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Observability and habitat characteristics of the mountain goat (oreamnos americanus blainville, 1816)… Foster, Bryan Richard 1982

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OBSERVABILITY AND HABITAT CHARACTERISTICS OF THE MOUNTAIN GOAT (Oreamnos americanus BLAINVILLE, 1816) IN WEST-CENTRAL BRITISH COLUMBIA by BRYAN RICHARD FOSTER B . S c , U n i v e r s i t y Of B r i t i s h Columbia, 1976 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF r MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES Department Of Animal Science We accept t h i s t h e s i s as conforming to the r e q u i r e d standard THE UNIVERSITY OF BRITISH COLUMBIA December 1982 © Bryan R i c h a r d F o s t e r , 1982 In p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of the requirements f o r an advanced•degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and study. I f u r t h e r agree that p e r m i s s i o n f o r e x t e n s i v e copying.of t h i s t h e s i s f o r s c h o l a r l y purposes may be granted by the Head of my Department or by h i s or her r e p r e s e n t a t i v e s . I t i s understood that copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n p e r m i s s i o n . Bryan R. F o s t e r Department of Animal Science The U n i v e r s i t y of B r i t i s h Columbia 2075 Wesbrook Pl a c e Vancouver, Canada V6T 1W5 Date: 20 December 1982 i i A b s t r a c t Mountain goat s p a t i a l d i s t r i b u t i o n , o b s e r v a b i l i t y and u t i l i z a t i o n of h a b i t a t types was i n v e s t i g a t e d from May, 1976 to August, 1978. Data were c o l l e c t e d by s y s t e m a t i c a l l y viewing mountain goats from the ground along a f i x e d t r a n s e c t . Mountain goats were c o n t a g i o u s l y d i s t r i b u t e d w i t h i n the study area, with 99.5 percent of a l l observed groups o c c u r r i n g on s o u t h e r l y - f a c i n g slopes of Maroon Mountain. Areas of monthly range use overlapped e x t e n s i v e l y . The h i g h e s t l e v e l s of contagion o c c u r r e d d u r i n g August and December and were c o r r e l a t e d with i n c r e a s e s i n group s i z e . K i d d i n g and r u t t i n g l o c a t i o n s were g e n e r a l l y clumped i n t h e i r d i s t r i b u t i o n s , but no geographic c o n c e n t r a t i o n s o c c u r r e d . The p o p u l a t i o n was estimated at 137 animals, comprising 90 females and 47 males. Mountain goats were most observable between February and May, however, only 42 percent of the p o p u l a t i o n was v i s i b l e , on average. R e p r e s e n t a t i v e p o r t i o n s of the female component ( i . e . >50% of the estimated t o t a l ) were observed d u r i n g 13 of 16 months examined, however, male r e p r e s e n t a t i o n was evident i n on l y 6 months f o r the same p e r i o d . Females were more observable than males because they o c c u r r e d i n l a r g e r groups, u t i l i z e d l e s s d e n sely-vegetated t e r r a i n at most times of the year, and spent more time a c t i v e . The observed mountain goats d i d not u t i l i z e b i o t i c or a b i o t i c h a b i t a t s e q u a l l y , nor i n constant p r o p o r t i o n s . D i r e c t s o u t h - f a c i n g , s c a r c e l y - v e g e t a t e d areas were predominantly used year round and c o n s i s t e d p r i m a r i l y of a broken rock s u b s t r a t e on b l u f f s o r o u t c r o p s b e t w e e n 35 a n d 41° s l o p e . D e c i d u o u s s h r u b l a n d s were h e a v i l y u t i l i z e d d u r i n g s p r i n g ' g r e e n - u p ' . D e n s e l y v e g e t a t e d a r e a s were u s e d by m o u n t a i n g o a t s d u r i n g p e r i o d s o f e x t r e m e ( h o t o r c o l d ) t e m p e r a t u r e and w i n d . Many g o a t s were l o c a t e d i n t h e a l p i n e d u r i n g t h e summer, b u t i n t h e s u b a l p i n e d u r i n g o t h e r s e a s o n s . N e a r l y h a l f o f a l l m o u n t a i n g o a t o b s e r v a t i o n s were l o c a t e d i n t h e v i c i n i t y o f t r e e l i n e . G o a t s d i d n o t a p p e a r t o p r e f e r h a b i t a t s warmed by a t m o s p h e r i c t h e r m a l i n v e r s i o n s d u r i n g w i n t e r . G r o u p c o m p o s i t i o n a f f e c t e d h a b i t a t s e l e c t i o n . M a l e - o n l y g r o u p s d i s p l a y e d f a r l e s s f r e q u e n t c h o i c e o f h a b i t a t t h a n t h e i r f e m a l e c o u n t e r p a r t s . T h i s may be a t t r i b u t e d t o t h e d o m i n a n t s t a t u s a n d h i g h e r p r o d u c t i o n r e q u i r e m e n t s o f r e p r o d u c t i v e f e m a l e s . M a l e - o n l y g r o u p s s p e n t more .time r e s t i n g t h a n f e e d i n g o r t r a v e l l i n g . P r o p o r t i o n a t e l y more t i m e was s p e n t r e s t i n g i n p r e f e r r e d h a b i t a t s . M i x e d - f e m a l e g r o u p s were o b s e r v e d t o s p e n d more t i m e f e e d i n g t h a n r e s t i n g o r t r a v e l l i n g . Some p r e f e r r e d h a b i t a t s were u s e d p r i m a r i l y f o r f e e d i n g , w h i l e o t h e r s were u s e d m a i n l y f o r r e s t i n g . I n l i g h t o f t h e p o t e n t i a l b i a s e s i n v o l v e d w i t h o b s e r v a t i o n t e c h n i q u e s , i t i s f e l t t h a t i n t e r - p o p u l a t i o n c o m p a r i s o n s o f p o p u l a t i o n s t r u c t u r e and r a n g e use s h o u l d be t r e a t e d w i t h c a u t i o n , p a r t i c u l a r l y when p o p u l a t i o n s o c c u p y d i f f e r e n t b i o t o p e s a n d have been s u b j e c t e d t o v a r y i n g l e v e l s o f h u n t i n g o r o t h e r f o r m s o f human d i s t u r b a n c e . The f u l f i l m e n t o f c u r r e n t h a r v e s t s t r a t e g i e s o f t h e s p e c i e s c a n n o t be m o n i t o r e d w i t h o u t more a c c u r a t e i n v e n t o r y t e c h n i q u e s . Table of Contents A b s t r a c t i i L i s t of Tables v i L i s t of F i g u r e s v i i Acknowledgements ix I. INTRODUCTION 1 II . STUDY AREA 4 1. SELECTION OF STUDY LOCATION 4 1 .1 R a t i o n a l e 4 1.2 Study Area Boundaries 6 2. PHYSIOGRAPHY AND GEOLOGY 8 3. CLIMATE 8 4. SOILS AND VEGETATION 9 I I I . METHODS 10 1. FIELD RECONNAISSANCE 10 2. SURVEY METHODOLOGY 10 2.1 A e r i a l Surveys 10 2.2 Ground Surveys 11 2.2.1 V e h i c u l a r Counts . 11 2.2.2 Backpacking Observations 12 3. POPULATION CHARACTERISTICS 12 3.1 D i s t r i b u t i o n And Movements 13 3.1.1 D i s t r i b u t i o n 13 3.1.2 Movements 15 3.2 P o p u l a t i o n S t r u c t u r e 16 3.2.1 P o p u l a t i o n S i z e 16 3.2.2 P o p u l a t i o n D e n s i t y 17 3.2.3 P o p u l a t i o n Composition 17 4. HABITAT CHARACTERISTICS 19 4.1 H a b i t a t Use 20 4.1.1 B i o t i c F a c t o r s 20 4.1.2 A b i o t i c F a c t o r s 24 4.2 H a b i t a t S e l e c t i o n 28 4.2.1 Data C o n s i d e r a t i o n s 28 4.2.2 B i o t i c F a c t o r s 33 4.2.3 A b i o t i c F a c t o r s 33 4.2.4 Mountain Goat A c t i v i t y Within P r e f e r r e d H a b i t a t s 34 IV. RESULTS 35 1. POPULATION CHARACTERISTICS AND OBSERVABILITY ....35 1.1 D i s t r i b u t i o n And Movements 35 1.1.1 D i s t r i b u t i o n 35 1.1.2 Movements 41 1.2 P o p u l a t i o n S t r u c t u r e 45 1.2.1 P o p u l a t i o n S i z e And Densi t y 45 1.2.2 P o p u l a t i o n Composition 45 1.3 Mountain Goat O b s e r v a b i l i t y 46 1.3.1 Census R e s u l t s 46 1.3.2 O b s e r v a b i l i t y Of Sexes 50 2. HABITAT CHARACTERISTICS 52 2.1 H a b i t a t Use 52 2.1.1 B i o t i c F a c t o r s 52 V 2.1.2 A b i o t i c F a c t o r s 69 2.2 H a b i t a t S e l e c t i o n 81 2.2.1 Data C o n s i d e r a t i o n s 81 2.2.2 B i o t i c F a c t o r s 84 2.2.3 A b i o t i c F a c t o r s 86 2.2.4 Mountain Goat A c t i v i t y Within P r e f e r r e d H a b i t a t s 92 V. DISCUSSION 96 1. HABITAT CHARACTERISTICS 96 1 .1 H a b i t a t Use 96 1.1.1 U n i v a r i a t e A n a l y s i s 96 1.1.2 M u l t i v a r i a t e A n a l y s i s 106 1.2 H a b i t a t S e l e c t i o n 108 1.2.1 E c o l o g i c a l F i n d i n g s 109 1.2.2 Concepts Of S e l e c t i o n 111 2. MOUNTAIN GOAT OBSERVABILITY 115 2.1 I n t r a s p e c i f i c I n f l u e n c e s 117 2.1.1 S o c i a l Behaviour 117 2.1.2 A c t i v i t y P a t t e r n s 118 2.2 I n t e r s p e c i f i c I n f l u e n c e s 119 2.2.1 B i t i n g I n s e c t s 119 2.2.2 Hunting 119 2.3 Weather 120 VI. CONCLUSIONS AND RECOMMENDATIONS 122 1. IMPLICATIONS FOR STUDY DESIGN 122 2. MANAGEMENT IMPLICATIONS 124 LITERATURE CITED 126 v i L i s t of Tables 1. Summary of s e x u a l l y dimorphic c h a r a c t e r i s t i c s used to c l a s s i f y mountain goats in the study area 18 2. Monthly d i s t r i b u t i o n s t a t i s t i c s of mountain goats i n h a b i t i n g Maroon Mountain, B.C 38 3. Mountain goat s e l e c t i o n of v e g e t a t i v e cover on Maroon Mountain (south slope) f o r male-only and mixed-female groups 85 4. Mountain goat s e l e c t i o n of aspect on Maroon Mountain (south slope) f o r male-only and mixed-female groups. 87 5. Mountain goat s e l e c t i o n of h i l l s i d e slope on Maroon Mountain (south slope) by male-only and mixed-female groups 89 6. Mountain goat s e l e c t i o n of 250-m a l t i t u d i n a l zones on Maroon Mountain (south slope) by male-only and mixed-female groups 91 7. P r e f e r r e d monthly h a b i t a t s used p r i m a r i l y f o r feed i n g and r e s t i n g by mixed-female groups on Maroon Mountain, B.C 94 v i i L i s t of F i g u r e s 1. Regional l o c a t i o n of the study area ( s t i p p l e d area d e p i c t s a l p i n e mountain goat h a b i t a t ) 5 2. Topographic d e t a i l of the study area ( d e l i m i t e d by the dashed 1 ine) 7 3. D i s t r i b u t i o n of s y s t e m a t i c a l l y observed mountain goats on Maroon Mountain, B.C., determined by v e h i c u l a r survey 36 4. D i s t r i b u t i o n ' o f neonate l o c a t i o n s i n the study area -1976 to 1978 40 5. D i s t r i b u t i o n of 1977 r u t t i n g l o c a t i o n s i n the study area 42 6. Observed movements of "female#2" and "bar-nosed female" on Maroon Mountain, B.C. (numbers i n d i c a t e sequence of s i g h t i n g s ) 43 7. Observed movements of "unicorn female" and " l e f t horn-stub female" on Maroon Mountain, B.C. (numbers i n d i c a t e sequence of s i g h t i n g s ) 44 8. Summary of mountain goat counts on Maroon Mountain and Mount Garland, B.C. ( h o r i z o n t a l bar=mean; v e r t i c a l bar=standard d e v i a t i o n ) 48 9. O b s e r v a b i l i t y of male and female components of the Maroon Mountain goat p o p u l a t i o n ( h o r i z o n t a l bar=mean; v e r t i c a l bar = standard d e v i a t i o n ) 51 v i i i 10. V e g e t a t i v e cover of key mountain goat range (south aspect) on Maroon Mountain, B.C 53 11. Summary of mountain goat use of v e g e t a t i v e cover on Maroon Mountain, B.C -. 61 12. Summary of mountain goat use of herbaceous understory types on Maroon Mountain, B.C ..64 13. Summary of mountain goat use of krummholz s t r a t a on Maroon Mountain, B.C 65 14. Summary of mountain goat use of deciduous shrub s t r a t a on Maroon Mountain, B.C 67 15. Summary of mountain goat use of f o r e s t woody shrub u n d e r s t o r i e s on Maroon Mountain, B.C 68 16. Summary of mountain goat use of landforms on Maroon Mountain, B.C : 70 17. Summary of mountain goat use of rock s u b s t r a t e s on Maroon Mountain, B.C 72 18. Summary of mountain goat use of aspects on Maroon Mountain, B.C 74 19. Summary of mountain goat use of h i l l s i d e slope on Maroon Mountain, B.C 76 20. Summary of mountain goat a l t i t u d i n a l use by 250-m c l a s s e s on Maroon Mountain, B.C 78 21. Summary of mean monthly temperature values f o r s i x weather s t a t i o n s on Maroon Mountain, B.C 80 i x Acknowledgements F i n a n c i a l support f o r t h i s study was pro v i d e d by grants awarded to my s u p e r v i s o r , Dr. David M. Shackleton, from the N a t i o n a l Science and E n g i n e e r i n g Research C o u n c i l of Canada, the Canadian W i l d l i f e S e r v i c e and the U n i v e r s i t y of B r i t i s h Columbia. My hearty thanks are a l s o extended to Dr. Shackleton f o r h i s i n t e r e s t and guidance i n the study and to a l l of my committee members f o r t h e i r c r i t i c a l review of t h i s manuscript. The U n i v e r s i t y of B r i t i s h Columbia awarded me a Summer Research S c h o l a r s h i p i n 1977, an A g r i c u l t u r a l Sciences Research S c h o l a r s h i p f o r 1977/78 and a Summer U n i v e r s i t y Graduate F e l l o w s h i p i n 1982. I was a l s o awarded a Canadian W i l d l i f e S e r v i c e W i l d l i f e B i o l o g y S c h o l a r s h i p f o r 1978/79. The B r i t i s h Columbia F i s h and W i l d l i f e Branch p r o v i d e d funding f o r p r e l i m i n a r y fixed-wing a e r i a l reconnaissance f l i g h t s and s e v e r a l hours of h e l i c o p t e r time f o r i n v e n t o r y , a f i e l d o f f i c e at T e r r a c e , B.C., camping and other f i e l d equipment, and three summer student p o s i t i o n s between 1977 and 1978. In p a r t i c u l a r , I thank Drs. David H a t l e r and D a r y l l Hebert (Regional W i l d l i f e B i o l o g i s t s ) , and Don Eastman ( W i l d l i f e Research C o o r d i n a t o r ) . S p e c i a l thanks are X given to Conservation O f f i c e r David T. Crack, who not only shared h i s o f f i c e with me and v o l u n t a r i l y s u p p l i e d l o g i s t i c support, but whose f a m i l y made t h e i r home as much mine as t h e i r own f o r the p e r i o d I conducted f i e l d r e s e a r c h a c t i v i t i e s . Thanks are a l s o extended to the Donations Committee of B r i t i s h Columbia F o r e s t Products L i m i t e d f o r t h e i r kind f i n a n c i a l support of t h i s study; to Mr. Bob Marsh of the p r o v i n c i a l C l i m a t o l o g y D i v i s i o n (now the A i r S t u d i e s Branch) for h i s guidance and supply of equipment f o r the i n s t a l l a t i o n of weather s t a t i o n s ; to Mr. James C a l d w e l l , of Malkan I n t e r n a t i o n a l C o n s u l t a n t s L t d . , f o r h i s expert demonstrations of photogrammetric procedures; and to the Research D i v i s i o n of the B r i t i s h Columbia F o r e s t S e r v i c e f o r the use of t h e i r i n v e n t o r y h e l i c o p t e r to conduct a mountain goat census d u r i n g the summer of 1976. Dr. F.L. B u n n e l l , of the U n i v e r s i t y of B r i t i s h Columbia, a l s o s u p p l i e d m e t e o r o l o g i c a l equipment f o r t h i s study. A d d i t i o n a l l y , I express my g r a t i t u d e to Mr. Dave Newman, base manager of Okanagan H e l i c o p t e r s L t d . , f o r h i s h e l p f u l n e s s i n s u p p l y i n g t h e i r B e l l 206B 'Jet Ranger' h e l i c o p t e r i n what I can only term rather unorthodox c o n d i t i o n s , but most- imp o r t a n t l y , f o r h i s maintenance of the e x p e r t i s e r e q u i r e d by p i l o t s who f l y i n mountainous t e r r a i n . x i I would l i k e to thank Laura Danby, my second summer a s s i s t a n t , i n 1978, who t o i l e d f o r 3 and 1/2 months with the drudgery a s s o c i a t e d with data p r e p a r a t i o n . T h i s t h e s i s c o u l d never have been completed without the s p e c i a l d e d i c a t i o n of Engel Rahs, my f i r s t summer a s s i s t a n t (1977), eventual c o - r e s e a r c h e r , and recent f i a n c e e . T h i s study i s as much hers as i t i s mine, and I thank her not only f o r her moral support and r e f r e s h i n g companionship i n the f i e l d , but a l s o f o r her perseverence with the p r e p a r a t i o n of the data. F i n a l l y , and e q u a l l y important, I would l i k e to acknowledge the success of resp e c t which developed between o u r s e l v e s and the g r i z z l y bears that i n c l u d e d p o r t i o n s of the study area as a p a r t of t h e i r home. A token i n v e s t i g a t i o n of some of our f i e l d s u p p l i e s was the o n l y alarming encounter. x i i D e d i c a t i o n Mr. Paul Otto Shulte Born - 15 A p r i l 1911 (Kien, Germany) Died - 22 December 1977 ( T e r r a c e , Canada) T h i s t h e s i s i s d e d i c a t e d to Paul Otto S h u l t 6 , one of the true p i o n e e r s of the T e r r a c e r e g i o n . He became a c q u a i n t e d with the study area d u r i n g a p e r i o d when stage coaches t r a v e r s e d Kitsumkalum v a l l e y and when t r a i l s and c a b i n s were c l e a r e d and c o n s t r u c t e d by hand. Mr. S h u l t 6 o f t e n e n l i g h t e n e d us with s t o r i e s of the a r e a , p a r t i c u l a r l y with h i s acute o b s e r v a t i o n s of w i l d l i f e - most notably of which were the white Kermode bear (Ursus americanus  kermodei) and the mountain goat. Paul S h u l t 6 d i e d of n a t u r a l causes on the 22nd day of December, 1977, while checking h i s t r a p l i n e near Maroon Creek. He never r e t u r n e d , to my sorrow, to r e c e i v e the Christmas g r e e t i n g s l e f t a w a i t i n g him. x i i i P o s t s c r i p t During the f a l l of 1978, the goats and the bears were once again p e r m i t t e d to r e c l a i m t h e i r v a l l e y , as t o r r e n t i a l r a i n s and f l o o d i n g washed away the l i n g e r i n g rememberances of Mr. Jack Garland and Mr. Paul S h u l t e , who i n 1938 e r e c t e d the miner's c a b i n (background of D e d i c a t i o n photo) i n which we l i v e d . V i r t u a l e l i m i n a t i o n of the Maroon Creek l o g g i n g road network ensued, thereby h a l t i n g (at l e a s t t e m p o r a r i l y ) almost a l l forms of d i s r u p t i n g human a c t i v i t y w i t h i n the v a l l e y . Thank you Rain - from a l l of us i n the v a l l e y . 1 I. INTRODUCTION Approximately 45 percent of the world's f r e e - r a n g i n g mountain goats (Oreamnos americanus) r e s i d e i n B r i t i s h Columbia (Johnson 1977, M i n i s t r y of Environment 1979). In the l a s t h a l f c entury, however, many goat p o p u l a t i o n s were exposed to a c c e l e r a t i n g hunting p r e s s u r e s through i n c r e a s e d v e h i c u l a r access (Quaedvlieg et a l . 1973, Phelps et a l . 1975). By the l a t e 1960's and the e a r l y 1970's, p r o v i n c i a l w i l d l i f e b i o l o g i s t s became aware of the consequences of gross mismanagement of the s p e c i e s through unregulated access (Demarchi p e r s . comm.). Harvest trends d e c l i n e d ( F o s t e r 1977) and some p o p u l a t i o n s were s e v e r e l y reduced or o b l i t e r a t e d (Phelps et a l . 1975). S t r i c t e r r e g u l a t i o n s were c r e a t e d , but were o f t e n unenforced. I n t e r a c t i o n s and c o n f l i c t s between mountain goats and v a r i o u s i n d u s t r i e s and r e c r e a t i o n i s t s are i n c r e a s i n g i n many areas of the p r o v i n c e . The need f o r more i n t e n s i v e ( i . e . watershed) management was r e c o g n i z e d f o r the s p e c i e s (Phelps et a l . 1975, Hebert 1978, Jamieson 1978), p a r t i c u l a r l y given that h a b i t a t v a r i a t i o n was i d e n t i f i e d as an important determinant of r e g i o n a l l y - s p e c i f i c c h a r a c t e r i s t i c s of mountain goat ecology (Fos t e r 1976, Hebert and T u r n b u l l 1977). Another e f f e c t of a c c e s s , l o s s of h a b i t a t , i s a l s o b e l i e v e d to have c o n t r i b u t e d to the severe r e d u c t i o n of some mountain goat p o p u l a t i o n s (Quaedvlieg et a l . 1973), yet no one i n B r i t i s h Columbia has documented range a t t r i b u t e s necessary f o r s u c c e s s f u l goat p r o d u c t i o n . P u b l i s h e d s t u d i e s of mountain goats i n t h i s p r o v i n c e p r i o r to 1976 are l i m i t e d to a taxonomic 2 examination of subspecies (McCrory 1965, Cowan and McCrory 1970), a study on the use of s a l t l i c k s (Hebert 1967, Hebert and Cowan 1971a), a n a t u r a l h i s t o r y study (Holroyd 1967), a b e h a v i o u r a l i n v e s t i g a t i o n (DeBock 1970), and an account of white muscle d i s e a s e (Hebert and Cowan 1971b). A l l f i e l d s t u d i e s were conducted i n the Kootenay r e g i o n of southwestern B r i t i s h Columbia. The need f o r l o c a l predevelopment s t u d i e s on mountain goat requirements has been p u b l i c i z e d only more r e c e n t l y ( H a t l e r 1978). At present, the s t a t e of Alaska i s l e a d i n g r e s e a r c h i n t h i s area as a r e s u l t of impending c o a s t a l l o g g i n g a c t i v i t i e s (Fox 1980, Fox and Taber 1981, Fox and Raedeke 1982, Schoen et a l . 1981, Smith 1981). These p r e l i m i n a r y s t u d i e s have confirmed that mountain goats r e q u i r e mature subalpine timber h a b i t a t , p a r t i c u l a r l y d u r i n g winter. Because moist c o a s t a l environments enable p r o d u c t i v e f o r e s t growth with h i g h commercial value (Resource P l a n n i n g Unit 1976), the advent of l o g g i n g poses a much more l i k e l y t h r e a t to l o s s of mountain goat h a b i t a t i n these regions than i t does in the d r i e r i n t e r i o r a r e a s . The broad purpose of t h i s study was to document both s p a t i a l and temporal p a t t e r n s of h a b i t a t use by mountain goats w i t h i n a densely populated r e g i o n of c o a s t a l w e s t - c e n t r a l B r i t i s h Columbia. The i n v e s t i g a t i o n was designed as an o b s e r v a t i o n a l study p r i m a r i l y due to inadequate funding f o r the p r o v i s i o n of r a d i o telemetry equipment. I t was f e l t t h a t the study area to be s e l e c t e d must o f f e r h i g h viewing c a p a b i l i t i e s , r e s u l t i n g i n two study o b j e c t i v e s . 3 One o b j e c t i v e of t h i s i n v e s t i g a t i o n was to document o b s e r v a t i o n a l c h a r a c t e r i s t i c s of the study p o p u l a t i o n which c o u l d a s s i s t i n determining (a) survey b i a s , and (b) the subsequent l e v e l of a n a l y s i s and degree of i n t e r p r e t a t i o n permissable under the main o b j e c t i v e . The hypothesis of the f i r s t o b j e c t i v e , presented i n i t s A l t e r n a t e (Ha) form, i s : • Mountain goats occur i n non-random p a t t e r n s of s p a t i a l d i s p e r s i o n . T h i s h y p o t h e s i s i n turn i s concerned with the main o b j e c t i v e of the study which was to i n v e s t i g a t e r e l a t i o n s h i p s of mountain goat h a b i t a t u t i l i z a t i o n . In t h i s study, ' h a b i t a t ' i s r e c o g n i z e d to c o n s i s t of both b i o t i c and a b i o t i c f a c t o r s . The h y p o t h e s i s formulated and t e s t e d under t h i s o b j e c t i v e , presented in i t s A l t e r n a t e (Ha) form, i s : • Mountain goats use some or a l l of the b i o t i c and a b i o t i c components of t h e i r range d i s p r o p o r t i o n a t e to t h e i r a v a i l a b i l i t y . The assumptions r e q u i r e d of t h i s a n a l y s i s , and t h e i r concomittant methods of accommodation, are d i s c u s s e d i n subsequent p o r t i o n s of t h i s t h e s i s . 