i n e (weathered i n t o s p i r e s and saw-toothed r i d g e s ) . Many of the mountains west of the Sawback range show Ca r b o n i f e r o u s and Devonian rock, w i t h the abrupt limestone c l i f f s r i s i n g to round or wedge-shaped summits. ( N a t i o n a l Parks A s s o c i a t i o n , I9k&ij.) From the many g l a c i e r s s t i l l e x i s t e n t w i t h i n the ar e a and from along the inter-mountain v a l l e y systems a r i s e the headwaters of the Bow, Red Deer, Clearwater and North Saskatchewan R i v e r s , a l l of which f l o w g e n e r a l l y eastward from the Park through the f o o t h i l l s of A l b e r t a . Two major t r i b u t a r i e s , the S i f f l e u r on the North Saskatchewan and the 10 Panther on the Red Deer, f o l l o w a somewhat s i m i l a r p a t t e r n . Within the c o n f i n e s o f the Park s e v e r a l r i v e r s of importance and c o u n t l e s s creeks c o n t r i b u t e to these major drainage s y -stems. Of p a r t i c u l a r note are the Cascade and Spray R i v e r s , and the F o r t y m i l e , Healy, Red E a r t h , Johnston and Pipestone Creeks f l o w i n g i n t o the Bow R i v e r , and the Mistaya, Howse and Alexandra R i v e r s f l o w i n g i n t o the North Saskatchewan. CLIMATE C l i m a t i c a l l y , the Park i s i n a comparatively low p r e c i -p i t a t i o n r e g i o n j u s t east of the Great D i v i d e , I t l i e s , however, a s t r i d e the d i v i d i n g l i n e between moderately heavy and v e r y l i g h t p r e c i p i t a t i o n a r e a s . The headwaters of the Bow, Red Deer, Clearwater and North Saskatchewan R i v e r s , the C o n t i n e n t a l D i v i d e i t s e l f , and west c e n t r a l areas g e n e r a l l y n o r t h to and i n c l u d i n g Bow Summit, r e c e i v e moderately heavy summer r a i n s and deep win t e r snows which encompass both moun-t a i n s and v a l l e y s . E a s t e r l y , the p r e c i p i t a t i o n becomes p r o -g r e s s i v e l y l e s s , w i t h comparatively l i g h t snow f a l l i n g i n the major v a l l e y s below the headwaters. Some e a s t e r l y p o r t i o n s of the Park experience near drought; the same c o n d i t i o n o b t a i n s i n the North Saskatchewan V a l l e y . The Spray V a l l e y i s t r a n s i t i o n a l , r e c e i v i n g c o n s i d e r a b l y more t o t a l moisture, winter and summer, than the adjacent Bow V a l l e y . 11 FLORA I t i s suggested by Clarke (1939:3) that immediately f o l l o w i n g the l a s t g l a c i a l r e t r e a t , the r e g i o n under study contained only two p l a n t a s s o c i a t i o n s , tundra and g r a s s l a n d , w i t h wet meadow i n t e r m e d i a t e . Then Engelmann spruce ( P i c e a Engelmannl) and A l p i n e f i r (Abies l a s l o c a r p a ) Invading from the west, i n t e r p o s e d themselves a t the J u n c t i o n of the two formations, and g r a d u a l l y pushed out to occupy the deep snow v a l l e y s of the west (some marshland areas s t i l l remain). From the northwest, white spruce ( P i c e a g l a u c a ) , w i t h p o p l a r (Populus tremuloides) as a f o r e r u n n e r , invaded the g r a s s l a n d a r e a of the east and pushed forward i n t o the mountains to meet the Engelmann a s s o c i a t i o n , l e a v i n g s m a l l areas of o r i -g i n a l p r a i r i e s t i l l e x tant. Thus today three major f l o r a l a s s o c i a t i o n s o c c u r — g r a s s l a n d , tundra ( a l p i n e ) and f o r e s t , the l a t t e r f a l l i n g i n t o three s u b d i v i s i o n s — b o r e a l , sub-alplne and montane. Green ( l 9 5 l ) s t a t e s t h a t there i s reason to b e l i e v e t h a t the Bow V a l l e y and I t s b o r d e r i n g mountain slo p e s supported dense stands o f dominant white spruce, crowded to the banks of the Bow R i v e r , f o r at l e a s t 110 years p r i o r to the t u r n of the Twentieth century. In the Lake Louise a r e a , however, i n 1S96 the major f l o r a l a s s o c i a t i o n from the Bow R i v e r to timber l i n e was Engelmann*s spruce w i t h an o c c a s i o n -a l b l a c k pine (Pinus ponderosa—was more probably Pinus a l b i o a u l l s ) . There were a few deciduous t r e e s and bushes 12 adjacent to the banks of the r i v e r s and open sphagnum swamps caused by morainal dams, and i n c l e a r i n g s c r e a t e d by snow avalanches. (Wilcox, 18>96:52>-6o.) Other p o s s i b l e e xceptions may have been s m a l l areas of o r i g i n a l g r a s s l a n d auch as s t i l l e x i s t a t M i l e 13 ( H i l l s d a l e ) west of B a n f f . O r i g i n of the Douglas f i r (Pseudotsuga t a x i f o l l a v a r . glauca) at low a l t i t u d e s i n the Bow V a l l e y i s not c l e a r . Lodgepole pine (Plnus c o n t o r t a v a r . l a t i f o l i a ) e x i s t s throughout the Park as a f i r e type, w i t h a tendency f o r such areas to r e v e r t to g r a s s l a n d a f t e r repeated or heavy burnings. In L5>90 a f i r e almost completely d e s t r o y e d the f o r e s t of the sl o p e s south of the Bow R i v e r from o p p o s i t e the V e r m i l i o n Lakes to beyond Baker Creek to the west. In 1903 f i r e d e s t r o y e d the f o r e s t from E a s t Gate to A n t h r a c i t e , n o r t h si d e of the Bow, and i n 1904- another f i r e burned the sl o p e s and v a l l e y f l o o r n o r t h of the Bow from V e r m i l i o n Lakes west to Baker Creek and n o r t h some d i s t a n c e up Johnston Creek. I t was f o l l o w i n g these f i r e s that aspen p o p l a r became a sub-s t a n t i a l p a r t of the f o r e s t a s s o c i a t i o n of t h i s area, and w i l l o w ( S a l i x Bebbiana Sarg.) took over s u i t a b l e v a l l e y f l o o r h a b i t a t s . In 1912 f i r e swept up both s i d e s of the Spray R i v e r from near Banff to about M i l e IS. The Panther R i v e r d i s t r i c t was burned i n 1926, and In 1928 the west slopes of Goat Range f e l l before f i r e . 1930 the area Cuthead toward the P a l l l s e r was burned; 193^ t n e Brewster Creek are a ; 1936 from F l i n t ' s Park toward the P a l l l s e r and Cascade 13 V a l l e y , and i n 19*4-0 the Howse R i v e r - N o r t h Saskatchewan R i v e r t r i a n g l e and the northeast slope o f Goat Range were burned. The f i r s t and second Kananaskis f i r e s , In 1926 and 1935-36 probably a f f e c t e d the southern reaches of the P a r k a I t i s c o n s i d e r e d probable that the main North Saskatchewan V a l l e y must have been burned w i t h i n the l a s t f i f t y y e ars, though no such f i r e i s recorded. C e r t a i n l y other f i r e s have oc c u r r e d i n the Park, but are not recorded o r remembered. But these burns, now a t v a r i o u s stages of g r a s s l a n d — l o d g e p o l e p i n e a s s o c i a t i o n and r e f o r e s t a t i o n , have p r o v i d e d the main b i g game f e e d i n g areas i n the past and s t i l l support the l a r g e r f a u n a l s p e c i e s — f o r a p o r t i o n of each year at l e a s t . They are complemented, and to some degree supplemented, by the many w i l l o w ( S a l i x ve.stlta Pursh.) areas p r o v i d e d by avalanche a c t i o n , and by wil l o w and other browse s p e c i e s now present i n or about creeks, r i v e r s and marshes. A l p i n e mea-dows, w i t h a s s o c i a t e d w i l l o w ( S a l i x saxlmontana Rydb.), p r o v i d e the t h i r d major source of food supply. FAUNA The h i s t o r y of the fauna of the Park i s u n f o r t u n a t e l y not w e l l recorded, a t l e a s t u n t i l v e r y recent times. Annual r e p o r t s on the Park made l i t t l e mention o f w i l d s p e c i e s u n t i l the l a t e 1930 ls. Th e r e f o r e , i t i s necessary to draw 14-r a t h e r broad i n f e r e n c e s from the few data a v a i l a b l e ; to b u i l d up a p i c t u r e of f a u n a l occurrence from which one can draw present comparisons and p o i n t up dynamic i n t e r -r e l a t i o n s h i p s . Comparatively l i t t l e i s known o f Rocky Mountain goat (Oreamnos amerlcanus missoulae) and c a r i b o u ( R a n g l f e r mon-ta n u s ) , e i t h e r p a s t or present, i n Banff Park. S i r George Simpson (184-7:118) r e p o r t e d sheep and goats along h i s route ( i n the Park) up to Simpson Pass i n 184-1. The r e c o r d of the E a r l of Southesk hunting i n through the Pipestone area i n 1859 would suggest the presence of goat, c a r i b o u or g r i z z l y bear (or a l l three) i n t h a t r e g i o n . Wilcox, In h i s r e p o r t p u b l i s h e d 1896 on h i s work i n the Lake Louise area, r e c o r d e d the presence of goat, b l a c k bear and mountain l i o n s . (Wilcox, 1896:58-60.) Caribou have been rec o r d e d o n l y as e a r l y as 1902, and as r e c e n t l y as 194-3 (Cowan, 194-3:13), 19^6 and pos-s i b l y 194-8 (Green, 194-8:7). The Park cannot be c o n s i d e r e d good c a r i b o u range, and i n l i g h t of present knowledge, t h i s s p e c i e s probably warrants l i t t l e c o n s i d e r a t i o n i n t h i s d i s -c u s s i o n of dynamic e c o l o g i c a l r e l a t i o n s h i p s . The same may be s a i d f o r the goat. The s l u t a t i o n might be completely changed when good data are a v a i l a b l e on both these s p e c i e s . Goat are p r e s e n t l y d i s t r i b u t e d throughout the Park i n a manner to suggest that most of the s u i t a b l e e c o l o g i c a l n iches are i n some degree u t i l i z e d . However, da t a are l a c k i n g 15 r e g a r d i n g t o t a l numbers, and c a r r y i n g c a p a c i t y of range i s unknown. Rocky Mountain bighorn (Ovls canadensis canadensis) have been indigenous to the Rockies and the f o o t h i l l s s i n c e e a r l i e s t r e c o r d s , and i n the U n i t e d S t a t e s used to extend east throughout the rugged and broken g r a s s l a n d s to the M i s s o u r i R i v e r and i n t o the Dakota badlands. With the advent of the white man t h e i r range was r e s t r i c t e d from t h i s p r o -bable optimum, and they were f o r c e d to a readjustment of h a b i -t a t whereby once mar g i n a l areas, or at best summer range, h i g h i n the mountains became home range. The s p e c i e s , so con-f i n e d to the east, was h a l t e d In t h e . w e s t e r l y movement where-ever deep snows o c c u r r e d a t or east of the C o n t i n e n t a l D i v i d e , and was f o r c e d to e x p l o i t a l p i n e meadows, and g r a s s l a n d s , the l a t t e r produced by the e v e r - i n c r e a s i n g burns t h a t h e r a l d e d the advent of the white men. The constant encroachment of f o r e s t s p e c i e s upon these burns, p l u s man's e f f o r t s a t f i r e c o n t r o l ( f i r s t o r ganized In 190.9 i n Banff Park (Canada, 1911:6)) would aggravate t h i s p r e c a r i o u s e x i s t e n c e . T h i s , i t would appear, has been the s i t u a t i o n i n Banff Park n o r t h of Simpson Pass ( C l a r k e , 19^2:3), where sheep have been recorded as e a r l y as 16 -^1 (Simpson, 1547:118). They are l a r g e l y c o n f i n e d to the e a s t e r n ranges, w i t h some i n v a s i o n of c e n t r a l areas n o r t h of and adjacent to the Bow V a l l e y . They were present i n c o n s i d e r a b l e numbers, p o s s i b l y maximum, u n t i l the l a t e 1 9 3 0 ,s when the p o p u l a t i o n s commenced 16 d w i n d l i n g . The same s i t u a t i o n a p p a r e n t l y o b t a i n s today, w i t h e x t i n c t i o n , though improbable, a p o s s i b i l i t y . There are no e a r l y r e c ords of mule deer (Odocoileus  hemionus hemionus) i n the Park a r e a . One can onl y surmise t h a t , s i n c e they are i n h a b i t a n t s of the b o r d e r l a n d between f o r e s t and g r a s s l a n d , they have f l o u r i s h e d or f a i l e d w i t h the s u c c e s s i v e changes of f l o r a l a s s o c i a t i o n s w i t h i n the Park. They w i l l most c e r t a i n l y have been i n the Bow V a l l e y s i n c e the f i r e s o f 1890 and 1903, as they are known to have been i n the f o o t h i l l s s i n c e e a r l i e s t r e c o r d . They are p r e s e n t l y found throughout the Park i n summer, from v a l l e y t o t i m b e r l i n e , but i n w i n t e r move to areas o f l i g h t snow—both w i t h i n and without the Park boundaries. The Bow V a l l e y i s the most important winter c o n c e n t r a t i o n a r e a w i t h i n the Park. W h i t e t a i l deer, (Odocoileus v i r g l n l a n u s macrourus) are scarce w i t h i n the Park today, and are not recor d e d i n any e a r l y d a t a . Cowan (194-3: 10) r e c o r d s a smal l band r e s i d e n t a l o n g the North Saskatchewan R i v e r , and a few o t h e r s along the B a n f f - J a s p e r highway from M i l e 92 to M i l e 102, A few summer occurrences have been noted In recent years, but w i n t e r r e s i d e n c e h a s n o t been e s t a b l i s h e d . F u r t h e r c o l o n i z a t i o n i s not c e r t a i n and w i l l p r o b a b l y be con-t r o l l e d by f u t u r e range c o n d i t i o n s . (Ibid; 1 0 . ) They can, i n the main, be ign o r e d i n t h i s study, s i n c e t h e i r i n f l u e n c e upon the f a u n a l complex i s minor. Moose ( A l c e s americana americana) are Indigenous to the area, but of recent o r i g i n i n most reaches o f the Park» In 17 1906, the Park Superintendent r e p o r t e d t h a t " ...There i s at present a good number of b i g game i n the park, c o n s i s t i n g c h i e f l y o f moose,, e l k , deer, bear, sheep, l y n x and goat, as w e l l as marten and beaver, b e s i d e s an u n l i m i t e d num-ber of game b i r d s , " (Green, 19^6:2)„ Since a t that time the Park encompassed 5,000 square m i l e s , (exact boundaries not known) i t i s pos-s i b l e the moose r e p o r t e d were i n the North Saskatchewan d r a i n -age a r e a . J« Naylor, r e t i r e d Park Warden, s t a t e s they were known i n t h a t r e g i o n l o n g before they were ever seen i n the lower reaches of the Park. Another p o s s i b i l i t y Is that they were i n the Ya-Ha-Tinda d i s t r i c t . However, the Park was reduced i n s i z e i n 1911, and i t was not u n t i l 1930 t h a t the Saskatchewan area was again I n c l u d e d . Thus, the 1916 r e p o r t , by Park Warden U. U., LaCasse, was the f i r s t r e c o r d of a moose i n the Park as then c o n s t i t u t e d , and the f i r s t r e p o r t e d o c c u r -rence i n the southern a r e a . T h i s animal came from the Kootenays over V e r m i l i o n Pass. The next r e c o r d was i n 1923, when a moose was seen by Wardens J . Naylor and W. O h i l d , near the abandoned beaver dams adjacent to A l t r u d e Creek and the Bow R i v e r . The same year, Warden P. G. Woodworth saw a moose while on p a t r o l i n t o the upper areas of the Park (Pipestone c o u n t r y ? ) . He c o n s i d e r e d i t a most remarkable s i g h t . From t h i s time on moose spread out over the Park. The Bow V a l l e y p o p u l a t i o n c o l o n i z e d , no doubt, i n a l l d i r e c t i o n s , but i t s e a s t e r n movement h a l t e d a t the confluence of the Cascade and Bow R i v e r s , where s u i t a b l e h a b i t a t ended. Moose IS" are p r e s e n t l y found along a l l Important water courses i n the Park. The Bow drainage system i s the major a r e a i n v o l v e d i n t h i s study. One cannot pass from c o n s i d e r a t i o n of the moose of the Park without some mention of the heaver. The two s p e c i e s have been a s s o c i a t e d e c o l o g i c a l l y throughout h i s t o r y , and the s i t u a t i o n here i s no e x c e p t i o n . I t i s d i f f i c u l t to escape the c o n c l u s i o n that n e i t h e r s p e c i e s c o u l d or would have c o l o n i z e d the area, other than t r a n s i t o r i l y , but f o r the f i r e s at the t u r n of the century. The aspen and p o p l a r a s s o c i a t i o n f o l -lowing the f i r e s c r e a t e d an environment a t t r a c t i v e to beaver. They, i n t u r n , c o n d i t i o n e d the environment to the needs o f moose. Thus commenced a sequence of e c o l o g i c a l events s t i l l seen today i n the lower Bow V a l l e y . The f i r s t beaver were observed i n 1921, near the mouth of A l t r u d e Creek, where they- had e s t a b l i s h e d a pond and lodge. Subsequent movement downstream took p l a c e , w i t h c o l o n i z a t i o n at Red E a r t h Creek, then Healy Creek, Sundance Creek and V e r m i l i o n Lakes and a r e a . U n s u i t a b i l i t y of h a b i t a t ended t h i s movement i n 1931 a t the j u n c t i o n of Cascade and Bow R i v e r s . About 1924-, p o p u l a t i o n p ressure i n the V e r m i l i o n Lakes a r e a was r e l i e v e d by e m i g r a t i o n up F o r t y m i l e Creek, but by 1934- t h a t area had been eaten out, and the p o p u l a t i o n d i s p e r s e d — p r o b a b l y to f a l l prey to coyotes (and p o s s i b l y to cougar). By 194-9, beaver i n the lower Bow V a l l e y were l a r g e l y c o n f i n e d to a small a r e a at V e r m i l i o n Lakes, where some r e s e r v e s of p o p l a r s t i l l remained. 