4 I I . STUDY AREA 1. SELECTION OF STUDY LOCATION 1.1 R a t i o n a l e The s e m i - c o a s t a l Nass Ranges was chosen as the study region due to i t s abundance of p r o d u c t i v e mountain goat h a b i t a t ( M i n i s t r y of Environment 1979). High mountain goat d e n s i t i e s are a t t r i b u t e d to the north-south 'herring-bone' c o n f i g u r a t i o n of the ranges, r e s u l t i n g i n numerous v a l l e y s l a y i n g east-to-west and o f f e r i n g numerous steep, s o u t h - f a c i n g mountain s i d e s with an abundance of a l p i n e h a b i t a t ( F i g u r e 1). F i v e p r o s p e c t i v e study s i t e s were i n i t i a l l y chosen a f t e r review of a F i s h and W i l d l i f e Branch p r e l i m i n a r y r e p o r t (Sumanik 1970) and subsequent examination of 1:50,000 topographic maps of the a r e a . A e r i a l reconnaissance of the Nass Ranges (96 km long by 46 km wide) was i n i t i a l l y undertaken by the author d u r i n g l a t e A p r i l of 1976. Maroon Mountain was chosen as the study s i t e f o r the f o l l o w i n g reasons; (1) A dense mountain goat p o p u l a t i o n of more than one hundred animals was known to i n h a b i t the mountain (Sumanik 1970). (2) I t was a c c e s s i b l e by (a) a l o g g i n g road network i n the v a l l e y to the south and to the west, and (b) a w e l l - d e f i n e d t r a i l to the a l p i n e on the northwest s i d e of the mountain. (3) The township of T e r r a c e was l o c a t e d l e s s than 40 km to the south, minimizing f e r r y time f o r fixed-wing and h e l i c o p t e r surveys, and time f o r o b t a i n i n g s u p p l i e s . 5 NASS RANGES (Hazelton Mountains) F i g u r e 1 - Regional l o c a t i o n of the study area ( s t i p p l e d area d e p i c t s a l p i n e mountain goat h a b i t a t ) . 6 (4) E x i s t i n g roads along Maroon Creek v a l l e y enabled e x c e l l e n t ground o b s e r v a t i o n c a p a b i l i t y of the e n t i r e south slope of Maroon Mountain. (5) Other secondary roads i n the adjacent Kitsumkalum v a l l e y enabled ground o b s e r v a t i o n north of the main mountain r i d g e and west of the main peaks from a d i s t a n c e of 3 to 8 km, t h e r e f o r e e l i m i n a t i n g the need to hike to the n o r t h - f a c i n g a l p i n e d u r i n g systematic sampling. (6) Two cabins were a v a i l a b l e f o r winter o b s e r v a t i o n ; one at t r e e l i n e on the n o r t h aspect and the other near the head of Maroon Creek v a l l e y i n the v a l l e y bottom. 1.2 Study Area Boundaries The proposed study area was separated by small secondary v a l l e y s to the north and to the south, and a l a r g e g l a c i a t e d v a l l e y and lake to the west. The e a s t e r n boundary was c o n s i d e r e d to be the d i v i d e of the Nass Ranges (Figure 2 ) . Telemetry s t u d i e s i n A l a s k a (Schoen et a l . 1981, Smith 1981, N i c h o l s i n p r e s s ) , Idaho (Kuck 1973) and Montana (Chadwick 1973, Rideout 1974) have shown that mountain goat home ranges are l e s s than the o u t l i n e d r e g i o n (75 km2) chosen as the study a r e a . A d d i t i o n a l l y , c o n v e r s a t i o n s with l o c a l l o g g e r s and other r e s i d e n t s suggested t h a t goats d i d not c r o s s the secondary v a l l e y s to the north and the south. 7 F i g u r e 2 - Topographic d e t a i l of the study area ( d e l i m i t e d by the dashed l i n e ) . 8 2. PHYSIOGRAPHY AND GEOLOGY Located 35.4 km north of the township of T e r r a c e (54° 47' n o r t h l a t i t u d e and 128° 39' west l o n g i t u d e ) , the study area l i e s w i t h i n the Skeena River drainage system. Maroon Mountain i s t y p i c a l i n landform of the Nass Ranges, c o n s i s t i n g of rugged t e r r a i n with a h i g h and narrow knife-edge r i d g e . The region was h e a v i l y g l a c i a t e d , with the higher peaks t y p i c a l l y being s c u l p t u r e d by c i r q u e g l a c i e r s ( H o l l a n d 1976). The three main peaks of Maroon Mountain are c l u s t e r e d and range between 2042 and 2066 m i n h e i g h t . The study area occurs w i t h i n a southern p o r t i o n of a metamorphosed v o l c a n i c and sedimentary rock b a t h o l i t h comprising the Bowser Group of the upper J u r a s s i c and Cretaceous age. D u f f e l and Souther (1964) d e s c r i b e r e g i o n a l geology i n more d e t a i l . 3. CLIMATE The nearest c o n t i n u o u s l y o p e r a t i n g government c l i m a t o l o g i c a l f a c i l i t y i s l o c a t e d at the T e r r a c e a i r p o r t , 50 km to the south. The o v e r a l l c l i m a t e v a r i e s s e a s o n a l l y from a moist, c o a s t a l , mesothermal environment to a d r i e r , more c o n t i n e n t a l - l i k e , p s e u d o - a r c t i c regime. T o t a l annual p r e c i p i t a t i o n averages 1301 mm, comprising 371.3 cm (29%) of u s u a l l y wet snow (water equivalent=371 mm). The annual temperature averages 6.0°C, with the h o t t e s t months being J u l y and August (x monthly temperature=16.2°C and 15.6°C, r e s p e c t i v e l y ) . Temperatures may reach 35°C d u r i n g the summer 9 and -25°C i n the w i n t e r . Mean annual s o l a r i n s o l a t i o n approaches 1375 hours, or 31% of the p o s s i b l e annual sunshine t o t a l . The s u n n i e s t month i s g e n e r a l l y August and the c l o u d i e s t i s most o f t e n December. Winds are g e n e r a l l y from the south d u r i n g snow-free months of the year ( A p r i l to October), however, f a l l and winter are the w i n d i e s t times of the year, with c o l d n o r t h e r l y winds. Monthly m e t e o r o l o g i c a l data are summarized in annual government r e p o r t s (Environment Canada 1976,1977,1978). 4. SOILS AND VEGETATION Maroon Mountain l i e s w i t h i n a h e a v i l y u t i l i z e d p o r t i o n of the newly e x p l o i t e d Skeena P u b l i c Sustained Y i e l d U n i t ( B r i t i s h Columbia F o r e s t S e r v i c e 1979). Spruce (P i c e a s p . ) , hemlock (Tsuga s p . ) , and f i r (Abies sp.) predominate the commercially v a l u a b l e timber s p e c i e s . Three b i o g e o c l i m a t i c zones ( K r a j i n a 1965) occur i n the study a r e a . The C o a s t a l Western Hemlock zone i s the wettest zone, o c c u r r i n g below 762 m above sea l e v e l . O r t h i c humo-ferric s o i l s are p r e v a l e n t i n t h i s zone ( V a l e n t i n e et a l . 1978). The Mountain Hemlock zone i s evident between 762 and 1554 m, r e t a i n i n g s i m i l a r s o i l types to the C o a s t a l Western Hemlock zone but having only one-half of i t s t o t a l annual p r e c i p i t a t i o n . The A l p i n e Tundra zone g e n e r a l l y occurs above 914 to 1554 m, depending on aspect, and has only o n e - t h i r d the p r e c i p i t a t i o n of the C o a s t a l Western Hemlock zone. D y s t r i c b r u n i s o l s and mini podzols predominate. 10 I I I . METHODS 1. FIELD RECONNAISSANCE P r e l i m i n a r y o b s e r v a t i o n s were conducted throughout the summer of 1976 (May to August), and f o r two weeks i n each of the months of October and December of that year. Observations recommenced i n May 1977 and continued f o r 16 c o n s e c u t i v e months, t e r m i n a t i n g i n e a r l y September 1978. In a l l , mountain goats were observed over three k i d d i n g p e r i o d s and three summers, one r u t t i n g p e r i o d and one e n t i r e w i n t e r . 2. SURVEY METHODOLOGY The study area was surveyed by one of two modes of t r a v e l : a i r and ground. A e r i a l surveys were designed to p r o v i d e b e t t e r data on p o p u l a t i o n s i z e , composition, and d i s t r i b u t i o n than were a v a i l a b l e from p r e v i o u s s t u d i e s (Sumanik 1970). Ground ( v e h i c u l a r and backpacking) surveys were conducted to o b t a i n l e s s c o s t l y data on both p o p u l a t i o n and h a b i t a t c h a r a c t e r i s t i c s . 2.1 A e r i a l Surveys Fixed-wing a i r c r a f t were used f o r two reconnaissance surveys. In each case, one t r a n s e c t was flown j u s t below t r e e l i n e , at approximately 1050 m, c o v e r i n g a b e l t 300 m wide. A h e l i c o p t e r was used f o r t r a n s p o r t and more i n t e n s i v e a e r i a l survey due to a d d i t i o n a l m a n e u v e r a b i l i t y , speed c o n t r o l and la n d i n g c a p a b i l i t i e s . However, funds were not s u f f i c i e n t to f u l f i l l the i n t e n s i v e survey methodology designed, t h e r e f o r e f l i g h t paths d i d not f o l l o w one s p e c i f i c contour, nor were they 11 maintained at a constant speed. Instead they zig-zagged over the study area sampling mountain goats p r o p o r t i o n a l to d e n s i t y ( i . e . from group to group), r a t h e r than c o v e r i n g a l l areas, or areas of no apparent use. Mountain goats were seldom sexed on these surveys, and only kid s and y e a r l i n g s were i d e n t i f i e d to age c l a s s . A d d i t i o n a l data were c o l l e c t e d on group s i z e and d i s t r i b u t i o n ( i n c l u d i n g e l e v a t i o n ) i n most cases. 2.2 Ground Surveys Two types of ground survey methods were used; v e h i c u l a r counts and backpacking o b s e r v a t i o n s . Observations of mountain goats were made with the a i d of 8X30 b i n o c u l a r s , a s p o t t i n g scope with 20X and 40X o c u l a r o b j e c t i v e s and a 20-40X zoom t e r r e s t r i a l , b i n o c u l a r t e l e s c o p e . Mountain goats were u s u a l l y observed from d i s t a n c e s g r e a t e r than 0.25 km, to e l i m i n a t e human d i s t u r b a n c e . A 35 mm camera was u t i l i z e d to document aspects of mountain goat ecology. 2.2.1 V e h i c u l a r Counts V e h i c u l a r surveys were conducted at approximately weekly i n t e r v a l s i n 1976, and biweekly i n 1977 and 1978. Weather and v i s i b i l i t y l e v e l s a f f e c t e d t i ming of surveys with respect to date of choice and time of day. A pickup truck was used d u r i n g snow-free months and a snowmobile d u r i n g winter. Mountain goats were l o c a t e d by systematic scanning of the t e r r a i n from f i x e d viewing l o c a t i o n s along a permanent road t r a n s e c t . Surveys were i n i t i a t e d at opposite ends of the t r a n s e c t each time i n order to reduce sampling b i a s . On 12 average, incomplete surveys covered one-half of the t r a n s e c t , and so were i n c l u d e d i n the a n a l y s i s of h a b i t a t r e l a t i o n s h i p s . O c c a s s i o n a l l y a survey had to be completed the f o l l o w i n g day, due to the onset of inclement weather, or dusk. When p o s s i b l e , o b s e r v a t i o n s were terminated l a t e i n the day a t a major geographic landmark and r e i n i t i a t e d e a r l y the subsequent morning. T h i s dusk-dawn c o n t i n u a t i o n o f t e n supported the b e l i e f t h a t l a r g e s c a l e movements by goats d i d not occur o v e r n i g h t . 2.2.2 Backpacking Observations Backpacking t r i p s were conducted at l e a s t once a month, g e n e r a l l y d u r i n g snow-free p e r i o d s o n l y . T h i s type of survey was c o n s i d e r e d to be a g e n e r a l , non-systematic reconnaissance, p r o v i d i n g supplemental i n f o r m a t i o n on mountain goat d i s t r i b u t i o n and range use. Backpacking survey data were not u t i l i z e d f o r s t a t i s t i c a l a n a l y s i s of p o p u l a t i o n estimates or h a b i t a t - r e l a t e d c h a r a c t e r i s t i c s of mountain goats, due to d i s t u r b a n c e f a c t o r s . 3. POPULATION CHARACTERISTICS A complete assessment of h a b i t a t s e l e c t i o n may r e q u i r e knowledge on the s t r u c t u r e of the population, under study (Oosenbrug and Theberge 1980). Thus i n f o r m a t i o n on p o p u l a t i o n d i s t r u b u t i o n and movements, s i z e , d e n s i t y and composition were c o l l e c t e d . 1 3 3.1 D i s t r i b u t i o n And Movements Pa t t e r n s of mountain goat d i s t r i b u t i o n and movements were recorded by g r i d l o c a t i o n d u r i n g a e r i a l and ground surveys. The northeast s e c t o r of the study area was sampled once a month du r i n g snow-free p e r i o d s by backpacking or h e l i c o p t e r surveys, and on a monthly b a s i s by h e l i c o p t e r only d u r i n g snowy c o n d i t i o n s (October through May). 3.1.1 D i s t r i b u t i o n The a n a l y s i s of mountain goat d i s t r i b u t i o n p a t t e r n s c o n s i d e r s n e i t h e r age nor sex c a t e g o r i e s , with the exception of k i d d i n g and r u t t i n g i n f o r m a t i o n , which do c o n s i d e r a d u l t males and females. a. Range Use Maps r e p r e s e n t i n g o v e r a l l and monthly cumulative range use by mountain goats were c o n s t r u c t e d f o r the p e r i o d May, 1977 through August, 1978. Mountain goat s i g h t i n g s i n 1976 were not recorded by g r i d l o c a t i o n but by approximate geographic f i x e s . Most mountain goat o b s e r v a t i o n s were p l o t t e d with an accuracy of ±10 m (one h a l f of the l e n g t h of one g r i d quadrant), with the exception of some homogenous (e.g. a l p i n e ) areas where an accuracy of ±30 m i s b e l i e v e d to be more r e a l i s t i c . A r e a l d e t e r m i n a t i o n of monthly range use was measured by dot g r i d (64 dots/km 2), based upon v e r t i c a l p r o j e c t i o n of monthly d i s t r i b u t i o n p a t t e r n s . 14 The m o d i f i e d minimum area polygon method (Harvey and Barbour 1965) was used to determine monthly range boundaries. b. S p a t i a l D i s p e r s i o n Monthly d i s p e r s i o n p a t t e r n s of mountain goats, o c c u r r i n g south of the main r i d g e only, were measured fo r degree of a g g r e g a t i o n . The purpose of t h i s e x e r c i s e was to determine i f range resources were being u t i l i z e d a t random ( i . e . p o i s s o n d i s t r i b u t i o n ) or i f c e r t a i n a t t r i b u t e s were r e s p o n s i b l e f o r uniform or clumped d i s t r i b u t i o n a l p a t t e r n s . An index of d i s p e r s i o n (I) ( E l l i o t 1971, p40) was used to d e s c r i b e monthly d i s p e r s i o n p a t t e r n s of mountain goats. Myers (1978) s t a t e d that t h i s index i s easy to compute and r e a d i l y understandable ( i n comparison to Green's and s t a n d a r d i z e d M o r i s i t a ' s c o e f f i c i e n t s ) . A d d i t i o n a l l y , ' I ' was found by Myers (1978) to be only weakly c o r r e l a t e d with d e n s i t y (or quadrat s i z e ) . The value ' I ' i s simply the (sample) variance-to-mean r a t i o . A contiguous g r i d system comprised of 129 400mx400m quadrats was superimposed over the g r i d maps of animal l o c a t i o n s and each quadrat was t a l l i e d by month and i n t o t a l . The quadrat s i z e used i s l a r g e r than that recommended by Greig-Smith (1964) fo r sampling p l a n t communities, because too many c e l l s with zero counts would have r e s u l t e d . Quadrat s i z e was chosen on the b a s i s of 15 mean d a i l y movements of mountain goats (x=400 m) i n a nearby study ( F o s t e r and Rahs 1981). Monthly v a l u e s of ' I ' i n excess of 3.0 were f i t t e d to a general negative binomial model (Petkau and Johnson, i n p r e s s ) . The c h i - s q u a r e s t a t i s t i c (X 2) was used to determine d i f f e r e n c e s between monthly d i s t r i b u t i o n s , at the 0.05 l e v e l of s i g n i f i c a n c e . Mountain goat k i d d i n g and r u t t i n g d i s t r i b u t i o n s were s i m i l a r l y c a l c u l a t e d , but small sample s i z e s and the l a r g e number of quadrats negated the use of Petkau and Johnson's ( i n p r e s s ) model. For these data, a X 2 value was c a l c u l a t e d ( E l l i o t 1971, p40) and r e d i s t r i b u t e d to the normal v a r i a b l e , 'd'. Agreement with a poisson d i s t r i b u t i o n was not r e j e c t e d (p>0.05) i f the a b s o l u t e value of 'd' was l e s s than 1.96. I f 'd' was g r e a t e r than 1.96, with a negative s i g n , a uniform d i s t r i b u t i o n was suspected. I f 'd' was g r e a t e r than 1.96 and p o s i t i v e , a contagious d i s t i b u t i o n was i m p l i e d . 3.1.2 Movements Mountain goats were i d e n t i f i e d f o r o b s e r v a t i o n by a nylon colour-coded neck c o l l a r and two b l a c k , r u b b e r i z e d 'Ritcey' ear-tags, each p a i n t e d with a white number, a f f i x e d subsequent to f r e e - r a n g i n g i m m o b i l i z a t i o n with 2.7 mg doses (1.1 cc) of Immobilon (etorphine h y d r o c h l o r i d e p l u s acepromazine - R e c k i t t and Colman, London, England). I n d i v i d u a l s were a l s o i d e n t i f i e d by morphological c h a r a c t e r i s t i c s , i n c l u d i n g broken horns, horn 1 6 d e f o r m i t i e s , s c a r s , molting p a t t e r n and pelage c o l o u r a t i o n . C e r t a i n i n d i v i d u a l s or groups which remained very c o h e s i v e , c o u l d be monitored f o r s e v e r a l c o n s e c u t i v e days, however, t h i s was not a common t r e n d . D a i l y movements were obtained by watching s e v e r a l groups of goats from dawn u n t i l dusk and r e c o r d i n g t h e i r l o c a t i o n s on a topographic map. I n d i v i d u a l goats were monitored over a longer p e r i o d by r e s i g h t i n g tagged and other u n i q u e l l y marked i n d i v i d u a l s . A 6 km2 p o r t i o n of the northeast corner of Mt. Garland, to the south, was covered d u r i n g each survey i n order to h e l p document the extent of immigration and emmigration to and from the study a r e a . 3.2 P o p u l a t i o n S t r u c t u r e Components of p o p u l a t i o n s t r u c t u r e were d e s c r i b e d i n terms of i t s s i z e , d e n s i t y , and composition. 3.2.1 P o p u l a t i o n S i z e P o p u l a t i o n s i z e estimates were d e r i v e d by the 'maximum cohort completion' method (Smith 1976, p35), i n which the maximum counts f o r each sex and age c l a s s were added together to o b t a i n a 'maximum observed count'. T h i s f i g u r e r e p r e s e n t s a minimum l i k e l y herd estimate. The technique was a p p l i e d to the v e h i c u l a r survey data o n l y . Information c o l l e c t e d on h e l i c o p t e r surveys was s t r a t i f i e d and compared to ground v e h i c u l a r survey areas, i n order to determine survey e f f i c i e n c y of the two techniques. P o p u l a t i o n estimates were d e r i v e d from i n d i c e s of ab s o l u t e p o p u l a t i o n coverage p r o v i d e d i n the l i t e r a t u r e (McCrory and Blood 1978, F o s t e r and Rahs 1981). These must be c o n s i d e r e d 17 as approximate, f o r t h e i r survey methods were o u t l i n e d i n c o m p l e t e l y . 3.2.2 P o p u l a t i o n D e n s i t y Mountain goat d e n s i t i e s were c a l c u l a t e d based upon c o r r e c t e d map a r e a . A r e a l d e t e r m i n a t i o n s were conducted with a dot g r i d (64 dots/km 2) and c o r r e c t e d by the mean h i l l s i d e slope of the area of mountain goat d i s t r i b u t i o n ( f o r more d e t a i l , see 4.2 H a b i t a t S e l e c t i o n ) . D e n s i t y was determined by d i v i d i n g the number of mountain goats observed i n t o the c o r r e c t e d survey a r e a . 3.2.3 P o p u l a t i o n Composition Mountain goats were c l a s s i f i e d by sex and age, on the b a s i s of horn morphology and behaviour, f o l l o w i n g the f i r s t summer of i n t r o d u c t o r y f i e l d work. One of f i f t e e n sex-age c a t e g o r i e s were recorded f o r each i n d i v i d u a l , based on three sex and f i v e age c l a s s e s . a. Sex As many of the c h a r a c t e r i s t i c s presented i n Table 1 as p o s s i b l e were used to i d e n t i f y males from females. Most sexing was a l s o estimated independently by a f i e l d a s s i s t a n t , and then compared. Where d i s c r e p a n c i e s o c c u r r e d between the two observers, or whenever sex c o u l d not be otherwise determined, the sex was designated as ' u n c l a s s i f i e d ' . 18 T a b l e 1 - Summary o f s e x u a l l y d i m o r p h i c c h a r a c t e r i s t i c s u s e d t o c l a s s i f y m o u n t a i n g o a t s i n t h e s t u d y a r e a . Sexual C h a r a c t e r i s t i c Male Female Reference symmetric horn curvature thick horns with bases nearly touching ' Roma n nos e 1 ad ul t c h a r a c t e r i s t i c s t r a i g h t e r horns, t y p i c a l l y 'kinked' near the t i p s narrow horns with bases well-spaced rostrum s t r a i g h t Casebeer et a l . 1950 Brandborg 1950,1955 Brandborg 1955 Hibbs 1965 Chadwick 1973 stretch stance urinary posture large s c r o t a l sac black, swollen o c c i p i t a l glands during the rut so i l e d rump, b e l l y and carpal j o i n t s during rut early spring molt - generally s o l i t a r y squatting urinary posture vulva and teats pink or flesh-coloured, swollen o c c i p i t a l glands no ' r u t t i n g - p i t ' a c t i v i t y barren females same as male other females molt during summer usually s o c i a l , frequently accompanied by young of the year DeBock 1970 Brandborg 1955 DeBock 19 70 Brandborg 1955 Geist 1964 Geist 1964 DeBock 1970 Brandborg 1955 Hebert 1967 Chadwick 1973 Brandborg 19 55 DeBock 19 70 Chadwick 1973 19 b. Age Age c a t e g o r i e s were d e f i n e d as; k i d , y e a r l i n g , subadult, a d u l t or u n c l a s s i f i e d . Kids and y e a r l i n g s were r e a d i l y d i f f e r e n t i a t e d on the b a s i s of body s i z e , horn growth (Fos t e r 1978) and conformation of the face (DeBock 1970; Smith 1976). In t h i s study, young mountain goats are c o n s i d e r e d k i d s u n t i l t h e i r f i r s t b i r t h d a y , and y e a r l i n g s u n t i l t h e i r second b i r t h d a y . A 'bi r t h d a y ' was c o n s i d e r e d to be the date of the f i r s t observed new-born of the year. I d e n t i f i c a t i o n of two-year o l d s was not attempted, due to v a r i a t i o n s i n horn growth i d e n t i f i e d by F o s t e r (1978), d e s p i t e the f a c t that other r e s e a r c h e r s c l a i m t o be capable of i d e n t i f y i n g t h i s cohort (e.g. DeBock 1970, Chadwick 1973, Thompson 1981). In t h i s study, two-year and o l d e r animals were c l a s s i f i e d , based p r i m a r i l y upon the width and the l e n g t h of the rostrum (see Smith 1976) and to a l e s s e r extent on horn s i z e . Subadults were c o n s i d e r e d to c o n s i s t of two and three-year o l d animals. Mountain goats estimated to be o l d e r than three years were c o n s i d e r e d a d u l t s . 4. HABITAT CHARACTERISTICS Important h a b i t a t v a r i a b l e s were i d e n t i f i e d d u r i n g the f i r s t summer's ad l i b , o b s e r v a t i o n s of the study p o p u l a t i o n . B i o t i c and a b i o t i c c h a r a c t e r i s t i c s of the h a b i t a t were assessed in terms of v e g e t a t i v e cover, p h y s i c a l s t r u c t u r e , and 20 microthermal v a r i a t i o n . V e g e t a t i v e cover of a p o i n t on a g r i d was recorded as the g r e a t e s t p o r t i o n of l a n d l y i n g w i t h i n that g r i d ' s quadrant (400 m 2). The terms ' a l p i n e ' and 'subalpine' r e f e r to areas above and below t r e e l i n e , r e s p e c t i v e l y . 4.1 H a b i t a t Use I n v e s t i g a t i o n of h a b i t a t use was conducted throughout the e n t i r e study area with the e x c e p t i o n of the northeast s e c t i o n (Ridges #1 and #2, see F i g u r e 2). V a r i a b l e s measured were; p l a n t cover, s u r f a c e e x p r e s s i o n , s u b s t r a t e , aspect, s l o p e , e l e v a t i o n , and temperature. V a r i a b l e s were c a t e g o r i z e d a p r i o r i and each i n t e r v a l was i n d i v i d u a l l y addressed i n the a n a l y s i s . Data were analyzed u s i n g s i n g l e c l a s s i f i c a t i o n l o g l i k e l i h o o d r a t i o (G) g o o d n e s s - o f - f i t t e s t s (Sokal and Rohlf 1980, p692) at the 0.05 l e v e l of s i g n i f i c a n c e . W i l l i a m ' s c o r r e c t i o n f o r c o n t i n u i t y was a p p l i e d to the G - s t a t i s t i c , f o r two-by-two comparisons on l y , i n order to o b t a i n a b e t t e r approximation to the c h i - s q u a r e d i s t r i b u t i o n (Sokal and Rohlf 1980, p 7 l 0 ) . A F o r t r a n program developed by M. G i l l i n g h a m ( F a c u l t y of F o r e s t r y , U n i v e r s i t y of B r i t i s h Columbia) and the author was used to c a l c u l a t e the G - s t a t i s t i c . 4.1.1 B i o t i c F a c t o r s a. V e g e t a t i v e Cover C l a s s i f i c a t i o n of b i o t i c landscapes i n the study area were based upon a p h y s i o g n o m i c - e c o l o g i c a l c l a s s i f i c a t i o n scheme o u t l i n e d by Mueller-Dombois and E l l e n b e r g (1974, p466). P l a n t formations were 21 d e l i n e a t e d from 1:15,840 black and white government a e r i a l photographs (Surveys and Mapping Branch 1974). Ground cover was d e f i n e d as the percentage of p o i n t s covered by the a e r i a l p a r t s of a p l a n t type under c o n s i d e r a t i o n , and was estimated v i s u a l l y . A v e g e t a t i o n map of approximately the same s c a l e was c o n s t r u c t e d of the area, i n 1976, with a K a i l p l o t t e r , u s i n g standard photogrammetric techniques (Slama 1980). Three f i e l d reconnaissance m i s s i o n s confirmed the f i n a l map d r a f t . P l a n t specimens were c o l l e c t e d d u r i n g the summer p e r i o d s , pressed and a i r - d r i e d , and l a t e r v e r i f i e d f o r i d e n t i f i c a t i o n at the U n i v e r s i t y of B r i t i s h Columbia herbarium. S c i e n t i f i c names were taken from the nomenclature of Hulten (1968), S c h o f i e l d (1969), H i t c h c o c k and C r o n q u i s t (1974) and Hale (1979). i . O v e rstory Assessment Six p l a n t formations were d e s c r i b e d on the b a s i s of h e i g h t and percent ground cover; Closed (non-giant) f o r e s t s are formed by t r e e s ranging from 5 to 50 m i n h e i g h t , with t h e i r crowns u s u a l l y i n t e r l o c k i n g . C l o s e d f o r e s t s are c h a r a c t e r i z e d by more than 50% ground cover. Open f o r e s t e d woodlands are formed by t r e e s at l e a s t 5 m t a l l , with most of t h e i r crowns not i n t e r l o c k i n g , comprising from 10 to 50% ground 22 cover. Scrub i s d e f i n e d i n terms of shrublands and t h i c k e t s , with p l a n t s ranging up to 4.6 m i n he i g h t . Shrublands range from 40 to 80% ground cover f o r deciduous types, and gr e a t e r than 50% fo r c o n i f e r o u s t y p e s . Deciduous scrub t h i c k e t s are c h a r a c t e r i z e d by ground cover exceeding 80%. Logging c l e a r c u t s are r e p r e s e n t a t i v e of on c e - f o r e s t e d , l o w - e l e v a t i o n h a b i t a t s now c l e a r e d of t h e i r commercially v a l u a b l e c o n i f e r o u s timber. Ground cover i s v a r i a b l e , depending upon the time s i n c e c l e a r i n g , but i s u s u a l l y g r e a t e r than 20%. P l a n t s are no higher than 5 m. T a l l - s e d g e Swamp i s f r e q u e n t l y f l o o d e d with fresh-water and commonly f o r long p e r i o d s . As a r u l e they are n a t u r a l by c r e a t i o n , however beavers (Castor canadensis) c r e a t e d the swamp i n t h i s study area. F o l i a g e i s f r e q u e n t l y t a l l e r than 30 cm with sedges dominating throughout. S c a r c e l y vegetated areas comprise i n t e r f l u v e s , v e r t i c a l rock f a c e s , and l i t t e r s l o p e s a l i k e , i n both the a l p i n e and s u b a l p i n e . These areas are g e n e r a l l y comprised of ground cover l e s s than or equal to 10%. P l a n t s are no higher than 0.9 m. Plan t formation c a t e g o r i e s were a d d i t i o n a l l y d e s c r i b e d on the b a s i s of growth form (e.g. 23 deciduous vs. c o n i f e r o u s ) , s p e c i e s composition, or both. Each formation may not represent e i t h e r a homogenous stand or a d i s t i n c t e n t i t y , but rat h e r one that may i n t e r g r a d e i n v a r i a b l e i n t e n s i t y with adjacent c a t e g o r i e s . P l a n t h e i g h t s were a s s i g n e d i n r e l a t i o n to the mean shoulder height of an a d u l t goat (x=0.92 m; Brandborg 1955). i i . Understory and S t r a t a C h a r a c t e r i s t i c s S e v e r a l understory c h a r a c t e r i s t i c s were s u b j e c t i v e l y d e s c r i b e d f o r the major p l a n t formations d u r i n g f i e l d c o n f i r m a t i o n of the p l a n t cover map. These i n c l u d e d herbaceous s t r a t a height and percent cover, scrub s t r a t a h e i g h t , and f o r e s t woody understory composition. Herbaceous c h a r a c t e r i s t i c s were d i v i d e d i n t o two types of cover (<50% or >50%), to d e s c r i b e each p l a n t formation i n both a l p i n e and subalpine environments. A l l s t r a t a were l e s s than 0.9 m, with the e x c e p t i o n of a s u b a l p i n e herbaceous meadow ( i . e . >50%) category, which sometimes exceeded 0.9 m. F o r e s t woody understory was d i v i d e d i n t o s i x types; c o n i f e r o u s , deciduous and mixed c o n i f e r o u s and deciduous t r e e (>5.0 m), and c o n i f e r o u s , deciduous and mixed c o n i f e r o u s and deciduous shrub (2.4 to 5.0 m). 24 Two krummholz s t r a t a were d e s c r i b e d ; a low stratum (upper a l t i t u d i n a l l i m i t of t r e e l i n e ) was d e s c r i b e d as shrubs l e s s than 1.2 m i n h e i g h t , and the high stratum (lower a l t i t u d i n a l l i m i t of t r e e l i n e ) c o n s i s t e d of t r e e s ranging from 1.2 to 4.6 m. Exposed deciduous shrublands and t h i c k e t s were a l s o d i v i d e d i n t o two s t r a t a ; a hi g h stratum (>1.0 m), and a low stratum (<1.0 m). 4.1.2 A b i o t i c F a c t o r s a. T e r r a i n C l a s s i f i c a t i o n i . Surface E x p r e s s i o n Surface e x p r e s s i o n d e s c r i b e s s u r f a c e forms that are not adequately shown ( p r i n c i p a l l y by contours) on the 1:50,000 topographic base map (Resource A n a l y s i s Branch 1978). E i g h t c a t e g o r i e s d e s c r i b e s u r f a c e e x p r e s s i o n ; rock outcrop, t a l u s , hummocky, s h e l f , r i d g e , peak, g u l l y or r a v i n e , and f a n . Slope p o s i t i o n (e.g. m i d - e l e v a t i o n , f o r e s t e d s l o p e , e t c . ) was not d e s c r i b e d . i i . Rock Substrate T h i s v a r i a b l e d e s c r i b e s i n o r g a n i c s u b s t r a t e t e x t u r e s on which mountain goats were observed (organic mats and snow-covered slo p e s were excluded). Rock s u b s t r a t e types were d e s c r i b e d in four c a t e g o r i e s ; broken c o n s o l i d a t e d (rugged 25 or crumbly) rock, smooth rock, s h a l e , and g r a v e l -s i l t s t o n e . Aspect Aspect was d i v i d e d i n t o e i g h t c l a s s e s r e l a t e d to exposure; n o r t h , n o r t h e a s t , e a s t , southeast, south, southwest, west and northwest. Aspects were determined by 45° a l l o t m e n t s of a compass p o i n t , s t a r t i n g at 337.5°. A n i n t h c l a s s ('no aspect') c o n s t i t u t e d the tops of r i d g e s and peaks and not merely f l a t areas (such as the 'Tea-Gardens' (see F i g u r e 2) north of the main r i d g e ) . Aspect at a p o i n t on a g r i d was recorded as the average aspect of the lan d l y i n g w i t h i n a 100 m r a d i u s from the p o i n t , r a t h e r than the immediate aspect of that p o i n t . Slope Slope i s d e s c r i b e d i n degrees ( r a t h e r than p e r c e n t ) , and was d i v i d e d i n t o eleven i n t e r v a l s of unequal s i z e r e l a t e d to steepness; <27°, 28-29°, 30-31°, 32-34°, 35-37°, 38-41°, 42-45°, 46-50°, 51-56°, 57-63°, and >63°. Slope was c a l c u l a t e d u s ing the t r i g o n o m e t r i c f u n c t i o n d e s c r i b e d by Thompson 1981, p 2 8 l ) . Map measurements were taken to the nearest 0.8 mm on an e n l a r g e d 1:50,000 topographic map. Each p o i n t of l o c a t i o n maintained the mid-point of the v e r t i c a l d i s t a n c e , which was h e l d constant at 91.4 m. 26 E l e v a t ion The 1928 m v e r t i c a l g r a d i e n t from r i d g e top to lake edge was assessed on two d i f f e r e n t o r d e r s . The f i r s t order of s e l e c t i o n u t i l i z e d t r e e l i n e as a geographic landmark. The upper l i m i t of t r e e growth (krummholz) was sampled every 500 m from the 1:50,000 base map. Two zones were subsequently d e f i n e d ; a l p i n e and s u b a l p i n e . The second order of s e l e c t i o n c l a s s i f i e d the a l t i t u d i n a l g r a d i e n t of the study area i n t o nine 250-m c l a s s e s ; <250 m, 251-500 m, 501-750 m, 751-1000 m, 1001-1250 m, 1251-1500 m, 1501-1750 m, 1751-2000 m, and >2000 m. E l e v a t i o n was c a l c u l a t e d to the nearest 15m, by superimposing the enla r g e d 1:50,000 (30.5 m i n t e r v a l ) contour map over the v e g e t a t i o n map with an ep i d i a s c o p e , using two government survey r e f e r e n c e p o i n t s i n the a l p i n e as c o n t r o l s f o r s c a l e . E l e v a t i o n of each c o n t r o l p o i n t was confirmed with a Thommen 2000 hand a l t i m e t e r and a l s o with the a l t i m e t e r i n a B e l l 206B 'Jet Ranger' h e l i c o p t e r . M i c r o c l i m a t e To assess annual microthermal v a r i a t i o n throughout the study area, s i x m e t e o r o l o g i c a l s t a t i o n s were e r e c t e d d u r i n g the summer of 1977 and monitored u n t i l the f o l l o w i n g summer: one was l o c a t e d i n the f o r e s t e d v a l l e y bottom, below goat ranges, at 725 m (stn #6); two were i n s t a l l e d on the lower l i m i t of 27 goat range - one i n a deciduous shrub r a v i n e at 879 m ( s t n #1); and the other i n a c o n i f e r o u s f o r e s t a t 893 m (stn #2); another was p l a c e d i n the krummholz, near t r e e l i n e , at 1433 m (stn #3); one sat on top of a g l a c i a l rock i n t e r f l u v e on the exposed, knife-edged, main r i d g e at 1692 m ( s t n #4); and the l a s t one was l o c a t e d on a weathered r i d g e i n the hummocky a l p i n e tundra at 1591 m (stn #5). A seventh weather s t a t i o n , monitored by the p r o v i n c i a l government, was l o c a t e d i n the extreme northwestern corner of the study area (Rosswood, B.C.) at an e l e v a t i o n of 152 m (see F i g u r e 2). Each r e c o r d i n g u n i t was i n s t a l l e d and maintained a c c o r d i n g to s p e c i f i c a t i o n s of the p r o v i n c i a l government Resource A n a l y s i s Branch (Climatology S e c t i o n 1974). A seven-day thermograph was used i n the v a l l e y - b o t t o m s t a t i o n (#6) and t h i r t y - d a y hygrothermographs were l o c a t e d at each of the remaining f i v e s i t e s . Each were i n s t a l l e d i n a Stevenson screen, s i t u a t e d 1.2 m above the ground on v e r t i c a l l y a d j u s t a b l e 3 m stands designed by the author. Two standard (15.2 cm diameter) p r e c i p i t a t i o n b a r r e l s , 'topped o f f with kerosene (to prevent e v a p o r a t i o n ) , and a t o t a l i z i n g annemometer accompanied each s t a t i o n . Square (5 cm) wire mesh was wrapped around each Stevenson screen to h e l p minimize damage c r e a t e d by porcupines ( E r e t h r i z o n dorsatum). 28 4.2 H a b i t a t S e l e c t i o n H a b i t a t components were examined only as s i n g l e independent v a r i a b l e s with respect to mountain goat u t i l i z a t i o n . A n a l y s i s of h a b i t a t s e l e c t i o n i s based upon use and a v a i l a b i l i t y of b i o t i c and a b i o t i c v a r i a b l e s . 4.2.1 Data C o n s i d e r a t i o n s S e v e r a l aspects of the s e l e c t i o n analyses were f i r s t c o n s i d e r e d . a. C a l c u l a t i o n of H a b i t a t A v a i l a b i l i t y A v a i l a b i l i t y was c o n s i d e r e d to i n c l u d e a g e n e r a l i z e d r e g i o n based upon the cumulative d i s t r i b u t i o n of mountain goats which were p l o t t e d on the base map. The e n t i r e h i l l s i d e was c o n s i d e r e d to be ' a v a i l a b l e ' to a goat ( i . e . from v a l l e y bottom creek to r i d g e - t o p ) . L a t e r a l boundaries c o n s i d e r e d i n c l u d e d two of the four major geographic b a r r i e r s d i s c u s s e d e a r l i e r as study area boundaries. A v a i l a b i l i t y of a h a b i t a t v a r i a b l e was measured in one of two ways; by d i r e c t measurement of area, or by random sampling f o r e s t i m a t i o n of p r o p o r t i o n s , i . D i r e c t A r e a l Measurement A r e a l estimates of a v a i l a b i l i t y were measured f o r p l a n t cover types and a l t i t u d i n a l zones by v e r t i c a l p r o j e c t i o n , using a dot g r i d (64 dots/km 2). C o r r e c t i o n of area f o r h i l l s i d e slope was determined by randomly sampling each 29 p l a n t and e l e v a t i o n category and a p p l y i n g the a p p r o p r i a t e c o r r e c t i o n f a c t o r , as d e s c r i b e d by Thompson (1981, p 2 8 l ) . Random sample s i z e s were determined using the bi n o m i a l p r o b a b i l i t y f u n c t i o n d e s c r i b e d i n Mendenhall (1971, p l 9 8 ) . The most c o n s e r v a t i v e values of 'p' and 'q' were used ( i . e . 0.5) because no estimate of p r o p o r t i o n s was a v a i l a b l e . A 90 percent l e v e l of confi d e n c e and 10 percent accuracy were s e l e c t e d f o r the de t e r m i n a t i o n of sample s i z e . Random sampling was i n some cases terminated prematurely i f i t e r a t i o n showed that the d e s i r e d l e v e l s of confi d e n c e and accuracy had been achieved, i i . E s t i m a t i o n of P r o p o r t i o n s by Random Sampling A 'non-mapping' technique (Marcum and Loftsgaarden 1980) was used f o r e s t i m a t i n g the p r o p o r t i o n s of slope and aspect c a t e g o r i e s . Estimates of h a b i t a t p r o p o r t i o n s were obtained by superimposing the goat l o c a t i o n g r i d system over an enlarged topographic map of Maroon Mountain and c l a s s i f y i n g a set of randomly d i s t r i b u t e d p o i n t s over the area of a v a i l a b i l i t y . Random c o o r d i n a t e s were generated u s i n g a computer program ( N i c o l 1981). The procedure f o r determining random sample s i z e f o r e s t i m a t i n g p r o p o r t i o n s f o l l o w e d that p r e v i o u s l y d e s c r i b e d f o r d e t e r m i n a t i o n of sample s i z e s f o r mean slope 3 0 (Mendenhall 1971). The mean slope of these values was then used to a d j u s t the area of a v a i l a b i l i t y w i t h i n each h a b i t a t category. Once c o r r e c t e d f o r h i l l s i d e s l o p e , p r o p o r t i o n s of a v a i l a b i l i t y were then r e c o n s t r u c t e d to equal the number of mountain goat o b s e r v a t i o n s f o r the time p e r i o d used i n the a n a l y s i s . T h i s d e s i g n , u t i l i z e d f i x e d marginal contingency t o t a l s f o r one c r i t e r i a ( i . e . h a b i t a t p r o p o r t i o n s ) and i s termed a 'Model I I ' contingency design (Sokal and Rohlf 1980, p735). The design was intended, i n p a r t , to a l l e v i a t e d i f f e r i n g e f f e c t s of frequency on the d i s t r i b u t i o n of the data. I t a l s o f a c i l i t a t e s computation of the t e s t s t a t i s t i c s i n v o l v e d . A v a i l a b i l i t y of s u r f a c e e x p r e s s i o n , s u b s t r a t e t e x t u r e , thermal s t r a t a and other c l i m a t o l o g i c a l data c o u l d not be a c c u r a t e l y assessed from maps or a e r i a l photographs and were t h e r e f o r e excluded from the a n a l y s i s of s e l e c t i o n . b. Other C o n s i d e r a t i o n s P r i o r to the a n a l y s i s of h a b i t a t s e l e c t i o n , each group of mountain goats was randomly subsampled f o r one ' r e p r e s e n t a t i v e ' i n d i v i d u a l , i n order to e l i m i n a t e b i a s e s i n v o l v i n g dependency of the data due to s o c i a l c o n s t r a i n t s . S e l e c t i o n of environmental a t t r i b u t e s 31 are b e l i e v e d to be i n f l u e n c e d by other c o n s p e c i f i c s (e.g. a nanny leads her k i d , the k i d does not choose where to be). Without subsampling, such an a n a l y s i s may be taken to i n d i c a t e h a b i t a t s e l e c t i o n by groups ra t h e r than by i n d i v i d u a l s , as presented i n the former s e c t i o n on H a b i t a t Use. A n a l y s i s using group data v i o l a t e s the requirement of independence i n s t a t i s t i c a l procedures. A d d i t i o n a l l y , two group types were d e f i n e d r a t h e r than a n a l y z i n g by sex a l o n e . Male groups c o n s i s t e d of male goats only, at any age. Mixed-female groups c o n s i s t e d of any composition having at l e a s t one female i n i t . Males o c c u r r e d i n both groups d e f i n e d , however, because they are subordinate to females, they were c o n s i d e r e d ' f o l l o w e r s ' i n the sense that i t i s the dominant animal that s e l e c t s i t s own environment (Chadwick 1973). A t h i r d group type may a l s o have c o n t a i n e d males and females ( ' u n c l a s s i f i e d group") but t h i s category was not used i n the a n a l y s i s . Two l i m i t a t i o n s were p l a c e d on d e f i n i n g a group; v i s u a l c o n t a c t , and p r o x i m i t y to c o n s p e c i f i c s . For example, two mountain goats may be v i s i b l e to each other when s e v e r a l hundred meters a p a r t , or they may be only 20 m apart but on the other s i d e of a r a v i n e , or v e g e t a t i o n s t r i p , or at extreme v e r t i c a l l i m i t s of a b l u f f ( i . e . on top and bottom), such that they are out of s i g h t of each o t h e r . N e i t h e r c o n d i t i o n 32 c o n s t i t u t e d a group. In the former case, the goats were aware of each other, and i n the l a t t e r case, they were not. The former s i t u a t i o n was r e g u l a t e d by a s s i g n i n g d i s t a n c e l i m i t a t i o n s , and f o r t h i s study, that boundary was a r b i t r a r i l y set at 100 m. V i s u a l c o n t a c t was the cue used to determine awareness between c o n s p e c i f i e s , and was evidenced by the o v e r t b e h a v i o u r a l responses of the goats (e.g. l o o k i n g up while g r a z i n g or t u r n i n g i t s head to s t a r e ) . Consequently, e s t i m a t i o n of group s i z e may not always represent p o i n t - i n - t i m e or 'scan' sampling (see Altmanri 1974). O c c a s i o n a l l y i n d i v i d u a l goats had to be watched for s e v e r a l minutes i n order to determine i f synchrony i n a c t i v i t y and d i r e c t i o n of movement appeared independent of c o n s p e c i f i e s . c. S t a t i s t i c a l A n a l y s i s Use and a v a i l a b i l i t y data were analyzed using a two-step technique; m u l t i n o m i a l contingency a n a l y s i s , using the l o g l i k e l i h o o d r a t i o (G) g o o d n e s s - o f - f i t t e s t (at the 0.05 l e v e l of s i g n i f i c a n c e ) , and subsequent a n a l y s i s of 90 percent simultaneous con f i d e n c e i n t e r v a l s (SCI), using the c o n s e r v a t i v e B o n f e r r o n i approach (Neu et a l . 1974, Marcum and Lofstgaarden 1980). The 90 percent SCI's were ( a r b i t r a r i l y ) chosen as a reasonable l e v e l of s i g n i f i c a n c e . By m a i n t a i n i n g a=0.l0, the standard normal d e v i a t e (Z) changes f o r each category ( k ) . As 33 a r e s u l t , i n d i v i d u a l c o n f i d e n c e i n t e r v a l s v a r i e d from 0.95 to 0.99 when 'k' e q u a l l e d 2 and 10, r e s p e c t i v e l y . 4.2.2 B i o t i c F a c t o r s a. V e g e t a t i v e Cover A l l s i x p l a n t formations o u t l i n e d i n the p r e v i o u s s e c t i o n on H a b i t a t Use were used i n the p r e f e r e n c e a n a l y s i s . Understory c h a r a c t e r i s t i c s were not assessed due to t h e i r small s c a l e . 4.2.3 A b i o t i c F a c t o r s a. Aspect S i x aspect i n t e r v a l s occur on the southern s l o p e s of Maroon Mountain; e a s t , southeast, south, southwest, and west. The r i d g e and main peaks ( i . e . 'no aspect') i n t e r v a l was not i n c l u d e d i n t h i s assessment because random sampling from a map was not p r e c i s e enough to d e l i n e a t e knife-edge r i d g e s or peak tops on Maroon Mountain. North, northeast and northwest aspects d i d not occur i n the area of a v a i l a b i l i t y , however, other micro-aspect s i t u a t i o n s probably e x i s t e d but were not recorded due to the 100 m r a d i u s r e s t r i c t i o n imposed d u r i n g data c o l l e c t i o n (see H a b i t a t Use). b. Slope Ten slope i n t e r v a l s occur on the southern s l o p e s of Maroon Mountain; <27°, 28-29°, 30-31°, 32-34°, 35-37°, 38-41°, 42-45°, 46-50°, 51-56°, and 57-63°. 34 However, random sampling d e s c r i b e d only the f i r s t seven i n t e r v a l s (0-45°), consequently the e i g h t h , n i n t h and tenth i n t e r v a l s (used by goats) and the 'eleventh' i n t e r v a l (>63°), unused by goats, were lumped together t o c r e a t e a ">46°' c l a s s f o r t h i s a n a l y s i s , c. E l e v a t i o n Although e i g h t a l t i t u d i n a l i n t e r v a l s (from 250 to 2250 m) were used by mountain goats, random sampling of a v a i l a b l e e l e v a t i o n bands documented one a d d i t i o n a l i n t e r v a l (<250 m). T h e r e f o r e , nine e l e v a t i o n i n t e r v a l s were used i n the pr e f e r e n c e a n a l y s i s . Because no random samples were l o c a t e d i n the upper or lowermost c a t e g o r i e s when sampling f o r mean slope c o r r e c t i o n s on the map (due to t h e i r small a r e a ) , the slope c o r r e c t i o n f a c t o r f o r the adjacent zone was used as a r e a l i s t i c c a l i b r a t i o n of these areas. 4.2.4 Mountain Goat A c t i v i t y Within P r e f e r r e d H a b i t a t s H a b i t a t types were c o n s i d e r e d to be p r e f e r r e d when an animal s e l e c t e d c e r t a i n d e s i r a b l e c h a r a c t e r i s t i c s w i t h i n that environment. T h i s a n a l y s i s i n v e s t i g a t e d the b i o l o g i c a l s i g n i f i c a n c e • of p r e f e r r e d h a b i t a t s by assessment of three a c t i v i t y p a t t e r n s ; f o r a g i n g , r e s t i n g , and t r a v e l . Foraging i n c l u d e d browsing and g r a z i n g p r a c t i c e s . Standing and ruminating animals were c o n s i d e r e d r e s t i n g . T r a v e l c o n s i s t e d p r i m a r i l y of animals walking or running. Animals at pl a y were recorded as e i t h e r walking or running. 35 IV. RESULTS 1. POPULATION CHARACTERISTICS AND OBSERVABILITY 1.1 D i s t r i b u t i o n And Movements Maroon Mountain appears to be a geographic ' i s l a n d ' with l i t t l e or no movement to and from the study area a p p a r e n t l y o c c u r r i n g . 1.1.1 D i s t r i b u t i o n a. Range Use Systematic v e h i c u l a r surveys covered approximately 73% of the study a r e a . These data showed that i n t e n s i t y of mountain goat use n o r t h and south of the c r e s t of the main r i d g e ( F i g u r e 3) was s i g n i f i c a n t l y d i f f e r e n t (G=1606.6;df=1). Slopes south of the main r i d g e comprised 29.4% (22.03 km2) of the study area, but were used by 99.5% of a l l groups recorded. Mountain goat o b s e r v a t i o n s ranged from r i d g e tops down to the v a l l e y - b o t t o m l o g g i n g roads. A 9 and 1/2 year o l d male was found dead i n one of the l o g g i n g c l e a r c u t s along Maroon Creek i n February, 1978. H e l i c o p t e r and backpacking surveys over the remaining 27% of the study area i n d i c a t e d that s l o p e s northeast of the main r i d g e were only o c c a s i o n a l l y u t i l i z e d d u r i n g l a t e summer, and never i n w i n t e r . Unless i n immediate p r o x i m i t y to escape t e r r a i n p r o vided by the main r i d g e , or H a l l Creek, mountain ON F i g u r e 3 - D i s t r i b u t i o n o f s y s t e m a t i c a l l y o b s e r v e d m o u n t a i n g o a t s on M a r o o n M o u n t a i n , B.C., d e t e r m i n e d by v e h i c u l a r s u r v e y . 