19 •Thus a c o l o n i z a t i o n , undoubtedly made p o s s i b l e by the f i r e s of 1S90, 1903 and 19G4, t h r i v e d , ate i t s e l f out and waned to but a few l o d g e s . As f o r the remainder of the Park, three new lodges were seen on the Spray and Bryant R i v e r s i n 19421, but no other f i r m l y e s t a b l i s h e d c o l o n i e s are known. I t i n e r a n t beaver may occur from time to time. The l a s t b i g game sp e c i e s to be considered, i s the e l k (Cervus canadensis n e l s o n l ) . I t has been noted (under moose) that e l k were r e p o r t e d w i t h i n the Park i n 1906. T h i s i s con-t r a r y to the statements of Mr. James Simpson and Mr. N.K. Luxton, both of whom s t a t e there were never e l k i n the Park, w i t h i n t h e i r memory, p r i o r to the f i r s t p l a n t i n g . Mr. Simpson, a t l e a s t , has been In the a r e a s i n c e before the t u r n of the century. I t i s p o s s i b l e that t h i s c o n f l i c t of o p i n i o n has a r i s e n because of the s h i f t i n boundaries that took p l a c e , but i t seems f a i r l y d e f i n i t e t h a t e l k have never been I n d i -genous to the Park, as now c o n s t i t u t e d , at l e a s t w i t h i n the memory of l i v i n g man. S i r G-eorge Simpson, however, d u r i n g h i s passage through the area i n 18>4l, r e c o r d e d t h a t one of h i s p a r t y k i l l e d a red deer two m i l e s from the Bow T r a v e r s e , where they c r o s s e d the Bow R i v e r (Simpson, 1847:122). He f u r t h e r r e p o r t e d f r e s h t r a c k s of r e d deer the second day down from Simpson Pass towards the Kootenay R i v e r ( l b i d | 1 2 1 ) . I t i s not known I f by t h i s he r e f e r r e d to e l k , but t h i s seems probable since the term red deer, as a p p l i e d to w h i t e t a i l deer, i s b e l i e v e d to be of more r e c e n t o r i g i n . C e r t a i n l y 20 e l k remains i n many areas along the f o o t h i l l country n o r t h of Morley, A l b e r t a , i n d i c a t e the s p e c i e s was common (and p l e n t i -f u l ) immediately east of the Park* These remains were r e p o r t e d to Mr. H. U. Green by Mr. George McLean, an educated Stoney Indian of venerable y e a r s . He s t a t e d t h a t i n some p l a c e s they covered s e v e r a l a c r e s , and he d i s c o u n t e d any theory of s l a u g h t e r by I n d i a n s . A g e n e r a l d i e - o f f through d i s e a s e and/or s t a r v a t i o n seems more pro b a b l e . T h i s whole s i t u a t i o n p a r a l l e l s the h i s t o r y of e l k i n the U n i t e d S t a t e s , where i t i s e a r l y recorded that e l k frequented the upper Rocky Mountain ranges i n summer, and where d i e - o f f s n e a r l y e x t i r p a t e d the s p e c i e s . E l k are known to have been i n the Canal Flats-Invermere country about the middle of the 19th century, and were prese n t i n some numbers to the 18J0*& when a severe w i n t e r almost e x t i r p a t e d the p o p u l a t i o n (from i n f o r m a t i o n g i v e n by Mr. M. Morigeau, Fairmont Springs, B. C.). They r e c o v e r e d slowly and by 1916-1920 were c o n s i d e r e d to be common a g a i n . They moved up and down the Kootenay drainage system, and undoubtedly approached, a t l e a s t , the summits o f the D i v i d e s . and the Banff Park border. I t i s not u n l i k e l y t h a t the e l k s i g h t e d a t Egypt Lake d u r i n g the summer of 194-5 (Green, 194-6: 4-) were from the Kootenay herd. Cowan a l s o r e c o r d s some i n s t a n c e s of e l k c r o s s i n g the west boundary o f the Park (Cowan, 194-3:14-). However, s i n c e i t was not u n t i l 1922 or 1923 that they were i n s u f f i c i e n t numbers to warrant an 'open 21 season i n B r i t i s h Columbia, i t i s u n l i k e l y t h a t p o p u l a t i o n p r e s s u r e has been s u f f i c i e n t to b r i n g about more than chance summer occupation of any p a r t of Banff Park. To the east of the Rockies e l k were p l e n t i f u l p r i o r to 12>91> and i t seems probable t h a t they were p r e s e n t i n most s u i t a b l e areas b o r d e r i n g the Park. The f i r e of 1890 should have p r o v i d e d o u t l e t f o r p r e s s u r e - i n d u c e d c o l o n i z a t i o n . But i n 1891-92 a most severe winter was experienced and i t i s recorded that i n Yellowstone N a t i o n a l Park 5,000 of an e s t i -mated 25,000 e l k d i e d of s t a r v a t i o n . Again i n 1898-99 u n u s u a l l y severe w i n t e r c o n d i t i o n s p r e v a i l e d , and the e l k along the e a s t e r n l i m i t s o f the Rockies became v i r t u a l l y ex-t i n c t , (Skinner, 1950:22.) Large gaps were l e f t i n a once almost continuous p o p u l a t i o n . I t i s b e l i e v e d that such a gap e x i s t e d immediately to the east of the Park, In 1916 arrangements were made by park a u t h o r i t i e s to o b t a i n e l k from the N a t i o n a l Park S e r v i c e of the U n i t e d S t a t e s , and In February, 1917> a t o t a l o f f i f t y - s e v e n e l k were r e c e i v e d from Yellowstone Park. In June, 1918, f o r t y - o n e were r e l e a s e d . In January of 1919 or 1920 an a d d i t i o n a l 206 were ob t a i n e d from the same source: twelve d i e d en r o u t e . F i f t y - t h r e e of these were r e l e a s e d a t Massive, twelve m i l e s west of Banff, and l 4 l a t D u t h i l l , e i g h t miles east of B a n f f . In a l l , 235 e l k were r e l e a s e d i n the Bow V a l l e y . (Green, 1946;2.) They f l o u r i s h e d , and soon began to migrate. Cowan (1943:14) g i v e s the f o l l o w i n g chronology of d i s p e r s a l as I n d i c a t e d by f i r s t 22 appearances i n the r e s p e c t i v e a r e a s : "Banff-Bow V a l l e y . Introduced 1917; Panther R i v e r -f i r s t seen 1927 (C. C. F u l l e r ) ; Snow Creek P a s s -f i r s t seen 1931 (W. C h i l d s ) ; Red Deer a t Yaha T i n d a Ranch, f i r s t seen 1933-34- (C„ Murphy); S a s k a t c h e w a n — f i r s t summering b u l l s 193° ( D a v i e s ) ; Indian Head c a b i n on Clearwater R i v e r — f i r s t seen 194-2 (Chas. P h i l l i p s ) ; L i t t l e P i p e s t o n e — f i r s t w i n t e r e d 194-2-4-3 (P. Woodworth); 9 m i l e s n o r t h of Lake Louise on J a s p e r H i g h w a y — f i r s t wintered 194-2-4-3 (P. Woodworth)," By 194-3 removal of s u r p l u s e l k was c o n s i d e r e d necessary and since t h a t date some e l k have ,been s l a u g h t e r e d each w i n t e r . In s p i t e of t h i s e l k continue to be p l e n t i f u l . Main perman-ent c o n c e n t r a t i o n s , determined by winter range, appear to be along the Bow V a l l e y from the E a s t Gate to L o u i s e , the Upper Spray V a l l e y , the Upper Cascade V a l l e y and the North Saskatchewan R i v e r . Other small l o c a l groups no doubt e x i s t . Nothing i s known of the advent of the major p r e d a t o r s i n the Park, except i n the case of wolves. Bear, both b l a c k (Eflarctos amerlcanus americanus) and g r i z z l y (Ursus h o r r i b i l -i s ) are present, the former i n some abundance. Simpson (184-7: 121) r e c o r d s bear t r a c k s on the south west slopes of Simpson Pass i n 184-1', but does not s t a t e f o r which s p e c i e s . Wilcox, i n I896, r e c o r d s b l a c k bear i n the Lake Lou i s e a r e a (Wilcox, 1896:58-60). Cougar ( F e l l s c o n c o l o r m l s s o u l e n s i s ) appear to be of r a r e occurrence i n the Park today, but were re c o r d e d a l s o by Wilcox i n 1896 ( I b i d ) . They were c o n s i d e r e d over-abundant by park a u t h o r i t i e s i n 1935, and Mr. E. R. Lee of Vancouver Island-w a a employed to hunt them. During the 23 p e r i o d 10 October, 1935, to 26 January, 1936, he k i l l e d nine c o u g a r — o n e at Massive and e i g h t a l o n g the Bow V a l l e y between Banff and East Gate* (From Government f i l e s . ) More r e c e n t l y , Green r e c o r d s a female and two k i t t e n s near the Aylmer Range i n 1946, t r a c k s of one a d u l t near Johnston Creek Range i n 19^6, and t r a c k s of two a d u l t s near Eisenhower Range i n 19*4-7 (Green, 19*4-9:35)* True s t a t u s o f the cougar i n the Park i s p r e s e n t l y unknown. The coyote (Canls l a t r a n s ) i s p l e n t i f u l l y r e p r e s e n t e d and has probably been present as long as the v a l l e y s have been semi-open and have supported deer. The t r a c k s of wolves (Canls lupus o c c i d e n t a l i s ) are r e c o r d e d by Simpson ( : 1 2 1 ) as o c c u r r i n g j u s t over Simpson Pass i n t o the Kooxtenay area, so presumably they ranged i n t o the a r e a now i n Banff Park. However, a c c o r d i n g ^ t o Cowan, they made t h e i r f i r s t appearance, i n r e c e n t times, i n the w i n t e r of 1942-43 when two entered the Park i n the Saskatchewan R i v e r area. In 1945 f i v e or s i x wolves hunted the Clearwater V a l l e y and denned j u s t o u t s i d e the Park. In 19*4-3 a s m a l l r e s i d e n t p o p u l a t i o n i n the Ya-Ha-Tinda a r e a worked al o n g the Red Deer R i v e r to Scotch Camp, and ranged up the Panther R i v e r to Windy Cabin. In 1946, however, evidence was seen that a p a i r had denned In the Panther R i v e r V a l l e y f o r a t l e a s t e i g h t y e a r s . (Cowan, 1947:145.) More r e c e n t l y , Warden E. Young r e p o r t s that In 194S there was a pack of f o u r t e e n wolves i n the Ya-Ha-Tinda Ranch a r e a . In 1944-45 three i n d i v i d u a l s passed through the Ghost River-Mlnnewanka Lake area, and i n 1943, 24-194-4- and 194-5 s i n g l e animals were seen at Bow Summit, Goat Creek, and i n the Bow V a l l e y e i g h t e e n m i l e s west of Banff ( i b i d ) . A c c o r d i n g to Mr. U. U. LaCasse, a r e t i r e d Park Warden, wolves came i n t o the Bow V a l l e y i n about 194-2-4-3 and a group became e s t a b l i s h e d up Johnston Canyon or Creek s h o r t -l y t h e r e a f t e r , o p e r a t i n g up to the P u l s a t i l l a Mountain a r e a and down i n t o the Bow V a l l e y . The pack f i n a l l y numbered nine a c c o r d i n g to h i s count. By 194-9 wolves were r e p o r t e d on a l l major game ranges i n the Park. In c o n s i d e r i n g , then, the g e n e r a l p i c t u r e of the f l o r a and fauna of the Park, i t i s seen t h a t today the major con-c e n t r a t i o n s of moose, e l k , sheep and mule deer are to be found, f o r p a r t of the year a t l e a s t , upon the same r a n g e s -ranges t h a t are i n the main the product of f i r e . The wolf and coyote, too, of n e c e s s i t y must frequent these same are a s . I t w i l l be seen, t h e r e f o r , that any one s p e c i e s or range i s s u b j e c t , i n some measure, to the impact of a complex i n v o l v i n g range, h e r b i v o r e and p r e d a t o r ; no one f a c t o r can be d i v o r c e d from the consequences of i t s b i o l o g i c a l whole. 25 ELK AND THEIR INTER-RELATIONSHIP WITH PARK BIOTA MOVEMENTS AND FOOD HABITS OF ELK As s t a t e d e a r l i e r , e l k p l a n t i n g s i n B a n f f Park were most s u c c e s s f u l , and from a slow b e g i n n i n g the p o p u l a t i o n s expanded u n t i l by 1936-37 ( a c c o r d i n g to v e r b a l r e p o r t s ob-t a i n e d i n 194-9) the ranges adjacent to the areas of l i b e r a -t i o n were a l r e a d y h e a v i l y overgrazed,, C o n d i t i o n s worsened u n t i l 194-3, when the f i r s t e l k s l a u g h t e r was c a r r i e d out. N o n - s e l e c t i v e removal has been c a r r i e d out every w i n t e r s i n c e . But i n 194-9 the mature aspens "chawed" (barked by . gnawing of e l k ) to h e i g h t s of seven f e e t , and the d e t e r i o r -ated c o n d i t i o n , or complete absence, of browse s p e c i e s bore mute testimony to past o v e r - p o p u l a t i o n by e l k , and gave c r e -dence to the thought t h a t a l l was not yet w e l l e c o l o g i c a l l y . Observations on e l k movements and f e e d i n g h a b i t s were begun on g May, 194-9, i n the Bow V a l l e y from Banff to Elsenhower, and continued d a i l y u n t i l l 6 May. I n d i v i d u a l o b s e r v a t i o n s l a s t e d from a few minutes to h a l f an hour, and were c a r r i e d out at v a r y i n g p e r i o d s of the day, though l a r g e l y d u r i n g l a t e a f t e r n o o n and evening. E l k numbering from one to t h i r t y were observed. During t h i s p e r i o d , o n l y one i n s t a n c e of browsing was noted; t h i s the c a s u a l browsing of dwarf b i r c h ( B e t u l a glandulosa) by one e l k (of a group of 26 t h i r t y ) d u r i n g the evening of 11 May, A male e l k was seen e a t i n g hark o f f a f a l l e n p o p l a r about 15:^5 hours on 10 May, During l 6 May, three b u l l e l k f e d i n the t y p i c a l moose h a b i -t a t of F i r s t and Second V e r m i l i o n Lakes area, but were d e f i -n i t e l y g r a z i n g . Seventeen May, twenty-six e l k were seen i n the 17 M i l e a r e a , f o u r o f them out i n the boggy, willow-type f l a t , but no f e e d i n g o b s e r v a t i o n s were p o s s i b l e . P r a c t i c a l l y a l l the a d u l t b u l l s had l e f t the w i n t e r range, and the females and young males were d i s p e r s e d i n t o s m a l l groups. From 17 May to 6 June, two hundred " e l k minutes" of o b s e r v a t i o n were accu-mulated; o n l y g r a s s and f o r b s were eaten by e l k . By the end of t h i s p e r i o d m i g r a t i o n to summer range was a p p a r e n t l y com-p l e t e d , and onl y a few females and an o c c a s i o n a l male remained on the winter range between Banff and Elsenhower. On 17 June, a cow e l k was seen south o f the lake i n the 17 M i l e F l a t r e g i o n . During the f i v e minutes i t was observed, I t searched out and thoroughly browsed a w i l l o w . Subsequent examination showed the ar e a to be completely grazed out, with food almost u n o b t a i n a b l e . I t seemed almost c e r t a i n t h at acute need f o r c e d t h i s c h o i ce of food upon the animal, which appeared to be f e e d i n g a c a l f . The same day a cow and c a l f were seen i n the t y p i c a l moose ar e a a t Muleshoe Lake, but were observed only to graze d u r i n g the ten minutes they were watched. On 25 June, i n the l a t e evening, two b u l l s and two cows were observed i n the 17 M i l e F l a t a r e a. The cOws were watched c l o s e l y f o r f o r t y minutes and were seen to browse 27 willows both c a s u a l l y and d e l i b e r a t e l y d u r i n g t h i s p e r i o d . F u r t h e r o b s e r v a t i o n s were d i f f i c u l t , due to darkness, but judging from the p o s i t i o n s o f the animals* heads, i t was con-s i d e r e d t h a t they browsed f i v e t o t e n per cent o f the time. Subsequent i n v e s t i g a t i o n i n d i c a t e d that sedges In the are a were s u f f i c i e n t l y h i g h t o i n v a l i d a t e p a r t o f t h i s d a t a . The f o u r t h o f J u l y , e i g h t cows and f o u r c a l v e s were observed g r a z i n g on the f l a t south of Second V e r m i l i o n Lake, and the same day ten a d u l t s and f o u r c a l v e s were seen g r a z i n g near 12 M i l e area west of Banff. On 25 J u l y , twenty e l k , b e l i e v e d to be females, were seen g r a z i n g between 10 M i l e and 11 M i l e i n l a t e evening. Three days l a t e r one cow was seen g r a z i n g i n t h i s a r e a . Throughout t h i s p e r i o d female e l k were known to be on the v a l l e y f l o o r of the Spray. Warden W. C h i l d r e p o r t e d 29 June t h a t he knew of a mixed herd between Goat Creek c a b i n and B a n f f . Cows were seen between Goat Creek c a b i n and 15 M i l e c a b i n on 25 J u l y , 29 J u l y and' 13 August, but no a c t u a l food o b s e r v a t i o n s c o u l d be made. A check of the area r e v e a l e d no evidence of rec e n t browsing of w i l l o w or dwarf b i r c h . Throughout t h i s p e r i o d , of course, e l k were observed upon the summer ranges of the Spray area. Observations by b i n o c u l a r i n d i c a t e d they were g r a z i n g . S i m i l a r l y , e l k were seen on the upper gra s s ranges o f the Cascade r e g i o n throughout the sum-mer. Warden E. C a r l e t o n r e p o r t e d that a herd o f twenty to t h i r t y e l k grazed f r e q u e n t l y around a swamp In F l i n t ' s Park 28 area d u r i n g the summer. On 13 August, t w e n t y - f i v e cows and c a l v e s were r e p o r t e d to be on the v a l l e y f l o o r about h a l f a mile i n s i d e Cascade gate, and on the 15th a mixed herd of 108 e l k was seen i n the wi l l o w f l a t on the v a l l e y f l o o r south of Cuthead. I t was not found p o s s i b l e to l a t e r v i s i t t h i s area to make food d e t e r m i n a t i o n s . E a r l y on 22 August, a herd of twenty e l k (two males, s i x c a l v e s and twelve cows.) were observed f o r twenty minutes o u t s i d e the c a b i n at H i l l s d a l e . They were seen to browse the short w i l l o w q u i t e d e l i b e r a t e l y . The two males were the most a v i d browsers, but the others browsed at l e a s t f i f t y per cent of the time. L a t e r examination of the approximately two acres concerned showed t h a t t o t a l a v a i l a b l e graze had been u t i l i z e d f o r t y to f i f t y per cent, and w i l l o w s t h i r t y to f o r t y per cent. Willows u t i l i z e d had f i f t y per cent o f the l e a d e r s eaten back from f o u r to ten i n c h e s . I t was noted at t h i s time that grasses and f o r b s were brown and v e r y dry, the r e s u l t of the dry season and normal f a l l c u r i n g . G e n e r a l l y , i t may be s a i d t h a t e l k were seen or r e p o r t -ed from almost every a r e a of the Park throughout the summer. On 22 August, seven b u l l s were seen, i n the upper reaches of Johnston Creek V a l l e y , moving r a p i d l y south toward the Bow— the f i r s t evidence of f a l l m i g r a t i o n f o r the r u t . The f i r s t b u g l i n g was r e p o r t e d about the middle of August but i t was not u n t i l 29 August t h a t i t was heard r e g u l a r l y d u r i n g the l a t e 29 evenings and throughout the n i g h t s . T h i s was l a t e r than the normal, a c c o r d i n g to competent o b s e r v e r s . No attempt was made to l i s t a l l p l a n t s p e c i e s u t i l i z e d by the e l k . W i l d l i f e l i t e r a t u r e i s r i c h w i t h s t u d i e s of e l k food h a b i t s , and the range problems pursuant t h e r e t o . Green (194-6:30-32) l i s t s the main g r a s s e s , shrubs and f o r b s of the Bow V a l l e y area, and notes those u t i l i z e d by e l k . Murie (1951:195-2525)» i n h i s monumental book on e l k , a l s o d e a l s In d e t a i l w i t h e l k foods and food h a b i t s . I t was noted, however, th a t shrubby c i n q u e f o i l ( P o t e n t l l l a f r u t l c o s a ) was completely untouched even i n h e a v i l y overbrowsed areas. Bearberry ( A r c t o s t a p h y l u s u v a - u r s l ) and b u f f a l o b e r r y (Shepherdia canaden-s i s ) showed l i t t l e evidence of e l k browsing, though c i t e d by both Green and Murie as e l k foods. Gaffney (194-1:444-) confirms t h i s l a c k of use of these three p l a n t s p e c i e s , except under extreme du r e s s . The v e r s a t i l i t y of the e l k i s so great that such minor v a r i a t i o n s i n food h a b i t are p r o b a b l y af l i t t l e importance, except where a f f e c t i n g p l a n t indices.. The obser-v a t i o n s r e g a r d i n g shrubby c i n q u e f o i l were c o n s i d e r e d s i g n i f i -cant i n t h i s l a t t e r r e s p e c t , s i n c e t h i s s p e c i e s f a l l completely to p o i n t up range c o n d i t i o n s i n B a n f f N a t i o n a l Park (see Murie, 1951:231). 30 VITAL STATISTICS OF ELK I t i s probable t h a t , s i n c e the 19^3 e l k s l a u g h t e r at l e a s t , b e t t e r r e c o r d has been kept of the Banf f Park e l k than of any other comparable herds on the c o n t i n e n t . I t was f e l t , t h e r e f o r e , t h a t l i t t l e f u r t h e r d a t a were needed, other than c a s u a l day to day counts as o p p o r t u n i t y p r o v i d e d . Such obser-v a t i o n s were somewhat spurred by e a r l y r e p o r t s of wardens that the c a l f crop was very low, and some e f f o r t to t a l l y Bow V a l l e y e l k was made. The re c o r d s kept from the time the f i r s t c a l v e s were seen (see Table I) I n d i c a t e d a cow to c a l f r a t i o o f 3:1. No attempt.was made to r e l a t e c a l v e s to t o t a l number of e l k seen. Warden W. Black estimated a ten per cent c a l f crop In the North Saskatchewan area, but gave no a c t u a l f i g u r e s . I t i s not thought that any d u p l i c a t i o n i n c a l f counts was made, but c e r t a i n l y some d u p l i c a t i o n i n y e a r l i n g counts o c c u r r e d . Twenty-four y e a r l i n g s i n a l l were seen, but h i g h e s t day counts f i x the f i g u r e at twelve. In a l l about s i x t y e l k remained on the Bow V a l l e y w i n t e r range throughout the summer. A c t u a l l y , l i t t l e r e l i a n c e i s p l a c e d upon these summer counts made on the lower ranges, since d r y cows may or may not be present, and c a l v e s are most d i f f i c u l t to see. Rather, the f i g u r e s g i v e n by Green (unpublished r e c o r d s ) f o r the y e a r s 1944-51 are co n s i d e r e d best i n d i c a t i o n of herd increment and are noted In Table I I . I t i s i n t e r e s t i n g to note t h a t the c a l f t o t o t a l herd r a t i o i n 1949 was 1:3. T i l i s confirms the 31 Table I . Sight r e c o r d s of e l k cows and c a l v e s i n Bow V a l l e y , Banff N a t i o n a l Park, i n 194-9. Date Number of a d u l t females seen Number of c a l v e s seen P l a c e Remarks 15 June 13 7 Muleshoe Lake Not ac c u r a t e due to d i s t a n c e 11 12 M i l e a r e a Observed c l o s e l y 17 June :.2 1 17 Mile, F l a t 1 l Muleshoe Lake 4- J u l y 8 4- Second V e r m i l i o n Lake Seen at d i s t a n c e 8 4- 12 M i l e area May have been more c a l v e s 18" J u l y 4- 4- 17 M i l e F l a t Observed c l o s e l y 25 J u l y 20 (#0 10-11 M i l e area In woods, may have been c a l v e s 28 J u l y 1 11 M i l e 13 August 1 1 Sheep L i c k , V e r m i l i o n Lakes 22 August 12 6 H i l l s d a l e Close count 32 b e l i e f t h a t many c a l v e s are missed i n a summer count. Table I I . E l k c a l f and y e a r l i n g counts (percentage of t o t a l herd) f o r Banff N a t i o n a l Park. Year Calves Percent o f t o t a l herd Y e a r l i n g s Percent o f t o t a l herd 19*44 23 13 19*4-5 21 11 19*4-6 11 -7 19*4-7 24 8 19*4-8 10 9 19*4-9 25 7 1950 8 16 1951 22 5 I t i s not intended to d i s c u s s f u r t h e r the i m p l i c a t i o n s of a 1:3 c a l f to t o t a l herd r a t i o . T h i s r a t e compares very f a v o r -a b l y w i t h the f i g u r e s g i v e n by Murle (1951:138-1*4-3). Cowan (19*4-3:16) and Green (19*4-6:10-11) a l s o have d i s c u s s e d the r e p r o d u c t i v e success at some l e n g t h . However, the c o n s i d e r -able l o s s , shown i n Table I I , from c a l v e s to y e a r l i n g s Is s i g n i f i c a n t , and as yet not s a t i s f a c t o r i l y e x p l a i n e d . The answer may w e l l f i t i n w i t h the g e n e r a l u n s a t i s f a c t o r y eco-l o g i c a l changes pursuant to e l k a c t i v i t i e s i n the Park. 33 ELK LOSS FACTOR Loss f a c t o r s are very w e l l d i s c u s s e d by G-reen i n h i s work on e l k of Banff Park (Green* 194-6:11-15). He covers mechanical d e s t r u c t i o n , p r e d a t o r s , c a l f and w i n t e r l o s s e s , and l o s s e s due to d i s e a s e , p a r a s i t i s m , a c c i d e n t and g e n e r a l f a c -t o r s . To b r i n g the l o s s e s due to n o n - s e l e c t i v e removal up to date, Green*s f i g u r e s ( i b i d . 12) are g i v e n i n Table I I I , w i t h a d d i t i o n s from h i s notes. I t i s i n d i c a t e d t h a t 1,4-59 e l k were removed by the s p r i n g of 195G» F u r t h e r r e d u c t i o n s have been made, but the f i g u r e s are not yet to hand. Table I I I . E l k removed by s l a u g h t e r from B a n f f N a t i o n a l Park. Year Number removed 194.3 - 4-4- 74-1944 - 4-5 200 194-5 - 4-6 352 19I4.6 - 4-7 321 194-7 - 4-g 257 1 194-8" - 4-9 93 194-9 r- 50 162 3**-Wolf p r e d a t i o n was c o n s i d e r e d important, and checks were made i n 19*4-9 to determine the occurrence of these p r e -d a t o r s . Warden W. Black saw two wolves i n the North Saskatchewan area i n August, and s t a t e d there were more i n h i s d i s t r i c t . He k i l l e d s i x i n 19*4-8. Wardens E. Young and J . Woledge r e p o r t e d a pack of f o u r t e e n was r e s i d e n t i n the YaHaTinda area i n 19*4-8, and was presumably s t i l l p r e s e n t . On 2 J u l y , a,female and three pups were seen by Warden E, C a r l e -ton near Cuthead i n the Cascade V a l l e y , and two pups were k i l l e d . On 15 August one wolf was seen i n the same ar e a . R e t i r e d Warden U. U. LaCasse s t a t e d that' there were nine wolves i n the Johnston Creek a r e a but d u r i n g the summer of 19*4-9 one was seen up the creek on 2*4- June (Woledge), f o u r were seen i n the same area 25 June (Young), two were seen near P u l s a t i l l a Summit on 20 J u l y (Woledge), and one comparatively f r e s h s c a t was seen on E d i t h Pass on 11 Augus t . During June two t o u r i s t s (women) r e p o r t e d two wolves seen from the h i g h -xiray near Johnston Creek. One o l d t r a c k , p o s s i b l y a w o l ^ s , was seen along. C a r r o t Creek d u r i n g the summer. .Warden W. C h i l d saw a wolf on 11 J u l y at 15 M i l e c a b i n i n the Spray V a l l e y . ' E l k l o s s to wolves was a major i n t e r e s t i n t h i s study. Opinion was d i v i d e d , among members bf the Park s t a f f , as to whether e l k or moose f e l l more r e a d i l y prey t o wolves. One warden f e l t there was no d i f f e r e n c e , but that the l e s s e r num-bers of moose made t h e i r l o s s e s more s i g n i f i c a n t . A l l f e l t 35 t h a t p r e & a t i o n upon e l k was s i g n i f i c a n t , however. Cowan s t a t e d i n 194-7 "that e l k c o n t r i b u t e d f o r t y - s e v e n per cent of the b i g game d i e t of wolves i n Banfif and J a s p e r Parks (Cowan, 194-7; 173)* T n e wolves have i n c r e a s e d i n Banff Park since the time of h i s s t u d i e s , so the g e n e r a l o p i n i o n r e g a r d -i n g p r e d a t i o n may be sound. However, no k i l l s d i r e c t l y a t t r i -b utable to wolves were seen i n 194-9; the c a r c a s s e s noted were beyond adequate i n t e r p r e a t i o n as to cause of death. T h i s p r e d a t o r - p r e y r e l a t i o n s h i p p o i n t s up a p o s s i b l e i n t e r e s t i n g s i t u a t i o n . E l k were p l a n t e d i n the Park p r i o r to the advent of the wolf, when no major c u r s o r i a l p r e d a t o r was p r e s e n t i n s i g n i f i c a n t numbers. In 1935 cougar c o n t r o l was c a r r i e d out over one of the main e l k c o n c e n t r a t i o n areas, and by 194-2-4-3, when the f i r s t wolves made t h e i r appearance, the e l k had a l r e a d y reached a p o p u l a t i o n h i g h r e q u i r i n g mechan-i c a l removal. No evidence has been seen i n the r e c o r d s , or throughout t h i s study, to suggest t h a t wolves at any time a f f e c t e d d e t r i m e n t a l l y the e l k p o p u l a t i o n (see a l s o Cowan, 194-7:172); r a t h e r the e f f e c t may have been b e n e f i c i a l . Well meant but u n o f f i c i a l k i l l i n g o f wolves has tended to perpe-tuate t h i s s i t u a t i o n . Gaffney (194-1:4:39), In a paper on e l k i n the F l a t h e a d , Montana, l i s t s the almost complete extermina-t i o n of cougajp i n that area as a c o n t r i b u t i n g f a c t o r to e l k o v e r p o p u l a t i o n . In the North Saskatchewan r e g i o n , however, where e l k f i r s t became f i r m l y e s t a b l i s h e d about 194-0-4-1 and wolves entered the area i n 194-2-4-3, the e l k herd seems to be 36 a d j u s t e d to i t s environment, and showing on l y moderate popu-l a t i o n i n c r e a s e . I t i s too presumptuous to s t a t e that wolves have been s o l e l y r e s p o n s i b l e f o r t h i s s i t u a t i o n ; but t h e i r presence has done no apparent harm and may indeed be b e n e f i -c i a l . At the present time, i n f a c t , any normal l o s s e s (other than by e p i z o o t i c s ) may be c o n s i d e r e d b e n e f i c i a l t o the e l k p o p u l a t i o n s g e n e r a l l y In the Park. ELK RANGES I t was apparent from the o u t s e t o f the 194-9 s t u d i e s that the e l k were t h e i r own worst enemies. In May i t w&a estimated, without p r e v i o u s knowledge of the range, that the B a n f f - H i l l s d a l e -Elsenhower a r e a had a grass c a r r y o v e r of about t w e n t y - f i v e per cent on the v a l l e y f l o o r . -This was i f u r t h e r u t i l i z e d b e f o r e the new growth c a r r i e d the s t i l l r e s i d e n t e l k , and f i n a l c a r r y o v e r g e n e r a l l y was about twenty per cent. T h i s f i g u r e , agreed w i t h Green's 194-9 r e p o r t on ranges, wherein a f i n a l c a r r y o v e r of twenty per cent was estimated ( t y p e w r i t t e n r e p o r t made a v a i l a b l e ) . H i l l s i d e s , p a r t i c u l a r l y south s l o p e s , showed v e r y heavy overuse. One such i n the H i l l s d a l e area showed n i n e t y - f i v e per cent p l u s usage f o r the f i r s t one t h i r d o f the slope, and one hundred per cent usage on up, wit h r o o t s and crowns of grass alone remaining. Hoof e r o s i o n v a r i e d from marked to s l i g h t a l o n g 37 the v a r i o u s s l o p e s . Areas of pin e g r a s s (Calamagrostis  rubescens) were r e l a t i v e l y untouched. On 12 May, a s i x f o o t wide 1 , 2 0 0 y a r d reconnaissance t r a n s e c t was made through H i l l s d a l e meadow from the cabin to the r a i l r o a d — t h u s i n c l u d i n g meadow and semi-open aspen ar e a s . Three hundred and eleven e l k p e l l e t groups, or one to every 7.7 square f e e t , were c o u n t e d . Only one dwarf b i r c h out of every hundred was found without a l l t e r m i n a l shoots eaten, a l l w i l l o w s were completely browsed, and no aspen r e p r o d u c t i o n ( s a p l i n g s or s e e d l i n g s ) was noted. On 13 May two s i m i l a r t r a n s e c t s were run, one from H i l l s d a l e c a b i n to Red E a r t h Creek road, and one from the road west to the f a r end of the south H i l l s d a l e meadow. On the f i r s t t r a n s e c t of 1 2 0 0 yards, through meadow, open aspen woods and 5 0 - 5 0 aspen and spruce, the count was again one group of p e l l e t s to every 7»7 square f e e t . Again a l l w i l l o w s were browsed completely and aspen r e p r o d u c t i o n was wanting. The second t r a n s e c t , of 1 6 0 0 yards, mainly through meadow, showed one group o f droppings to every 2.4- square f e e t . Square metre quadrates were c l i p p e d i n the n o r t h and south H i l l s d a l e meadows i n May, as were s e v e r a l a l s o a l o n g the highway to t e s t e l k pressure i n wooded and/or pine grass areas. A comparable set was c l i p p e d i n August, a f t e r comple-' t i o n of summer growth. The o r i g i n a l quadrates i n the n o r t h meadow were a l s o r e c l l p p e d to t e s t the e f f e c t s of one hundred per cent w i n t e r u t i l i z a t i o n upon the ensuing year's growth. 3* T o t a l s r e g i s t e r e d are shown i n Table IV. A l l weights are f o r a i r - d r i e d c o n d i t i o n . The f i g u r e s shown f o r the p l o t s a l o n g the highway confirm the b e l i e f t h a t deep woods and p i n e grass a s s o c i a t i o n s were r e l a t i v e l y unused by e l k f o r f e e d i n g areas, since the f a l l c l i p s s t i l l i n c l u d e d a major p o r t i o n of the past w i n t e r ' s c a r r y o v e r . I f the o r i g i n a l e stimates of s p r i n g c a r r y o v e r were ac c u r a t e (as there i s reason to b e l i e v e they were) then the growth i n 1949 was down bad l y . T h i s was b e l i e v e d to be the case, judged from v i s u a l o b s e r v a t i o n . The y i e l d from the r e - c l i p p e d p l o t s p o i n t e d up the harm to be done by d r a s t i c o v e r - u t i l i z a t i o n . C l a r k e and a s s o c i a t e s , i n an e c o l o g i c a l and g r a z i n g c a p a c i t y study on the p r a i r i e s , recom-mend a f o r t y - f i v e per cent c a r r y o v e r of the p r i n c i p a l forage s p e c i e s ( C l a r k e et a l , 1942:20). Hunter, i n Colorado, b e l i e v e s t h at a twenty per cent c a r r y o v e r of g r a s s e s and weeds i s s u f f i c i e n t (Hunter, I945i23g>). The range o b s e r v a t i o n s made i n Banff Park suggest that Hunter's f i g u r e a l l o w s no margin of s a f e t y , and that t h i r t y per cent to f o r t y per cent c a r r y -over at l e a s t i s to be d e s i r e d . Late r a i n s i n J u l y brought some r e l i e f to the ranges, but by l a t e August the grass areas were s t i l l i n u n s a t i s f a c t o r y c o n d i t i o n . G e n e r a l l y , t h i s s i t u a t i o n a p p l i e d to a l l grass ranges from Banff to Eldon; some areas were poorer than at H i l l s d a l e . The shrubs i n the H i l l s d a l e meadow a r e a were e x t r e -mely overbrowsed. Average h e i g h t of dwarf b i r c h was f i f t e e n to e i ghteen i n c h e s , and of willows twelve to eighteen inches,. 39 Table IV, Data on square metre quadrates c l i p p e d i n H i l l s d a l e area, Banff N a t i o n a l Park'. Area Number S p r i n g Comparative Per cent Estimated of p l o t s c l i p f a l l c l i p s p r i n g per cent c l i p p e d (gms.) (gms.) c a r r y o v e r s p r i n g i n terms c a r r y o v e r of f a l l growth Along highway South meadow North meadow 7 10 10 10 476.5 732.8 999.1 2116.0 1023.4 1788.1 1007.1 65.0. 47.2 20 60.5 ; 20 r e - c l i p * * 0 r i g l n a l ten p l o t s i n North meadow r e - c l i p p e d to t e s t e f f e c t of one hundred p e r cent u t i l i z a t i o n . Most w i l l o w s appeared to be n e a r i n g e x t i n c t i o n — a s i t u a t i o n u n i v e r s a l - t h r o u g h o u t the area Banff to E l d o n . Wet or semi-wet ground willows showed b e t t e r growth, w i t h h e a v i l y browsed l i v e stems to h e i g h t s o f three to f o u r f e e t ; a l l growth above t h i s was k i l l e d . I t was estimated t h a t browse u t i l i z a t i o n was e i g h t y per cent to one hundred per cent. I t was noted, by p e l l e t o b s e r v a t i o n s , t h a t except i n a few r a r e i n s t a n c e s there appeared to be a d i s t i n c t d i v i d i n g l i n e between areas of e l k and moose u t i l i z a t i o n . T h i s l i n e o c c u r r e d a t the edge of a l l moist or boggy a r e a s . Thus i n 40 lowland meadows the u s u a l overbrowsed c o n d i t i o n common to e l k range would p r e v a i l , yet small boggy w i l l o w patches would be completely untouched by e l k j though h e a v i l y browsed by moose. T h i s was p a r t i c u l a r l y noted i n the 10 to 12 M i l e area west of Banff and i n the. 17 M i l e meadow. In the l a t t e r case, the a r e a completely surrounding the boggy meadow, and i n c l u d -i n g the l a k e , was d e s t r u c t i v e l y browsed by e l k yet the ea s t e r n two t h i r d s o f the meadow., covered g e n e r a l l y by w i l -lows up to two f e e t i n he i g h t and over two to three a c r e s by w i l l o w s f i v e f e e t In h e i g h t , was almost untouched, except by moose. E l k t r a c k s were seen p a s s i n g through t h i s a r e a but no d i r e c t evidence of browsing was seen. Perhaps the most s t a r t l i n g e f f e c t o f e l k upon t h i s range was the almost complete removal of aspen r e p r o d u c t i o n throughout the Bow r e g i o n . Aspen s e e d l i n g s and s a p l i n g s were to a l l i n t e n t s and purposes completely absent from the imme-d i a t e v i c i n i t y of a l l favored graze areas, and r e p r o d u c t i o n was inadequate to pre s e r v e the stands. I t was noted t h a t e l k browsing grew p r o g r e s s i v e l y l e s s as one moved back from " p r e f e r r e d " g r a z i n g a r e a s . Check p l o t s showed that s i x t y -s i x per cent o f the aspens immediately a d j a c e n t to the H i l l s -d a l e meadow were "chawed", many of them to h e i g h t s o f seven f e e t . Two hundred yards back o n l y f i f t y per cent were, "chawed". August checks i n d i c a t e d t h a t young s e e d l i n g s d i d get up to a he i g h t of s e v e r a l inches by f a l l , but they presumably were eaten completely back again each w i n t e r and s p r i n g . 4-1 Overpopulation by e l k brought about the o r i g i n a l aspen de-t e r i r o a t i o n i n c o n j u n c t i o n w i t h extreme range overuse. I t would appear that even reduced herds now remove a l l seed-l i n g s , p r e b a b l y i n c i d e n t a l l y while g r a z i n g . In the Queens Park a r e a a c o n s i d e r a b l e stand of aspens, a p p a r e n t l y "chaw-ed" a t an age when they were Just too b i g to be broken down, was observed to be going down under the w i n t e r snows. L i t t l e regrowth was seen i n t h i s area and i t i s e v i d e n t i t w i l l not s u r v i v e . Re-examination o f the browse s p e c i e s i n August i n d i -cated n e a r l y one hundred per cent r e c o v e r y from the s p r i n g c o n d i t i o n . However, t h i s was s t i l l far,below the p o t e n t i a l . To check t h i s , d ata were d e r i v e d from the Index p l o t about ten m i l e s west of Banff, and noted i n Green's work on e l k (194-6:27) as P l o t number 1, Sub-area A, U n i t area 2 — f e n c e d i n A p r i l , 194-4-, when u t i l i z a t i o n of dwarf b i r c h and w i l l o w was e i g h t y p e r c e n t . The area e n c l o s e d mature aspen, so was t y p i c a l of the area. When checked, the o u t s i d e area showed the t y p i c a l c o n d i t i o n — d w a r f b i r c h and w i l l o w up to f o u r t e e n -twenty inches i n h e i g h t and aspen r e p r o d u c t i o n n i l . W i t h i n the p l o t , f i v e dwarf b i r c h clumps averaged s i x f e e t In h e i g h t and w i l l o w s averaged three to f i v e f e e t . Aspen r e p r o d u c t i o n was as f o l l o w s : s i x t e e n s a p l i n g s averaging seven f e e t , n ine averaging three to f o u r f e e t and f i f t e e n f i r s t year s e e d l i n g s . I t i s not known i f Hunter (194-5:238) Is a g a i n b o r d e r i n g the c r i t i c a l p o i n t i n range usage when he recommends a f i f t y per cent c a r r y o v e r of c u r r e n t years browse growth. But the com-p a r i s o n s noted above c l e a r l y i n d i c a t e the degrading e f f e c t of e i g h t y per cent to one hundred per cent browse u t i l i z a t i o n . Before l e a v i n g the d i s c u s s i o n o f t h i s e,lk range, some mention must be made of another f a c t o r working i n the n o r t h H i l l s d a l e meadow. Ground s q u i r r e l s ( C l t e l l u s columblanus  columbianus) were found to be most numerous. Invas i o n by l a r g e rodents Is a commonly accepted index of over-grazed range; t h i s a r e a proved no e x c e p t i o n . Whether these ground s q u i r r e l s w i l l e v e n t u a l l y b e n e f i t the range by e a t i n g rhe u n d e s i r a b l e p l a n t s p e c i e s , as i s suggested by Bond (194-5:233), i s a moot, q u e s t i o n . By f a l l I t was observed that every "gopher" hole was surrounded by an area eaten completely b a r e — f o r an average r a d i u s of three f e e t . I t was estimated i n 194-9 that u n l e s s they were c o n t r o l l e d , the c a r r y i n g capa-c i t y of the n o r t h meadow would be lowered s i g n i f i c a n t l y . Under present circumstances, t h i s would, I t Is b e l i e v e d , outweight any p o s s i b l e good achieved. The range from Banff to the East Gates was checked, and the s i t u a t i o n found to be e s s e n t i a l l y the same as that a l r e a d y d e s c r i b e d — w i t h r e c o v e r y even l e s s than at H i l l s d a l e , , D e s t r u c t i v e browsing of w i l l o w growth was found at the entrance to the gorge l e a d i n g to the C a r r o t Creek summer range. T h i s , however, was b e l i e v e d to be due to a b u i l d up of e l k p r i o r to a ru s h through the gorge, or p o s s i b l y because of de l a y engen-dered by unfavorable c o n d i t i o n s w i t h i n i t . T h i s 4-3 i n t e r p r e t a t i o n was strengthened by the f a c t t h a t w i l l o w s ( S a l i x subcoerulea P i p e r = S a l i x Drummondlana B a r r a t v a r , subcoerulea P i p e r ) and b i r c h ( B e t u l a pumlla L. v a r . glandu-l l f e r a Regel.) on up the creek n o r t h of the gorge were p l e n -t i f u l and i n good c o n d i t i o n — t h e y showed o n l y moderate browsing by moose ( p e l l e t d i a g n o s i s ) . The C a r r o t Creek sum-mer range showed the d e s t r u c t i v e e f f e c t of "chawing" upon the aspens, though graze c o n d i t i o n s were f a i r . T h i s suggested t h a t t h i s area was used, on o c c a s i o n at l e a s t , as a w i n t e r range. The Spray R i v e r e l k ranges, were i n s p e c t e d s e v e r a l , times d u r i n g June, J u l y and August. They were g e n e r a l l y i n e x c e l l e n t c o n d i t i o n , with the south-east s e c t i o n of the upper range showing d e t e r i o r a t i o n by l a t e summer. Some of the r i d g e s of sharp repose a l s o showed d e c l i n e i n range c o n d i t i o n . However, s i n c e t h i s v a l l e y r e c e i v e s more p r e c i p i t a t i o n than the Bow area, there was no reason seen to c o n s i d e r the ar e a overpopulated. The v a l l e y f l o o r c a r r i e d o n l y a l i m i t e d amount of forage, but no i n d i c a t i o n of summer browsing by the then r e s i d e n t e l k was seen. However, heavy wintter browsing of w i l l o w on the low, open f l a t s , w i t h u t i l i z a t i o n e i g h t y per cent to n i n e t y p e r cent and average w i l l o w h e i g h t eighteen to twenty-four inches, was noted. I t was i m p o s s i b l e to d e t e r -mine whether t h i s was the work of e l k or moose, but i t was probably caused by both. Regrowth i n August appeared to be e x c e l l e n t , i n terms of the s p r i n g c o n d i t i o n . 