3 7 goats were observed to u t i l i z e h i l l s i d e s n orth of the main r i d g e f o r p e r i o d s of a few hours at the most. T r a v e l i n these areas was g e n e r a l l y at a f a s t walk, due to the g e n t l e - s l o p i n g nature of the a r e a . Seasonal ranges overlapped with v a r y i n g i n t e n s i t y . The l e a s t o v e r l a p i n monthly ranges had a c o e f f i c i e n t of a r e a l a s s o c i a t i o n of 0.18 (see T a y l o r 1977, p177), between areas used i n August and November, 1977. Monthly range s i z e s were s m a l l e s t d u r i n g p e r i o d s of hot summer temperatures ( J u l y , 1977 and 1978) and l a r g e s t d u r i n g s p r i n g thaw and the a s s o c i a t e d 'green-up' ( A p r i l to June). Non-systematic surveys d u r i n g the summer of 1976 showed range s i z e to be much l a r g e r , i n the absence of p r e d a t o r s . b. S p a t i a l D i s p e r s i o n Mountain goats were determined to be non-randomly d i s t r i b u t e d over the s l o p e s south of the main r i d g e , t h e r e f o r e r e j e c t i n g the N u l l Hypothesis (G=32.3;df=128). Each a n a l y s i s by month a l s o r e j e c t e d the N u l l Hypothesis, however, mountain goat d i s p e r s i o n p a t t e r n s d u r i n g the months of May and November, 1977, d i d not s t a t i s t i c a l l y f i t the 'general' negative binomial model used i n the a n a l y s i s (Table 2). Mountain goats tended to be most aggregated i n summer (June through August), when food r e s o u r c e s were more abundant. However, two months in p a r t i c u l a r (August and December, 1977) d i s p l a y e d s i g n i f i c a n t l y higher 38 Table 2 - Monthly d i s t r i b u t i o n s t a t i s t i c s of mountain goats inhabiting Maroon Mountain, B.C. Date Range Use Number o f Goats p e r Q u a d r a t Goodness-of - f i t t o N e g a t i v e B i n o m i a l (km 2) Mean Sample ,—, V a r i a n c e (x) , (s^) Index o f D i s p e r s i o n ( I ) a Maximum L i k e l i h o o d E s t i m a t e ( k ) b L ° 9 C h i -L i k e l i h o o d S q u a r e <j.f . E s t i m a t e ( 0 ) c 1977 May S. 4 1. 38 7. 14 5. 17 0. 183 7. 558 23. .248* 7 Jun 10. 8 2. 05 18. .80 9. 17 0. 178 11. 545 4 . 124 9 J u l 4 . 1 1. 03 8. .42 8. 17 0. 080 12. ,860 4. ,851 5 Aug 8. 3 1. 97 23. ,98 12. 17 0. 125 15. .799 13. , 888 8 Sep 8. 6 1. 76 15. . 36 8. .73 0. 250 7. , 039 11. ,832 8 Oct 5. 5 1. 06 5. .06 4. .80 0. 171 6. . 199 4. . 879 6 Nov 8. .1 1. 59 8. .42 5. . 30 0. .187 8. . 515 32. .173* 8 Dec 5. , 4 1. 03 17. .05 16. , 55 0. .066 15. .587 0, .609 4 1970 Ja n 6. , 6 1. ,17 6, . 36 5. .44 0. , 209 5. ,608 8. .410 7 Feb 13. .2 1. .78 9 , . 80 5, .51 0. ,299 5. .931 7. .494 9 Mar 12. . 3 1. . 74 7 , . 46 4. . 29 0. . 370 4 , .714 9. . 241 8 Apr 14, . 5 1. .46 4, .91 3, .36 0. .520 2, .803 4, .176 7 May 9. , 6 1. . 73 11, . 54 6 .67 0. . 248 6. .957 9, .487 8 Jun 9, . 2 1. .48 9 .99 6, .75 0, . 171 8, . 634 13, . 179 7 J u l 2 . 6 0, .75 4 . 88 6, . 51 0, .075 10. . 041 5 . 275 4 Aug 7. . 8 1, . 02 10 . 80 10 . 59 0, . 119 8 . 526 6 .040 6 a 2 , — I = s / x b k (c l u m p i n g v a r i a b l e ) i s c a l c u l a t e d by t h e Newton-Raphson i t e r a t i o n p r o c e d u r e ( B i s h o p , F i e n b e r g and H o l l a n d 1975, p84) c 6 = x / k * s i g n i f i c a n t a t p<0.05 39 l e v e l s of mountain goat a g g r e g a t i o n than a l l o t h e r r e m a i n i n g months (T a b l e 2 ) . August, 1978 showed a s i m i l a r t r e n d , but was not s i g n i f i c a n t . Low l e v e l s of a g g r e g a t i o n were noted throughout the w i n t e r ( J a n u a r y t o A p r i l ) , most n o t a b l y i n the month of A p r i l , a t the onset of s p r i n g 'green-up'. There was no apparent r e l a t i o n s h i p between d i s p e r s i o n p a t t e r n and range s i z e (Spearman's t=-3.7;df=14), however, a p o s i t i v e c o r r e l a t i o n o c c u r r e d between clumpedness and group s i z e (t=3.4;df=14). O v e r a l l , k i d d i n g (and accompanying p o s t - p a r t u m i s o l a t i o n ) a r e a s were a l s o d e t e r m i n e d t o be c o n t a g i o u s l y d i s t r i b u t e d (d=7.3). In 1976, however, neonate l o c a t i o n s were randomly d i s t r i b u t e d (d=0.6). The e x i s t e n c e of a n n u a l ' k i d d i n g grounds' was d i s m i s s e d on the b a s i s of a l a c k i n g e o g r a p h i c a l c o n c e n t r a t i o n ( F i g u r e 4 ) . C e r t a i n e n v i r o n m e n t a l a t t r i b u t e s may be common f o r each s i t e , however t h i s was not examined. S i m i l a r l y , d u r i n g the r u t (November and e a r l y December), b o t h t e n d i n g a d u l t males and a d u l t females were non-randomly d i s t r i b u t e d (males: d=2.8; f e m a l e s : d=17.2). Both d i s t r i b u t i o n s a r e b e l i e v e d t o be as much a r e s u l t of a r e a l c o n c e n t r a t i o n due t o d e s c e n d i n g s n o w l i n e s and i n c r e a s i n g snowpack a t t h i s time of y e a r , as they a r e of s o c i a l f a c t o r s . The e x i s t e n c e of a n n u a l ' r u t t i n g grounds' was d i s m i s s e d on the b a s i s of F i g u r e 4 - D i s t r i b u t i o n o f n e o n a t e l o c a t i o n s i n t h e s t u d y a r e a - 1976 t o 1978. 41 a lac k i n g e o g r a p h i c a l c o n c e n t r a t i o n ( F i g u r e 5 ). 1.1.2 Movements Only one mountain goat, a 3.5 y e a r - o l d female, was captured and permanently marked (August, 1977). 'Female#2' was r e s i g h t e d 32 times over a 12 month p e r i o d ( F i g u r e 6). D e s p i t e numerous attempts, f r e e - r a n g i n g i m m o b i l i z a t i o n of more mountain goats was found to be l o g i s t i c a l l y too d i f f i c u l t and time consuming. However, permanent i d e n t i f i c a t i o n was p o s s i b l e f o r seven other mountain goats w i t h i n the p o p u l a t i o n . Six of these were i d e n t i f i a b l e by unique (damaged) horn c h a r a c t e r i s t i c s and one o l d nanny had a scar across her rostrum. Each i d e n t i f i a b l e goat was always r e s i g h t e d south of the main r i d g e of Maroon Mountain ( F i g u r e s 6 and 7), c o n f i r m i n g the importance of t h i s mountainside i n r e l a t i o n to the e n t i r e study area. Three of these animals ('female#2', 'unicorn female' and ' l e f t horn-stub female') d i s p l a y e d pronounced l a t e r a l s h i f t s i n range use of a t l e a s t 5 to 7 km d u r i n g l a t e June and e a r l y J u l y . No obvious movements of t h i s nature were otherwise recognized, even though these same goats were r e p e a t e d l y s i g h t e d i n the more open a l p i n e zone ( u s u a l l y along or near the knife-edge r i d g e t o p ) . No m i n e r a l l i c k was d i s c o v e r e d , which c o u l d p o s s i b l y have e x p l a i n e d t h i s movement by females. Figure 6 - Observed movements of "female#2" and "bar-nosed female" on Maroon Mountain, B .C. (numbers indicate sequence of s ight ings). F i g u r e 7 - O b s e r v e d movements of " u n i c o r n f e m a l e " and " l e f t h o r n - s t u b f e m a l e " on Maroon M o u n t a i n , B.C. (numbers i n d i c a t e s e q u e n c e of s i g h t i n g s ) . 45 1.2 Po p u l a t i o n S t r u c t u r e 1.2.1 P o p u l a t i o n S i z e And D e n s i t y The minimum Maroon Mountain goat p o p u l a t i o n s i z e was p r e d i c t e d to be 137 mountain goats, based upon the maximum count, which was obtained d u r i n g a h e l i c o p t e r survey. McCrory and Blood (1978) estimated that h e l i c o p t e r s miss 26 to 30% of the a c t u a l p o p u l a t i o n d u r i n g a e r i a l surveys, t h e r e f o r e the Maroon p o p u l a t i o n may range as high as 185 to 196 mountain goats. Based upon the minimum known p o p u l a t i o n s i z e of 137 animals, the mountain goat d e n s i t y was 9.59 goats/km 2, over the area of observed d i s t r i b u t i o n . The p r e d i c t e d Maroon Mountain p o p u l a t i o n s i z e (n=l85 to 196 goats) approximated a d e n s i t y of between 12.96 and 13.73 animals/km 2. A e r i a l d e t e r m i n a t i o n s were s i g n i f i c a n t l y d i f f e r e n t when c o r r e c t e d f o r h i l l s i d e slope (x slope=34°'correct ion factor=1.20). T h e r e f o r e , d e n s i t y v a l u e s were reduced to 10.79 and 11.40 mountain goats/km 2. 1.2.2 P o p u l a t i o n Composition Neonates were f i r s t observed on May 26, 1976, May 24, 1977 and May 19, 1978. Sex-age r a t i o s were not assessed f o r the 1976 data because a d u l t female c l a s s i f i c a t i o n was b i a s e d towards 'goats with k i d s ' and because males c o u l d not be d i s t i n g u i s h e d from females with a hig h degree of c e r t a i n t y . The maximum kid/100 a d u l t female r a t i o was 73.7, i n 1977. The maximum number of y e a r l i n g / 1 0 0 a d u l t females was 52.7, i n 1978. These data suggest a f i r s t - y e a r m o r t a l i t y r a t e of 28.5%. The maximum 46 a d u l t sex r a t i o was 52.6 males/100 a d u l t females. A c t u a l p r o p o r t i o n s of the Maroon Mountain p o p u l a t i o n composed of males and females were c a l c u l a t e d from the maximum survey count, r e g a r d l e s s of age, a p p o r t i o n i n g the ' u n c l a s s i f i e d ' segment by the c l a s s i f i e d sex r a t i o , and adding the two f i g u r e s f o r each sex t o g e t h e r . In t h i s manner, i t was estimated that the maximum number of males and females i n the Maroon Mountain p o p u l a t i o n were 47 and 90, r e s p e c t i v e l y . Most of the males were c l a s s i f i e d as a d u l t s (75.6%). Subadults comprised 3.9%, y e a r l i n g s 13.4% and k i d s 0.2% of the c l a s s i f i e d males. S i m i l a r l y , most females were a l s o c l a s s i f i e d as a d u l t s (86.1%). Subadults comprised 3.9%, y e a r l i n g s 9.7% and k i d s 0.3% of the c l a s s i f i e d females. Kids were the most d i f f i c u l t to sex and, t h e r e f o r e , r e t a i n e d the highest- p r o p o r t i o n of u n c l a s s i f i e d animals. 1.3 Mountain Goat O b s e r v a b i l i t y V a r i a t i o n s i n mountain goat o b s e r v a b i l i t y were examined by census r e s u l t s , comparison of survey technique and mountain goat sex. 1.3.1 Census R e s u l t s Censuses i n c l u d e d a 6 km2 area on Mt. Garland south of Maroon Mountain, i n which on average 8 percent of censused goats were seen. Although not used elsewhere, they have been i n c l u d e d i n the f o l l o w i n g d i s c u s s i o n of census techniques to maintain complete survey data. 47 A e r i a l Counts Four h e l i c o p t e r surveys were conducted i n a 'Jet Ranger' d u r i n g May, August, December, 1976 and J u l y , 1977. T o t a l counts ranged from 70 to 149 mountain goats ( F i g u r e 8), producing a mean count of 118.8 animals (SD=35.9). Snow-free months (May to August) p r o v i d e d the highest counts (114 to 142), with the lowest v a r i a t i o n (SD=18.5). However mountain goat o b s e r v a b i l i t y by r e p l i c a t i o n with h e l i c o p t e r was not constant (G=35.1;df=3). Two f i x e d wing a i r c r a f t surveys were conducted ( A p r i l , 1976, i n a de H a v i l l a n d 'Beaver'; August, 1977 i n a Cessna 180), d u r i n g which only 14 mountain goats and one lone i n d i v i d u a l were observed, r e s p e c t i v e l y . C o r r e c t i n g these counts by m u l t i p l y i n g with a f a c t o r of 5.0, to cover the v e r t i c a l g r a d i e n t of the study area, r e s u l t e d i n counts of 70 and 5 animals, or a mean of 37.5 mountain goats, which i s f a r l e s s than the p r e v i o u s a e r i a l survey technique and much more v a r i a b l e (SD=46.0 g o a t s ) . Mountain goat o b s e r v a b i l i t y between the two fixed-wing surveys was a l s o s i g n i f i c a n t l y d i f f e r e n t (G=67.2;df=1). V e h i c u l a r Counts Many v e h i c u l a r surveys were not completed due to changing environmental v a r i a b l e s d u r i n g the course of each survey, which averaged between 6 and 12 hours i n d u r a t i o n . However, a t o t a l of 63 completed road F i g u r e 8 - Summary of m o u n t a i n g o a t c o u n t s on Maroon M o u n t a i n and Mount G a r l a n d , B.C. ( h o r i z o n t a l b a r - m e a n ; v e r t i c a l b a r = s t a n d a r d d e v i a t i o n ) . 49 surveys were accomplished over 24 months of o b s e r v a t i o n between A p r i l , 1976 and August, 1978 (F i g u r e 8). The average number of goats observed by road survey was 63.14 (SD=27.4). The p r o p o r t i o n of the t o t a l (estimated) p o p u l a t i o n was not constant throughout the year (G=812.1;df=62), ranging from 12.6% to 85.4% of the t o t a l p o s s i b l e number observable. As expected, maximum v e h i c u l a r counts g e n e r a l l y o c c u r r e d d u r i n g the p e r i o d of mountain goat p a r t u r i t i o n . Late f a l l - e a r l y winter and summer p e r i o d s g e n e r a l l y p r o v i d e d the lowest counts. During the f i r s t two y e a r s , 59.7 and 57.7 goats were seen on average, but i n 1978, a mean of 74.1 goats were seen. Mean counts d i d not d i f f e r s i g n i f i c a n t l y between years ( K r u s k a l - W a l l i s 1 Hc=8.1;df=2). c. Comparison of Surveys Fixed-wing surveys were inadequate f o r c o n c l u s i v e comparisons. H e l i c o p t e r surveys were s t a t i s t i c a l l y d i f f e r e n t from ground surveys. A p a i r - w i s e comparison of h e l i c o p t e r and follow-up v e h i c u l a r counts i n d i c a t e s a s i g n i f i c a n t d i s p a r i t y between the two techniques (G=49.8;df=3). H e l i c o p t e r censuses p r o v i d e d 1.9 times g r e a t e r accuracy than ground censuses, on average. The l e v e l of p r e c i s i o n f o r each of the two methods appeared s i m i l a r ( a i r SD=20.7; ground SD=18.6), when each of the four h e l i c o p t e r surveys were compared to 50 t h e i r r e s p e c t i v e follow-up v e h i c u l a r surveys, however, h e l i c o p t e r surveys were c o n s i d e r e d more p r e c i s e because of t h e i r p r o p o r t i o n a t e l y reduced v a r i a t i o n i n r e l a t i o n to t o t a l numbers. 1.3.2 O b s e r v a b i l i t y Of Sexes The average number of males and females were not e q u a l l y observable throughout the year (Wilcoxon's T=0.0;n=16), nor were they c o n s i s t e n t l y seen throughout the sampling p e r i o d (males: G=405.5,df=37; females: G=345.7,df=37). R e p r e s e n t a t i v e p r o p o r t i o n s ( i . e . >50%) of the male segment of the p o p u l a t i o n were only observable d u r i n g 6 of the 16 months sampled ( F i g u r e 9). They were seen l e a s t d u r i n g the p e r i o d of mountain goat p a r t u r i t i o n ( l a t e May-early June), and on i n t o the summer ( J u l y -September). Males were most f r e q u e n t l y observed d u r i n g l a t e winter (February and March) and e a r l y s p r i n g 'green-up' ( A p r i l and May), and s e c o n d a r i l y d u r i n g the rut (November). Females were c o n s i d e r a b l y more v i s i b l e than males, being represented i n 13 of the 16 months sampled ( F i g u r e 9 ) . S i m i l a r to males, they were l e a s t v i s i b l e d u r i n g p a r t s of the summer, and a l s o i n l a t e f a l l , d u r i n g and a f t e r the r u t t i n g p e r i o d . 51 NUMBER OF SURVEYS 1 3 3 4 3 2 3 1 2 2 2 2 2 3 2 3 100 (0 Ul < 5 cc UJ m 5 => z z < Ul s 30 maximum est imated number (n=47) 1 ± (50%) M J J A S 0977) O N D J F M MONTH A M C1978) J J A 100 CO Ul _ l < 2 u. O (0 2 Z Z 4 Ul 2 (50%) MONTH F i g u r e 9 - O b s e r v a b i l i t y of male and female components of the Maroon Mountain goat p o p u l a t i o n ( h o r i z o n t a l bar=mean; v e r t i c a l bar=standard d e v i a t i o n ) . 52 2. HABITAT CHARACTERISTICS 2.1 H a b i t a t Use 2.1.1 B i o t i c F a c t o r s a. V e g e t a t i v e Cover D e s c r i p t i o n Six p l a n t formations, comprising f i f t e e n c a t e g o r i e s of v e g e t a t i v e cover, were d e s c r i b e d on the south aspect of Maroon Mountain and are d e p i c t e d i n F i g u r e 10. These i n c l u d e : C l o s e d (non-giant) f o r e s t s . F i v e c a t e g o r i e s are d e s c r i b e d w i t h i n the t e x t , three of which are c o n i f e r o u s ; (1) The Tsuga-Abies (Hemlock-Fir) f o r e s t i s the major cover type on n o r t h , e a s t , and w e s t - f a c i n g s l o p e s , from as low as 150 m above sea l e v e l , on the shores of Kitsumkalum Lake, to the 1252 m (mean) t r e e l i n e region of Maroon Mountain. Tsuga s p e c i e s occur from v a l l e y bottom to t r e e l i n e , whereas Abies g r a n d i s and moreso ' A_^  l a s i o c a r p a become i n c r e a s i n g l y abundant above 610 m. The understory c o n s i s t s p r i m a r i l y of cryptogams such as Hylocomium splendens, Pleurozium  s c h r e b e r i , R h y t i d i a d e l p h u s l o r e u s , and R h y t i d i o p s i s  robusta, forming a dense mat. M e n z i e s i a f e r r u g i n e a and s e v e r a l Vaccinium s p e c i e s occur i n the upper subalpine f o r e s t understory, where the crown cover i s s l i g h t l y l e s s dense. (2) The P i c e a - A b i e s ( S p r u c e - F i r ) f o r e s t occurs on F i g u r e 10 - V e g e t a t i v e cover of key mountain goat, range (south a s p e c t ) on Maroon Mountain, B.C. 53 54 s o u t h - f a c i n g slopes and v a l l e y - b o t t o m s . Picea glauca and P_j_ s i t c h e n s i s are most abundant on slop e s l e s s than 762 m, but Abies a m a b i l i s and Tsuga h e t e r o p h y l l a are a l s o common. Moss, r i p a r i a n , L y s i c h i t u m and Oplopanax a s s o c i a t i o n s are a l s o common, on the v a l l e y -f l o o r . Above 762 m, Pi c e a s p e c i e s are q u i c k l y r e p l a c e d by Tsuga mertensiana. Abies a m a b i l i s . i s r e p l a c e d by A_^  l a s i o c a r p a , which becomes the dominant t r e e s p e c i e s above 1067 m. A cryptogamic mat of moss sp e c i e s s i m i l a r i n composition to the Tsuga-Abies f o r e s t predominates, and a few widely s c a t t e r e d shrubs, mentioned i n the pre v i o u s f o r e s t type, are l o c a t e d w i t h i n the subalpine f o r e s t understory. P t i l i u m c r i s t a - c a s t r e n s i s and s e v e r a l P y r o l a s p e c i e s are a d d i t i o n s to t h i s u nderstory. (3) The Pinus-Populus (Pine-Aspen) f o r e s t occurs o n l y on the d r i e r , rocky s l o p e s of the south-aspect of Maroon Mountain, at the entrance to Maroon Creek v a l l e y . Pinus c o n t o r t a i s the p r i n c i p a l t r e e s p e c i e s . Secondary t r e e s are Populus tremuloides and Tsuga  h e t e r o p h y l l a . A r c t o s t a p h y l o s u v a - u r s i , Chimaphila  umbellata, Cornus canadensis, Linnaea b o r e a l i s , and P y r o l a a s a r i f o l i a compose a scant, but more d i v e r s e , u n d e r s t o r y . A heavy cryptogam i n f l u e n c e i s e v i d e n t , c o n s i s t i n g of P e l t i g e r a apthosa, Pleurozium s c h r e b e r i , P o l y t r i c h u m juniperinum, and P t i l i u m c r i s t a -c a s t r e n s i s . T h i s area was burned sometime duri n g the 55 1920's or 1930's. (4) A f o u r t h f o r e s t category i s a mixed f o r e s t of deciduous t r e e s with some c o n i f e r o u s t r e e s . O c c u r r i n g p r i m a r i l y on the south aspect, on o l d and u n d i s t u r b e d rock l i t t e r s l o p e s , the Populus-Alnus-Thuja (Apsen-Alder-Cedar) f o r e s t c o n t a i n s a v a r i e t y of t r e e s p e c i e s . B e t u l a p a p y r i f e r a , P i c e a g l a u c a , Populus  tremuloides and Thuja p i i c a t a are the dominant t r e e s , w i t h Alnus rubra and A_^  s i n u a t a becoming abundant under more r i p a r i a n i n f l u e n c e . Acer and Alnus shrub forms comprise the t a l l e r understory s p e c i e s . D r y o p t e r i s a u s t r i a c a and Gymnocarpium d r y o p t e r i s are abundant low-form ( f e r n ) s p e c i e s , i n a d d i t i o n to a l a r g e v a r i e t y of mosses (mentioned p r e v i o u s l y ) which c a r p e t the rocky f l o o r . (5) The l e a s t abundant f o r e s t cover d e s c r i b e d i s the sub-montane deciduous f o r e s t , u s u a l l y found i n small patches on g e n t l y - s l o p i n g h i l l s i d e s or on a l l u v i a l f a n s . The Populus-Alnus-Acer (Cottonwood-Alder-Maple) f o r e s t forms two types. Populus tremuloides i s the main t r e e s p e c i e s of the f i r s t type, with Athyrium  f i l i x - f e m i n a and Gymnocarpium d r y o p t e r i s common i n the understory, i n a d d i t i o n to a v a r i e t y of f o r b s . The second type i s comprised of Populus t r i c h o c a r p a as the major t r e e s p e c i e s . C h a r a c t e r i s t i c a l l y on a l l u v i a l f ans, Acer and Alnus shrub forms occur i n abundance. The low-form understory s p e c i e s are s i m i l a r l y d i v e r s e 56 in both types. Open f o r e s t e d woodlands. Three c o n i f e r o u s woodland c a t e g o r i e s were re c o g n i z e d . Each i s merely a d i v e r s i f i c a t i o n of the three c o n i f e r o u s f o r e s t s d e s c r i b e d p r e v i o u s l y . The i n f l u e n c e of rocky s u b s t r a t e and i n c r e a s e d l i g h t f i l t r a t i o n i s shown by a d i v e r s e a r r a y of shrubby p l a n t s such as Acer, Alnus, and Vaccinium s p e c i e s . Graminoid s p e c i e s of A g r o s t i s and Calamagrostis are commonly a s s o c i a t e d with the l a r g e r shrub forms, as are cryptogam, S a x i f r a g a and Sedum s p e c i e s with the more open rocky a r e a s . Scrub. Three scrub c a t e g o r i e s occur, c o n s i s t i n g of two shrublands and one shrub t h i c k e t . (1) The deciduous shrubland i s c h a r a c t e r i z e d by shrubs which are clumped and di v i d e d , from each other by a grass stratum ( s i m i l a r to a scrub p a s t u r e ) . Two types occur, determined by rocky i n f l u e n c e ; one with a t a l l stratum and the other with a low stratum. The t a l l stratum, or Alnus type grows to 4.6 m i n height and i s comprised almost s o l e l y of clumps of Alnus s i n u a t a , i n t e r s p e r s e d by such graminoid s p e c i e s as Agropyron  caninum, A g r o s t i s e x a r a t a , A. scabra, Calamagrostis  canadensis, Festuca o v i n a , Poa a l p i n a and Trisetum  spicatum. A c h i l l e a m i l l e f o l i u m , Rosa nutkana, Carex and L u z u l a , s p e c i e s and other herbs are a l s o abundant. The low stratum, or Rubus type, grows to 1m i n h e i g h t , comprised of Heracleum lanatum, Rubus 57 p a r v i f l o r u s , S m i l a c i n a s t e l l a t a and Veratrum v i r i d e . Few graminoid s p e c i e s occur i n t h i s sub-type. ( 2 ) A second ( c o n i f e r o u s ) shrubland, or 'krummholz', i s composed mostly of c r e e p i n g or lodged needle-leaved phanerophytes such as Abies l a s i o c a r p a and some Tsuga  mertensiana. Shrub forms are grouped and d i v i d e d from each other by e i t h e r exposed rock, cryptogamic or herbaceous l a y e r s , or a l l three s t r a t a types i n mosaic f a s h i o n . The undergrowth may be d i v e r s e , c o n s i s t i n g of many a l p i n e and subalpine shrub, herb and moss-l i c h e n s p e c i e s r e q u i r i n g a wide range of moisture g r a d i e n t s . Shrubs such as Empetrum nigrum, Rubus  pedatus, P h y l l o d o c e , S a l i x and Vaccinium s p e c i e s a l l i n t e r m i x , along with A c h i I l e a m i l l e f o l i u m , A r t e m i s i a  n o r v e g i c a , Carex and Luzula s p e c i e s , S i l e n e a c a u l i s , and Festuca ovina and Poa graminoid s p e c i e s . Broad-l e a f e d f o r b s are not abundant, however numerous cryptogam s p e c i e s occur, p a r t i c u l a r l y P l a t i s m a t i a  g l a u c a , Pleurozium s c h r e b e r i , and A l e c t o r i a s p e c i e s . ( 3 ) The deciduous shrub t h i c k e t i s more or l e s s dense scrub with or without a s i g n i f i c a n t l y reduced herbaceous undergrowth, in c o n t r a s t to the deciduous shrubland. Deciduous t h i c k e t s occur p r i m a r i l y on seepage slopes below c l i f f rock or along r a v i n e s . P l a n t s p e c i e s , when present, are e s s e n t i a l l y the same as those d e s c r i b e d of the deciduous shrubland, only i n d i f f e r i n g p r o p o r t i o n s ; i . e . l e s s herbs and more woody 58 p l a n t s . Logging C l e a r c u t . Logged areas north of Maroon Creek ranged from one to seven years of age, as of the summer of 1978. Each c l e a r c u t was slash-burned a f t e r l o g g i n g o p e r a t i o n s were completed. Succession progressed i n order of cryptogam, shrub, and herb s t r a t a , r e s p e c t i v e l y . Dense cryptogam cushions of Dicranum, Mn i um, Pogonatum and P o l y t r i c h u m moss s p e c i e s q u i c k l y invaded the more r e c e n t l y c l e a r e d areas of organic s u b s t r a t e , i n a d d i t i o n to a host of other bryophytes. Alnus rubra, A. s i n u a t a and other woody c o l o n i z e r shrubs such as Aruncus S y l v e s t e r , Chimaphila umbellata, Ribes l a x i florum, Rosa nutkana, Sambucus racemosa, Sorbus s i t c h e n s i s , and Cornus, Rubus, S a l i x and Vaccinium s p e c i e s e s t a b l i s h e d themselves soon afterwards, growing i n p r o f u s i o n . A g r o s t i s , Arabics, Carex , C i r s i um, E p i l o b i u m , Mimulus, Phleum and S m i l a c i n a s p e c i e s dominate i n the o l d e r c l e a r c u t s (>2 years of age), along with A c h i l l e a  m i l l e f o l i u m , . A n a p h a l i s margaritacea, Lupinus  l a t i f o l i u s , Montia s i b i r i c a , and T i a r e l l a t r i f o l i a t a . Every c l e a r c u t had been r e p l a n t e d with P i c e a t r e e s e e d l i n g s p r i o r to the f a l l of 1978. T a l l - s e d g e Swamp. Open bodies of water, c r e a t e d by now-abandoned beaver dams support l a r g e homogenous stands of Carex and Equisetum in t e r m i x e d . Where orga n i c m a t e r i a l has f i l l e d i n more r a p i d l y , a few 59 small P i c e a s i t c h e n s i s and P_^  qlauca snags stand, l e s s than 5 m i n h e i g h t . A few v a l l e y - b o t t o m graminoid s p e c i e s a l s o grow on these higher a r e a s . The beaver dams themselves are covered with young shrub growth, such as Alnus and S a l i x s p e c i e s , along with a v a r i e t y of herbs. S c a r c e l y vegetated a r e a s . Subalpine and a l p i n e c a t e g o r i e s are re c o g n i z e d . P l a n t s may be s c a t t e r e d or absent. (1) In the s u b a l p i n e , chasmophytic v e g e t a t i o n dominated, with permanent p l a n t s rooted i n f i s s u r e s of rocks or w a l l s . Many c o l o n i z e r shrubs occured, i n a d d i t i o n to r e t a r d e d growth forms of both c o n i f e r o u s and deciduous t r e e s . S i l e n e • a c a u l i s , Sedum and Senec i o s p e c i e s , i n a d d i t i o n to many graminoid s p e c i e s , occured on well-manured s u r f a c e rock. Dicranum, Mnium, P e l t i g e r a , and Po l y t r i c h u m cryptogam s p e c i e s a l s o abound. (2) A l p i n e i n t e r f l u v e s , rocks and scree supported permanent herbs or half-woody, p r o s t r a t e p l a n t s with i n f r e q u e n t occurrence. Much of the v e g e t a t i v e cover comprised cryptogam s p e c i e s . A l p i n e slopes on the south aspect of Maroon Mountain were g e n e r a l l y s p a r s e l y - v e g e t a t e d , u s u a l l y with l e s s than 10% ground cover. No w e l l - d e f i n e d p l a n t communities dominated an area l a r g e enough to map. T h e r e f o r e , s c a r c e l y vegetated rock, small herbaceous meadows, and 60 cryptogamic mats of l i c h e n and bryophyte s p e c i e s were recognized but not d e p i c t e d on the v e g e t a t i o n map (see F i g u r e 10). b. Use of V e g e t a t i v e Cover i . O verstory Assessment Only three ' f o r e s t ' c a t e g o r i e s were used to d e s c r i b e mountain goat use ( c o n i f e r o u s , deciduous and mixed). References to c o n i f e r o u s f o r e s t s i n c l u d e d a l l three c o n i f e r o u s f o r e s t c a t e g o r i e s ( i . e . Hemlock-Fir, S p r u c e - F i r , and Pine-Aspen). 'Woodlands' were c o n s i d e r e d as a s i n g l e category ( i . e . c o n i f e r o u s ) o n l y , due to an absence of deciduous and mixed c o n i f e r o u s - d e c i d u o u s formations i n the study area. Mountain goats d i d not u t i l i z e v e g e t a t i v e cover c a t e g o r i e s i n equal p r o p o r t i o n s (G=7680.4;df=9). They were observed to use twelve of the f i f t e e n p l a n t formation c a t e g o r i e s on Maroon Mountain p r e v i o u s l y d e s c r i b e d . These i n c l u d e two s c a r c e l y vegetated areas ( a l p i n e and s u b a l p i n e ) , krummholz, the three c o n i f e r o u s f o r e s t and woodland c a t e g o r i e s , mixed f o r e s t s , deciduous shrublands, and deciduous shrub t h i c k e t s . The s c a r c e l y - v e g e t a t e d rock category, i n c l u d i n g both a l p i n e and subalpine subtypes, r e c e i v e d more use (69.4%), o v e r a l l , than the remaining c a t e g o r i e s combined (Figure 11). 97 131 1 0 0 CO Z o < > cc III CO CD U» O z -Ul (J cc Ul a. 2961 Mean • • V e g e t a t i v e C o v e r T a l l - s e d g e S w a m p l a g g i n g C l e a r c u t D e c i d u o u s Fo r e s t M i x e d F o r e s t D e c i d u o u s S h r u b l a n d D e c i d u o u s S h r u b T h i c k e t C o n i f e r o u s W o o r i l a n d C o n i I e ro us Fo re s t K r u m m h o I z S c a r c e l y V e g e t a t e d A r e a s CTi MONTH F i g u r e 11 - Summary o f m o u n t a i n g o a t u s e o f v e g e t a t i v e c o v e r on Maroon M o u n t a i n , B.C. 62 Deciduous shrublands were the second-most used p l a n t cover (10.5%). Krummmholz, c o n i f e r o u s f o r e s t s , c o n i f e r o u s woodlands and deciduous shrub t h i c k e t s were used i n f r e q u e n t l y (6.3%, 5.9%, 4.0%, and 3.4%, r e s p e c t i v e l y ) . Mountain goats were only s p o r a d i c a l l y observed i n the mixed f o r e s t cover types (0.2%). No mountain goats were observed i n the deciduous f o r e s t , l o g g i n g c l e a r c u t s , or the t a l l - s e d g e swamp. U t i l i z a t i o n of v e g e t a t i v e cover was dependent upon time of year (G=5790.1;df=75). S c a r c e l y vegetated areas were used p r o p o r t i o n a t e l y more than any other category i n every month (n=16). However, i n May, 1978, mountain goat use of deciduous shrub areas exceeded that of the s c a r c e l y vegetated rock type ( F i g u r e 11). Deciduous shrublands and t h i c k e t s were a l s o used e x t e n s i v e l y from September through November, and again from February through A p r i l . C o n i f e r o u s timber areas were used p r i m a r i l y d u r i n g the f a l l and winter, whereas the krummholz zone r e c e i v e d most of i t s use d u r i n g the snow-fr e e p e r i o d s , i i . Understory C h a r a c t e r i s t i c s Sample s i z e s f o r some c h a r a c t e r i s t i c s were low i n c e r t a i n months. T h e r e f o r e , one should c o n s i d e r the f o l l o w i n g i n t e r p r e t a t i o n b e a r i n g 63 t h i s i n mind. Herbaceous understory types were not u t i l i z e d e q u a l l y by mountain goats (G=2133.5;df=3). Monthly d i f f e r e n c e s accounted f o r part of t h i s v a r i a t i o n (G=608.0;df=45). Averaged over the study p e r i o d , the most common understory used by mountain goats (68.8%) was comprised of l e s s than 50% herbaceous cover ( F i g u r e 12), and was t y p i c a l l y a s s o c i a t e d with rocky s u b s t r a t e s . T h i s type was u t i l i z e d most through a l l months with the ex c e p t i o n of December, when herbaceous meadows (low stratum type) were e x c l u s i v e l y used in sub a l p i n e areas. T h i s low stratum type was used mostly from September through December, and again from February through May. The a l p i n e (low stratum) and subalpine (high stratum) herbaceous meadows were predominantly u t i l i z e d d u r i n g snow-free p e r i o d s of the year (June through August). The o v e r a l l d i v i s i o n of use of p l a n t s t r a t a w i t h i n the krummholz zone (Figure 13) was s i m i l a r (G=0.8;df=1), with 52.5% of the mountain goats observed u t i l i z i n g the high stratum. S e a s o n a l l y , however, use of the two s t r a t a v a r i e d s i g n i f i c a n t l y (G=94.6;df=15). The higher stratum was used mostly i n the f a l l , winter and s p r i n g , f o r what appeared to be thermal c o v e r . The lower 11 153 231 120 240 206 132 75 12 16 42 77 116 216 185 97 130 2048 100 CO z o < > cc Ul to CO z Ul u cc Ul a. so M jTTTTTmt mm. J A S O N D J (1977) F M A M J (1978) Mean • Understory < 50% Herbaceous (Alpine & Subalpine) > 50% Herbaceous (Alpine) > 50% Herbaceous & > 0.9m (Subalpine) > 50% Herbaceous & <0.9m (Subalpine) ON M O N T H F i g u r e 12 - Summary of m o u n t a i n g o a t u se of h e r b a c e o u s u n d e r s t o r y t y p e s on Maroon M o u n t a i n , B.C. n 11 43 19 24 54 24 6 9 27 17 20 24 20 38 0 9 345 S t r a t i 100 | | High ( l . 2 - 4 . 6 m ) low (< 1.2 m) CO < > SC Ul (0 CD O SO < Ul u cc Ul a. ° M J J A S O N D J F M A M J J A Mean C1977) (1978) MONTH F i g u r e 13 - Summary o f m o u n t a i n g o a t use of krummholz s t r a t a on Maroon M o u n t a i n , B.C. 66 stratum was u t i l i z e d the h e a v i e s t d u r i n g l a t e s p r i n g and throughout the summer. Although the low woody stratum may hinder t r a v e l due to i t s d e n s i t y , i t a l s o a s s i s t s mountain goats i n removal of t h e i r shedding h a i r . Often the stunted t r e e s are clumped and o f f e r a l u s h herb mat s u i t a b l e f o r f o r a g i n g . In the deciduous shrub cover category ( F i g u r e 14), mountain goats were most f r e q u e n t l y observed (71.5%) i n areas o f f e r i n g the higher v e g e t a t i o n s t r a t a , as t h i s type was c h a r a c t e r i s t i c of a d i v e r s e a r r a y of herbaceous understory m a t e r i a l (G=64.4;df=1). The lower stratum g e n e r a l l y o f f e r e d l i t t l e forage other than the shrubs themselves. Use of each s t r a t a was v a r i a b l e throughout the year (G=29.1;df=15), with no apparent t r e n d . F o r e s t woody shrub understory types were a l s o not used i n equal p r o p o r t i o n s (G=281.1;df=3). Use of each type v a r i e d throughout the year (G=80.6;df=45) as w e l l (Figure 15), however, c o n i f e r o u s t r e e s were u t i l i z e d most, except d u r i n g p a r t u r i t i o n (May and June). C o n i f e r o u s shrub u n d e r s t o r i e s were used p r o p o r t i o n a t e l y more d u r i n g the k i d d i n g p e r i o d , but were a l s o h e a v i l y u t i l i z e d d u r i n g w i n t e r . N e g l i g i b l e use of the mixed ( c o n i f e r o u s -n 23 21 3 100 0 24 39 40 2 CO Z o *-> rz Ul CO to z Ul o cc Ul a. SO 19 40 29 71 8 M A S 0977) M A M (1978) 324 Mean Shrub Strata | | High (> 1.0m ) FTi] low (< 1.0m) ON MONTH F i g u r e 14 - Summary of m o u n t a i n g o a t use o f d e c i d u o u s s h r u b s t r a t a on Maroon M o u n t a i n , B.C. M A S (1977) N M A M (1978) Mean MONTH F i g u r e 15 - Summary o f m o u n t a i n g o a t use of f o r e s t woody s h r u b u n d e r s t o r i e s on Maroon M o u n t a i n , B.C. 69 deciduous) timber and mixed shrub understory types was e v i d e n t . 2.1.2 A b i o t i c F a c t o r s a. T e r r a i n C l a s s i f i c a t i o n i . S urface E x p r e s s i o n O v e r a l l a n a l y i s showed that mountain goats u t i l i z e d landforms i n unequal p r o p o r t i o n s (G=1114.8;df=6). Rock b l u f f s , outcrops or k n o l l s were used s i g n i f i c a n t l y more (42.4%) than any other t e r r a i n f e a t u r e d e s c r i b e d ( F i g u r e 16). They g e n e r a l l y o f f e r e d u n r e s t r i c t e d viewing of the immediate surroundings and f r e q u e n t l y served as bedding a r e a s . Benches, ledges, and s h e l v e s were used l e s s f r e q u e n t l y (19.1%) by goats, as were r a v i n e s (14.5%), r i d g e s (10.4%), and t a l u s s l o p e s (9.6%). Peaks (2.7%) and hummocky (tundra) areas (1.3%) were u t i l i z e d to a much l e s s e r e x t e n t . Mountain goat use of topographic f e a t u r e s were a l s o dependent upon time of year (G=l080.7;df=90). Rock b l u f f s , outcrops and k n o l l s were used throughout the year, but with a preponderance of use d u r i n g the f a l l and winter p e r i o d s (November t o March) when snow covered most of the ground. A s h i f t to usage of steep r a v i n e rock w a l l s o c c u r r e d i n l a t e winter and II 80 140 49 106 95 56 144 69 91 91 100 73 118 57 53 105 1427 Landlorm O M J J A S O N D J F M A M J J A Mean (1977) (1978) MONTH F i g u r e 16 - Summary of m o u n t a i n g o a t u s e o f l a n d f o r m s on Maroon M o u n t a i n , B.C. 71 e a r l y s p r i n g (March to May), subsequent to the peak in use of rock b l u f f s , outcrops and k n o l l s . Both • c a t e g o r i e s have important snow-shedding f e a t u r e s b e l i e v e d to a t t r a c t mountain goats f o r f e e d i n g . Benches, ledges and s h e l v e s were f r e q u e n t l y u t i l i z e d by mountain goats d u r i n g the p e r i o d of p a r t u r i t i o n i n 1977, however, steep r a v i n e w a l l s and r i d g e - t o p s were used by the goats d u r i n g the same p e r i o d i n 1978. These d i f f e r e n c e s are thought to r e s u l t from v a r y i n g temperature and snow c o n d i t i o n s between the two years ( i . e . more snow i n 1978). i i . Rock Substrate Summary of the t o t a l mountain goat use of rock s u b s t r a t e s shows d i s p r o p o r t i o n a t e l y higher use of broken rock (87.7%), throughout the year (G=3579.2;df=3) ( F i g u r e 17). Slopes of loose shale were the second most used s u b s t r a t e by goats (only 6.7%), s i m i l a r i n degree to use of g r a v e l slopes (4.6%). Smooth rock s u r f a c e s were u t i l i z e d only 0.9% of the time. Temporal v a r i a t i o n i n the use of s u b s t r a t e types a l s o o c c u r r e d (G=492.9;df=45). The broken rock category was c o n s i s t e n t l y used more than a l l other c a t e g o r i e s throughout the e n t i r e study p e r i o d . Shale s l o p e s were u t i l i z e d p r i m a r i l y d u r i n g the l a t e s p r i n g and e a r l y summer (June n 85 172 85 160 162 85 115 45 103 171 191 123 116 163 80 119 1975 Rock Substrata 100 CO z o 4 > cc ui (0 ea O »0 z Ul u cc Ul GL D i r t , G r a v e l Smooth Rock jj Sha le | | B roken Rock M A S 0977) M A M 0978) Mean MONTH F i g u r e 17 - Summary o f m o u n t a i n g o a t u se o f r o c k s u b s t r a t e s on Maroon M o u n t a i n , B.C. 7 3 through August) but to a much l e s s e r extent i n 1977, when mountain goats used d i r t - g r a v e l s l o p e s p r o p o r t i o n a t e l y more than f o r the same p e r i o d i n 1978. The former type i s g e n e r a l l y used f o r t r a v e l and bedding d u r i n g hot temperature extremes. The l a t t e r category was p r i m a r i l y used f o r ' dusting' by mountain goats, g e n e r a l l y d u r i n g high summer temperatures. Aspect O v e r a l l assessment of aspect u t i l i z a t i o n by mountain goats suggests that each category i s not used in equal p r o p o r t i o n (G=7566.0;df=8) . D i f f e r e n c e s i n use were accounted f o r by monthly e f f e c t s (G=520.4;df=120) (Figure 18). Between e a r l y May, 1977 and l a t e August, 1978, 96% of a l l mountain goats s y s t e m a t i c a l l y observed on Maroon Mountain o c c u r r e d on s o u t h e r l y - f a c i n g slopes between 112° and 247°. N o r t h e r l y a s p e c t s were r a r e l y used by mountain goats and almost e x c u l s i v e l y d u r i n g c o n d i t i o n s of high ambient temperatures i n the snow-free months of June through September. An i n c i d e n t a l mountain goat s i g h t i n g on a north aspect i n l a t e May probably arose from ' e x p l o r a t o r y ' t r a v e l over tundra areas s u b j e c t e d to recent snow-melt c o n d i t i o n s . Slope Mountain goats u t i l i z e d s l o p e s ranging from 16° M O N T H F i g u r e 18 - Summary of m o u n t a i n g o a t u se o f a s p e c t s on Maroon M o u n t a i n , B . C . 75 to 62° d i s p r o p o r t i o n a t e l y (G=3546.5;df=l0). However, n e a r l y 84% of a l l mountain goats observed u t i l i z e d i n c l i n e s between 30° and 45° (Figure 19). T h i r t y -e i g h t percent of the goats were observed i n the 38-41° i n t e r v a l c l a s s . Temporal v a r i a t i o n s o c c u r r e d i n the use of slope i n t e r v a l s (G=866.2;df=150). The 38-41° i n t e r v a l was predominantly u t i l i z e d throughout the year. Steeper slope c l a s s e s (42-45°,46-50° , 51-56°, and 57-63°) were used by mountain goats p r i m a r i l y d u r i n g months when abundant snow cover was present (November and February) or when the ground was exposed by shedding or m e l t i n g snow (March through June). More g e n t l e grades (<27°) were used p r o p o r t i o n a t e l y more d u r i n g the summer p e r i o d s ( J u l y to September), when snow-free c o n d i t i o n s enabled occupancy of a l p i n e r i d g e and tundra areas. d. E l e v a t i o n T r e e l i n e was estimated to occur at 1292 m n o r t h of the main r i d g e , and at 1194 m south of the main r i d g e , averaging 1252 m throughout the e n t i r e study a r e a . The a l p i n e zone ranges v e r t i c a l l y over 734 m on the southern s l o p e s of Maroon Mountain. The subalpine zone ranges v e r t i c a l l y over n e a r l y h a l f as much again (1057 m). O v e r a l l mountain goat use was s i g n i f i c a n t l y d i f f e r e n t between 250-m i n t e r v a l e l e v a t i o n c l a s s e s n 178 264 133 251 225 137 205 133 151 229 224 186 223 191 97 131 2958 deg rees MONTH F i g u r e 19 - Summary of m o u n t a i n g o a t use o f h i l l s i d e s l o p e on Maroon M o u n t a i n , B.C. 77 (G=2174.7;df=8). However, more than 60% of the mountain goats observed o c c u r r e d i n a l p i n e h a b i t a t ( F i g u r e 20). The two 250-m i n t e r v a l s adjacent to t r e e l i n e (1001-1250 m and 1251-1500 m) accounted f o r 45% of a l l mountain goat o b s e r v a t i o n s . No mountain goats were observed below 250 m. Temporal v a r i a t i o n s i n the use of 250-m a l t i t u d i n a l zones were a l s o apparent (G=2281.2;df=120). Use of higher e l e v a t i o n s i n c r e a s e d d u r i n g months of snowpack r e c e s s i o n and extreme sunny (warm) temperatures (June through September). A f t e r t h i s p e r i o d , a l t i t u d i n a l use dropped s t e a d i l y , to w e l l below t r e e l i n e i n November, with the onset of stormy weather and str o n g n o r t h e r l y winds. Most of the mountain goats u t i l i z e d s u b a l p i n e h a b i t a t s from October t o May, except i n January. e. M i c r o c l i m a t e Weather s t a t i o n s were monitored f o r up to twelve months, r e c o r d i n g temperature, humidity, wind speed, and p r e c i p i t a t i o n l e v e l s whenever p o s s i b l e . U n f o r t u n a t e l y only the temperature data were worthy of a n a l y s i s ( e x c l u d i n g the government maintained f a c i l i t y at Rosswood, B.C.), due to instrument f a i l u r e s and other v a r i o u s mishaps. The 'main r i d g e ' weather s t a t i o n (#4) was blown over by str o n g n o r t h e r l y winds i n October and was not recovered u n t i l March. A n a l y s i s of the study s i t e temperature data n 178 264 133 251 227 137 205 133 151 229 225 1 86 223 1 91 97 1 31 2961 meters MONTH F i g u r e 20 - Summary of m o u n t a i n g o a t a l t i t u d i n a l u s e by 250-m c l a s s e s on Maroon M o u n t a i n , B.C. 79 (Figure 21) r e v e a l e d s e v e r a l important c l i m a t o l o g i c a l f e a t u r e s on mountain goat ranges. Mean monthly temperatures were s i g n i f i c a n t l y d i f f e r e n t between s t a t i o n s (Hc=13.6;df=6). Of the seven weather s t a t i o n s assessed, s t a t i o n #1 ( ' W a t e r f a l l ' : 879 m) was the warmest, d u r i n g the s p r i n g months ( A p r i l to June). At t h i s time, the c o l d e s t l o c a t i o n s were the a l p i n e s t a t i o n s ( s t a t i o n #4, at 1692 m, and s t a t i o n #5, at 1591 m). Throughout the summer p e r i o d , Rosswood recorded the h i g h e s t mean monthly temperature v a l u e s . During e a r l y f a l l (October), the head of Maroon v a l l e y and the v a l l e y bottom a c t e d as ' c o l d - s i n k s ' , always being i n the shade due to the low angle of the sun. A l p i n e s t a t i o n s a l s o maintained c o l d temperatures at t h i s time, with the ' t r e e l i n e ' s t a t i o n being the warmest l o c a t i o n . During November, temperatures dropped, becoming i n d i c a t i v e of winter, p a r t i c u l a r l y at s t a t i o n #5. However, i n December, a pronounced thermal i n v e r s i o n had formed a warm a i r mass above t r e e l i n e with a l p i n e s t a t i o n s #3, #4(?) and #5 approaching mean monthly temperatures up to 17°C warmer than lower e l e v a t i o n a r e a s . In winter (January to March), however, the a l p i n e r i d g e s a p p a r e n t l y c o o l e d down to temperatures s i m i l a r to that of the other s t a t i o n s , but the ' t r e e l i n e ' s t a t i o n remained much warmer than the others f o r the e n t i r e three-month p e r i o d . 80 20 i o o Ul E CC Ul CL 2 UJ • > - i x »-z o 5 z < UJ 2 15 10 H •10 15 O . - O ROSSWOOD I152>> 8TN. NO. 6 I 7 2 5 a l »• — » STN . NO. 1 I 8 7 9 - ) STN. NO. 2 ( 8 0 3 - I • STN . NO. 3 (1433-1 » — • STN . NO. 8 11591B! a... A STN. NO. 4 ( 1 « 9 2 a l A S O N 0977) M A M J C1978) MONTH F i g u r e 21 - Summary of mean monthly temperature values f o r s i x weather s t a t i o n s on Maroon Mountain, B.C. Note thermal i n v e r s i o n s as shown by s t a t i o n no.3 i n December, January, and February. 81 F o r e s t e d areas ( s t a t i o n #2) and adjacent non-f o r e s t e d areas ( s t a t i o n #1) were not s i g n i f i c a n t l y d i f f e r e n t i n t h e i r mean monthly temperature regimes (T=21;n=12). However, f o r e s t e d areas were noted to be s l i g h t l y warmer than neighbouring exposed areas from September to February, with the e x c l u s i o n of November. From March through August, f o r e s t e d areas were, on average, 4°C c o o l e r than exposed areas. 2.2 H a b i t a t S e l e c t i o n T h i s a n a l y s i s i s presented to determine i f h a b i t a t s were being used by mountain goats d i s p r o p o r t i o n a t e to t h e i r a v a i l a b l i l i t y . S e l e c t i o n d e s c r i b e s a circumstance, not an a c t . ' P r e f e r r e d ' h a b i t a t s are those i n which mountain goats were found to use a p a r t i c u l a r h a b i t a t category more than i n p r o p o r t i o n to i t s a v a i l a b i l i t y . 'Avoidance' r e f e r s to the converse ( i . e . goat use i s s i g n i f i c a n t l y l e s s than i n p r o p o r t i o n to i t s a v a i l a b i l i t y ) . ' S e l e c t i o n ' r e f e r s to the circumstances of avoidance and p r e f e r e n c e ; nothing e l s e i s i m p l i e d by the use of these three terms. 