44 Quite a d i f f e r e n t s i t u a t i o n was ev i d e n t on the Cascade ranges. These areas, c h i e f l y invaded h i g h - a l t i t u d e sheep ranges, were seen to be i n poor c o n d i t i o n throughout the sum-mer. I n c i p i e n t e r o s i o n was q u i t e e v i d e n t . Since many of these ranges are on steep.westerly s l o p e s of the P a l l i s e r Range, e r o s i o n i s to be expected a f t e r o n l y moderate o v e r -g r a z i n g . T h i s , however, made the s i t u a t i o n no l e s s acute. Considerable browsing of the w i l l o w s on the Cascade v a l l e y f l o o r was noted, and q u e s t i o n i n g e l i c i t e d the i n f o r m a t i o n that i n the f a l l of 194-8 e l k were found on the v a l l e y f l o o r d u r i n g the s l a u g h t e r — t h e f i r s t time t h i s had been noted. Obviously, browsing must have o c c u r r e d s i n c e graze s p e c i e s were l i m i t e d . No s e r i o u s harm was yet evident i n 194-9,but as noted e a r l i e r , e l k i n some numbers were a g a i n on the v a l l e y f l o o r — t h i s time In August. I t must be a n t i c i p a t e d t h a t i f t h i s behaviour continues, s e r i o u s harm to the browse s p e c i e s must c e r t a i n l y ensue. A check on 29 August, of the F l i n t ' s Park and Windy ranges to the n o r t h showed them to be brown and i n poor c o n d i t i o n , with much bare ground showing. G e n e r a l l y , the e n t i r e range area of the Cascade gave the impression of low c a r r y i n g c a p a c i t y and impaired v i t a l i t y . The North Saskatchewan e l k ranges were g i v e n scanty check. Since the range l i e s i n a low p r e c i p i t a t i o n area, no l u s h growth was expected. However, the s e v e r a l short v i s u a l checks i n d i c a t e d no d e t e r i o r a t i o n of range. Low wil l o w growth on some of the r i v e r f l a t s had been h e a v i l y browsed, 4-5 probably by both e l k and moose. I t Is not b e l i e v e d that t h i s I n d i c a t e d any problem. Aspen growth and r e p r o d u c t i o n appeared to be normal on the e l k range, and "chawing" was n e g l i g i b l e . G e n e r a l l y , the range showed no s i g n s of over-p o p u l a t i o n by e l k . In c o n c l u s i o n , the e l k have shown a remarkable a b i l i t y to p i o n e e r i n t o and to occupy p r a c t i c a l l y a l l summer g r a z i n g ranges i n the Park. With the e x c e p t i o n of the North Saskatchewan range, where they have not yet reached a peak, and p o s s i b l y the Spray range, they have subsequently abused these summer ranges to the p o i n t where p r o g r e s s i v e . r a n g e d e t e r i o r a t i o n i s an accomplished f a c t . MOVEMENTS AND FOOD HABITS OF MOOSE S p e c i a l emphasis was l a i d upon the study of the moose p o p u l a t i o n i n the Bow V a l l e y from Banff west to the 17 M i l e meadow, and c o n s i d e r a b l e time was spent o b s e r v i n g movements and food h a b i t s . Check s t u d i e s were made i n s e v e r a l other areas, f o r comparison purposes and to d e t e r -mine, i f p o s s i b l e , i n f l u e n c e of presence or absence of e l k upon these f a c t o r s . O bservations commenced 85 May, 194-9, and on 12 May t h e . f i r s t moose were seen, two cows and one new-born c a l f , i n the V e r m i l i o n Lakes area. By 27 May the minimum number 46 ( l e s s c a l v e s ) known to have been i n the Banff to .17 M i l e area was twenty-one. Y e a r l i n g s c o n s t i t u t e d n i n e t e e n per cent of the t o t a l d u r i n g t h i s p e r i o d (see Table V I ) , but the l a s t y e a r l i n g disappeared IS May. I t was remarked by. Green t h a t the cows d r i v e the y e a r l i n g s away Just p r i o r to c a l v i n g , and these o b s e r v a t i o n s seemed to bear out the f a c t . T h i s was f u r t h e r shown by f a c t that l a t e i n June Warden W. C h i l d saw a dry cow s t i l l accompanied by a y e a r -l i n g , while on 4- J u l y Green noted a cow, w i t h a c a l f , v i g o r o u s l y a t t a c k i n g a y e a r l i n g t h a t a p p a r e n t l y claimed r e l a t i o n s h i p . By 18" June the p o p u l a t i o n i n the V e r m i l i o n Lakes-7 M i l e Beaver Dam-Muleshoe Lake a r e a alone was eighteen ( l e s s c a l v e s ) and on 20 June seventeen were c o u n t e d — f o u r -teen of them cows. By 2 J u l y there were n i n e t e e n moose i n the a r e a — f o u r of them d e f i n i t e l y y e a r l i n g s and three doubt-f u l . Thus the y e a r l i n g s had r e t u r n e d to the summer marshes and now c o n s t i t u t e d a t l e a s t twenty-one per cent of the t o t a l . On 3 J u l y there were f o u r t e e n moose pre s e n t ; by 8 J u l y only seven were seen. I t became apparent that the f l u c t u a t i o n s i n numbers bore g r e a t e r s i g n i f i c a n c e than mere i n a c c u r a c y i n o b s e r v a t i o n . This, was the more obvious s i n c e two grey females were known to have been present i n e a r l y May, m i s s i n g i n l a t e May and present a g a i n by mid-June. By 8 J u l y they were again absent. I t was a l s o n o t i c e d t h a t the moose d i v i d e d t h e i r f e e d i n g *7 time, i n the v a l l e y , between wet v e g e t a t i o n and dry w i l l o w and dwarf b i r c h . T h i s was d i s c u s s e d w i t h S p e c i a l Warden H, U. Green, and i t was concluded t h a t the moose must be moving i n and out of the Bow V a l l e y . I t appeared that they came i n from dry feed areas, s a t i a t e d themselves on wet food, and then r e t u r n e d to the upper d r y feed a r e a s . F u r t h e r i n -v e s t i g a t i o n showed that a p a r t y of g e o l o g i s t s , while a t the top of the c o l on Mt. Norquay on k J u l y , had seen a b u l l moose on top of the r i d g e (coming out of F o r t y m i l e Creek) and moving down toward V e r m i l i o n Lakes. On 10 J u l y a cow and c a l f were r e p o r t e d moving past the f o o t o f the s k i l i f t on Mt. Norquay—presumably i n or out of F o r t y m i l e . On 11 J u l y , a b u l l moose was f o l l o w e d f o r some f o u r hundred yards up the draw from Second V e r m i l i o n Lake toward the c o l p r e v i o u s l y mentioned. Subsequent examination of E d i t h Pass r e v e a l e d that the game t r a i l i n t o the F o r t y m i l e was w e l l used, wit h r e c e n t movement by moose In both d i r e c t i o n s apparent. No i n d i c a t i o n of any movement over the pass n o r t h of H i l l s d a l e was seen, but d i s c u s s i o n s w i t h Warden E. Royle from Healy Creek i n d i c a t e d that moose came down that v a l l e y , f e d i n the Bow marshes, and r e t u r n e d south up the creek a g a i n . As a r e s u l t of t h i s apparent s i t u a t i o n , the Waterfowl Lakes a r e a was s t u d i e d to determine i f t h i s movement might be common to other s e c t i o n s of the Park. Reports of e a r l i e r years i n d i c a t e d a l a r g e moose p o p u l a t i o n i n t h i s a r e a , I t being not uncommon to see up to ten animals pres e n t at one time. However, only three cows and one c a l f were seen d u r i n g s e v e r a l v i s i t s throughout the summer. I t was noted, and con-firm e d by Warden R. H a s k e l l , t h a t they d i d not stay permanent-l y a t j t h e Upper Lake, but came and went p e r i o d i c a l l y . Exam-i n a t i o n of the a r e a i n d i c a t e d l i t t l e movement to the n o r t h , along the M i s t a y a R i v e r toward the Howse R i v e r , but w e l l -worn t r a i l s south up to Mistaya Lake area were e v i d e n t . One cow was f o l l o w e d along t h i s route u n t i l , she reached the h i g h r ground east of M i s t a y a Lake. i t seemed, then, that here, as i n the Bow, a movement to and from wet v e g e t a t i o n was t a k i n g p l a c e . A number of samples of wet and underwater v e g e t a t i o n had been c o l l e c t e d at the V e r m i l i o n Lakes; a s i m i l a r c o l l e c t i o n was made here (see Table V ) . I n q u i r y e l i c i t e d the i n f o r m a t i o n that pond-weeds were once most p l e n t i f u l In Upper Waterfowl Lake; now they were extremely s c a r c e . I t was o n l y by wading or rowing downstream from a f e e d i n g animal t h a t samples ( l o s t from the mouth) were obt a i n e d . These proved to be two of the same spe-c i e s found i n the Bow V a l l e y . I t seemed probable that pond-weeds once drew moose to the area, but subsequent s c a r c i t y o f such foods, p l u s overbrowsing of adjacent w i l l o w growth, had d i s c o u r a g e d s i g n i f i c a n t summer m i g r a t i o n s and occasioned the seeming p a u c i t y of moose i n the area. In other words, there were probably Just as many moose as ever but they no longer congregated i n the one a r e a . F o l l o w i n g t h i s the Spray V a l l e y , from G-oat Creek to 4-9 Table V. Pond p l a n t s c o l l e c t e d from some moose summer ranges i n Banff N a t i o n a l Park i n 194-9? Second V e r m i l i o n Lake and a r e a — c o l l e c t e d d u r i n g J u l y Equisetum f l u v i a t i l e L. Equisetum p a l u s t r e L» HIppuris v u l g a r i s L. Myriophyllura exalbescens Fern. Potamogeton r i c h a r d s o n i l (Benn) Rydb. Potamogeton p e c t i n a t u s L. Ranunculus c l r c i n a t u s S i b t h . Second V e r m i l i o n Lake and a r e a — c o l l e c t e d d u r i n g August Ranunculus p u r s h i i R i c h . S a g g i t a r i a l a t i f o l i a W i l l d . (deep water form) Sparganum a n g u s t l f o l i u m Michx. U t r i c u l a r i a v u l g a r i s L» v a r . americana Gray Upper Waterfowl L a k e — C o l l e c t e d d u r i n g J u l y Drepanocladus sp. (moss) Potamogeton p e c t i n a t u s L. Seven M i l e Beaver D a m — c o l l e c t e d d u r i n g August Chara sp. E l e o c h a r i s a c i c u l a r i s R. and S. H i p p u r l s v u l g a r i s L. Potamogeton a m p l i f o l i u s Tuck. Potamogeton panoramitanus B i v . v a r . major G. F i s c h . Potamogeton p e c t i n a t u s L. Sparganum a n g u s t l f o l i u m Michx. U t r i c u l a r l s v u l g a r i s L. v a r . americana Gray V a l l i s n e r i a americana ? Michx. (i n t r o d u c e d ) ^ I d e n t i f i c a t i o n s by Dr. T. M. C. T a y l o r of U n i v e r s i t y of B r i t i s h Columbia. 50 the 15 M i l e c a b i n , was checked. A well-u s e d t r a i l was found along the w e s t e r l y s i d e of the v a l l e y , a p p a r e n t l y l e a d i n g to Sundance Pass (and the Bow V a l l e y ) . I t showed every evidence of f r equent use by moose,*and a h e a v i l y browsed s t r i p con-f i n e d t o a few f e e t on e i t h e r s i d e o f the t r a i l I n d i c a t e d t r a n s i e n t u t i l i z a t i o n . Thus i t seemed most probable t h a t Spray V a l l e y moose conformed to the p a t t e r n and p e r i o d i c a l l y f e d i n the V e r m i l i o n Lakes area. A s i m i l a r t r a i l and browse c o n d i t i o n was found along the Cascade R i v e r south of Cuthead. There seemed l i t t l e doubt that here again the same p a t t e r n obtained, w i t h the moose p o s s i b l y m i g r a t i n g to the Bow V a l l e y f o r wet v e g e t a t i o n . A reconnaissance of the Graveyard a r e a i n the n o r t h e r n s e c t i o n o f the Park i n d i c a t e d that moose f e d there i n the swamps and sloughs In summer. Animals were seen almost com-p l e t e l y submerged, f e e d i n g on pondweeds. P l a n t s p e c i e s appeared to be the same as i n the Bow area, though no c o l -l e c t i o n s were made f o r c o n f i r m a t i o n . Few o b s e r v a t i o n s were made i n t h i s d i s t r i c t , but past moose counts i n d i c a t e d move-ments s i m i l a r to the Bow and Waterfowl ar e a s . Observations i n the areas of Johnston and Baker Creeks, and Boom and Helen Lakes, i n d i c a t e d moose were prese n t d u r i n g the summer. No evidence o f r e g u l a r summer movements was noted i n the Johnston and Baker Creek areas, but r e g u l a r movement between Boom and A l t r u d e Lakes was i n d i c a t e d . S i m i l a r l y , i t seemed probable that movement was t a k i n g p l a c e between Helen Lake 51 and Hector Lake areas. Observations of winter movements of the moose were not, of course, o b t a i n a b l e d u r i n g t h i s study. But from i n q u i r y i t appeared that most of the moose d i s a p p e a r from the swamps w i t h the f i r s t appearance of i c e . Most of the wardens claimed that they go to the many t r i b u t a r y streams and v a l l e y s to winter; the o l d b u l l s going o f t e n to the summits or passes. There w i l l o w ( S a l i x v e s t l t a Pursh. and S. subcoerulea P i p e r = S. Drummondlana B a r r a t v a r . subcoe-r u l e a (Piper)) would be p l e n t i f u l , but snow f r e q u e n t l y deep. A few moose remain i n the w i l l o w brakes, around V e r m i l i o n Lakes, made a v a i l a b l e as the t o u r i s t season ends. Toward s p r i n g , too, l a r g e numbers of moose have been seen i n the 17 M i l e meadow area. However, Mr. LaGasse estimated the t o t a l p o p u l a t i o n from Banff to Lake L o u i s e , e x c l u s i v e of t r i b u t a r y v a l l e y s , at about f o r t y , and maintained that the l a r g e number ( r e p o r t e d l y f o r t y moose) seen i n the 17 M i l e meadow i n 1942 or k-J was not normal and so not i n d i c a t i v e of the l o c a l s i t u a t i o n . I t i s p o s s i b l e that the advent of the f i r s t wolf might have caused t h i s e f l u x t o the v a l l e y , or unusual snows might have occ u r r e d ; u n f o r t u n a t e l y no r e c o r d s are a v a i l a b l e f o r check. I t was not p o s s i b l e , from check of Wardens' d i a r i e s , to d e t e c t any s i g n i f i c a n t d i f -f e r e n c e i n the number of s i g h t o b s e r v a t i o n s of moose d u r i n g winter or summer, except on the summer c o n c e n t r a t i o n areas a l r e a d y mentioned. I t remains to be determined then, i f 52 any major win t e r m i g r a t i o n takes p l a c e , or i f moose merely d i s t r i b u t e themselves more evenly throughout the dry food areas d u r i n g that season. During the summer, s e v e r a l hundred "moose minutes" were spent o b s e r v i n g the food eaten by moose i n the Bow V a l l e y r e g i o n . They were observed to browse e n t i r e l y , e a t i n g w i l l o w s , dwarf b i r c h , b l a c k and white p o p l a r (Populus b a l s a m l f e r a L. and P. tremuloldes Michx.), and v a r i o u s other low shrubs, p l u s o c c a s i o n a l lodgepole pine t i p s , u n t i l the equisitum (Equlsitum f l u v i a t i l e L. and E. p a l u s t r e L.) became evident along the edge of the V e r m i l i o n Lakes about 26 May. T h i s l a t t e r food, then, was eaten a v i d -l y , i n c o n j u n c t i o n w i t h p e r i o d s of browsing willows ( S a l i x  Bebblana Sarg.). By 1 June pondweeds became a v a i l a b l e (see Table V i i ) , and remained the major a t t r a c t i o n of the V e r m i l i o n Lakes and 1 M i l e Beaver Dam area throughout the summer. P e r i o d i c browsing of w i l l o w continued throughout. No evidence of other than c a s u a l g r a z i n g of d r y grasses and f o r b s was seen, and t h i s appeared to be mainly a c c i d e n t a l . However, by 18 June some new sedge growth (Carex spp.) was being eaten, and l a t e r some slough grass (Beokmannia  Syzlgachne (Steiid) Fern.) was grazed s l i g h t l y i n the v i c i n i t y of the Beaver Pond. No attempt was made to determine r e l a -t i v e times spent, f e e d i n g upon the d i f f e r e n t s p e c i e s , but moose were seen to f e e d i n the V e r m i l i o n Lakes f o r over an hour at a time. At Upper Waterfowl Lake, a cow moose was observed f e e d i n g i n the g l a c i e r - f e d waters f o r j u s t over 53 two hours. When o b s e r v a t i o n s concluded the end of August, underwater and/or marsh v e g e t a t i o n was s t i l l b e i n g consumed i n c o n j u n c t i o n w i t h w i l l o w . Checks were made of Boom Lake, Cirque Lake and E d i t h Pass areas between 27 J u l y and l 6 August. I t was i n t e r e s t i n g to note that the on l y f r e s h browsing evident was upon F a l s e A z a l i a ( M e n z l e s i a f e r r u g l n e a Hook.). Willow and dwarf b i r c h showed moderate winter browsing, but the F a l s e A z a l i a appeared to be the f a v o r e d browse of the f l u i d summer p o p u l a t i o n s . VITAL STATISTICS OF MOOSE Observations on r e p r o d u c t i o n were made c h i e f l y i n the Bow V a l l e y area from Banff to Eisenhower. However, r e c o r d s were kept of a l l p e r s o n a l o b s e r v a t i o n s , and of r e l i a b l e r e p o r t s r e c e i v e d from v a r i o u s other r e g i o n s of the Park. The f i r s t c a l f seen was born 12 May i n the. V e r m i l i o n Lakes area, and the l a s t newborn c a l f was found 22 June. During t h i s pesripd f o u r -teen c a l v e s ( i n c l u d i n g one set of twins) were noted, while s i x t e e n cows were known to be i n the area. At l e a s t one of the three d r y cows was not yet of bree d i n g age. One a d d i t i o n -a l cow was known to be i n the 17 M i l e meadow a r e a a t t h i s time and i t i s b e l i e v e d she was dry. Thus there were f o u r t e e n c a l v e s f o r s i x t e e n cows or a 1:1.4 c a l f to cow r a t i o . 