2.2.1 Data C o n s i d e r a t i o n s The data were f i r s t c o n s i d e r e d i n f i v e d i f f e r e n t ways before the a n a l y s i s of s e l e c t i o n was undertaken. a. Area of A v a i l a b i l i t y Based upon the cumulative d i s t r i b u t i o n of mountain goats presented i n F i g u r e 3, and known movements of s e v e r a l i n d i v i d u a l s ( F i g u r e s 6 and 7), 82 the area ' a v a i l a b l e ' to goats was c o n s i d e r e d to i n c l u d e the e n t i r e s o u t h e r l y - f a c i n g slope of Maroon Mountain, from r i d g e - t o p down to Maroon Creek i t s e l f . The summering areas north of the main r i d g e were not a v a i l a b l e to the goats i n 1977 or 1978 due to the common occurrence of p r e d a t o r s (Cards lupus, Gulo qulo and Ursus a r c t o s ) . C o r r e c t i o n For H i l l s i d e Slope A r e a l c a l c u l a t i o n s and random sample e s t i m a t i o n of p r o p o r t i o n s of a v a i l a b i l i t y showed no s i g n i f i c a n t d i f f e r e n c e i n the p r o p o r t i o n s of p l a n t cover (Wilcoxon's T=16.0;n=10), aspect (T=5.0;n=5), slope (T=l7.0;n=8), and 250-m i n t e r v a l s (T=14.5;n=9) a f t e r c o r r e c t i n g f o r h i l l s i d e s l o p e , although i n each case the a b s o l u t e area of each category was o b v i o u s l y l a r g e r . The c o r r e c t e d data were i n c l u d e d i n the p r e f e r e n c e a n a l y s i s . Surface e x p r e s s i o n , s u b s t r a t e t e x t u r e and p l a n t understory v a r i a b l e s were not used. Random Subsampling From Groups O r i g i n a l l y i t was planned to subsample groups i n order to s t a t i s t i c a l l y analyse independent data by reducing b i a s e s r e s u l t i n g from s o c i a l c h o i c e s r a t h e r than h a b i t a t c h o i c e s . However, I d i d not use the independent (subsampled) data i n the f i n a l a n a l y s i s , as i t was f e l t that s e x - r e l a t e d a t t r i b u t e s ( i . e . group types) played a more s i g n i f i c a n t r o l e i n goat ecology, 83 and i n s u f f i c i e n t data were a v a i l a b l e to subsample f o r both group s i z e and group type on a monthly b a s i s . However, the i m p l i c a t i o n s of random subsampling of one animal per group as opposed to using a l l i n d i v i d u a l s i n the a n a l y s i s were ex p l o r e d . S i g n i f i c a n t d i f f e r e n c e s i n h a b i t a t use oc c u r r e d f o r some v a r i a b l e s . Use of p l a n t cover (G=20.3;df=10) and e l e v a t i o n at 250-m i n t e r v a l s (G=70.5;df=8) were s i g n i f i c a n t l y d i f f e r e n t between the subsampled and the o v e r a l l data, but aspect (G=11.4;df=8) and slope (G=17.8;df=10) showed no s i g n i f i c a n t d i f f e r e n c e s . I n f l u e n c e of S o c i a l S t r u c t u r e Group type was a l s o chosen as an important f a c t o r i n the p r e f e r e n c e a n a l y s i s because 33.2% of a l l males were a s s o c i a t e d with female goats. I t seemed l o g i c a l , i n view of the f a c t that males are l e s s a g g r e s s i v e than females (Chadwick 1973, Rideout 1974), that male use of h a b i t a t a t t r i b u t e s was determined by the dominant sex, whom they followed (when i n t h e i r company). Under t h i s same r a t i o n a l e , when used f o r group subsampling, only males i n 'male-only' groups represented true s e l e c t i o n of h a b i t a t a t t r i b u t e s by each sex. Sample Repre s e n t a t i o n A f u r t h e r r e s t r i c t i o n p l a c e d on the data p r i o r to a n a l y s i s was adequate r e p r e s e n t a t i o n of the absolute 84 numbers of male and female components of the p o p u l a t i o n . From F i g u r e 9, i t was apparent that the sampled p r o p o r t i o n s of males and females d i d not exceed 50% of each component f o r 10 and 3 months, r e s p e c t i v e l y , of the 16 months sampled. Because an assessment of s e l e c t i o n based upon a non-r e p r e s e n t a t i v e p r o p o r t i o n of a sex would be m i s l e a d i n g , the a n a l y s i s was r e s t r i c t e d to those months i n which each sex was represented by more than 50% of i t s t o t a l estimated number ( i . e . November, 1977 and January to May, 1978 f o r males; May, June, August to November, 1977 and January to July', 1978 f o r females). 2.2.2 B i o t i c F a c t o r s a. V e g e t a t i v e Cover Mixed-female groups were more s e l e c t i v e f o r v e g e t a t i v e cover than male-only groups (Table 3). Only i n May, 1978 was a p l a n t cover type not used e q u a l l y with respect to i t s a v a i l a b i l i t y by male-only groups, whereas f o r the same p e r i o d , mixed-female groups used v e g e t a t i v e cover d i s p r o p o r t i o n a t e l y i n every month but January. Male-only groups p r e f e r r e d deciduous shrub t h i c k e t s d u r i n g May, 1978, but mixed-female groups showed an i n d i f f e r e n c e t o t h e i r use. Mixed-female groups p r e f e r r e d the s c a r c e l y vegetated areas i n a l l months but September and October of 1977, 8 5 Table 3 - Mountain goat s e l e c t i o n of v e g e t a t i v e cover on Maroon Mountain (south slope) f o r male-only and mixed-female groups. Date Log Likelihood Number o f Goats Male -only Group Selection 1 Ratio Scarcely Vegetated Areas Kruamholz Coniferous Forest Coniferous Woodland Deciduous Shrub Thicket Deciduous Shrubland Mixed Forest Deciduous Forest Logging Clearcut Swamp 1977 May-Oct ro m n m B n m m m n n SI Nov 9.3963 16 0 (0) 0 0 0 0 (0) (0) (0) (0) Dec m n o m m m in m Ca ID n m 197B Jan 11.696B 12 0 0 (0) 10) (0) (0) (0) (0) (0) (0) Feb 11.3037 40 0 0 0 0 0 0 0 (0) (0) (0) Mar 16.3164 41 0 0 0 0 0 0 (0) (0) (0) (0) Apr 12.6631 38 0 0 0 0 0 0 0 (0) (0) (0) Hay 21.2146* .42 0 0 0 0 + 0 (0) (0) (0) (0) Jun-Aug m HI to D m m m 01 D D m Date Log Likelihood Number of Goats Mixed-female Group Selection 1 Ratio Scarcely Vegetated Areas Krunmhols Coniferous Forest Coniferous Woodland Deciduous Shrub Thicket Deciduous Shrubland Mixed Forest Deciduous Forest Logging Clesrcut Swsmp 1977 May 66.0977 151 * 0 - 0 0 + (-» (0! (-) <0| Jun 95.8730* 233 0 - 0 0 0 - (0) (-) (0) Jul in tn Dl m m ID m m tn m n m Aug 91.8657* 217 + 0 - <-> <-) - (-) (0) (-) (0) Sep 2.8833 179 0 0 0 0 0 0 (0) (0) (0) (0) Oct 27.2944* 106 0 0 0 0 0 + 0 (0) (-) (0) Nov 72.9258* 150 • - - 0 0 0 (0) (-) (0) Dec m D m B n B m • m B B B 1978 Jan 4.6045 128 0 0 0 (0) (0) (0) (0) (0) (0) (0) Feb 88.6914* 172 + - 0 0 {-) 0 (-1 (0) <-) (0) Mar 68.2544* 159 + 0 - 0 0 0 0 (0) (-) (0) Apr 59.9629* 118 + 0 - 0 0 0 0 ID) (-) 10) May 27.7710* 164 0 (-) - 0 0 • (-1 (0) (-) (0) Jun 17.9751* 164 + 0 - - (-) 0 (-1 10) (-) 10) Jul 82.8145* 80 • (-) (0) (0) 0 10) (0) (0) (0) Aug at D * II in n Ol m • B • B 1 - preferred, or used i n greater proportion than available; *0> - no eel action, or used l a equal proportion to availability! '-' • avoided, or used in less proportion than available. Bracketed syabols indicate no use. m - non-representative data (i.a. leas than 50* of tha estimate sa* eoaponant i n tha population) • significant at p<0.05 86 and January and May of 1978, followed s e c o n d a r i l y by pr e f e r e n c e f o r deciduous shrublands i n May, October, November, 1977 and May, 1978. Mixed-female groups a l s o avoided more cover c a t e g o r i e s (e.g. f o r e s t e d areas) than d i d male-only groups. Although the a n a l y s i s f r e q u e n t l y showed deciduous f o r e s t s , l o g g i n g c l e a r c u t s , and the t a l l - s e d g e swamp to be used i n p r o p o r t i o n to t h e i r a v a i l a b i l i t y , these c a t e g o r i e s r e c e i v e d no use at a l l by e i t h e r group type ( i . e . they were a v o i d e d ) . T h i s suggests an i n a b i l i t y of Marcum and Loftgaarden's (1980) technique to d i f f e r e n t i a t e use and a v a i l a b i l i t y where low sample s i z e s occur or small areas e x i s t (deciduous f o r e s t s and the t a l l - s e d g e swamp comprised 1.41 and 0.67% of the study area, r e s p e c t i v e l y ) at the given p r o b a b i l i t y l e v e l . 2.2.3 A b i o t i c F a c t o r s a. Aspect Contingency t e s t s of independence on mountain goat use and a v a i l a b i l i t y of aspect are presented i n Table 4. Male-only groups were l e s s s e l e c t i v e of aspect c a t e g o r i e s than were female groups. Male-only groups used aspect c a t e g o r i e s d i s p r o p o r t i o n a t e to t h e i r a v a i l a b i l i t y i n February only, p r e f e r r i n g south exposures and a v o i d i n g southwest exposures. Mixed-female groups d i s p l a y e d s i m i l a r s e l e c t i o n of aspect 87 T a b l e 4 - M o u n t a i n g o a t s e l e c t i o n o f a s p e c t on Maroon M o u n t a i n ( s o u t h s l o p e ) f o r m a l e - o n l y and m i x e d - f e m a l e g r o u p s . Log L i k e l i h o o d Ratio Number of Goats Male-only Group S e l e c t i o n * Southeast Southwest 1977 May-Oct Nov Dec 1978 Jan Feb Har Apr May Jun-Aug 7.2407 17.3677* 6.3071 8.0205 5.4019 12 40 40 38 41 (0) (0) (0) (0) (0) (0) (0) 0 (0) (0) Date Log L i k e l i h o o d Ratio Number of Goats M i x e d - f e m a l e Group S e l e c t i o n * South S o u t h e a s t Southwes t East Wust 1977 May 21 .6896* 149 + - 0 (0) 0 Jun 30 .3149* 232 + 0 - 0 (-) J u l m m m m m m m Aug 27 .0449* 206 0 - 0 (0) 0 Sep 31 .5552* 165 + 0 - 0 (0) Oct 13 .3237* 106 0 - (0) (0) Nov 25 .1196* 150 + 0 - (0) 0 Dec m m m m m m m 1978 Jan 12 .5278* 128 0 0 - (0) (0) Feb 45 .6958 172 + 0 - (0) (0) Mar 30 .2554* 159 + 0 - (0) (0) Apr 36 .3213* 118 + 0 - (0) (0) May 19 .8711* 161 + 0 0 (0) (0) Jun 34 .5952* 161 + 0 - 0 (0) J u l 32 .1929* 69 + 0 - 0 0 Aug m m m m m m RI ' + ' = p r e f e r r e d , or used i n g r e a t e r p r o p o r t i o n than a v a i l a b l e ; '0' » no s e l e c t i o n , or used i n equal p r o p o r t i o n to a v a i l a b i l i t y ; '-' - avoided, o r used i n l e s s p r o p o r t i o n than a v a i l a b l e . Bracketed symbols i n d i c a t e no use. in *> non-representative data ( i . e . l e s s than 50% of the estimated sex component i n the popu l a t i o n s i g n i f i c a n t at p<0.05 88 d u r i n g February, however they a l s o u t i l i z e d aspect d i s p r o p o r t i o n a t e to i t s a v a i l a b i l i t y i n a l l other months as w e l l . In each case, south a s p e c t s were g e n e r a l l y p r e f e r r e d and southwest aspects g e n e r a l l y avoided. Western a s p e c t s were a l s o avoided i n June, 1977, and southeast f a c i n g slopes were avoided i n May and August, 1977. b. Slope Mixed-female groups were more s e l e c t i v e of slope c a t e g o r i e s than were male-only groups (Table 5 ) . Mixed-female groups s e l e c t e d at l e a s t two slope c a t e g o r i e s i n each month of the a n a l y s i s except f o r August, 1977. Male-only groups s e l e c t e d only one category i n three of the s i x months analy s e d . In November and March, male-only groups p r e f e r r e d the 38-41° slope c l a s s . G e n t l e slopes (<27°) were avoided i n May. Although contingency a n a l y s i s of the February data concluded that s e l e c t i o n o c c u r r e d i n t h i s month, none was h i g h l i g h t e d by the SCI a n a l y s i s . Mixed-female groups p r e f e r r e d the 38-41° slope c l a s s i n almost every month. A d d i t i o n a l l y , the 35-37° slope c l a s s was p r e f e r r e d i n June, 1977. Although used p r i m a r i l y d u r i n g the summer, g e n t l e r s l o p e s (<29°) were avoided f o r the most p a r t . Female groups a l s o avoided the steeper s l o p e s (>42°) throughout most of the year. 89 T a b l e 5 - M o u n t a i n g o a t s e l e c t i o n o f h i l l s i d e s l o p e on Maroon M o u n t a i n ( s o u t h s l o p e ) by m a l e - o n l y a n d m i x e d - f e m a l e g r o u p s . Log Male-only Group S e l e c t i o n Ratio Goats < 2 7 ° 28-29° 30-31° 32-34° 35-37° 38-41° 42-45° >4 6° 1977 May-Oct m m m m m m m m m Nov 15 7711* IE (0) (0) (0) 0 0 + 0 0 Dec m m m m m m m m m m 1978 Jan 9 0934 12 0 0 0 (0) 0 0 (0) (0) Feb 25 3066* 40 0 10) 0 (0) 0 0 0 (0) Mar 17 9121* 41 0 0 0 0 0 + 0 0 Apr 13 5117 38 0 (0) (0) 0 0 0 0 0 May 24 6040* 42 (-) (0) 0 0 0 0 0 0 Jun-Aug m m m m m m m m m m Mixed-female Group S e l e c t i o n Log Number Date L i k e l i h o o d Ratio Of Goats <27° 28-29° 30-31° 32-34° 35-37° 38-41° 42-45° >46 1977 May 68 4316 151 - 0 0 - 0 + 0 0 Jun 91 9209* 233 - 0 0 0 + + 0 -J u l m m m m m m m m m m Aug 1 6886 217 0 0 0 0 0 0 (0) (0) Sep 73 9569* 177 0 (-) 0 0 0 + - (-) Oct 86 1724* 106 (-) (0) 0 0 0 - 0 Nov 60 5420* 150 - (01 0 - 0 * 0 0 Dec m m ID m m m m m m 1978 Jan 59 4756* 128 - (0) 0 0 0 + 0 (-) Feb 33 7539* 172 - (-) 0 0 0 + 0 0 Mar 63 2915* 158 0 0 0 0 + - 0 Apr 41 0298* 118 - (0) 0 0 0 + 0 0 May 67 1953* 164 - 0 0 0 0 + 0 0 Jun 42 3711* 164 - (-) 0 0 0 + 0 0 J u l 22 6504* 80 0 0 0 0 0 0 0 (-) Aug ID m m m m n m m m m o '+' • preferred, or used in greater proportion than available; *0' - no selection, or used in equal proportion to availability; '-' • avoided, or used in less proportion than available. Bracketed symbols Indicate no use. . non-representative data (i.e. less than 50% of the estimate sex component in the population) significant at p<O.0S 9 0 c. E l e v a t i o n Male-only groups p r e f e r r e d a l p i n e (above t r e e l i n e ) areas d u r i n g January and the subalpine (below t r e e l i n e ) zone from March through May. Mixed-female groups e x h i b i t e d s i m i l a r p r e f e r e n c e s d u r i n g January and May, 1977, however, from February through A p r i l , they u t i l i z e d both zones i n equal p r o p o r t i o n to t h e i r a v a i l a b i l i t y . In other months, not represented by male-only groups, mixed-female groups g e n e r a l l y p r e f e r r e d the a l p i n e zone. In a more d e t a i l e d s e l e c t i o n a n a l y s i s of mountain goat a l t i t u d i n a l use (Table 6), mixed-female groups were more s e l e c t i v e f o r c e r t a i n 250-m e l e v a t i o n bands than were the male-only groups. D i s p r o p o r t i o n a t e use of e l e v a t i o n by the male groups was evident only i n May, 1978, when the subalpine (751-1250 m) i n t e r v a l was p r e f e r r e d . Mixed-female groups, however, d i s p l a y e d s e l e c t i o n i n nine of the t h i r t e e n months ana l y s e d . In November, when male-only groups d i d not s e l e c t f o r any p a r t i c u l a r e l e v a t i o n , mixed-female groups p r e f e r r e d e l e v a t i o n s about or j u s t below t r e e l i n e (751-1000 m) and avoided higher e l e v a t i o n s . In winter (January and March), female groups d i s p l a y e d a tendency to occupy a l p i n e areas near t r e e l i n e and avoided subalpine areas i n January, whereas the male-only groups showed no p r e f e r e n c e . Female groups e x h i b i t e d a high l e v e l of s e l e c t i o n i n May, 1977 but 91 T a b l e 6 - M o u n t a i n g o a t s e l e c t i o n o f 250-m a l t i t u d i n a l z o n e s on Maroon M o u n t a i n ( s o u t h s l o p e ) by m a l e - o n l y a n d m i x e d -f e m a l e g r o u p s . M a l e - o n l y Group S e l e c t i o n 1 Log Number S u b a l p i n e A l p i n e L i k e l i h o o d of — •••• 1 • 1 . . R a t i o Goat a 164-250m 251-500m 501-7 50m 7 51- 1000m 1001- 1250m 1251-150 0m 150I-17 50tr, 17 51 -2000m 2 001-206 6rr. Oct m m m m m m m m m m m Nov 14 . 3419 18 (0) (0) 0 0 0 (0) 10) 0 (0) Dec m m w> m m m m m m m m 1978 Jan 11. .4149 12 10) (0) (0) (0) 0 0 0 0 10} Feb 7. .7847 40 (0) 0 0 0 0 0 0 0 10) Mar 12. 7B37 41 (0) 0 0 0 0 0 (0) 0 (0) Apr 11. 8216 38 (0) 0 0 0 0 0 0 0 (0) May 17. 0195* 42 (0) 0 0 + 0 0 0 (0) (0) Aug m m m tn m m m m m m m Mixed-female Group S e l e c t i o n 1 Date Log L i k e l i h o o d R a t i o Number of Goats S u b a l p i n e A l p i ne 164-250m 251-500m 501-75Om 751-1000m 1001-1250m 1251-1500m 1501-1750m 1751-2000m 2001-2066n 1977 flay f>b. 10M* 151 (0) ( -) - 0 0 • - - (0) Ju.» 9.B340 23 J (0) (0) 0 0 0 0 0 0 (0) J u t m m m m m m m m m m m Aug 281.8228* 217 (0) <-> (-) I-) - - • 0 (0) Sop 31.8140* 179 (0) (-) I-) - - • <-) (0) Oct 66.3701* 106 (0) (0) 1-) - • - (-} (0) Nov 51.6216* 154 (0) 10) 0 + 0 - (-) (0) Dec m m m n m m m m m m 1978 Jan 97.7725* 128 10) 10) - - (-) • • 0 (0) Feb 9.3682 172 (0) 0 0 0 0 0 0 0 (0) Mar 19.8135* 1 59 (0) 0 0 0 0 • 0 0 10) Apr 16.7046* 118 (0) (0) 0 0 0 0 0 0 (0) May 5.0640 164 (0) 0 0 0 0 0 (0) (0) (0) Jun 3.3667 164 10) 0 0 0 0 0 0 0 (0) J u l 27.2134* 80 (0) (0) <-> - (-) (-) 0 • (0) Aug a m ra m m m m m B in n '+' > preferred, or used in greater proportion than available] '0' • no selection, or used in equal proportion to availability) '-' - avoided, or used in leaa proportion than available. Bracketed syabols indicate no use. m - non-representative data (i.e. less than 50% of the estimate sex component in tha population) * significant at p<0.05 92 t o t a l i n d i f f e r e n c e d u r i n g May, 1978. They a l s o showed i n d i f f e r e n c e s to use of e l e v a t i o n zones d u r i n g other months of the s p r i n g ( i . e . A p r i l and June, 1978), and tended to p r e f e r higher e l e v a t i o n s (>1250 m) d u r i n g the summer p e r i o d . 2.2.4 Mountain Goat A c t i v i t y Within P r e f e r r e d H a b i t a t s a. O v e r a l l A c t i v i t y P a t t e r n s The most f r e q u e n t l y observed mountain goat a c t i v i t i e s were feeding (48.8%) and r e s t i n g (43.0%). T r a v e l comprised only 8.3 percent of the o b s e r v a t i o n s . P r o p o r t i o n s of feeding and r e s t i n g d i d not vary s i g n i f i c a n t l y by month (G=5.1;df=5). A c t i v i t y p a t t e r n s v a r i e d , however, by group type. Male-only groups tended to r e s t (48.4%) and t r a v e l (11.8%) more than mixed-female groups (resting=42.1%; t r a v e l = 8 . 2 % ) ; thus i t f o l l o w e d that male-only groups fed (39.9%) l e s s than female groups (49.7%). When the p r o p o r t i o n s of f e e d i n g were assessed by group type over the same p e r i o d ( i . e . November, 1977 and January to May, 1978), no s i g n i f i c a n t d i f f e r e n c e s by month were observed (males: G=8.7;df=5; females: G=1.8;df=5), however, p r o p o r t i o n s of mixed-female group a c t i v i t i e s d i d a l t e r s i g n i f i c a n t l y d u r i n g other times of the year (G=87.0;df=13). 93 b. A c t i v i t y P a t t e r n s Within P r e f e r r e d H a b i t a t s Data were i n s u f f i c i e n t to s t a t i s t i c a l l y compare a c t i v i t y p a t t e r n s of both male-only and mixed-female groups f o r each h a b i t a t category, on a monthly b a s i s , t h e r e f o r e , d e s c r i p t i v e s t a t i s t i c s (% use) only were used to d e s c r i b e p r e f e r r e d h a b i t a t s . The p r o p o r t i o n s of r e s t i n g , f e e d i n g , and t r a v e l e x h i b i t e d by male-only groups were s i m i l a r i n a l l h a b i t a t s , however there was a tendency f o r males to r e s t more i n p r e f e r r e d h a b i t a t s . R e s t i n g predominated (x=60.0%), followed by fe e d i n g (x=31.1%) and t r a v e l (x=8.9%). For mixed-female groups, however, most p r e f e r r e d h a b i t a t c a t e g o r i e s (54.2%) were used p r i m a r i l y f o r feeding (x=57.0%; r e s t i n g , x=37.5%; t r a v e l , x=5.5%), while others (41.7%) were used mainly f o r r e s t i n g (x=52.3%; fe e d i n g , x=36.5%; t r a v e l , x=11.2%). The most p r e f e r r e d h a b i t a t s were f e e d i n g areas o c c u r r i n g mainly i n s o u t h - f a c i n g , deciduous shrublands between 1001 and 1500 m, on s l o p e s of 38 to 41°. Other p r e f e r r e d h a b i t a t s were s c a r c e l y vegetated rock areas, and e l e v a t i o n s between 750-1000 and 1501-1750 m (Table 7 ) . The most p r e f e r r e d h a b i t a t s where r e s t i n g mainly o c c u r r e d were s c a r c e l y vegetated rock areas with a d i r e c t southern exposure. In June, 1977, these h a b i t a t s o c c u r r e d on slo p e s between 35 and 41°, and i n J u l y , 1978, these h a b i t a t s were l o c a t e d w i t h i n the 1751 to 2000 m e l e v a t i o n i n t e r v a l . Other p r e f e r r e d 94 T a b l e 7 - P r e f e r r e d m o n t h l y h a b i t a t s u s e d p r i m a r i l y f o r f e e d i n g and r e s t i n g by m i x e d - f e m a l e g r o u p s on Maroon M o u n t a i n B . C . Prcdomi nant A c t i v i t y Feeding Pl a n t Cover Aspect Slope E l e v a t i o n P l a n t Cover Aspect Slope ( ° ) (in) ( ° ) E l e v a t i o n (m) 1977 May Deciduous South 38-41 Shrubland S c a r c e l y Veg. Rock S c a r c e l y Veg. Rock South 35-37 38-41 1251-1500 J u l Aug S c a r c e l y Veg. Sock Sep Oct Deciduous Shrubland 1001-1250 1251-1500 S c a r c e l y Veg. Rock Deciduous Shrubland 1001-1250 Dec 1976 Jan 38-41 1251-1500 1501-1750 Apr May S c a r c e l y Veg. Rock Deciduous Shrubland 38-41 1251-1500 S c a r c e l y Veg. Rock S c a r c e l y Veg. Rock Aug S c a r c e l y Veg. Rock S c a r c e l y Veg. Rock 95 h a b i t a t s were deciduous shrublands and e l e v a t i o n s between 1000 and 1500 m (Table 7). 96 V. DISCUSSION 1 . HABITAT CHARACTERISTICS Most mountain goat s t u d i e s , i n c l u d i n g t h i s one, were based on d i r e c t o b s e r v a t i o n of the animals, p r i m a r i l y because t h i s method i s c o s t - e f f e c t i v e and causes minimum d i s t u r b a n c e to the animals. The i m p l i c a t i o n s of o b s e r v a t i o n a l techniques are c o n s i d e r e d where p o s s i b l e i n the i n t e r p r e t a t i o n of the data, and are more s p e c i f i c a l l y addressed i n S e c t i o n 2. 1.1 H a b i t a t Use S e v e r a l r e s e a r c h e r s have d e s c r i b e d mountain goat h a b i t a t use (e.g. H j e l j o r d 1971, Rideout 1974, McFetridge 1977, Smith 1976, B u r l e i g h 1978, Fox 1978, Thompson 1981, Schoen et a l . 1981), however, each view the term ' h a b i t a t ' to c o n s i s t of a d i f f e r e n t set of environmental v a r i a b l e s or v a r i a b l e c a t e g o r i e s . U n f o r t u n a t e l y , no s t a n d a r d i z e d g u i d e l i n e s have been followed which allow comparison of h a b i t a t v a r i a b l e s ( i n c l u d i n g boundaries f o r t h e i r d e l i n e a t i o n ) . In an attempt to overcome t h i s major problem, some (e.g. B u r l e i g h 1978, Shea 1980) have r e c e n t l y adopted r e g i o n a l l y s p e c i f i c h a b i t a t c l a s s i f i c a t i o n s used by the United S t a t e s Department of A g r i c u l t u r e . 1.1.1 U n i v a r i a t e A n a l y s i s Mountain goat h a b i t a t r e l a t i o n s h i p s have been i n v e s t i g a t e d mainly by u n i v a r i a t e a n a l y s i s of use i n f o r m a t i o n . The most common v a r i a b l e s addressed have been p l a n t cover, aspect, s l o p e , and e l e v a t i o n , with d i s t a n c e to escape t e r r a i n becoming 97 i n c r e a s i n g l y c o n s i d e r e d . S t i l l other f a c t o r s , such as i n t e r s p e c i f i c and i n t r a s p e c i f i c c o m p e t i t i o n and c l i m a t e are a l s o c o n s i d e r e d important. These l a t t e r f a c t o r s are addressed i n c i d e n t a l l y i n t h i s s e c t i o n but are d i s c u s s e d more f u l l y i n S e c t i o n 2.)1 a. V e g e t a t i v e Cover Mountain goats u t i l i z e a v a r i e t y of p l a n t formations throughout the year, due to t h e i r annual v e r t i c a l movements between a l p i n e and subalpine areas ( K l e i n 1953). Heterogeneity of goat ranges was f i r s t q u a n t i f i e d ( s u b j e c t i v e l y ) by Smith (1981) i n t o c a t e g o r i e s of low, medium and h i g h . Nearly 60% of h i s o b s e r v a t i o n s o c c u r r e d i n areas with high p l a n t v a r i a b i l i t y , and o f t e n i n a s s o c i a t i o n with old-growth f o r e s t . Smith (1976) e a r l i e r concluded that mountain goat use of v e g e t a t i v e types was most evenly d i s t r i b u t e d i n November, May and June, when snow depths i n f l u e n c e d goat movements the most ( f a l l descent and s p r i n g a s c e n t ) . The r e s u l t s from my study are comparable with Smith's (1976) data; October, May and June r e c e i v e d the most evenly d i s t r i b u t e d use of p l a n t cover types. In many s t u d i e s , however, mountain goat use of v e g e t a t i v e cover appears to be p o s i t i v e l y a s s o c i a t e d with more open h a b i t a t s . For example, most s t u d i e s , i n c l u d i n g t h i s , d e s c r i b e high l e v e l s of mountain goat use of a l p i n e tundra, broken rock and boulder f i e l d s 98 (Hibbs 1965, Chadwick 1973, Rideout 1974, McFetridge 1977, Adams 1981, Thompson 1981). Although mountain goats have been observed to use mature timber and shrubby areas i n a number of s t u d i e s as w e l l (e.g. Hebert 1967, Kuck 1973, Singer 1975, Smith 1976, McFetridge 1977, B u r l e i g h 1978), the authors d i d not conclude that these cover types were h e a v i l y u t i l i z e d nor e s s e n t i a l to mountain goats. Rideout (1974) found s i g n i f i c a n t use of f o r e s t e d areas using t e l e m e t r y , but only r e c e n t l y have mature c o n i f e r o u s f o r e s t s been re c o g n i z e d as an important component of mountain goat h a b i t a t (Shea 1980, Smith 1981, Fox and Raedeke 1982). In t h i s study, mountain goats were b e l i e v e d to use f o r e s t e d areas to a v o i d b i t i n g i n s e c t s and d i r e c t sun d u r i n g the summer, and c o l d temperatures and d r i v i n g winds i n the win t e r . Snow depth, hunting, and s o c i a l behaviour may a l s o r e s u l t i n g r e a t e r use of timbered areas by mountain goats (see S e c t i o n 2 ) . Deciduous shrub areas were c o n s i d e r e d by Shea (1980) to c o n s t i t u t e important f o r a g i n g areas f o r mountain goats d u r i n g the s p r i n g . Shrub areas ( i n c l u d i n g shrublands and t h i c k e t s ) c o n s t i t u t e d the second-most important p l a n t cover type i n t h i s study, and were o f t e n used f o r bedding as w e l l as f o r f o r a g i n g (see Table 7). Mountain goats were commonly seen to stand on t h e i r h i n d l e g s to eat a l d e r l e a v e s on high shrubs. 99 Burned areas were a l s o noted to be favoured by mountain goats (McFetridge 1977, F o s t e r and Rahs 1981), however t h e i r use appeared to drop with i n c r e a s i n g d i s t a n c e from escape t e r r a i n . b. Aspect Peck (1972), Rideout (1974), Smith (1976), Stevens (1979) and Thompson (1981) concurred that s o u t h - f a c i n g a s p e c t s were favoured by mountain goats and that i n c r e a s e d use of t h i s aspect appeared to be a f u n c t i o n of i n c r e a s i n g snow depth on other a s p e c t s . The c o n c l u s i o n s from t h i s study are s i m i l a r . I t should be p o i n t e d out, however, that numerous windswept, snow f r e e areas occurred on the tops of a l p i n e k n o l l s and r i d g e s on the north aspect of Maroon Mountain, but i n t e n s e n o r t h e r l y winds, h i g h s n o w - d r i f t b u i l d up i n l o w - l y i n g areas and l a c k of cover p r e c l u d e d use of these areas by mountain goats d u r i n g w i n t e r . Southern exposures r e c e i v e d more i n s o l a t i o n , were warmer, and had a snowline higher i n e l e v a t i o n , and l e s s accumulated snow than any other exposure, which al l o w s l e s s r e s t r i c t e d t r a v e l and enables forage to be more r e a d i l y a v a i l a b l e d u r i n g winter and s p r i n g (Schoen et a l . 1981). N i c h o l s (pers. comm.), however, s t a t e d t h at t y p i c a l l y deep autumn s n o w f a l l s i n Alaska c o u l d i s o l a t e mountain goats wherever they happened to occur at the time, f o r the e n t i r e w i n t e r . N o r t h e r l y a s p e c t s were g e n e r a l l y c o n s i d e r e d by 100 Rideout (1974), Smith (1976) and Thompson (1981) to be used p r i m a r i l y d u r i n g summer, f o r thermoregulatory c o n t r o l and a c c e s s i b i l i t y of new p l a n t growth. During h i g h summer temperatures, Stevens (1979) observed 80% of the goats i n her study area on n o r t h - f a c i n g a s p e c t s and found that they u t i l i z e d southern exposures d u r i n g c o o l e r temperatures i n summer. In c o n t r a s t , Chadwick (1973) and Kuck (1973) found l i t t l e d i s p r o p o r t i o n a t e use of exposure by mountain goats d u r i n g summer, which may have been due to the d i s t r i b u t i o n of s u i t a b l e h a b i t a t . During t h i s study, a higher i n c i d e n c e of goat use of northern aspects on Maroon Mountain o c c u r r e d only d u r i n g the summer of 1976, when u n l i k e subsequent y e a r s , l a r g e p r e d a t o r s were absent. However, on Mount Garland, to the south, mountain goats were f r e q u e n t l y observed on north and e a s t e r l y - f a c i n g s l o p e s throughout the year. c. Slope Surface i n c l i n a t i o n i s reco g n i z e d as an important component of mountain goat h a b i t a t (Fox 1980, Schoen et a l . 1981, Fox and Raedeke 1982). The use of h i l l s i d e slope by mountain goats v a r i e s c o n s i d e r a b l y between s t u d i e s (see McFetridge 1977, Fox 1978, Stevens 1979). U n f o r t u n a t e l y , few r e s e a r c h e r s u t i l i z e d standard slope c l a s s e s , h i n d e r i n g a c c u r a t e comparisons. 101 On Maroon Mountain, l e s s than 5% of the goats were observed on s l o p e s g r e a t e r than 45°, i n part because of the l i m i t a t i o n s i n d i s c r i m i n a t i o n of the mapping method (<91 v e r t i c a l meters) f o r e s t i m a t i n g s l o p e . Although other work shows s i m i l a r slope use, v a r i a t i o n does occur. For example, Smith (1976) c o n s i s t e n t l y observed mountain goats on slopes g r e a t e r than 45° d u r i n g winter and s p r i n g , and on s l o p e s l e s s than 45° i n summer and f a l l . In c o n t r a s t , Thompson (1981) observed 70% of h i s summer goat s i g h t i n g s on sl o p e s l e s s than 15°, however, goat use was a f f e c t e d by a r t i f i c i a l s a l t l i c k s . R ecently, Schoen et a l . (1981) and Fox and Raedeke (1982) have suggested that d i s t a n c e to the nearest c l i f f ( i . e . escape t e r r a i n ) was the s i n g l e -most important a t t r i b u t e d e f i n i n g mountain goat h a b i t a t throughout the year. T h i s v a r i a b l e was not examined i n t h i s study because the e n t i r e south slope of Maroon Mountain was c o n s i d e r e d to c o n s t i t u t e escape t e r r a i n . C e r t a i n l y no animal c o u l d have been more than 200 m from the nearest c l i f f face i n t h i s a r e a. Steep h i l l s i d e s o f f e r both p r o t e c t i o n from p r e d a t o r s and w i n t e r i n g forage areas. In t h i s study, more than 80% of the o b s e r v a t i o n s each month c o n s i s t e n t l y o c c u r r e d on s l o p e s between 30 and 45°, except d u r i n g J u l y and August, 1978, when g e n t l e r s l o p e s were used p r o p o r t i o n a t e l y more. Although some 1 02 use of more g e n t l y - s l o p i n g areas was documented in the summer of 1977 and 1978, i t was apparent that wolves (Canis lupus) i n h i b i t e d mountain goat use of such areas i n these y e a r s , i n c o n t r a s t to 1976 when no l a r g e p r e d a t o r s were observed and the more g e n t l y -s l o p i n g areas were more commonly used. S i m i l a r responses i n range u t i l i z a t i o n by D a l l ' s sheep (Ovis  d a l l i d a l l i ) were observed by Murie (1944) i n the presence and absence of wolves. Fox (1980) c o n s i d e r e d predator avoidance to be more i n f l u e n t i a l than even forage a v a i l a b i l i t y f o r mountain goat h a b i t a t use. In t h i s study, use of steep areas on key ranges o c c u r r e d mainly d u r i n g l a t e hunting season (October) and may have been a response to gunshots and i n c r e a s e d v e h i c u l a r a c t i v i t y a long the v a l l e y - b o t t o m . Use of steeper areas was a l s o observed d u r i n g the rut (November), p o s s i b l y i n response to l o g g i n g o p e r a t i o n s a l s o i n the v a l l e y - b o t t o m . F a l l i n g t r e e s sounded remarkably l i k e r i f l e shots when they crashed onto other f a l l e n t r e e s d u r i n g the c o l d weather. Both sounds of g u n f i r e and f a l l i n g t r e e s were observed to c r e a t e alarm and d i s t r a c t i o n i n both males and females d u r i n g t h e i r c o u r t s h i p behaviour. Responses to these s t i m u l i were even g r e a t e r than to l a t e winter and s p r i n g avalanches. Smith (1976) and Kuck (1977) i n d i c a t e d that snow-shedding c h a r a c t e r i s t i c s of steep s l o p e s were the 103 primary determinant f o r u t i l i z a t i o n of winter ranges by mountain goats. Adams (1981) found that goats u t i l i z e d steeper areas d u r i n g winters with g r e a t e r s n o w f a l l . P e r l a and M a r t i n e l l i (1978) summarized the occurrence of major s l a b avalanches and found that 96% o c c u r r e d on i n c l i n a t i o n s between 35 and 45°. T h i s corresponds with the observed winter use of slope c a t e g o r i e s i n t h i s study (84% between 30-45°). However, more than 80% of a l l goats observed o c c u r r e d on i n c l i n e s between 30 and 45° slope year-round ( s p r i n g , summer and f a l l of 1977, and w i n t e r , 1978). Although steep snow-free s l o p e s may be f r e q u e n t l y used i n winter, there was no evidence from t h i s study to suggest that these areas were used d i s p r o p o r t i o n a t e l y more than any other time of the year. I t t h e r e f o r e seemed l i k e l y , as Schoen et a l . (1981) suggested, that the primary f u n c t i o n of steep t e r r a i n was to serve as an i n n a c c e s s i b l e r e t r e a t from p r e d a t o r s . d. E l e v a t i o n Many r e s e a r c h e r s d e s c r i b e v a r i a t i o n i n a l t i t u d i n a l use e x h i b i t e d by mountain goats between summer and winter, but few have q u a n t i f i e d these v e r t i c a l movements. Some (e.g. Rideout 1974, Smith 1976) have observed a p o s i t i v e a s s o c i a t i o n between mountain goat d i s t r i b u t i o n s and annual snow p a t t e r n s , while others (e.g. Thompson 1981) i n f e r r e d that summer use of high e l e v a t i o n s was r e l a t e d to p l a n t phenology. 104 Smith (1978) summarized a l t i t u d i n a l s i g h t i n g s of goats by month and found t h a t from December to May, they used low e l e v a t i o n s , and from June to November, they occupied higher a l t i t u d e s . S i m i l a r o b s e r v a t i o n s have been made by o t h e r s (e.g. Anderson 1940, Brandborg 1955, Holroyd 1967, Chadwick 1973, Kuck 1973, Rideout 1974). However, Casebeer e t a l . (1950), L e n t f e r (1955), Chadwick (1973) and Vaughan (1975) f r e q u e n t l y observed goats at low e l e v a t i o n s and on windblown r i d g e - t o p s d u r i n g w i n t e r . Vaughan (1975) noted t h a t most goats i n h i s "study area occupied windblown r i d g e - t o p s at very high a l t i t u d e s , due to few, s o u t h - f a c i n g c l i f f s at lower e l e v a t i o n s , however, Hebert and T u r n b u l l (1977) determined that i n c r e a s e d snow d e n s i t y (wetness) determined s u b a l p i n e w i n t e r i n g l o c a t i o n s of goats. In t h i s study, while mountain goats were widely d i s t r i b u t e d a l t i t u d i n a l l y throughout the year, two prominent p a t t e r n s of v e r t i c a l ascent and descent were apparent over the twelve month p e r i o d . A t y p i c a l summer ascent (May to August) o c c u r r e d and was b e l i e v e d t o be c o r r e l a t e d t o r e c e d i n g snowline and the p a t t e r n of p l a n t growth. A f a l l descent ( l a t e September to November) appeared to be a f u n c t i o n of i n c r e a s i n g s n o w f a l l and st r o n g n o r t h e r l y winds, p a r t i c u l a r l y i n November. The l o c a t i o n of k i d d i n g and 105 r u t t i n g a c t i v i t i e s a l s o seemed to be a f u n c t i o n of snow d i s t r i b u t i o n and concomitant l o c a t i o n of a d u l t females. In January, a l p i n e areas were h e a v i l y used, due to the a v a i l a b i l i t y of wind-blown, snow-free areas on exposed s l o p e s . Subalpine e l e v a t i o n s were again used more as snow accumulated over the winter and then d u r i n g s p r i n g 'green-up', p r i o r to the l a t e s p r i n g e a r l y summer ascent t o a l p i n e areas. Another p o s s i b l e e x p l a n a t i o n f o r the descent d u r i n g l a t e December may have been that the animals were too warm i n the a l p i n e thermal i n v e r s i o n zone with t h e i r long winter pelage. I t i s not d e f i n i t e l y known i f use of lower e l e v a t i o n s , such as l o g g i n g c l e a r c u t s and some rocky outcrops along Maroon Cr-eek, would have o c c u r r e d i n the absence of human a c t i v i t y i n the v a l l e y - b o t t o m . Assessment of monthly v e r t i c a l movement allowed documentation of only broad c y c l e s of a l t i t u d i n a l use, and these o c c u r r e d mainly d u r i n g the w i n t e r . At t h i s time, mountain goats were o f t e n observed i n the most rugged and barren a l p i n e h a b i t a t s imaginable. These areas seemed to be u n f i t f o r any animal to l i v e out the winter. However, more i n t e n s i v e winter o b s e r v a t i o n s showed that c y c l i c a l a l t i t u d i n a l p a t t e r n s were e x h i b i t e d by many mountain goats every 3 to 10 days. T h i s frequency range appeared c o r r e l a t e d with the occurrence of high pressure systems. I f s u i t a b l e windswept areas were a v a i l a b l e i n the a l p i n e , goats 106 would sometimes stay three or four days, u n t i l the onset of the next low p r e s s u r e d i s t u r b a n c e . If the goats, on moving up to the a l p i n e , c o u l d not f i n d s u i t a b l e f o r a g i n g areas, they u s u a l l y r e t u r n e d to t r e e l i n e or subalpine areas the very same or next day as t h e i r i n i t i a l upward movement. 1.1.2 M u l t i v a r i a t e A n a l y s i s Four s t u d i e s of mountain goat h a b i t a t use have been conducted to date, i n which m u l t i v a r i a t e a n a l y s e s have been employed. As w i l l be seen i n the d i s c u s s i o n below, they vary i n t h e i r f i n d i n g s , and much of t h i s v a r i a t i o n appears due to the f a c t t h at they have i n c l u d e d d i f f e r e n t v a r i a b l e s i n t h e i r models, as much as to the f a c t that they were c a r r i e d out i n q u i t e d i f f e r e n t study areas. Schoen et a l . (1981) used three i n t e r v a l - l e v e l h a b i t a t v a r i a b l e s ( e l e v a t i o n , s l o p e , and d i s t a n c e to escape t e r r a i n ) to study mountain goat h a b i t a t u t i l i z a t i o n i n A l a s k a , by d i s c r i m i n a n t f u n c t i o n a n a l y s i s . They i d e n t i f i e d one v a r i a b l e which c o u l d account f o r 100% of the v a r i a t i o n i n goat s i g h t i n g s . T h i s was d i s t a n c e to the nearest c l i f f . In a p r e l i m i n a r y t e s t of t h e i r model, 100% of the independent o b s e r v a t i o n s were a c c u r a t e l y p r e d i c t e d . But, i n another Alaskan study, Fox and Raedeke (1982) showed that a g r e a t e r number of v a r i a b l e s were important. In t h i s case, four v a r i a b l e s ( e l e v a t i o n , timber volume, sl o p e , and d i s t a n c e from c l i f f s ) accounted f o r 74% of the goat o b s e r v a t i o n s and 100% of the areas not i n h a b i t e d by mountain goats. However, 1 07 n o m i n a l l y - s c a l e d v a r i a b l e s (e.g. aspect, p l a n t cover, landform, e t c . ) were not assessed i n e i t h e r study due to r e s t r i c t i o n s of the s t a t i s t i c a l a n a l y s i s . I t i s b e l i e v e d that i n c l u s i o n of nominal v a r i a b l e s may enhance the model's power, although a d i f f e r e n t a n a l y s i s technique i s r e q u i r e d . McFetridge (1977) conducted a m u l t i v a r i a t e a n a l y s i s of n o m i n a l l y - s c a l e d a t t r i b u t e data on mountain goat h a b i t a t r e l a t i o n s h i p s i n A l b e r t a , between May and October o n l y . T h i s a n a l y s i s c o n s i d e r e d p l a n t cover, a s p e c t , s l o p e , e l e v a t i o n and d i s t a n c e to escape t e r r a i n . McFetridge (1977) concluded that the most s i g n i f i c a n t r e l a t i o n s h i p of use o c c u r r e d with a l l of the v a r i a b l e s i n c l u d e d i n the model, and was a p p l i c a b l e mostly to nursery groups, r a t h e r than a l l goat o b s e r v a t i o n s . His August data y i e l d e d the best r e l a t i o n s h i p s , although the 'di s t a n c e to r o c k - g r a v e l ' v a r i a b l e d i d not have a s i g n i f i c a n t e f f e c t on h a b i t a t r e l a t i o n s h i p s . T h i s v a r i a b l e c o n s i s t e d of s u r f i c i a l rock, g r a v e l , shale and s o i l , and was noted to be u s u a l l y steep and un s t a b l e , however, i t d i d not c o n s t i t u t e c l i f f - l i k e escape t e r r a i n . Shea (1980) developed mountain goat h a b i t a t s u i t a b i l i t y index models f o r s p r i n g , summer and winter ranges i n Alaska and c o n s i d e r e d aspect, p l a n t cover and d i s t a n c e to escape t e r r a i n as the most important c r i t e r i a i n p r e d i c t i n g seasonal range use. However, the models have not been t e s t e d . 108 1.2 H a b i t a t S e l e c t i o n The use of h a b i t a t i s a f f e c t e d by numerous s o c i a l and environmental f a c t o r s , t h e r e f o r e one would expect temporally d i s p r o p o r t i o n a t e use of environmental a t t r i b u t e s , with respect to h a b i t a t a v a i l a b i l i t y . However, the i m p l i c a t i o n s of such d i s p r o p o r t i o n a t e use o f t e n c r e a t e s a cloudy i s s u e . T h i s may be due i n l a r g e p a r t , to s e v e r a l c onnotations of the terms 'preference' and ' s e l e c t i o n ' which have been abused and used m i s l e a d i n g l y by v a r i o u s a u t h o r s . For example, McCrory et a l . (1977) and B u r l e i g h (1978) d i s c u s s e d ' h a b i t a t s e l e c t i o n ' from a p u r e l y s u b j e c t i v e assessment. H j e l j o r d (1971) and ot h e r s r e f e r r e d to ' h a b i t a t p r e f e r e n c e ' based only on the measurement of usage. In each case, the t e r m i n o l o g i e s used misrepresented the comparisons undertaken, even i n the q u a n t i f i c a t i o n of usage. One of the f i r s t mountain goat s t u d i e s e x t e n s i v e l y a d d r e s s i n g the concepts of use and a v a i l a b i l i t y was conducted by Adams (1981). However, h i s comparisons were f o r the summer p e r i o d o n l y , and were based on the forage r a t i o (Stoddart e t a l . 1975) which excluded any form of s t a t i s t i c a l c o n s i d e r a t i o n . Thompson (1981), Schoen et a l . (1981) and Fox and Raedeke (1982) d e s c r i b e d mountain goat h a b i t a t u t i l i z a t i o n by contingency a n a l y s i s of use and a v a i l a b i l i t y . However, few authors have shown complete p r o c e d u r a l documentation and c o n s i d e r a t i o n of sampling requirements. Thompson (1981) and Fox and Raedeke (1982) only p a r t i a l l y o u t l i n e d t h e i r methodology f o r c a l c u l a t i n g h a b i t a t a v a i l a b i l i t y , i n c l u d i n g d e l i n e a t i o n of t h e i r boundaries. Only Thompson (1981) c o r r e c t e d a v a i l a b i l i t y data, obtained by 109 v e r t i c a l map p r o j e c t i o n , f o r h i l l s i d e s l o p e . None of the authors a p p a r e n t l y c o l l e c t e d t h e i r data i n a systematic f a s h i o n , thus r a i s i n g the q u e s t i o n of i t s v a l i d i t y . 1.2.1 E c o l o g i c a l F i n d i n g s The l e v e l of h a b i t a t s e l e c t i o n by the two groups being compared was c o n s i d e r e d to be the sum of t h e i r p r e f e r e n c e s and avoidances. T h e r e f o r e , the group type with the l e a s t number of h a b i t a t s used i n equal p r o p o r t i o n to t h e i r a v a i l a b l i l i t y was c o n s i d e r e d to be 'more s e l e c t i v e ' . I t i s l e f t open to d i s c u s s i o n whether one group type i s more (or l e s s ) s e l e c t i v e than the other group type, when, f o r example, group 'A' p r e f e r r e d 6 h a b i t a t s and avoided 2 of them, while group 'B' p r e f e r r e d 2 h a b i t a t s and avoided 6 of them. The a n a l y s i s used i n t h i s study confirmed p r e v i o u s suggestions t h a t : a) sexual d i f f e r e n c e s occurred i n the s e l e c t i o n of h a b i t a t by mountain goat group types, and b) mixed-female groups were more s p e c i a l i z e d , or s e l e c t i v e , than male-only groups. Fox and Raedeke (1982) d i d not c o n s i d e r the e f f e c t of sex d u r i n g t h e i r i n v e s t i g a t i o n , however Schoen et a l . (1981) concluded that s i g n i f i c a n t d i f f e r e n c e s i n h a b i t a t use o c c u r r e d between sexes f o r the g r e a t e r p o r t i o n of the v a r i a b l e s assessed. U n f o r t u n a t e l y , n e a r l y h a l f of the seasonal r e l a t i o n s h i p s examined in t h e i r a n a l y s i s , v i o l a t e d the c h i - s q u a r e requirements concerning expected f r e q u e n c i e s . They concluded, however, that d u r i n g winter, females were d i s t r i b u t e d at higher e l e v a t i o n s on steeper, more broken and r o c k i e r s l o p e s , c l o s e r to c l i f f s than 110 were males. In the present study, male-only groups s e l e c t e d f o r slope and aspect h a b i t a t c a t e g o r i e s s i m i l a r to those s e l e c t e d by mixed-female groups, but at d i s i m i l a r e l e v a t i o n s and i n d i f f e r e n t v e g e t a t i v e cover c a t e g o r i e s . Male-only groups s e l e c t e d f o r more dense v e g e t a t i v e cover i n s u b a l p i n e e l e v a t i o n s than d i d mixed-female groups. Sex d i f f e r e n c e s i n h a b i t a t s e l e c t i o n have a l s o been documented f o r example i n bighorn sheep (Ovis canadensis) (Shank 1979a, M a r t i n e l l i and Hudson 1981, T i l t o n and W i l l a r d 1982). Schoen et a l . (1981) hypothesized that females may be more s e l e c t i v e than males, choosing areas of g r e a t e r s e c u r i t y from p r e d a t i o n and areas of g r e a t e r forage a v a i l a b i l i t y . The f i n d i n g s from t h i s study support part of t h i s h y p o t h e s i s . Mixed-female groups were found more f r e q u e n t l y to s e l e c t steep, rocky, s o u t h - f a c i n g a l p i n e h i l l s i d e s d u r i n g summer and winter and s i m i l a r l y c h a r a c t e r i z e d s ubalpine h i l l s i d e s i n the f a l l , than were male-only groups. One would expect, a p r i o r i t h a t whatever c o n d i t i o n s were s e l e c t e d , food resources would be an important component in the s e l e c t i o n p r o c e s s , however, Kuck (1977) showed that mountain goat s e l e c t i o n f o r steep, rocky areas i n winter was not i n response to a b e t t e r food resource, but to snow-shedding c h a r a c t e r i s t i c s . 