54 One cow and c a l f and one cow and y e a r l i n g were r e -p o r t e d i n the Spray V a l l e y . Two cows and c a l v e s were r e p o r t e d i n the Johnston Lake area, and two cows and c a l v e s were seen i n the Cascade a r e a . Three cows were seen a t Upper Waterfowl Lake s e v e r a l times but on l y one c a l f was noted; i t i s b e l i e v e d two of the cows were d r y . On the Howse R i v e r two cows were seen and both had c a l v e s . In two t r i p s past the Graveyard area f o u r cows and two c a l v e s were seen. I t was p o s s i b l e the same cow and c a l f were seen each time so the count had to be taken two cows and one c a l f . I t i s seen, however, t h a t the lowest c a l f - c o w r a t i o noted was 1:3. I t was not known what the summer movements o f dry cows were, but from o b s e r v a t i o n s of m i g r a t i o n s to pondweed areas t h e r e was no reason to b e l i e v e that they segregated themselves or avoided the l a k e s . There-f o r e the calf-cow r a t i o s observed were taken as a r e l i a b l e i n d i c a t i o n of the c a l f crop. For the same reason, i t was b e l i e v e d that the sex and age counts made i n the V e r m i l i o n Lakes area were a r e l i a b l e index of the composition of the moose p o p u l a t i o n i n the Park. By 27 May twenty-one moose ( l e s s calves), were known to have been i n the a r e a . By 18 June the p o p u l a t i o n p r e s e n t was e i g h t -een p l u s t en c a l v e s . T h e r e a f t e r o n l y day r e c o r d s were kept except i n the case of c a l v e s , and major counts are shown i n Table VI. As noted e a r l i e r , by 27 May the y e a r l i n g s ( t h r e e males and one female p o s s i b l e ) made up n i n e t e e n per cent of the t o t a l ( l e s s c a l v e s ) . On 2 J u l y the y e a r l i n g s c o n s t i t u t e d at l e a s t twenty-one per cent of the p o p u l a t i o n , w i t h a sex 55 Table VI. C l a s s i f i c a t i o n , by age and sex, of moose known t o be i n the V e r m i l i o n Lakes -17 M i l e meadow area, Banff N a t i o n a l Park, from May to J u l y , 1949. Date A d u l t Males Ad u l t Females Young Males Young Y e a r l i n g s Females Calves Unde-t e r -mined 27 May 2 7 7 2(1) 3(4) 4 l g June 3 12 2 10 1 2 J u l y 4 g (2) 7(5) 4 3 J u l y 3 g 3 6 4 J u l y 6 6 1 3 g J u l y 2 2 3 2 Obrackets indicate possible numbers where sex determinations i n doubt. r a t i o of f i v e males to two females, with two of the males pos-s i b l y two year o l d s . On 3 J u l y the y e a r l i n g r a t i o was one male to two females; on g J u l y two males to one female. Thus an approximate r a t i o of three males to two females was obser-ved i n y e a r l i n g s . Young males and females were recorded s e p a r a t e l y In e a r l y counts but lumped i n w i t h a d u l t s as the summer prog r e s s e d . J u l y f i g u r e s were c o n s i d e r e d most I n d i c a -t i v e f o r a d u l t sex r a t i o s , s i n c e d u r i n g the c a l v i n g season the cows p a r t i c u l a r l y s e l e c t e d the v a l l e y bottom. I t Is b e l i e v e d the r a t i o ran very c l o s e to 1:1. -In a l l , there was n o t h i n g 56 seen to i n d i c a t e an unbalanced or d e c l i n i n g p o p u l a t i o n . Green, from study of Wardens' r e t u r n s and p e r s o n a l observa-t i o n s , estimates the 1950 moose population, as 24-7. T h i s f i g u r e compares f a v o r a b l y w i t h i n t e r p r e t a t i o n o f Wardens* r e p o r t s f o r e a r l i e r y e a r s , and i n d i c a t e s a more or l e s s s t a t i c c o n d i t i o n s i n c e 194-5 (as opposed to the s i t u a t i o n o u t -sid e the Park i n A l b e r t a where moose have d e c l i n e d ) , (Green, unpublished r e p o r t . ) Cowan, i n h i s e a r l i e r s t u d i e s , found moose to be d e c l i n i n g . I t i s c o n s i d e r e d probable that moose in the Park reached a maximum p o p u l a t i o n d e n s i t y i n the l a t e 1 9 3 ° fs and then d e c l i n e d s l i g h t l y to a steady l e v e l about 194-5. MQOSE LOSS FACTORS Loss f a c t o r s w i t h r e s p e c t to moose have been ade-qu a t e l y covered by Cowan (194-3:18-19 and 1944:73) . The w i n t e r t i c k (Dermacentor a l b l p l c t u s ) appears to be of some importance. No s e r i o u s c o n d i t i o n was seen i n 194-9, hut from c o n v e r s a t i o n w i t h other persons i t appeared t h a t these t i c k s have caused, or have c o n t r i b u t e d t o , c o n s i d e r a b l e losses,, Predator l o s s e s d i d not appear s i g n i f i c a n t (see a l s o Cowan, 194-7:167), and QtUy two probable wolf k i l l s were noted. However, Wardens maintained that wolf k i l l s d i d occur, and 57 as s t a t e d e a r l i e r , o p i n i o n d i f f e r e d as to s e v e r i t y of preda-t i o n compared to t h a t upon e l k . I t should be a g a i n noted, however, t h a t s i n c e the moose p o p u l a t i o n was f a r below that of the e l k , n u m e r i c a l l y equal p r e d a t l o n upon both would have f a r g r e a t e r s i g n i f i c a n c e with r e s p e c t to the moose. Warden P. G-. Woodworth s t a t e d that he found f i v e dead moose i n the L i t t l e Pipestone country e a r l y In the s p r i n g o f 194-9, but he c o u l d not say they had been k i l l e d by wolves. Some l o s s e s of c a l v e s occur due to drowning, when the cows swim or wade acr o s s l a k e s too deep and too wide f o r t h e i r o f f s p r i n g . There Is an obvious l o s s , shown by the v i t a l s t a t i s t i c s , from c a l f to y e a r l i n g age. T h i s i s a t t r i b u t a b l e , i t i s b e l i e v e d , to n o n - s u r v i v a l of the f i r s t w i n t e r . I t was c o n s i d e r e d i n 194-9 t h a t the Park supported, i n most areas, the maximum num-bers of moose d e s i r a b l e f o r good management. I t seems probable, then, that w i n t e r c o m p e t i t i o n f o r food m i t i g a t e s a g a i n s t the c a l v e s to the p o i n t where, d e s p i t e h i g h r e p r o d u c t -i v e success, the p o p u l a t i o n i s o n l y able to m a i n t a i n the steady l e v e l noted. Thus the l o s s f a c t o r s , p r e s e n t l y not c r i t i c a l , p o i n t up a s i t u a t i o n of p o s s i b l e f u t u r e concern should range c o n d i t i o n s worsen. MOOSE RANGES I t was not p o s s i b l e to adequately d e a l w i t h the moose range from Banff to the 7 M i l e Beaver Dam a r e a i n the 58 Bow B a i l e y without c o n s i d e r i n g ' t h e combined e f f e c t s of beaver a c t i v i t y , w a t e r k i l l and moose browsing. Without doubt, the beaver, i n the p a s t , c o n t r i b u t e d l a r g e l y to the d e s i r a b l e sum-mer h a b i t a t and food found i n t h i s a r e a . However, t h e i r con-t i n u e d a c t i v i t i e s were co n s i d e r e d to be somewhat of a mixed b l e s s i n g . They d o u b t l e s s maintained the e x i s t e n t water l e v e l s , and were t h e r e f o r i n s t r u m e n t a l i n r e t e n t i o n o f the e x c e l l e n t underwater v e g e t a t i o n ; as w e l l as the wet area willows t h a t s t i l l p e r s i s t e d along l a k e and marsh edges. At the same time, they had maintained water l e v e l s such that l a r g e s e c t i o n s of w i l l o w brakes, i n the T h i r d V e r m i l i o n Lake and 7 M i l e Beaver Dam areas, had been unquestionably w a t e r - k i l l e d , thus removing these s i g n i f i c a n t areas from use by moose. I t has a l r e a d y been p o i n t e d out t h a t the summer moose p o p u l a t i o n was migra-t o r y or t r a n s i e n t , and much l a r g e r than the area could s u s t a i n over any p e r i o d of time. Such animals d i v i d e d t h e i r time b e t -ween wet v e g e t a t i o n and browse. By August the w i l l o w s i n the immediate v i c i n i t y of the l a k e s and marshes showed n i n e t y to one hundred per cent u t i l i z a t i o n , w i t h c o n s i d e r a b l e d e s t r u c t -i v e browsing of the major brakes. Wet v e g e t a t i o n s t i l l con-t i n u e d to be e x c e l l e n t . The f l o o d k i l l of w i l l o w s In the g e n e r a l area, then, c o n t r i b u t e d to the c o n c e n t r a t i o n of these t r a n s i e n t animals, and no doubt hastened the t r e n d toward u l t i m a t e d e s t r u c t i o n of the browse s p e c i e s . In the Muleshoe area, i n June, f i v e cows and f o u r c a l v e s were p r e s e n t — w o r k i n g the s m a l l green runs i n 59 c o n j u n c t i o n with, w i l l o w browsing. On 3 J u l y no moose were seen; and browse was found to be n i n e t y to one hundred per cent u t i l i z e d ; browsing animals c o u l d no l o n g e r s u r v i v e t h e r e . F a r t h e r west, i n the v i c i n i t y of the 7 M i l e Dam, w i l -low height i n the open f l a t s was twelve to eighteen i n c h e s , and a l l growth was h e a v i l y browsed. T a l l w i l l o w brakes along the stream edges were n i n e t y to one hundred per cent u t i l i -zed, and no l i v e stems over f o u r f e e t i n h e i g h t s u r v i v e d . Normal h e i g h t f o r the open f l a t w i l l o w s would be a t l e a s t f o u r to f i v e f e e t , and f o r those i n the brakes ten to twelve f e e t and h i g h e r . The F o r t y m i l e , Johnston, Baker and Boom Creek areas bore e x c e l l e n t w i l l o w growth, and browse g e n e r a l l y was good. No check was made of the Healy Creek and Red E a r t h Creek ar e a s . At the 17 M i l e meadow, e a r l y i n v e s t i g a t i o n i n d i -c a t e d moderate to heavy winter u t i l i z a t i o n of browse s p e c i e s . However, i n the main a r e a summer w i l l o w growth and f r u i t i n g were e x c e l l e n t . T h i s was due to complete l a c k of summer usage, a s i t u a t i o n b e l i e v e d due to complete absence of e q u i -situm or pondweed i n the nearby lake and to the d e s t r u c t i v e browsing by e l k of surrounding a r e a s . The area l e a d i n g to Helen Lake was i n v e r y good c o n d i t i o n , and the Bow Summit I t s e l f showed e x c e l l e n t browse c o n d i t i o n s . West of Upper Waterfowl Lake, i n c l u d i n g up to and around Cirque Lake, moose range was a g a i n e x c e l l e n t . At the Waterfowl Lakes, however, the c o n d i t i o n was s i m i l a r to that at V e r m i l i o n Lakes. Willows were h e a v i l y to d e s t r u c t i v e l y 6o browsed, except where p r o t e c t e d by the road camp ( t h i s de-monstrated summer browsing s i n c e the camp was onl y open d u r i n g the summer). The degree of u t i l i z a t i o n decreased as one moved p r o g r e s s i v e l y f a i t h e r from the l a k e , a g a i n demon-s t r a t i n g summer usage. South up the M i s t a y a R i v e r and t r i -b utary creeks browsing was heavy. As e a r l i e r mentioned, however, underwater v e g e t a t i o n was n e g l i g i b l e except a t the entrance of the Mis t a y a R i v e r i n t o Upper Waterfowl Lake. C a r r y i n g c a p a c i t y g e n e r a l l y , then, was low. The Howse R i v e r area showed i n d i c a t i o n s o f e i g h t y to n i n e t y per cent browse u t i l i z a t i o n i n meadows when i n s p e c -te d e a r l y i n the summer. Due to the d r y season recove r y was not one hundred per cent by August, but i t was s u f f i c i e n t to i n d i c a t e good range condition,, Browse c o n d i t i o n s appeared to be good i n the Graveyard area, and a l s o a t Owen Creek. In the l a t t e r a r e a aspens had about f i f t y p e r cent of the l e a d e r s removed i n some small l o c a l i t i e s . The Cascade moose areas g e n e r a l l y were i n good con-d i t i o n . As s t a t e d e a r l i e r , some d e t e r i o r a t i o n of browse spe c i e s was seen on the Cascade V a l l e y f l o o r south o f Cuthead, but i t was not yet s i g n i f i c a n t . In the Spray V a l l e y c o n d i -t i o n s were not good. South and east of the 15 M i l e c a b i n the main t a l l w i l l o w brakes were h e a v i l y browsed g e n e r a l l y . Browsing was heavy along the t r a i l l e a d i n g t o Sundance Pass. I t was c o n s i d e r e d t h a t t h i s area was c a r r y i n g the maximum numbers concomitant w i t h continued good range management. 61 T h i s was c o n s i d e r e d t r u e of a l l major moose ranges i n the Park, i n p a r t i c u l a r the summer c o n c e n t r a t i o n a r e a s . Only minor creeks and s l i d e areas i n d i c a t e d p o s s i b l e underpopula-t i o n . MOOSE IN RELATION TO ELK L i t t l e i n d i c a t i o n was seen of any d i r e c t c o n f l i c t between moose and e l k , p o s s i b l y due to the f a c t that e l k use of browse areas i s l a r g e l y c o n f i n e d to the w i n t e r while these s t u d i e s were c a r r i e d out d u r i n g the summer. I n q u i r y e l i c i t e d no i n s t a n c e s of p h y s i c a l c o n f l i c t , however, and r e p o r t s were r e c e i v e d of the two s p e c i e s l i v i n g compatably i n c e r t a i n areas. Some observed i n s t a n c e s of e l k browsing have a l r e a d y been noted?; and i t was c o n s i d e r e d s i g n i f i c a n t t h a t i n the main, Bow V a l l e y e l k c o n c e n t r a t i o n areas browse s p e c i e s were h e a v i l y o v e r - u t i l i z e d or k i l l e d , w h i le aspen r e p r o d u c t i o n was n e g l i g i b l e . I t cannot be doubted t h a t c o n s i d e r a b l e a r e a was thus removed.from use by moose. However, In view of the s o l i t a r y h a b i t s of the moose, i t i s u n l i k e l y t h a t t h i s s i t u a -t i o n was a major f a c t o r i n l i m i t i n g the moose p o p u l a t i o n . The tendency of e l k to g r a d u a l l y p r e s s a g a i n s t the moose swamp areas, though, would seem to i n d i c a t e that t f such areas d r i e d up g r a d u a l l y , e l k would take them over, e l i m i -n a t i n g moose or a l t e r i n g t h e i r m i g r a t i o n p a t t e r n s . T h i s 62 l a t t e r e f f e c t may a l r e a d y have o c c u r r e d , s i n c e no moose were seen to move from Johnston Creek through H i l l s d a l e to the V e r m i l i o n Lakes d u r i n g the summer. Too, the 17 M i l e meadow area was completely unused by moose d u r i n g the summer, pos-s i b l y due to the d e s t r u c t i v e a c t i v i t i e s of the e l k i n the surrounding a r e a . Mention has a l r e a d y been made of the s i t u a t i o n i n the Cascade V a l l e y . In 194-9 t n e movement of e l k to the v a l l e y f l o o r had not yet r e s u l t e d i n s e r i o u s browse d e t e r i o r a t i o n . There i s l i t t l e doubt, however, that i f t h i s new behaviour p a t t e r n p e r s i s t s , the browse w i l l soon be e l i m i n a t e d and the main v a l l e y f l o o r made untenable f o r moose. A more complex e f f e c t Is seen i n the removal of aspen growth and r e p r o d u c t i o n by e l k . In the Bow V a l l e y t h i s i s c o n t r i b u t i n g to the eventual e x t i r p a t i o n of the beaver, which i n t u r n w i l l remove the pondweed areas and hasten r e t u r n of dry c o n d i t i o n s d e t r i m e n t a l to the moose. I t must be r e c o g -n i z e d , then, that e l k are a f f e c t i n g the normal p l a n t s u c c e s s i o n of the area, and w i l l i n t u r n u n f a v o r a b l y i n f l u e n c e the moose. The f u l l s i g n i f i c a n c e o f t h i s i n f l u e n c e may not yet be appa-r e n t . 63 MOVEMENTS AND FOOD HABITS OF MULE DEER Since the study i n 1949 was p a r t i c u l a r l y d i r e c t e d to moose s t u d i e s , l i t t l e attempt was made to determine move-ments and food h a b i t s of the mule deer, beyond r e c o r d i n g o b s e r v a t i o n s . Deer, o r deer s i g n s , were noted throughout the summer i n a l l s e c t i o n s of the Park. Cowan (1943:11) s t a t e s t h a t most of the summer deer p o p u l a t i o n migrates out of the Park i n the autumn, and w i n t e r s at the lower l e v e l s of the f o o t h i l l s . Thus the major problem o f wint e r range i s l i m i t e d i n the Park. However, the Bow V a l l e y , which maintains a c e r t a i n r e s i d e n t p o p u l a t i o n throughout the summer, p r e s e n t l y p r o v i d e s a w i n t e r i n g ground f o r a d i s t a n c e of some seven m i l e s from Banff to the ar e a of the 7 M i l e Beaver Dam. Edwards, i n h i s s t u d i e s on the Malheur N a t i o n a l F o r e s t ranges i n Oregon, s t a t e s t h a t the deer spend n e a r l y f o u r months on a l a r g e w i n t e r assembly a r e a before moving to the wint e r con-c e n t r a t i o n area f o r the two c r i t i c a l months February to A p r i l (Edwards, 194-2:213). I f t h i s s i t u a t i o n o b t a i n s to any degree i n Banff Park, then a much l a r g e r a r e a than that a l r e a d y d e s c r i b e d may be v i t a l to w i n t e r s u r v i v a l o f deer. I t was most n o t i c e a b l e that w h i l e some few deer were seen throughout the Bow area, the main summer deer popu-l a t i o n was found In c l o s e p r o x i m i t y to Banff i t s e l f , or those areas where t o u r i s t t r a f f i c had, a p p a r e n t l y , r e s t r i c t e d the d e s t r u c t i v e a c t i o n of e l k . They were a l s o found i n the deep 64-to semi-open woods surrounding the l a k e s and swamps—areas l i t t l e f requented by e l k and h e l d to be t y p i c a l l y moose h a b i t a t . Thus f i f t y - o n e s i g h t r e c o r d s were noted f o r the area from Banff to the 7 M i l e Beaver Dam, while o n l y e i g h t -een were noted f o r the e n t i r e area from 1 M i l e to the Junc-t i o n of the Lake L o u i s e - J a s p e r H i g h w a y s — a d i s t a n c e of n e a r l y t h i r t y m i l e s . Yet a l l t h i s l a t t e r a r e a should have been s u i t a b l e deer t e r r i t o r y ; most of i t should have been w i n t e r as w e l l as summer range. Most o b s e r v a t i o n s were made be f o r e the end of the f i r s t week i n June, s i n c e by t h a t time most deer appeared to have migrated. They r e t u r n e d by the middle of August but time d i d not permit o f worthwile o b s e r v a t i o n s . No attempt was made to determine the food h a b i t s of the deer i n the Park. Studies of deer foods and food h a b i t s are numerous and exhaustive i n d e t a i l . I t can be s t a t e d g e n e r a l l y t h a t deer are browsers by p r e f e r e n c e but do eat grasses and f o r b s i n q u a n t i t i e s v a r y i n g from season to season. Use of these l a t t e r two forage types v a r i e s a l s o from a b a s i c minimum to maxima dependent upon range c o n d i t i o n . Percentage of t o t a l d i e t made up by any s i n g l e s p e c i e s depends upon a v a i l a b i l i t y . Cowan (1944:4-9-52) g i v e s some data on deer food h a b i t s i n Jas p e r Park, and i t was assumed, f o r pur-poses of the study, that the food h a b i t s were e s s e n t i a l l y the same i n Banff Park. Aspen and w i l l o w were prese n t i n the area where the g r e a t e r p o p u l a t i o n count was made, but through-out most of the remaining semi-open areas of the Bow V a l l e y 6 5 w i l l o w and aspen were unimportant f o r deer s i n c e the two s p e c i e s had been v i r t u a l l y e l i m i n a t e d by e l k , Shepherdla was present but not s u f f i c i e n t l y abundant to be of s i g n i -f i c a n t v a l u e . Bearberry was abundant and p r o b a b l y the c h i e f support of deer throughout the winter ranges shared w i t h the e l k . VITAL STATISTICS OF MULE DEER Time d i d not permit attempting an a c t u a l f u l l count of deer i n the Bow V a l l e y d u r i n g the summer study, but s i g h t counts were t a l l i e d . The h i g h e s t s i n g l e count f o r one day was s i x d e e r i T h i s was i n d i r e c t c o n t r a s t to the s i t u a -t i o n as r e p o r t e d i n the e a r l i e r years of the Park. Hewitt ( 1 9 2 1 : 2 3 8 ) quotes a warden's r e p o r t of e a r l y 1 9 1 9 which s t a t e d that seventy-one sheep and twelve deer were seen near Massive ( H i l l s d a l e a r e a ? ) — p r e s u m a b l y i n one day. He f u r t h e r quotes the Superintendent as r e p o r t i n g i n A p r i l of the same year t h a t along the motor road f o r ten m i l e s west of Banff 375 sheep, 10 goats and 16 deer were seen ( i b i d : 2 3 7 ) . Mr. LaCasse r e p o r t e d to me that p r i o r to about 1 9 3 6 - 3 7 willows used to be p l e n t i f u l and In e x c e l l e n t c o n d i t i o n throughout the H i l l s d a l e area, and that deer and sheep were p l e n t i f u l . A check of the d i a r i e s of Warden G. P. Woodworth f o r 194-2 66 r e v e a l e d that the two l a r g e s t d a i l y s i g h t r e c o r d s f o r the e n t i r e area Banff to Lake Louise to the D i v i d e , were f i v e and nine deer. These were d u r i n g the month of May, so com-parable to the e a r l i e r counts, as w e l l as those of 194-9. Other counts by Woodworth f o r 194-2 were h i g h e r but not con-s i d e r e d u s e f u l f o r comparison as most of the deer were undoubtedly seen i n the Lake Louise d i s t r i c t * However, a g e n e r a l check of the r e c o r d s seemed to I n d i c a t e that the deer p o p u l a t i o n remained p r a c t i c a l l y s t a t i c from t h a t year to 1949. T h i s i s i n t e r p r e t e d to mean that removal of e l k by s l a u g h t e r s i n c e 194-3 has f a i l e d to produce s i g n i f i c a n t i n c r e a s e i n deer; r e s u l t a n t range improvement has not been r e f l e c t e d i n the deer p o p u l a t i o n to date. In a summer study as c a r r i e d out i n 194-9, i t was not p o s s i b l e to make any i n t e l l i g e n t estimate of f e c u n d i t y of deer, or to determine increment. The does h a b i t u a l l y hide t h e i r fawns while they themselves feed or move about, f r e q u e n t l y a c o n s i d e r a b l e d i s t a n c e away. The fawns remain cached f o r the g r e a t e r p a r t of each d a y — e v e n a f t e r a t t a i n -i n g the age of c o n s i d e r a b l e a c t i v i t y . Too, most of the does had d i s p e r s e d from the Bow V a l l e y c o n c e n t r a t i o n areas p r i o r to fawning. Thus not more than h a l f - a - d o z e n fawns were seen throughout the summer, altho u g h every day was spent i n the f i e l d (not i n search of d e e r ) . As p r e v i o u s l y s t a t e d , the deer were re-a p p e a r i n g i n the v a l l e y by mid-August, but v a l i d doe-fawn counts were not p o s s i b l e before the f i r s t of 67 September when the investigation ended. Figures given by Green (unpublished report) for 194-8, In the Bow Valley, i n -dicate that yearlings made up approximately 13.6 per cent of the t o t a l herd. Comparison of t h i s figure with the 17 per cent and 12 per cent given by Cowan fo r 194-3 (194-3:11-12) would seem to indicate that there has been l i t t l e change i n the situation since the e a r l i e r date. Robinette and Olsen (194-4-:l60-l6l) found i n central Utah that yearlings c o n s t i -tuted 27o4- per cent of the t o t a l herd. Herd increment i n the Bow Valley, then, appears to be low. MULE DEER LOSS FACTORS Loss factors i n the deer population of Banff Park remain r e l a t i v e l y unknown. Cowan (194-3:12 and 194-4-:33-34-) has dealt with t h i s problem i n both Banff and Jasper Parks, and has discussed i n p a r t i c u l a r parasites and diseases. It i s not desired to comment further on t h i s aspect of the s i t u a t i o n . However, i t was noticed that there was a r e l a t i v e l y heavy population of coyotes present i n Banff Park i n 194-9, and one cannot doubt that a number of fawns must annually f a l l prey to these predators. Further, the s i g n i f i c a n t wolf population i n the Park must have some effect upon the deer. These l a t t e r , unable to compete adequately with elk for food, would be expected to succumb more readi l y to predation than the elk. 6g Thus l o s s e s would he g r e a t e r than would be expected on the b a s i s of chance encounter alone. Loss f a c t o r s g e n e r a l l y would be aggravated i n importance due to the p a u c i t y and poverty of the prey p o p u l a t i o n . MULE DEER RANGES Deer summer g e n e r a l l y throughout the Park, and are so s c a t t e r e d as to make range d e t e r m i n a t i o n s d i f f i c u l t i f not i m p o s s i b l e . Thus np attempt was made, i n 19^9, to take p a r t i c u l a r cognizance of deer summer range c o n d i t i o n s . Browse was p l e n t i f u l a l o n g the minor streams and v a l l e y s , s l i d e s and small openings, and i t was c o n s i d e r e d t h a t the d i s p e r s e d deer p o p u l a t i o n would experience no d i f f i c u l t y i n coming through the summer i n good c o n d i t i o n . S i n c e ^ o n l y major win t e r range w i t h i n the Park i s i n the Bow V a l l e y , most empha-s i s was p l a c e d upon t h i s a r e a i n the study. I t has a l r e a d y been, s t a t e d t h a t browse s p e c i e s were almost completely want-ing throughout the wint e r range a r e a . Some improvement i n graze s p e c i e s was noted, as compared t o the s i t u a t i o n p r i o r to the implementation of an e l k removal program i n 19^3, but graze c o n d i t i o n s g e n e r a l l y were s t i l l f a r below normal. I t was c o n s i d e r e d that the e n t i r e w i n t e r range was marginal and p r e c a r i o u s f o r deer. 69 MULE DEER IN RELATION TO ELK The i n f l u e n c e of e l k on deer has been a b l y d i s -cussed by C l i f f i n h i s paper presented to the North American W i l d l i f e Conference i n 1939 ( C l i f f , 1939:560-569). I t Is r e a l i z e d that the range foods of the Oregon t e r r i t o r y d i s -cussed v a r y c o n s i d e r a b l y from those of the Banff area, but the same b a s i c r e l a t i o n s h i p s e x i s t and one cannot escape the same c o n c l u s i o n — t h a t e l k f l o u r i s h a t the expense of deer. C l i f f found t h a t over a 17'-year p e r i o d ( i n c l u d i n g a p e r i o d of i n t e n s e range overuse) e l k showed an average annual i n -crease of twenty-three per cent compared to 14-,4- per cent f o r the mule deer. He estimated that one e l k r e p l a c e d three deer i n h i s a r e a . No data are a v a i l a b l e f o r the Bow V a l l e y but i t i s not b e l i e v e d the replacement r a t i o has been so h i g h . The y e a r l i n g percentage? a l r e a d y g i v e n would seem to i n d i c a t e t h a t deer should have been i n c r e a s i n g more r a p i d l y than e l k . I t i s b e l i e v e d , however, t h a t these f i g u r e s do not r e f l e c t the t r u e s i t u a t i o n as the h i g h deer y e a r l i n g counts were mainly f o r areas not g r e a t l y frequented by e l k . I t i s b e l i e v e d t h a t upon the h e a v i l y used e l k ranges (once e x c e l l e n t deer ranges) counts would show an extremely low s u r v i v a l r a t e of deer and a s t a t i c or f a i l i n g t o t a l p o p u l a t i o n — a s compared to an i n c r e a s i n g e l k p o p u l a t i o n . C l i f f a l s o found (Ibid:567-568) t h a t the s i z e and weight of the mule deer decreased as c o m p e t i t i o n with e l k 70 i n c r e a s e d . Stoddart and Rasmussen (194-5:254) found i n Utah t h a t deer on overhrowsed range produced an annual fawn crop of IS. 7 per cent of the female herd compared to 75 per cent and 77 percent i n other s e c t i o n s of the S t a t e . Deer of a l l ages from the overhrowsed area averaged about 70 per cent of the weight of the other deer, c a l c u l a t e d from k i l l d a t a . They s t a t e t h a t "There i s no q u e s t i o n that an underfed'deer not only i s smaller than normal deer but a l s o i t s c a p a c i t y to reproduce i s l e s s " . T h i s has been f u r t h e r confirmed by Morton and Cheatum (1946) and Cheatum and Severinghaits ( l 9 5 Q ) i n t h e i r work on w h i t e t a i l deer i n New York S t a t e . There are no data as yet a v a i l a b l e to prove or d i s p r o v e such a s i t u a t -i o n i n Banff Park. But the thought p r e s e n t s i t s e l f t h a t here, w i t h much of the once e x c e l l e n t w i n t e r range now marginal, and with the main winter deer p o p u l a t i o n s of the Bow V a l l e y r e -s t r i c t e d to extremely l i m i t e d areas, the same c o n d i t i o n c o u l d v e r y w e l l o b t a i n . Mule deer are known to be t e n a c i o u s i n t h e i r adherence to home ranges and customary f e e d i n g grounds. I t i s c o n s i d e r e d u n l i k e l y that the v a r i o u s p o p u l a t i o n s i n v o l -ved i n c o m p e t i t i o n w i t h e l k i n Banff Park have moved to new w i n t e r areas from the summer ranges. Rather i t i s more p r o -bable t h a t they have s u f f e r e d the d e b i l i t a t i n g and degrading e f f e c t s of impoverished range c o n d i t i o n s . There was ho reason to b e l i e v e t h a t e l k were a f f e c t -i n g the deer of the North Saskatchewan V a l l e y . Browse c o n d i -t i o n s i n t h a t area were s t i l l good. P r a c t i c a l l y a l l other 7 1 deer of the Park w i n t e r o u t s i d e i t s boundaries, so n o t h i n g was l e a r n e d of t h e i r w i n t e r i n g c o n d i t i o n s . However, C l a r k , i n h i s r e p o r t on the Park i n 1939» gave i t as h i s o p i n i o n that the Park was f u l l y stocked w i t h deer ( C l a r k , 1939:9). No data were observed i n t h i s study to suggest that such was not the case. Thus i t must be presumed that any encroach-ment of e l k upon deer range w i t h i n the Park would r e s u l t In c o n d i t i o n s unfavorable to deer, w i t h a subsequent downward tr e n d i n t h e i r numbers. There can be no q u e s t i o n that some such encroachment has o c c u r r e d . MOVEMENTS AND FOOD HABITS OF BIG-HORN SHEEP . The ecology and l i f e h i s t o r y o f the bighorn sheep of Banff N a t i o n a l Park have r e c e n t l y been extremely w e l l covered by G-reen ( 1 9 4 - 9 ) . Most of the i n f o r m a t i o n , then, g i v e n f o r t h i s s p e c i e s has been drawn from h i s work. However, c e r t a i n p e r s o n a l i d e a s and o b s e r v a t i o n s have been i n c o r p o r a t e d i n t o t h i s b r i e f r e c a p i t u l a t i o n . A c c o r d i n g to G-reen (1949:20), the b i g h o r n of Banff Park are not c o n f i n e d to any one range w i t h i n the r e g i o n , but move, i n bands of v a r y i n g numbers, from range to r a n g e -o f t e n remaining i n each l o c a l i t y o n l y f o r a few days. W e l l t r a v e l l e d t r a i l s j o i n most of the ranges, and are used u n t i l 72 w e l l i n t o the winte r , or u n t i l deep snow l i m i t s movement. Dis t a n c e s t r a v e l l e d vary from l e s s than f i v e to more than twenty-tvro mile 3 . I t was r e p o r t e d i n 194-9 that there had been some movement of sheep from the Aylmer range area to the V e r m i l i o n l i c k , but no evidence wgs a c t u a l l y seen of wide-spread sheep movements,. These sheep are d i u r n a l i n t h e i r a c t i v i t y , and can u s u a l l y be seen upon the ranges—-feeding, l o a f i n g and r e s t i n g . They have so l o s t t h e i r f e a r of humans i n the Park that the ewes and young males can o f t e n be r e a d i l y a p p r o a c h e d — e s p e -c i a l l y at l i c k s . The rams, i n separate bands i n summer, tend to remain more wary and s u s p i c i o u s . From o b s e r v a t i o n s made i n 194-9, i t can be s a i d t h a t the f l o c k s appeared to f o l l o w no set r o u t i n e . They appeared at the V e r m i l i o n l i c k and again disappeared w i t h apparent l a c k of r e g u l a r i t y or reason. The numbers appearing v a r i e d from day to day, al t h o u g h some of the same i n d i v i d u a l s appeared to be presen t s u c c e s s i v e times. There appears to be l i t t l e seasonal m i g r a t i o n among the Banff Park sheep. Rather they appear to move somewhat i n -d i s c r i m i n a t e l y about t h e i r ranges, tending to f a v o r s o u t h e r l y and xvesterly s l o p e s . In w i n t e r they frequent areas where snow i s l e a s t heavy or has blown c l e a r . These a g a i n are mainly s o u t h e r l y and w e s t e r l y slopes which are most exposed to the p r e v a i l i n g winds. Cowan ( 194-3:4—5) d i s c u s s e s sheep winter ranges i n some d e t a i l . Sheep are b a s i c a l l y g r a z e r s but do consume smal l 73 q u a n t i t i e s of browse. Cowan (1944:51) found i n Jasper Park t h a t grasses and sedges c o n s t i t u t e d e i g h t y - t h r e e per cent of-the t o t a l food consumed, f o r b s ten per cent and shrubs o n l y seven per cent. I t may be presumed that e s s e n t i a l l y the same s i t u a t i o n o b t a i n s i n Banff Park. Feeding p e r i o d s a l t e r n a t e w i t h r e s t i n g and l o a f i n g p e r i o d s . During the summer, at l e a s t , c o n s i d e r a b l e t i m e . i s spent a t m i n e r a l " l i c k s " . At such times the bands o f sheep work the d i r t of the " l i c k " , n i p o c c a s i o n a l sedges and grasses i n the v i c i n i t y , p o s s i b l y d r i n k i f water i s nearby, and g e n e r a l l y g i v e the appearance of r e s t i n g . Free water i n the immediate v i c i n i t y i s not, a p p a r e n t l y , an e s s e n t i a l to good sheep range. Escape cover appears to be a main c o n s i d e r a t i o n when sheep are g r a z i n g i n a c l o s e l y c o n f i n e d a r e a . I t i s not known j u s t how much t h i s need d i c t a t e s food c h o i c e . VITAL STATISTICS OF BIG-HORN SHEEP I t i s extremely, d i f f i c u l t to a r r i v e a t any d e f i n i t e c o n c l u s i o n s r e g a r d i n g sheep p o p u l a t i o n s i n Banff Park without a d e t a i l e d study. They wander over s e v e r a l ranges, fb»m l a r g e or s m a l l f l o c k s at random and break and reform w i t h a c e r t a i n amount of . s h u f f l i n g , thus making ac c u r a t e counts most d i f f i c u l t . Lambs may or may not accompany ewes to the l i c k s . 7 4 S p e c i a l marking techniques would he i n v a l u a b l e In the study of such p o p u l a t i o n s . Throughout the summer of 194-9, d a i l y counts were made of sheep i n the Bow V a l l e y . Undoubtedly c o n s i d e r a b l e d u p l i c a t i o n o c c u r r e d . However, the t o t a l count f o r the sum-mer was 278 sheep of a l l ages. T h i s aggregate was made up of ages and sexes as f o l l o w s : 63 a d u l t rams, 4-2 a d u l t ewes, 27 y e a r l i n g s , 21 lambs and 125 unaged and unsexed. In p e r c e n t -ages t h i s worked out to 22.7 per cent a d u l t rams, 15.1 per cent a d u l t ewes, 9.7 per cent y e a r l i n g s , 7.5 per cent lambs and 4-5.0 per cent unaged and unsexed. Green (194-9:30) g i v e s composition of aggregate counts made from A p r i l to September, f o r the f i v e years 194-4- to 1948 i n c l u s i v e . These show r e l a -t i v e s t a b i l i t y i n sex and age r a t i o s . F i g u r e s f o r 1948 are l 6 , l per cent rams (over two y e a r s ) , 60.8 per cent ewes (over two y e a r s ) , 3r° P e r cent rams and ewes two y e a r s , 3»^ per cent rams and ewes y e a r l i n g and 16.5 per cent lambs. Comparison of h i s f i g u r e s w i t h the 1949 f i g u r e s i n d i c a t e s c e r t a i n r a t h e r great d i s c r e p a n c i e s , but Green shows no unaged and unsexed group so the f i g u r e s are not s t r i c t l y comparable, e s p e c i a l l y f o r ewes, young rams and y e a r l i n g s . Great d i f f i c u l t y was experienced i n 1949 i n d i s t i n g u i s h i n g between two year o l d rams and ewes of v a r i o u s ages. T h i s was g r a d u a l l y overcome with experience, but h a s t y or d i s t a n t o b s e r v a t i o n s were s t i l l u n c e r t a i n . T h i s c e r t a i n l y caused i n a c c u r a c y i n counts. How-ever,, i t i s b e l i e v e d that the g r e a t e r percentage of the 75 unaged-unsexed group was ewes, which would b r i n g the t o t a l somewhat i n l i n e w i t h Green's f i g u r e s . Y e a r l i n g counts were f e l t to be j u s t f a i r l y a c c u r a t e , but compared f a v o r a b l y with the 194-8 f i g u r e s f o r the Dormer f l o c k (Green, 1948:11). Ad u l t rams and lamb counts were c o n s i d e r e d to be q u i t e accu-. r a t e . I t i s b e l i e v e d that Green's f i g u r e s f o r rams are low, and not i n d i c a t i v e of the true s i t u a t i o n . They were compiled from counts taken e n t i r e l y a t the V e r m i l i o n l i c k , where a d u l t rams are i n f r e q u e n t l y seen. I t i s d i f f i c u l t to s t a t e i f the ,1949 f i g u r e s i n d i -cate any s i g n i f i c a n t change i n p o p u l a t i o n composition. Green (194-9;31) shows a p r o g r e s s i v e d e c l i n e i n lamb m o r t a l i t y from 17o3 per cent i n a 194-4 to 13.6 per cent i n 194-7, which sug-g e s t s an i n c r e a s i n g p o p u l a t i o n . I t i s p o s s i b l e that lamb s u r v i v a l to y e a r l i n g age improved s u b s t a n t i a l l y the w i n t e r of 194-8-4-9, so a c c o u n t i n g f o r the h i g h percentage o f y e a r l i n g s seen d u r i n g the study. However, no p l a u s i b l e reason f o r the drop i n lambs i n 194-9 i s apparent, u n l e s s the r a p i d d e t e r i o r -a t i o n of the upland ranges f i n a l l y made i t s e l f f e l t through reduced f e r t i l i t y or f e c u n d i t y . A f o l l o w - u p study i s neces-sary to p r o v i d e f u r t h e r data i n t h i s r e g a r d . One other pos-s i b i l i t y i s lamb l o s s to p r e d a t o r s , as coyotes appeared to be a c t i v e i n 194-9. U n f o r t u n a t e l y there are no comparative d a t a w i t h which to check t h i s p o s s i b i l i t y . Aggregate counts at the V e r m i l i o n l i c k i n d i c a t e d l i t t l e change i n p o p u l a t i o n . Table VII shows the counts f o r 76 the years 1944-4-9 i n c l u s i v e . Table VII.. Humber of counts and aggregate t o t a l s of bighorn sheep seen a t V e r m i l i o n l i c k , Banff N a t i o n a l Park, f o r 194-4 to 1949 i n c l u s i v e . Year Number of Counts Aggregate T o t a l 1944 7 29 444-1945 35 445 1946 35 . 