111 1.2.2 Concepts Of S e l e c t i o n S e l e c t i o n does not simply imply preference ( i . e . g r e a t e r use than a v a i l a b i l i t y ) , but a l s o i n c l u d e s avoidance responses. Although mixed-female groups s e l e c t e d v a r i o u s h a b i t a t a t t r i b u t e s throughout the year, they were a l s o much more s e l e c t i v e than males i n the sense that they avoided n e a r l y twice as many a v a i l a b l e h a b i t a t types than they p r e f e r r e d . The l e s s s e l e c t i v e ( i n both d i r e c t i o n s ) male-only groups, t h e r e f o r e may f i t i n t o the s t r a t e g y of ' g e n e r a l i s t s ' who can optimize t h e i r use of h a b i t a t c o n d i t i o n s by g e n e r a l l y u s i n g h a b i t a t types as they encounter them. Thus they show l i t t l e avoidance or p r e f e r e n c e f o r any p a r t i c u l a r h a b i t a t . a. Order of S e l e c t i o n If an a n a l y s i s of s e l e c t i o n i s to i n v o l v e examining the p r o p o r t i o n s of use r e l a t i v e t o a v a i l a b i l i t y , then i t i s e s s e n t i a l to determine what i s a v a i l a b l e to the animal. Johnson (1980) d e s c r i b e d four h y p o t h e t i c a l o r d e r s of s e l e c t i o n r e l a t i n g to an animal's p r e f e r e n c e s . He r e f e r s to f i r s t order s e l e c t i o n as the p h y s i c a l or g e o g r a p h i c a l range of a s p e c i e s , w i t h i n which second order s e l e c t i o n may determine the home range of an i n d i v i d u a l . T h i r d order s e l e c t i o n p e r t a i n s to the usage of v a r i o u s h a b i t a t components w i t h i n the home range, and f i n a l l y , f o u r t h order s e l e c t i o n can determine the procurement of food items from those a v a i l a b l e at t h a t s i t e . The concept of a v a i l a b i l i t y i s o f t e n q u i t e 1 12 a r b i t r a r y , based upon components deemed by the i n v e s t i g a t o r to be a v a i l a b l e to the animal. However, each order i s r e l a t e d to a higher order, not by the observer, but by the animal. For example, an animal's home range occurs i n a c e r t a i n l o c a t i o n because that animal has a l r e a d y 'decided' to l i v e i n that p a r t i c u l a r p a r t of i t s range d i s t r i b u t i o n . Consequently, c o n c l u s i o n s drawn from such a n a l y s e s may be i n c o n s i s t e n t . Johnson (1980) proposed a method of ranking a v a i l a b l e home range data, to e l i m i n a t e c l a s s i f i c a t i o n e f f e c t s , however no unbiased s t a t i s t i c a l method i s a v a i l a b l e f o r comparing usage data to constant p r o p o r t i o n s of a v a i l a b i l i t y , such as attempted i n t h i s study. b. I n t e r p r e t i n g S e l e c t i o n i . A r t e f a c t s of S e l e c t i o n If an animal i s not observed to occur w i t h i n c e r t a i n h a b i t a t s , then i t i s g e n e r a l l y assumed that these areas are avoided. However, n e i t h e r Marcum and Lof t s g a a r d e n ' s (1980) SCI technique nor E v e r i t t ' s (1977) a n a l y s i s of contingency r e s i d u a l s (used by Schoen at a l . 1981) f o r determining h a b i t a t s e l e c t i o n were s u f f i c i e n t l y s e n s i t i v e to c o n s i d e r zero use of small areas or p r o p o r t i o n s to be avoided. T h i r t y - f o u r percent of the a l l h a b i t a t s r e c e i v e d no use by mountain goats i n t h i s study, and of those, n e a r l y h a l f 113 (47%) were c l a s s i f i e d as avoided i n the SCI a n a l y s i s ; the remainder were c o n s i d e r e d to be used i n equal p r o p o r t i o n to t h e i r a v a i l a b i l i t y . T h i s problem i s not addressed i n other s t u d i e s , but i s c e r t a i n l y apparent (e.g. Schoen et a l . 1981). In a d d i t i o n to the problem of zero use, v a r i a t i o n s i n s e l e c t i o n between sexes impose f u r t h e r l i m i t a t i o n s . For example, given an imbalanced sex r a t i o of two females f o r every male, such as the Maroon Mountain p o p u l a t i o n , there was p r o p o r t i o n a t e l y l e s s chance f o r males to occupy as many h a b i t a t s as females because there are fewer of them. When the data were c o r r e c t e d , by i d e n t i f y i n g zero-use h a b i t a t s as avoided, before conducting the s e l e c t i o n a n a l y s i s , no change i n the r e l a t i v e d i f f e r e n c e s between the sexes r e s u l t e d . However, d i f f e r e n t i n t e r p r e t a t i o n s would have occ u r r e d between the two sexes. For example, males c o u l d have been mistakenly c o n s i d e r e d to be more s e l e c t i v e than females, because they avoided more h a b i t a t types, which was, i n a c t u a l f a c t , not the case. There appears to be no immediate s o l u t i o n to t h i s problem. Development of a new pre f e r e n c e index based upon p r o p o r t i o n s should a l l e v i a t e the problem of d i s p a r a t e sex r a t i o s . 1 1 4 A d d i t i o n a l l y , b i a s e s may have a r i s e n because there were fewer males than females. However, females would be expected to s e l e c t l e s s h a b i t a t s because they g e n e r a l l y o c c u r r e d i n l a r g e r groups, of which there were fewer i n number. T h i s confounding e f f e c t was not addressed i n the a n a l y s i s , but warrants f u t u r e c o n s i d e r a t i o n , i i . P r eference and Avoidance H a b i t a t s i d e n t i f i e d by s e l e c t i o n a n a l y s i s as e i t h e r p r e f e r r e d or avoided should not be taken b l i n d l y at face v a l u e , nor should h a b i t a t s used i n p r o p o r t i o n to t h e i r a v a i l a b i l i t y be ignored. Animals presumably make d e c i s i o n s to some extent as to the region where they r e s i d e , what h a b i t a t s they use, what p l a n t s they eat, e t c . , but whether these ' d e c i s i o n s ' are b i o l o g i c a l l y meaningful remains to be t e s t e d . An a n a l y s i s of s e l e c t i o n cannot d i c t a t e whether p r e f e r r e d areas are e s s e n t i a l and avoided areas unimportant or dangerous. For example, h a b i t a t A (e.g. a small grove of t r e e s ) may be more p r e f e r r e d than h a b i t a t B (e.g. a s p a r s e l y vegetated, rocky h i l l s i d e ) ; but h a b i t a t A may be used f o r comfort only (e.g. shade) and not e s s e n t i a l f o r s u r v i v a l , whereas h a b i t a t B i s b i o l o g i c a l l y e s s e n t i a l f o r food and escape t e r r a i n . Although mixed-female groups e x h i b i t e d more 1 15 i n s t a n c e s of s e l e c t i o n than males, t h i s does not n e c e s s a r i l y mean that females u t i l i z e d more 'optimal' areas than male-only groups, f o r the l a t t e r may be a b l e to e x p l o i t t h e i r environment more e f f i c i e n t l y as a g e n e r a l i s t . They a l s o have l e s s demands than the females (e.g. pregnancy, l a c t a t i o n ) and o f f s p r i n g (e.g. growth, development). 2. MOUNTAIN GOAT OBSERVABILITY Mountain goat s t u d i e s have g e n e r a l l y been o b s e r v a t i o n a l i n nature (e.g. DeBock 1970, Vaughan 1975, McFetridge 1977, Wright 1977, Stevens 1979, Adams 1981). A d d i t i o n a l l y , most s t u d i e s have implemented non-systematic sampling and have been conducted i n s e v e r a l b i o t o p e s , ranging from northern b o r e a l f o r e s t escarpments and subalpine canyons to t y p i c a l a l p i n e - s u b a l p i n e mountainous c o n d i t i o n s . Consequently, each p o p u l a t i o n s t u d i e d has the p o t e n t i a l to o f f e r unique v i s i b i l i t y b i a s e s , i n a d d i t i o n to s p e c i f i c c h a r a c t e r i s t i c s of p o p u l a t i o n demography and range u t i l i z a t i o n . I t i s c l e a r from t h i s study that o b s e r v a t i o n a l b i a s e s o c c u r r e d d u r i n g systematic sampling of the Maroon Mountain goat p o p u l a t i o n , which l i m i t the comparison and d i s c u s s i o n of p o p u l a t i o n and range c h a r a c t e r i s t i c s of t h i s herd with other mountain goat s t u d i e s . The l a r g e v a r i a t i o n i n sex and age r a t i o s shown by D r i v e r et a l . (1978), between two months of the same year, supports t h i s f i n d i n g . N o n - v i s i b l e p o r t i o n s of the 116 p o p u l a t i o n c o u l d have been accounted f o r by one or more of the f o l l o w i n g reasons: a) mountain goats c o u l d have occupied e q u a l l y v i s i b l e h a b i t a t s , s i m i l a r i n p r o p o r t i o n s to those sampled, but i n non-observable areas w i t h i n the study area (even on the south s l o p e , every square meter of h i l l s i d e was not v i s i b l e ) ; b) mountain goats c o u l d have been l o c a t e d i n more densely vegetated ( i . e . n o n - v i s i b l e ) areas w i t h i n the study area; or c) mountain goats may have emigrated from the study area, i n which case e i t h e r a) or b) c o u l d be t r u e . The f i r s t p o s s i b i l i t y was s u b s t a n t i a t e d o n l y i n 1976, and then by ad l i b , o b s e r v a t i o n s of mountain goats on Ridges #1 and #2, where both s o l i t a r y males and nursery groups were p e r i o d i c a l l y observed, d u r i n g the summer o n l y . However, mountain goats were not observed i n these areas d u r i n g the p e r i o d of systematic o b s e r v a t i o n s . During t h i s p e r i o d , the presence of wolves appeared to r e s t r i c t mountain goat range use south of the main r i d g e . The t h i r d p o s s i b i l i t y was a l s o r e j e c t e d ; f i r s t l y because i n t e n s i v e b e h a v i o u r a l o b s e r v a t i o n s (>1OOhrs/month) were conducted on the southern slope of Maroon Mountain, i n a d d i t i o n to the time spent d u r i n g v e h i c u l a r surveys, and no d i r e c t i o n a l movements of animals going over the main r i d g e or out along Ridge #3 were observed. However, ad l i b , o b s e r v a t i o n s in 1976 o n l y , d i d document l i m i t e d mountain goat use of Ridge #3 d u r i n g the summer. Secondly, surveys of adjacent areas ( i . e . areas south of the Hanging V a l l e y , Mount Garland and Wesach Mountain) d i d not show any n o t i c e a b l e i n f l u x i n numbers of animals d u r i n g 1 1 7 the study p e r i o d . T h i r d l y , ground t r u t h i n g showed no i n t e r -mountain movement. The only i n d i c a t i o n s of low e l e v a t i o n use by mountain goats were one set of t r a c k s on the Maroon Creek l o g g i n g road i n January, 1977, an a d u l t male w i n t e r i n g a c r o s s from the l o g g i n g road near the mouth of Maroon Creek v a l l e y i n 1978, and a dead male found i n a c l e a r c u t along Maroon Creek i n February, 1978. In each case, these animals or t h e i r s i g n , were observed w i t h i n the study area and i n months i n which the m a j o r i t y of the estimated p o p u l a t i o n was most v i s i b l e (see F i g u r e 9) . Consequently, b) , u t i l i z a t i o n of l e s s v i s i b l e areas w i t h i n the study area, i s l e f t as the only p o s s i b l e a l t e r n a t i v e . S e v e r a l i n t e r s p e c i f i c and i n t r a s p e c i f i c f a c t o r s , i n a d d i t i o n to weather, are b e l i e v e d to a f f e c t mountain goat o b s e r v a b i l i t y . 2.1 I n t r a s p e c i f i c I n f l u e n c e s 2.1.1 Soc i a l Behaviour Males accounted f o r much of the v a r i a t i o n i n mountain goat o b s e r v a b i l i t y due to s e x - s p e c i f i c d i f f e r e n c e s i n h a b i t a t s e l e c t i o n , r e s u l t i n g from female a g g r e s s i o n . Competitive e x c l u s i o n has been the most popular theory governing sexual d i f f e r e n c e s i n mountain goat h a b i t a t s e l e c t i o n (Chadwick 1973,1977, Kuck 1977, Smith 1976). I t was p o s t u l a t e d that sex-s p e c i f i c d i f f e r e n c e s i n h a b i t a t s e l e c t i o n would become more apparent with i n c r e a s i n g p o p u l a t i o n s i z e , because resources would become more l i m i t i n g . Under these c o n d i t i o n s , males should tend to u t i l i z e more timbered or brushy h a b i t a t s of 118 marginal value than females with young. Conversely, males may be expected to u t i l i z e s i m i l a r h a b i t a t s as females, i f the p o p u l a t i o n was d e c l i n i n g due to hunting, f o r example, in which case resource a v a i l a b i l i t y would not be a f f e c t e d . Other s o c i a l f a c t o r s a l s o a f f e c t o b s e r v a b i l i t y , as noted by B a i l e y and Johnson (1977). For example, males are g e n e r a l l y s o l i t a r y or occur i n s m a l l e r groups than females (Chadwick 1973, 1977), and t h e r e f o r e may be more e a s i l y missed by the observer. A d d i t i o n a l l y , males are more l i k e l y to be p l a c e d i n the ' u n c l a s s i f i e d ' category than females because fewer c h a r a c t e r i s t i c s are a v a i l a b l e to i d e n t i f y males i n comparison to females (e.g. maternal b e h a v i o u r ) . 2.1.2 A c t i v i t y P a t t e r n s In t h i s study, males were more l e t h a r g i c than females ( i . e . they r e s t e d more than they fed, d i u r n a l l y ) . Males were f r e q u e n t l y observed to bed f o r p e r i o d s of s e v e r a l hours, i n c o n s t r a s t to the two-hour c y c l i c a l p a t t e r n s d e s c r i b e d f o r females (Chadwick 1973). R e s t i n g animals were c o n s i d e r e d more d i f f i c u l t to see because of t h e i r reduced v i s i b l e s u r f a c e area and because they were s t a t i o n a r y . P a r t r i d g e (1979) s t a t e d that d i f f e r e n c e s i n t i m i n g of a c t i v i t y may c o n t r i b u t e to e c o l o g i c a l s e g r e g a t i o n . In mountain goats, Rideout (1974) documented that both sexes show a c t i v i t y peaks at dawn, i n the l a t e morning, and j u s t b efore s u n r i s e . However, males d i s p l a y e d l e s s a c t i v i t y i n the a f t e r n o o n , and an e a r l i e r peak i n the evening. Stevens (1979) observed a d u l t males n o c t u r n a l l y f e e d i n g , on a l e a s t two o c c a s i o n s . 119 McFetridge (1977), Fox (1978) and Stevens (1979) noted that mountain goats u t i l i z e d c e r t a i n h a b i t a t types f o r d i f f e r e n t purposes. I n a c t i v i t y i n densely vegetated h a b i t a t s poses a much more s e r i o u s o b s e r v a t i o n a l b i a s than i n a c t i v i t y i n s p a r s e l y vegetated a r e a s . 2.2 I n t e r s p e c i f i c I n f l u e n c e s 2.2.1 B i t i n g I n s e c t s B i t i n g i n s e c t s were f r e q u e n t l y observed to d i s t u r b m o l t i n g males and barren females d u r i n g the s p r i n g , and o f t e n s t i m u l a t e d t h e i r disappearance i n t o h e a v i l y timbered a r e a s , which were noted to support much sparser d e n s i t i e s of b i t i n g i n s e c t s . Thus, these animals would be l e s s v i s i b l e at t h i s time of year, whereas females accompanied by young, r e t a i n i n g p r o t e c t i v e winter c o a t s (Hebert 1967, McFetridge 1977), remained i n open h a b i t a t s . Because females with kids molted l a t e r i n summer, they were able to o b t a i n r e l i e f from b i t i n g i n s e c t s by occupying exposed, breezy s i t e s i n the a l p i n e , such as r i d g e - t o p s and small peaks. These h a b i t a t s were r e a d i l y v i s i b l e . 2.2.2 Hunting The observed number of a d u l t males/100 a d u l t females dropped i n August, September and October, a f t e r a steady i n c r e a s e from May. T h i s was observed to c o i n c i d e with the hunting season on Maroon Mountain. Although both sexes were open to hunting, males were g e n e r a l l y the most s o u g h t - a f t e r (however there were p r o p o r t i o n a t e l y l e s s of them s h o t ) . I t i s 1 20 b e l i e v e d that most of the males u t i l i z e d densely vegetated h a b i t a t s d u r i n g t h i s p e r i o d . Swenson (1982) s i m i l a r l y noted that mixed-grass p r a i r i e mule deer i n c r e a s e d t h e i r use of upland timbered cover types d u r i n g hunting season. In another study, Douglas (1971) monitored the response of chamois (Rupicapra  r u p i c a p r a ) and red deer (Cervus elaphus) to an i n t e n s i v e e r a d i c a t i o n campaign. S h o r t l y a f t e r the hunting of females ceased, i t was apparent that both sexes were exceedingly shy and wary, stayed c l o s e to f o r e s t cover and fed i n the open mainly at n i g h t . 2.3 Weather Fox (1978) concluded that the e f f e c t s of weather on mountain goat a c t i v i t y and h a b i t a t use c o u l d s i g n i f i c a n t l y i n f l u e n c e survey r e s u l t s . During t h i s study, i t was noted that hot summer temperatures induced mountain goats to use dense timber and shrub areas f o r p r o t e c t i o n from d i r e c t sun. Stevens (1979) noted t h a t the c o n i f e r o u s canopy was used almost e x c l u s i v e l y d u r i n g the h o t t e s t part of the summer. Stormy, winter weather was a l s o observed to s t i m u l a t e mountain goat use of densely vegetated areas, f o r p r o t e c t i o n from strong winds and d r i v i n g snow. Hebert and T u r n b u l l (1977) a t t r i b u t e d winter use of timbered areas i n c o a s t a l B r i t i s h Columbia to the c h a r a c t e r i s t i c dense snow-pack of the area, i n c o n t r a s t to i n t e r i o r areas, where dry, powder snows are f r e q u e n t l y blown f r e e from a l p i n e r i d g e s . S e v e r a l c l i m a t i c c o n d i t i o n s c r e a t e animal i n a c t i v i t y which a l s o reduces o b s e r v a b i l i t y . Both Fox (1978) and Stevens (1979) 121 observed that goats were more i n a c t i v e on sunny days than on cloudy days. In t h i s study, goats were observed to stand f o r long p e r i o d s on c o l d winter days. Malecheck and Smith (1976) noted the same response by c a t t l e . They a l s o c o n s i d e r e d wind v e l o c i t y to be i n v e r s e l y r e l a t e d to d a i l y t r a v e l , s i m i l a r to the e f f e c t of snow depth (Smith 1976). 1 22 VI. CONCLUSIONS AND RECOMMENDATIONS 1. IMPLICATIONS FOR STUDY DESIGN In o b s e r v a t i o n a l s t u d i e s , a n a l y s i s of open h a b i t a t s may only y i e l d a p a r t i a l understanding of mountain goat ecology. A confounding problem, noted a l s o by Rounds (1981), may occur with deciduous shrub and t r e e cover, i n which animal o b s e r v a b i l i t y f o r the observer v a r i e s s e a s o n a l l y between and w i t h i n cover types. Systematic sampling methods can help minimize b i a s e s , or at l e a s t h e l p to keep t h e i r departures c o n s t a n t , however, the u l t i m a t e goal i s not s o l e l y p r e c i s i o n but a l s o accuracy. The use of a more e f f i c i e n t technique i s recommended. For example, h e l i c o p t e r surveys g e n e r a l l y r e s u l t i n higher counts than ground or fixed-wing surveys ( B a l l a r d 1975,1977), but they may a l s o a f f e c t an animal's a c t i v i t y and p o s s i b l y even h a b i t a t use (Fos t e r and Rahs, i n p r e s s ) . An a l t e r n a t i v e s o l u t i o n to the problem of v i s i b i l i t y would be to use remote-sensing d e v i c e s a f f i x e d to i n d i v i d u a l animals. Using t h i s technique, Schoen et a l . (1981) d i s c o v e r e d that c e r t a i n mountain goat i n d i v i d u a l s are ' a l p i n e d w e l l e r s ' , while others spend most of t h e i r l i f e i n the f o r e s t . However, i t may be necessary to o b t a i n r e p r e s e n t a t i v e samples from a l l sex and age c l a s s e s i f d i f f e r e n t i a l l e v e l s of s e l e c t i o n commonly occurs f o r mountain goats as has been documented f o r mountain sheep (Shank 1979a, M a r t i n e l l i and Hudson 1981, T i l t o n and W i l l a r d 1982). A few telemetry s t u d i e s have been conducted with mountain goats (e.g. Chadwick 1973, Rideout 1974, Schoen et a l . 1 23 1981, N i c h o l s , i n p r e s s ) , but these have c o n c e n t r a t e d on only a small f r a c t i o n of the a c t u a l p o p u l a t i o n , which p r e c l u d e s documentation of p o p u l a t i o n demography and sex-age d i f f e r e n c e s i n mountain goat ecology. Radio-telemetry s t u d i e s are not without other disadvantages e i t h e r . In a d d i t i o n to e x c e s s i v e c o s t s i n v o l v e d with t r a p p i n g , tagging and equipment f o r m o n i t o r i n g mountain goat movements, the problem of s i g n a l bounce may become e x c e s s i v e i n rugged and rocky mountain goat h a b i t a t . Furthermore, the accuracy of t r i a n g u l a t i o n f i x e s may a l s o leave something to be d e s i r e d , p a r t i c u l a r l y when a s s e s s i n g s m a l l - s c a l e h a b i t a t r e l a t i o n s h i p s i n patchy environments. For example, Denton (1973) r e p o r t e d a mean e r r o r polygon of 4.3 ha i n a western Montana e l k study using a e r i a l t r a c k i n g , with only 88 percent of the approximation areas a c t u a l l y i n c l u d i n g the e l k or the t e s t t r a n s m i t t e r . Although a e r i a l t r a c k i n g has the advantages of speed and a c c e s s i b i l i t y of even the l a r g e s t , most remote and rugged study areas, i t may be r e l a t i v e l y i n n a c c u r a t e i f v i s u a l c o n t a c t i s not made (Denton 1973). However, some s p e c i e s , such as mountain goats and mountain sheep, seem more alarmed by a i r c r a f t at long d i s t a n c e s (Shank 1979b). B i g g i n s and P i t c h e r (1978) compared the e f f i c i e n c y of v i s u a l ( f l i g h t g r i d ) and t e l e m e t r i c techniques f o r mule deer (Odocoileus hemionus) and concluded that the two methods produced s i g n i f i c a n t l y d i f f e r e n t p a t t e r n s of d i u r n a l h a b i t a t use. A d d i t i o n a l l y , n o c t u r n a l h a b i t a t use was impossible to assess o b j e c t i v e l y by v i s u a l techniques and was s i g n i f i c a n t l y 124 d i f f e r e n t than d i u r n a l use f o r instrumented deer. Mountain goats are known to be a c t i v e n o c t u r n a l l y (Chadwick 1973, Rideout 1974, Stevens 1979). 2. MANAGEMENT IMPLICATIONS The e f f e c t s of harvest or h a b i t a t m a n i p u l a t i o n on mountain goat p o p u l a t i o n demography cannot be determined u n t i l p o p u l a t i o n parameters are f i r s t a c c u r a t e l y documented. For example, at the recommended 5 percent a d u l t mountain goat harvest l e v e l o u t l i n e d by the B r i t i s h Columbia M i n i s t r y of Environment (1979), one would have to be more than 95% c o n f i d e n t of the accuracy of survey i n f o r m a t i o n c o l l e c t e d , i n order to document the e f f e c t s of t h i s h arvest s t r a t e g y . At t h i s time, there i s no technique a v a i l a b l e with the p r e c i s i o n and accuracy r e q u i r e d f o r such i n t e n s i v e management of the s p e c i e s . For example, the seven systematic h e l i c o p t e r t r a n s e c t surveys i n Van Drimmelin's (1982) mountain goat study c o n t a i n e d t o t a l counts averaging 31% from the estimated p o p u l a t i o n mean at the 95% p r o b a b i l i t y l e v e l . Using t h i s i n f o r m a t i o n as t e s t data, i t was estimated that more than 250 surveys, at a t o t a l c o s t exceeding $66,000.00 (1/2 hour of h e l i c o p t e r time per survey, at $500.00/hour) would have been r e q u i r e d to be 95% c o n f i d e n t that the t o t a l count obtained was w i t h i n 5% of the p o p u l a t i o n mean. S a t i s f y i n g s t a t i s t i c a l sampling requirements would be i n c r e a s i n g l y more d i f f i c u l t to achieve f o r s m a l l e r - s i z e d p o p u l a t i o n s ( i . e . <100 animals) due to the l a r g e v a r i a t i o n a s s o c i a t e d with m i s s i n g s e v e r a l animals. S i m i l a r l y , l e s s dense 125 p o p u l a t i o n s ( i . e . <10 goats/km 2) may a l s o i n c r e a s e survey v a r i a t i o n . The M i n i s t r y of Environment (1979) r e c o g n i z e d the need f o r development of new and s t a n d a r d i z e d i n v e n t o r y techniques f o r mountain goats, however i t i s the recommendation of t h i s study to c o n s i d e r the s u b j e c t a p r i o r i t y i s s u e . Management g u i d e l i n e s should not be implemented without the a b i l i t y to monitor t h e i r e f f e c t . 1 26 LITERATURE CITED Adams, L.G. 1981. Ecology and p o p u l a t i o n dynamics of mountain goats, Sheep Mountain-Gladstone Ridge, Colorado. M.S. T h e s i s , Colorado State U n i v e r s i t y , F o r t C o l l i n s , Colorado. 189pp. Altmann, J . 1974. O b s e r v a t i o n a l study of behaviour: sampling methods. Behaviour 49:227-267. Anderson, N.A. 1940. Mountain goat study. Washington Dept. Game, B i o l . B u l l . 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