54o 1947 35 571 1948 4-9 527 1949 29 297 G-reen made the 1944—4-8 counts from A p r i l to September (Green, 1949l3°)• T n e 1949 counts were made from May to August i n c l u -s i v e . I t Is not known whether the 1949 count r e p r e s e n t s a drop In p o p u l a t i o n , or merely r e f l e c t s the s h o r t e r p e r i o d covered. There can be l i t t l e doubt t h a t the sheep p o p u l a t i o n of the Park has dropped v e r y c o n s i d e r a b l y i n the past ten or more years. As a l r e a d y noted, a Park Warden e a r l y i n 1919 counted 71 sheep near Massive, and the Superintendent r e p o r t e d i n A p r i l of the same year that 375 sheep were seen w i t h i n ten mi l e s of Banff, west along the highway. Mr. U. U. LaCasse 77 s t a t e d that i n comparatively recent years sheep used to be p l e n t i f u l i n the Massive area. Green (Ibid:28) i n d i c a t e s that twenty years ago 300 to 400 sheep were observed y e a r l y on the Sawback-Vermlllon range. H i g h e s t count from 1942 to 1948 was l 4 7 . The s i t u a t i o n i n the Cascade a r e a was seen to be even more s e r i o u s . A Park Warden r e p o r t e d f i f t y - t w o sheep on Cuthead mountain e a r l y i n 1919 (Hewitt, 1921:238). Park Warden E. Stenton s t a t e d that i n 1927 or 1928 he and Mr. James Simpson counted between 600 and 700 sheep on the P a l l l s e r range. I t was not uncommon to count 250 sheep d u r i n g a t r i p along the P a l l l s e r and Cuthead ranges en route to Windy Cabin. By 1949 one was l u c k y to see twenty head on such a t r i p . A c c o r d i n g to Green ( i b i d : 2 8 ) , the h i g h e s t count on the P a l l l s e r from 1942 to 1948 was f o r t y - t h r e e . During the s e v e r a l t r i p s made through t h i s a r e a i n the 1949 study, no sheep was, .'seen on the two main ranges. F u r t h e r evidence of d e c l i n e i n p o p u l a t i o n i s seen i n the g r a d u a l disappearance of sheep from the areas south of the Bow R i v e r . A c c o r d i n g to Green ( l b i d : 2 S ) o l d Park f i l e s make mention of one hundred sheep south of the Bow i n 19l4. Whether some of these were summer migrants or not i s not known. Green presumes they were a l l Included i n a permanent p o p u l a t i o n on Goat Range, but i t i s known that there were and s t i l l are sheep on the Mt. Bourgeau Range. Cowan, i n 1943, suggests a l i b e r a l estimate of the Goat Range po p u l a -t i o n would be f i f t e e n (Cowan, 1943:6). As f a r as c o u l d be 72 determined i n 1949, the l a s t of these had been e x t i r p a t e d . .The Mt. Bourgeau f l o c k s t i l l numbers about t w e n t y - f i v e . I t i s i mpossible to escape the c o n c l u s i o n , then, as both Cowan and. G-reen have, noted, t h a t the b i g h o r n sheep of the Park have d e c l i n e d g r e a t l y and, i n s p i t e of the apparent d e c l i n e i n lamb m o r t a l i t y , may not yet be h o l d i n g t h e i r own at the r e -duced l e v e l . BIG-HORN SHEEP LOSS FACTORS Loss f a c t o r s , r e l a t i n g both t o annual l o s s e s and to p r o g r e s s i v e p o p u l a t i o n d e c l i n e , have been thoroughly d i s c u s -sed; by Cowan (1943:6-8, 1944:16-20 and 59-63) and Green (1949:30-31 and 34-49). It' Is perhaps r e p e t i t i o u s to r e c a p i -t u l a t e i n any d e t a i l t h e i r o b s e r v a t i o n s and f i n d i n g s . I t i s d e s i r e d , however, to l a t e r p o i n t up c e r t a i n f e a t u r e s of these p o s s i b l e l o s s f a c t o r s and to t h i s end some r e p e t i t i o n i s neces-sary. Actinomycosis has been found to be o f g e n e r a l o c c u r -rence i n the sheep of Jasper Park and presumably then too i n Banff Park f l o c k s where i t s presence i s known,. T h i s d i s e a s e i s c o n s i d e r e d more c h r o n i c than acute, but i t can k i l l through the medium of m a l n u t r i t i o n of the h o s t . Two s p e c i e s of lungworm are present i n bighorn, the h a i r lungworm 79 ( P r o t o s t r o n g y l u s s t i l e s i ) ana the thr e a d lungworm ( P l c t y o -caulus v l v i p a r u s ) . Both these p a r a s i t e s induce a pneumonic c o n d i t i o n , and while r a r e l y d i r e c t l y f a t a l , they can and do cause d e b i l i t y and lowered v i t a l i t y which cause the host to f a l l prey to p r e d a t o r s , m a l n u t r i t i o n or some o t h e r more s e r -ious d i s e a s e such as Haemorrhagic s e p t i c e m i a (caused by P a s t e u r e l l a o v i s e p t l c u s ) . P o t t s (1938:896) b e l i e v e s t h a t s e pticemia was r e s p o n s i b l e f o r the g r e a t l o s s e s i n sheep i n Rocky Mountain N a t i o n a l Park, Colorado, but suggests t h a t L y mphadinitis caseosus (caused by Corynebacterlum pyogenes) f o l l o w i n g lungworm i n f e s t a t i o n may be e q u a l l y important. Stomach worms of the genus O s t e r t a g l a were found by Cowan i n every sheep examined In the Parks; here a g a i n the same f a c t o r of d e b i l i t a t i o n , through blood l o s s and secondary i n f e c t i o n , i s p r e s e nted. S e v e r a l other I n t e r n a l p a r a s i t e s are known to occur i n sheep but t h e i r importance has yet to be demonstrated. E x t e r n a l p a r a s i t e s , i n the form of the wint e r t i c k (Perma-centor a l b i p i c t u s ) and the Rocky Mountain t i c k (Dermacentor  andersoni) are known to occur i n Banff sheep, but no s e r i o u s e f f e c t has yet been shown. I t should be noted, however, that again these p a r a s i t e s f o s t e r a lowered v i t a l i t y of the host. G-reen, i n p a r t i c u l a r , has g i v e n d e t a i l e d i n f o r m a t i o n r e g a r d i n g p r e d a t o r s i n Banff and t h e i r e f f e c t upon the bighorn (Green, 1 9 ^ 9 : 3 ^ - W . I t i s not d e s i r e d to enlarge go upon h i s work, hut he l i s t s cougars, wolves and coyotes as main probable p r e d a t o r s . As f a r as cougars are concerned, i t must be remarked that d u r i n g the f o u r months spent i n study i n the Park i n 1949, not one cougar t r a c k was i d e n t i f i e d a l -though many t r a i l s , of both horses and game, were covered on f o o t by the i n v e s t i g a t o r . One d o u b t f u l t r a c k , not new, was seen along C a r r o t Creek but i t i s b e l i e v e d that i t was an o l d wolf t r a c k s i n c e i t was s l i g h t l y l o n g e r than wide. Thus i t Is b e l i e v e d t h a t G-reen h i t very c l o s e to the t r u t h when he s t a t e d t h a t one cougar c o u l d have been r e s p o n s i b l e f o r a l l the s i g h t o b s e r v a t i o n s he has noted. Or i t may be t h a t the Park i s on the t r a v e l route of the o c c a s i o n a l i t i n e r a n t cougar, inasmuch as the s i g h t o b s e r v a t i o n s extend over s e v e r a l y e a r s . I t i s doubted that the Park c o n t a i n s any s i g n i f i c a n t r e s i d e n t cougar p o p u l a t i o n . Wolves were noted to be p r e s e n t i n some numbers i n 1949, as were coyotes too. With the l a t t e r i t was d i f f i c u l t to determine exact numbers since i n d i v i d u a l animals wandered g r e a t l y and were d i f f i c u l t to r e - i d e n t i f y . Two coyotes were | seen s t a l k i n g a group of lambs and ewes near the V e r m i l i o n l i c k . The two separated and attempted to cut the sheep o f f from nearby excape t e r r a i n , but the f i n a l outcome was f o r e -s t a l l e d by a shot f i r e d a t one coyote. I t was apparent, though, t h a t some harassment at l e a s t was t a k i n g p l a c e . g l D i s c u s s i o n of the ranges w i l l be g i v e n i n subsequent paragraphs. But i t i s apt to p o i n t out here t h a t much of the best sheep range i n the Park i s composed of open slopes a c o n s i d e r a b l e d i s t a n c e from escape t e r r a i n . E x c e l l e n t v i s i -b i l i t y a f f o r d s some p r o t e c t i o n , but i t i s p o s s i b l e f o r c u r -s o r i a l p r e d a t o r s to harass f l o c k s and so c o n f i n e them, upon occ a s i o n , to the l e s s s u i t a b l e and marginal p o r t i o n s of t h e i r range. Murie (1944:14-2) has noted t h i s s i t u a t i o n w i t h wolves and sheep d u r i n g h i s i n v e s t i g a t i o n s i n Mt. McKinley Park i n A l a s k a . As w i l l be noted l a t e r , then, the combination o f d e b i l i t a t i n g p a r a s i t i s m and d i s e a s e , harassment by p r e d a t o r s f o r c i n g use of marginal range, and range abuse and overuse by competitive h e r b i v o r e s could w e l l c o n s t i t u t e environmental r e s i s t a n c e beyond the a b i l i t y of the bigh o r n to withstand. While i t seems h i g h l y probable that the o r i g i n a l l a r g e f l o c k s were h e a v i l y decimated by e i t h e r Haemorrhagic s e p t i c e m i a or verminous broncho-pneumonia, (Cowan, 194-3?7), i t may w e l l be th a t they are being f u r t h e r decimated or h e l d a t a dangerous low by the combination of f a c t o r s h e r e i n d e s c r i b e d . 82 BIG-HORN SHEEP RANGES Green (1949:18-19) g i v e s e l e v e n ranges f©r bighorn sheep i n Banff Park, d e s c r i b e d as Eisenhower, Johnston Creek, Sawback, V e r m i l i o n , Aylmer, C a r r o t Creek, P a l l i s e r , Dormer, Bare Mountain, Red Deer and Mount Wilson, a r e a s . He i n c l u d e s the Upper Cascade R i v e r V a l l e y to F l i n t ' s Park w i t h i n the P a l l i s e r area; the Panther R i v e r and Snow Creek w i t h i n the Bare Mountain area, and T y r r e l l , McConnell and D i v i d e Creeks w i t h i n the Red Deer area. The map a c r o s s page shows these ranges as o u t l i n e d by him. He has, however, overl o o k e d s e v e r a l other sheep ranges, i n c l u d i n g the Bourgeau Range, Sulphur Mountain (summer range only) and the Mt. Drummond-Cyclone Mountain a r e a (exact s t a t u s unknown). These concur f a i r l y w e l l w i t h the areas noted by Cowan (194-3?4-5) w i t h the e x c e p t i o n t h a t by 194-9 no sheep, were known to be r e s i d e n t south of the Bow R i v e r other than on the Bourgeau Range. Thus the Goat Range f l o c k has been e l i m i n a t e d . Cowan d e a l s more p a r t i c u l a r l y w i t h w i n t e r ranges, as being the acute problem, but s i n c e the w i n t e r ranges a r e , i n the main, p a r t of the l a r g e r ranges d e s c r i b e d by Green i t i s c o n s i d e r e d s u i t a b l e to d e a l w i t h the more g e n e r a l a r e a s . The Mount Wilson and Red Deer areas were not examined i n 194-9, and the Bare Mountain and Dormer areas were observed c u r s o r i l y from a d i s t a n c e . However, from knowledge of the 23 e l k s i t u a t i o n i n t h e Mount Wilson area, i t was assumed t h a t i t had not yet s u f f e r e d from the dep r e d a t i o n s of e l k and was s t i l l i n good c o n d i t i o n . The Red Deer, Bare Mountain and Dormer Ranges were u t i l i z e d by e l k i n summer (G-reen, 194-9; 32-33) and i t was- cons i d e r e d , as Judged by other ranges, t h a t t h i s c o m p e t i t i o n would be to the detriment of the ranges and the sheep. A l l the other sheep ranges were h e a v i l y used by e l k at some time of the year, and were c o n s i d e r e d to be over-u t i l i z e d to some degree. E l k were seen h i g h on the Elsenhower,, Johnston Creek and Sawback sheep ranges d u r i n g the summer, and i t was deduced from p e l l e t o b s e r v a t i o n s t h a t they had been on some of the lower l e v e l s of the V e r m i l i o n Range. The C a r r o t Creek Range was h e a v i l y grazed, w i t h areas showing the t y p i c a l e l k u t i l i z a t i o n c o n d i t i o n of browse k i l l i n g and aspen d e t e r i o r -a t i o n . The P a l l l s e r range showed i n c i p i e n t e r o s i o n , from both hooves and g r a z i n g , and poor graze c o n d i t i o n s . T h i s s i t u a t i o n was aggravated by the extreme drought of e a r l y summer i n 194-9. The Aylmer Range i s a summer e l k range, and i t was seen to be i n j u s t f a i r c o n d i t i o n . Thus i t i s seen that o f a l l the sheep ranges i n Banff Park, p o s s i b l y only one, a t Mount Wilson, has escaped the e l k . I t was p o i n t e d out i n the h i s t o r y of the Park t h a t p r a c t i c a l l y a l l ranges but a l p i n e meadows have been c r e a t e d by f i r e s . C e r t a i n l y p r a c t i c a l l y a l l the w i n t e r sheep ranges 84-are f i r e c r e a t e d . Thus normal f o r e s t s u c c e s s i o n , e x t e n s i o n and regrowth c o n s t a n t l y exert p r e s s u r e upon a l r e a d y r e s t r i c -ted ranges. Recent f i r e s have opened up some new ranges, and Cowan suggested i n 194-3 that approximately the same t o t a l amount o f f r a n g e has remained a v a i l a b l e f o r the p a s t t h i r t y or f o r t y y e a r s . However, Park a u t h o r i t i e s are making every e f f o r t to p e r f e c t f i r e p r e v e n t i o n and f i r e f i g h t i n g techniques, and i t seems probable t h a t f o r e s t p r e s s u r e w i l l f u r t h e r l i m i t the sheep over the forthcoming years. BIGHORN SHEEP IN RELATION TO ELK The encroachment of e l k upon sheep range has- Just been d i s c u s s e d . No d i r e c t s t u d i e s have been c a r r i e d out to d e t e r -mine s i m i l a r i t i e s i n food h a b i t s of the two s p e c i e s i n Banff Park, but P i c k f o r d and Reid, i n t h e i r s t u d i e s i n t o c o m p e t i t i o n between e l k and domestic l i v e s t o c k i n Oregon, found t h a t the food p r e f e r e n c e s were very s i m i l a r , and they concluded that competition was keen. They found t h a t a small group of p l a n t s p e c i e s p r o v i d e d the major bulk of food f o r both s p e c i e s , and estimated that 1.25 e l k r e p l a c e d f i v e domestic sheep ( P i c k f o r d and Reid, 194-3:328-332). I t i s b e l i e v e d t h a t s i m i l a r i t y o f food h a b i t i n the Park has l e d to d i r e c t c o m petition between these two s p e c i e s . The v i g o r and a g g r e s s i v e n e s s of the e l k 85 herds, p l u s e x c e p t i o n a l v e r s a t i l i t y i n food h a b i t , has l e d to almost complete o c c u p a t i o n of a l l major sheep r a n g e s — to the detriment of the l e s s a g g r e s s i v e sheep. These l a t t e r have been f o r c e d , through l a c k of f o o d or f o r as yet unknown p s y c h o l o g i c a l reasons, more and more to the edges, of the ranges, where l i v i n g i s marginal and p r e c a r i o u s . I t Is not known i f sheep d i s l i k e e l k and w i l l not range f r e e l y over common ground, but i t was noted that ranges frequented by e l k were l i t t l e used by sheep. T h i s was p a r t i c u l a r l y seen on some p o r t i o n s of the V e r m i l i o n and Sawback ranges. However, i t was r e p o r t e d t h a t on the Dormer range both s p e c i e s s t i l l f eed over the same range. Thus i t was c o n s i d e r e d more p r o -bable that i t was a matter of sheer p h y s i c a l i n a b i l i t y of sheep to compete w i t h e l k f o r food. In any event, the sheep are being slowly but s u r e l y pushed o f f much of the choice range. One can only speculate on what has happened to them In the p a s t . I t seems probable t h a t the e p i z o o t i c s b e l i e v e d to have decimated the f l o c k s p r i o r to the f i r s t i n v e s t i g a t i o n s i n the 194-0's were the r e s u l t of sudden v i r u l e n c e of the disease i n c o n j u n c t i o n with h i g h p o p u l a t i o n s c o n t a i n e d w i t h i n r a t h e r r e s t r i c t e d ranges. T h i s c o n c e n t r a t i o n would be p a r t i -c u l a r l y e v ident i n the winter time. Since t h i s major d e c l i n e , worsening range c o n d i t i o n s would not o n l y prevent adequate recovery; they might w e l l c o n t r i b u t e d i r e c t l y to a p r o g r e s s i v e g6 d e c l i n e i n the s p e c i e s . I t has been suggested that e l k c a r r i e d d i s e a s e onto the ranges where sheep then f e l l v i c t i m s to i t . There seems to be l i t t l e to s u b s t a n t i a t e t h i s r e a s o n i n g . I t i s f e l t that the s i t u a t i o n i s one of a l e s s v i g o r o u s s p e c i e s f a l l i n g before the r-igdr^s of heavy environmental p r e s s u r e . I t was p o i n t e d out that most of the d i s e a s e s s u f f e r e d by sheep are of the d e b i l i t a t i n g type, and that p r e d a t o r s may tend to harass them i n t o marginal areas. These f a c t o r s , combined wit h the g e n e r a l range poverty and marginal e x i s t e n c e t h r u s t upon them by e l k , make- the s i t u a t i o n p r e c a r i o u s to the p o i n t where any minor change i n the balance might mean heavy l o s s or e x t i r p a t i o n . I t has been s t a t e d that l a c k of, adequate food p r e d i s p o s e s domestic stock to d i s e a s e , and t h a t most i n s t a n c e s o f m o r t a l i t y i n deer were the r e s u l t of m a l n u t r i -t i o n , through range d e p l e t i o n and secondary i n f e c t i o n . ( T a y l o r and Hahn, .1947:321.) There appears to be l i t t l e reason to doubt that the bighorn sheep of Banff Park are f a i l i n g b efore j u s t such a combination of circumstances. They may be expected to f a l l easy prey to s e p t i c e m i c , a c t i -nomycotic and bronchial-pneumonic e p i z o o t i c s . g6 SUMMARY DISCUSSION OF DYNAMIC  INTER-RELATIONSHIPS OF ELK WITH PARK BIOTA There has been o u t l i n e d a s i t u a t i o n wherein the r e g i o n now known as Banff N a t i o n a l Park has passed from the g r a s s -lands of the immediate p o s t - g l a c i a l e r a to a now h e a v i l y wooded p r i m a r i l y c o n i f e r o u s btfome. Deciduous types* i n p a r t i c u l a r aspen, are found along watersheds, i n some s e m i - f o o t h i l l areas, and i n t e r m i x e d w i t h Lodgepole pine as a f i r e t y p e. Willows and other shrubs grow i n low r e g i o n s , s l i d e areas, along creeks and i n n e a r - a l p i n e to a l p i n e a s s o c i a t i o n s . A l p i n e mea-dows s t i l l e x i s t , and w i t h burned-over areas and a few t r u e p r a i r i e r e g i o n s make up the major g r a z i n g components. Record of mammalian fauna Is scanty, except f o r v e r y recent times. But i t has been seen how deer, sheep and goats have been indigenous f o r as long as r e c o r d s permit a p p r a i s a l , w i t h deer and sheep In p a r t i c u l a r dependent, f o r p o p u l a t i o n suecess, upon f i r e type v e g e t a t i o n produced through n a t u r a l 87 causes. Into t h i s biome then came the beaver-moose complex, of very r e c e n t o r i g i n but nonetheless i n n a t u r a l e c o l o g i c a l sequence. F i r e s produced s a t i s f a c t o r y beaver h a b i t a t ; bea-ver a c t i v i t y p r o v i d e d s u i t a b l e summer h a b i t a t f o r moose. Predators have probably been present f o r as l o n g as h e r b i -vorous prey were a v a i l a b l e . The n a t u r a l dynamic aspects of the s i t u a t i o n become apparent. While any one of the mammalian s p e c i e s above-mentioned might s u r v i v e i n l i m i t e d numbers i n the c o n i f e r o u s climax a s s o c i a t i o n , a l l are dependent upon f i r e and the r e -s u l t a n t s e r a i stages i n the f l o r a f o r u l t i m a t e o c c u p a t i o n and success ( i n t e r p r e t e d as long term p o p u l a t i o n s u r v i v a l at optimum l e v e l ) . T h e i r f o r t u n e s are i n t i m a t e l y a s s o c i a t e d w i t h f i r e a c t i o n and i n t e r - s p e c i f i c c o m p e t i t i o n f o r food. Any f a c t o r a f f e c t i n g the balance of a v a i l a b l e range or normal p o p u l a t i o n movement or i n c r e a s e must n e c e s s a r i l y have some immediate r e p e r c u s s i o n s upon the whole e c o l o g i c a l complex. Thus the N a t i o n a l Park p o l i c y of f i r e p r e v e n t i o n c o u l d not f a i l to p l a c e c e r t a i n s t r e s s e s upon a l l these indigenous popu-l a t i o n s . The beaver have eaten themselves out and t h e i r p o p u l a t i o n s have waned. Subsequent d r y i n g up of beaver dam s i t e s and swamps has l i m i t e d summer moose range. I t seems probable that t h i s would be a n a t u r a l e c o l o g i c a l sequence o f events i n so l i m i t e d a beaver h a b i t a t , but f i r e c o n t r o l c o u l d not f a i l to quicken t h i s p r o g r e s s i o n . S i m i l a r l y , the ranges m of deer and sheep have been c u r t a i l e d and contained, and sheep i n p a r t i c u l a r have been a f f e c t e d . I t i s b e l i e v e d that the major f i r e s of the e a r l y 1900's, i n c o n j u n c t i o n w i t h t o t a l , p r o t e c t i o n o f the sheep, opened up o p p o r t u n i t i e s f o r maximum i n c r e a s e . More adequate f i r e c o n t r o l i n l a t e r y e a r s , combined w i t h f o r e s t encroachment and regrowth, l i m i t e d the p o s s i b i l i t y of subsequent expansion. Thus the s i t u a t i o n developed where maximum but con t a i n e d p o p u l a t i o n s f e l l b e fore e p i z o o t i c ^ The t o u r i s t a c t i v i t y of the N a t i o n a l Park has most c e r t a i n l y a f f e c t e d the normal e c o l o g i c a l sequence of events. Any w i l d s p e c i e s s u b j e c t e d to o b s e r v a t i o n by l a r g e numbers of people can s c a r c e l y be expected to ma i n t a i n a completely normal h a b i t p a t t e r n . As e a r l y as 1399 there were 7,3&9 v i s i t o r s to Banff Park (Canada, 1907:3); by 1950-51 the f i g u r e stood at 459,273 (Canada, 1950-51 :14). T o u r i s t trade too means roads, t r a f f i c , d i s p o n i n g o f c e r t a i n ranges, and an o v e r a l l p o l i c y d i r e c t e d to entertainment of the p u b l i c . As e a r l y as 1907 there were two s t r e t c h e s of road t o t a l l i n g n i n e t y - s i x m i l e s i n the Park (Canada, 1907:4). The now p r o -posed Trans-Canada highway through the wint e r range of one major e l k herd i n the Park, and through the c h i e f summering range of the moose, w i l l be a major blow to these s p e c i e s and emphasizes again the i n c o m p a t i b i l i t y o f w i l d e r n e s s con-cept w i t h t o u r i s t t r a d e . But without any doubt, the 89 g r e a t e s t impact upon the e c o l o g i c a l p i c t u r e of Banff Park was c r e a t e d by the Importation of e l k to p r o v i d e one more a t t r a c t i o n f o r the p u b l i c . P r i m a r i l y , three v i t a l po.ints were overlooked i n the s t o c k i n g program. The f i r s t was the f a i l u r e ' i n the b e g i n n i n g to a p p r e c i a t e the v i g o r o u s behaviour and v o r a c i o u s and v e r s a -t i l e f e e d i n g h a b i t s of the e l k . The second was that the f i r s t e l k were wintered over i n what might' normally have been con s i d e r e d s p r i n g or summer r a n g e — a t l e a s t i f t h e i r p a t t e r n f o l l o w e d that of the r e g i o n from which they were obtained. In Yellowstone Park they are known to migrate 150 to 200 m i l e s out to the w i n t e r ranges (Gaffney, 1941:4-36) and i t might w e l l have been expected that they should migrate from t h i s p l a n t i n g area to the adjacent f o o t h i l l s where range was more abundant. But, a c c o r d i n g to Murie (1951;64-), the migra-t o r y h a b i t s of e l k are c o n d i t i o n e d by h a b i t , and having once spent a w i n t e r i n any g i v e n l o c a l i t y they w i l l r e t u r n to i t — u s u a l l y by the same r o u t e . Thus the c o n d i t i o n s of these p l a n t i n g s tended to e s t a b l i s h the p a t t e r n f o r the f u t u r e . T h i s s i t u a t i o n was aggravated by the f a c t t h a t one e l k p l a n t was made west of Banff townsite. T h i s meant t h a t the main avenue of escape f o r any surplus p o p u l a t i o n was cut o f f once the m i g r a t i o n p a t t e r n was e s t a b l i s h e d to the west. From the beginning, then, the stage was set f o r a problem of overpopu-l a t i o n . The t h i r d f a c t o r to be c o n s i d e r e d i s t h a t a major 90 h e r b i v o r e was p l a n t e d i n an area.where no major p r e d a t o r e x i s t e d i n s i g n i f i c a n t numbers. Some cougar presumably were present but i n 1935, when e l k were a l r e a d y nearIng a c r i t i c a l h i g h , c o n t r o l measures were i n s t i t u t e d and nine cougar were removed from one of the major e l k problem a r e a s . I t was not u n t i l s e r i o u s e l k o v e r p o p u l a t i o n was an accomplished f a c t t h a t wolves f i r s t appeared i n the Park. These pre d a t o r s were then powerless to a f f e c t the continued i n c r e a s e of the prey. In the North Saskatchewan V a l l e y alone, where prey and pre d a t o r were f i r s t recorded w i t h i n a few years of one an-other, the e l k appear to be i n harmony w i t h the c o e x i s t e n t b i o t a . Tte p r e d a t o r - p r e y r e l a t i o n s h i p has been f u r t h e r aggra-vated by misguided though well-meant wolf c o n t r o l . The S t a f f of the Park have .the f a i r l y t y p i c a l a n t i p a t h y to these p r e d a t o r s , and c a r r y out c o n t r o l measures c o n t i n u o u s l y as op p o r t u n i t y p e r m i t s — b y t a c i t understanding of a u t h o r i t y i f not by a c t u a l order. C e r t a i n of the game s p e c i e s , p a r t i c u l a r -l y deer and sheep, may r e q u i r e some p r o t e c t i o n but the over-a l l e f f e c t of these a c t i o n s has been the r e d u c t i o n of a much needed c o n t r o l upon the e l k . T h i s has probably worsened a bad s i t u a t i o n . I t has been p o i n t e d out that one cannot p o i n t the f i n g e r d i r e c t l y at e l k as a f a c t o r d e t r i m e n t a l to the wel-f a r e of moose i n the Park. However, they have unquestionably 91 a s s i s t e d In the overbrowsing of c e r t a i n probable moose areas, they have c o n t r i b u t e d to overbrowsing of main avenues of approach to the moose summer swamp areas, and they have com-p l e t e l y removed aspen r e p r o d u c t i o n and c r i p p l e d willow and dwarf b i r c h growth over much of the Bow V a l l e y . T h i s l a t t e r a c t i o n has c o n t r i b u t e d to the e v e n t u a l "eat-out" by beaver, and has l e s s e n e d any hope of r e p o p u l a t i o n by them. T h i s i n t u r n w i l l hasten the e v e n t u a l d e c l i n e of moose i n the Park. A f u r t h e r p o s s i b i l i t y of c o n f l i c t has been d i s c u s s e d , the use of the Cascade V a l l e y f l o o r by e l k i n w i n t e r . I t seems apparent t h a t , while no major c o n f l i c t i s yet apparent, the e l k are e x e r t i n g a steady pressure a g a i n s t moose every p l a c e they c o e x i s t . I t i s suggested t h a t the moose, more s o l i t a r y by nature, w i l l l o s e i n t h i s c o m p e t i t i o n . The e l k have unquestionably i n f l u e n c e d the deer popu-l a t i o n of the Park by t h e i r d e s t r u c t i o n of browse s p e c i e s throughout most of the Bow-Valley r e g i o n . Deer have been f o r c e d to marginal areas adjacent to towns, Wardens 1 cabins and to the deep woods adjacent to swamps. In these l a t t e r areas they may be competing on a minor s c a l e w i t h moose, adding to the d e s t r u c t i v e use a l r e a d y d e s c r i b e d . I t seems p o s s i b l e that the e l k , I f u n c o n t r o l l e d , w i l l e v e n t u a l l y con-s i g n the deer to the r o l e of a r a r e or o c c a s i o n a l s p e c i e s In the Park, I t i s perhaps w i t h r e s p e c t to bighorn sheep that the 92 e l k have been most harmful. Confined to l i m i t e d areas, but reproducing w i t h great v i g o r , the e l k invaded every sheep range i n the Park (with one p o s s i b l e exception) to some de-gree. They have d r i v e n them from the bottom l a n d s and have overgrazed many of t h e i r winter ranges. They have now com-menced the same i n v a s i o n of the sheep summer ranges. T y p i c a l of t h i s s i t u a t i o n i s the P a l l i s e r Range. I t i s not known i f r, heavy use of a range by e l k makes i t repugnant to sheep, or i f they are Just unable to compete p h y s i c a l l y f o r food, but they have been slowly and s u r e l y f o r c e d i n t o m arginal a r e a s , and t h e i r numbers have decreased. I t has been suggested that the major l o s s of sheep was due to one or more e p i d e -m i c s — p r o b a b l y haemorrhagic s e p t i c e m i a . P o t t s (193^^96) suggests t h a t t h i s d i s e a s e i s not so h i g h l y contagious as b e l i e v e d , but r a t h e r t h a t i t i s endemic but dormant i n most sheep herds. He b e l i e v e s that h i g h herd l o s s i s due to simultaneous exposure to bad weather and m a l n u t r i t i o n . I t i s suggested that i f t h i s d i s e a s e i s present i n the Banff Park, any sudden i n c r e a s e i n v i r u l e n c e might w e l l prove f a t a l to f l o c k s s u f f e r i n g d e b i l i t a t i n g p a r a s i t i s m and impo-v e r i s h m e n t — t h e l a t t e r l a r g e l y a t t r i b u t a b l e to e l k . The s i t u a t i o n i s aggravated by the encroachment of f o r e s t growth v i g o r o u s l y p r o t e c t e d from f i r e . Under e x i s t i n g circumstances i n the Park i t seems p o s s i b l e that bighorn sheep, even more than deer, may be doomed to r e v e r t to the s t a t e of a r a r e 93 s p e c i e s e x i s t i n g upon marginal h a b i t a t . L i t t l e mention has been made of goat and c a r i b o u w i t h -i n the Park but some comment seems d e s i r a b l e . One can only, speculate upon the p o s s i b l e e f f e c t of e l k upon these s p e c i e s . But o b s e r v a t i o n s made i n 194-9 of the a g i l i t y o f e l k , combined w i t h the knowledge of t h e i r v e r s a t i l i t y of food h a b i t , would l e a d one to b e l i e v e that they may, i f u n c o n t r o l l e d , feed upon some c h o i c e r ranges used by goat. Since goat at a l l times appear to l e a d a somewhat marginal e x i s t e n c e , any i n t r u s i o n upon t h e i r range by e l k , or d i s p l a c e d bighorn, would be d e t r i m e n t a l to them. S i m i l a r l y , any hope of i n c r e a s e i n c a r i -bou would be o b v i a t e d by any movement of e l k i n t o t h e i r mar-g i n a l ranges. E x c e s s i v e range use and i n c i p i e n t e r o s i o n i n the Park, both due to e l k , may have more f a r r e a c h i n g consequences than the e l i m i n a t i o n of competitive f a u n a l s p e c i e s . Skinner, while d i s c u s s i n g the e f f e c t s of o v e r p o p u l a t i o n by e l k , has noted t h i s same problem. He s t a t e s t h a t almost permanent changes take p l a c e i n the s o i l i f p l a n t r e t r o g r e s s i o n (due to range overuse) i s allowed to c o n t i n u e . Trampling becomes p r o g r e s s i v e l y more severe and humus i n the top s o i l i s r e -duced. A b i l i t y to absorb and r e t a i n moisture i s l e s s e n e d and f i n a l l y e r o s i o n by r a i n , snow and wind takes p l a c e . And he p o i n t s out t h a t while p l a n t r e d u c t i o n i s f a i r l y easy to c o r r e c t , s o i l breakdown i s most d i f f i c u l t to a r r e s t . I t may o f t e n take 94 l o n g e r than a l i f e t i m e * (Skinner, 1950;23.) Thus i t w i l l be seen that the c o n d i t i o n s of t r a m p l i n g and e r o s i o n , so o b v i o u s l y apparent i n 1949, may w e l l a l t e r f l o r a l s u c c e s s i o n beyond the l i m i t s of present v i s i o n , thus i n f l u e n c i n g the e n t i r e ecology o f a d e l i c a t e l y balanced b i o t a . One cannot escape the c o n c l u s i o n that the i n t r o d u c -t i o n of t h i s major, v i g o r o u s and a g g r e s s i v e h e r b i v o r e i n t o the Park must a f f e c t the normal dynamic complexes to be found among the b i o t a . Not only have the mammalian species been i n f l u e n c e d but o t h e r c l a s s e s , too. B i r d s p e c u l i a r to a deciduous h a b i t a t have been l i m i t e d by the d e s t r u c t i o n o f aspen and unde r s t o r y s p e c i e s . B i r d l i f e i n the wil l o w t h i c k e t s must have been a d v e r s e l y a f f e c t e d . One can o n l y surmise the changes w i t h r e s p e c t to i n v e r t e b r a t e l i f e as a r e s u l t of the d e s t r u c t i v e v e g e t a t i v e changes, b o t h above and below ground, that have been caused by e l k . But i n the com-bined changes wrought upon both v e g e t a t i v e c h a r a c t e r and f a u n a l composition of the Park one reads the l e s s o n that man i s , i n f a c t , a member of the complex b i o t i c community where-ever he e x i s t s , and h i s a c t i o n s i n f l u e n c e the immediate dynamic t r e n d of e v o l u t i o n — o f t e n f a r beyond h i s p e r c e p t i o n . 95 BIBLIOGRAPHY V A l l a n , J . A, 1913 Bond, R. M, 194-5 Canada 1907 and 1911 194-4-1950-51 Rocky Mountains (Bankhead to Golden). Ottawa. Department of Mines. G e o l o g i c a l Survey. T r a n s c o n t i n e n t a l E x c u r s i o n C I, Guide Book No. 8, P a r t I I . pp. 167-199, maps and t a b l e . Range Rodents and P l a n t Succession. Trans. 10th N. Am. W. Conf. pp. 229-234-. Department of the I n t e r i o r . N a t i o n a l Parks Report. Government P r i n t i n g Bureau, Ottawa. Department of Mines and Resources. Report of Lands, Parks and F o r e s t Branch. King's P r i n t e r , Ottawa. F i s c a l year ending March 31st. Department of Resources and Development. Report of f o r f i s c a l year ending 1951. Ottawa. King's P r i n t e r and C o n t r o l l e r of S t a t i o n e r y . Cheatum, E. L. and C. W. Severinghaus. V a r i a t i o n s i n F e r -1950 t i l i t y of W h i t e - t a i l e d Deer R e l a t e d to Range C o n d i t i o n s . Trans. 15th N. Am. W. Conf. pp. 170-189. C l a r k e , C. H. D. W i l d l i f e I n v e s t i g a t i o n i n Banff N a t i o n a l 1939 Park, 1939. Ottawa. Department of Mines and Resources. N a t i o n a l Parks Bureau. 26 pp. (mimeo.). ........ Wild L i f e I n v e s t i g a t i o n s i n Banff and Jasper 194-2 N a t i o n a l Parks i n 194-1. Ottawa. Department of Mines and Resources. N a t i o n a l Parks Bureau. 21 pp. (mimeo.). 96 C l a r k e , S. E., J . A. Campbell, and J . B. Campbell. 1942 C l i f f , E. P. 1939 An E c o l o g i c a l and G r a z i n g C a p a c i t y Study o f the Native Grass P a s t u r e s i n Southern A l b e r t a , Saskatchewan and Manitoba. Dominion o f Canada, D i v i s i o n of Forage Crops, Department of A g r i c u l t u r e . Pub. No. 73$, Tech. B u l l . 44. 31 PP. R e l a t i o n s h i p between E l k and Mule Deer i n the  Blue Mountains of Oregon^ Trans. 4th N. Am. W. Conf. pp. 5bO-5b9. Cowan, I . McT. Report on Game C o n d i t i o n s i n Banff, Jasper 1943 and Kootenay Parks. N a t i o n a l Parks Bureau, Ottawa. 72 pp. (Mimeo.). 1944 1947 Edwards, 1942 Gaffney, 1941 Green, H, 1946 Report of W i l d l i f e S t u d i e s i n Jas p e r , Banff and Yoho N a t i o n a l Parks 1944 and P a r a s i t e s , Diseases and I n j u r i e s of,Game Animals i n the Rdcky Mountain N a t i o n a l Parks.1942-1944. N a t i o n a l Parks Bureau. Ottawa. 84 pp., 23 f i g s . , maps. (Mimeo.). The Timber Wolf i n the Rocky Mountain N a t i o n a l Parks o f Canada. R e p r i n t from Canadian J o u r n a l of Research, D, 25:139-174. October, 1947. P u b l i s h e d by the n a t i o n a l Research C o u n c i l of Canada. 0. T. Survey o f wint e r deer range, Malheur N a t i o n a l  F o r e s t , Oregon. Jo u r . W. Mgt. b(J):210-220. W. S.; The E f f e c t s of Winter E l k Browsing, South Fork  of the F l a t h e a d R i v e r , Montana. J o u r . W. Mgt. £ W : 4 2 7 - 4 5 3 . . . U. The E l k of Banff N a t i o n a l Park. 32 pp. (Mimeo.). 1942 1949* Occurrence of Game and O^her Mammals. Banff N a t i o n a l Park. 18 pp.. ( t y p e w r i t t e n ) . The Bighorn Sheep of Banff N a t i o n a l Park. N a t i o n a l Parks and H i s t o r i c S e r v i c e . Develop-ment S e r v i c e s Branch, Ottawa. 53 PP», 15 p l a t e s . (Mimeo.). 1951 H i s t o r y and Occurrence of Moose i n Banff N a t i o n a l Park. Unpublished Manuscript. 97 Hewitt., Co Cordon* The Conservation of the Wild L i f e of 1921 Canada* Charles S c r i b n e r ' s Sons, New York. 344 pp., i l l u s . , p l a t e s , maps, c h a r t s . Hunter. G-. N. Methods of Determining Trends i n B i g Game 1945 Numbers and Range C o n d i t i o n s . Trans 10th N. Am. W. Conf., pp. 234-241. Morton. Glenn H., and E. L. Cheatum. Regiona l D i f f e r e n c e s 1946 In Breeding P o t e n t i a l 0 f i n i t e - t a i l e d Deer i n New York. Jour. W. Mgt. 10(3):242~248. " Murie, A. The Wolves of Mount McKinley. Washington, 1944 U. S. Government P r i n t i n g O f f i c e , U. S. N a t i o n a l Park S e r v i c e , Fauna S e r i e s No. 5. 23S pp., i l l u s . ( i n c l . maps, t a b l e s ) . Murie, 0. J * The E l k . o f North America. The Stackpole Company, 1951 H a r r i s b u r g , Penn. and the W i l d l i f e Management I n s t i t u t e , Washington, D. C. 376 pp., 29 i l l u s , , 32 f i g s . N a t i o n a l Parks A s s o c i a t i o n . N a t i o n a l Parks of Canada. N a t i o n a l 1948 Parks Magazine. N a t i o n a l Parks A s s o c i a t i o n , Washington, D. C, 22(94):4o. N a t i o n a l Parks of Canada. The N a t i o n a l Parks of Canada. 1932 Department of Mines and Resources, Lands, Parks and F o r e s t Branch, N a t i o n a l Parks Bureau, Ottawa; 64 pp., i l l u s . , f o l d . , c o l . maps. P l c k f o r d , G. D. and E. H. Rei d . Competition o f E l k and 1943 Domestic L i v e s t o c k f o r Summer Range Forage. Jour. W. Mgt. 2(3^32S-332. P o t t s , M. K. Observations on Disease of Bighorn i n Rocky 1932 Mountain N a t l o n a l T a r k . Trans. 3rd N. AmT W. Conf. pp. 893-897. Robinette, W. L e s l i e , and Orange A. Olsen. S t u d i e s of the 1944 P r o d u c t i v i t y of Mule Deer In C e n t r a l Utah. Trans. 9th N. Am. W. Conf, pp.156-161. Simpson, S i r George. N a r r a t i v e of a journey around the world, 1847 d u r i n g the.years 1841 and 1842. London, H. Colburn. V o l . 1. 438 pp., 1 map. Skinner, C. K. Problems of Surplus E l k In Yellowstone Park. 1950 Wyoming Wil d L i f e . Wyoming Game and F i s h . " Commission, Cheyenne, Wyoming, XIV(6):22-26. 9S Stoddart, L. A, and D. I . Rasraussen. B i g Q-ame—Range L i v e -194-5 stock Competition on Western Ranges. Trans. 10th N. Am. W. Conf. pp. 251-256. T a y l o r . W. P. and H, C. Hahn. D i e - o f f s among the White-Stalled 1947 Deer i n the Edwards P l a t e a u of Texas. Jour, W. Mgt, 11(4):317-323. Warren, P. S. Banff Area, A l b e r t a . G e o l o g i c a l Survey o f 1927 Canada. G e o l o g i c a l S e r i e s . Memoirs 153, No.134, 94 pp., i l l u s . Wilcox, W. D. Lake L o u i s e , i n the Canadian Rocky Mountains. 1896" Excerpt from the J o u r n a l s and Proceedings of The Royal G e o g r a p h i c a l Society,..,!,. S a v i l e Row. F r a n c i s Edwards, 83 High Street,' 1 Marylebone W. 64 pp. W i l l i a m s , M. B. Through the Heart of the Rockies and 1921 S e l k i r k s . P u b l i s h e d under the d i r e c t i o n of S i r James Lougheed, M i n i s t e r of the I n t e r i o r , Ottawa. 105 pp* 3 append., 3 maps. Kootenay N a t i o n a l Park and The Banff-Windemere 1929 Highway, King's P r i n t e r , Ottawa. 45 pp., p l a t e s and maps. 

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