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Ground-truth and large-scale 70 mm aerial photographs in the study of reindeer winter rangeland, Tuktoyaktuk… Sims, R. A. 1983

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GROUND-TRUTH AND LARGE-SCALE 70 mm AERIAL PHOTOGRAPHS IN THE STUDY OF REINDEER WINTER RANGELAND, TUKTOYAKTUK PENINSULA AREA. N.W.T. by RICHARD ALLAN SIMS B.Sc.Hon., Lakehead U n i v e r s i t y , 1974 M.Sc, U n i v e r s i t y of Manitoba, 1977 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES Dept. of Fores try/Remote Sensing 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 A p r i l , 1983 ©Richard A. Sims, 1983 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements for an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y available for reference and study. I further agree that permission for extensive copying of t h i s thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. I t i s understood that copying or publication of t h i s thesis for f i n a n c i a l gain s h a l l not be allowed without my written permission. Department of F o r e s t r y  The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date DE-6 (3/81) ABSTRACT Reindeer (Eangifev tarandus tarandus L.) winter rangeland i n the Tuktoy-aktuk P e n i n s u l a area, N.W.T., was s t u d i e d using a g r o u n d - t r u t h / l a r g e - s c a l e (1:1,400-1:3,400) remote sensing program. Ground-truth of v e g e t a t i o n , s o i l s and general environment was conducted at 112 r e p r e s e n t a t i v e s i t e s l o c a t e d throughout the study area. Two-way i n d i -c a t o r species a n a l y s i s (TWINSPAN) of v e g e t a t i o n cover by 420 pla n t taxa assigned s i t e s among four b r o a d l y - d e f i n e d 'vegetation groups'. The v e g e t a t i o n groups could be considered as ecosystemic u n i t s since they are a l s o d i f f e r -e n t i a t e d by a range of s i t e parameters, i n c l u d i n g slope p o s i t i o n c l a s s e s , general cover features measured i n 10 m x 10 m p l o t s , mineral s o i l t e x t u r e c l a s s e s , the occurrence of organic s o i l s and ice-wedge polygons, and c e r t a i n s o i l p h y s i c a l and chemical parameters. Lichens are of p a r t i c u l a r • i m p o r t a n c e as the winter d i e t mainstay f o r the r e i n d e e r , and d i f f e r e n c e s among v e g e t a t i o n groups are r e f l e c t e d by dominant l i c h e n t a xa, and l i c h e n ground cover, biomass and standing crop estimates. Lichen cover at s i t e s ranged up to 89.3% and, fo r s i t e s where l i c h e n cover >_20%, standing crop ranged from 194.4 to 6,377.6 kg.ha -*. Larg e - s c a l e c o l o u r - i n f r a r e d (CIR) 70 mm stereo photographs were acquired throughout the study area along 44 f l i g h t l i n e s , and a t o t a l of 1,469 photo-frames were i n t e r p r e t e d and i n v e n t o r i e d . Data were summarized according to 7 reindeer management zones defined w i t h i n the study area. A l l Ice-wedge polygons are common t e r r a i n features i n p o o r l y - d r a i n e d l o c a -t i o n s and are estimated to cover 23.4% of the land s u r f a c e . T e r r a i n d i s t u r b -ance by v e h i c l e s was observed i n 19.0% of the photo-frames, but accounted f or g e n e r a l l y low cover which ranged from 0.5% to 2.0% f o r d i f f e r e n t r e i n d e e r man-agement zones. Lichen Types were recognized as the a i r - p h o t o i n t e r p r e t e d e q u i v a l e n t s o f three of the four v e g e t a t i o n groups. W i t h i n Lichen Types i d e n t i f i e d on photo-frames, m i c r o d e n s i t o m e t r i c measurements were made of l i c h e n patches. Quanti-t a t i v e study of the measurements helped confirm s e p a r a b i l i t y of the Lichen Types; L i n e a r D i s c r i m i n a n t Function a n a l y s i s provided an 81.1% c o r r e c t r e -assignment of 296 sets of measurements among Lichen Types. Percent l i c h e n cover and l i c h e n standing crop were summarized acc o r d i n g to the rei n d e e r management zones. Measured on the photo-frames, l i c h e n cover v a r i e d among zones from 1.48% to 14.24% of land area. Using l i c h e n biomass measurements from ground-truth s t u d i e s , l i c h e n standing crop ranged from 39.6 kg.ha~i to 572.0 kg.ha~i f o r d i f f e r e n t zones. These estimates allowed the study area's w i n t e r c a r r y i n g c a p a c i t y to be t e n t a t i v e l y determined at 20,373 r e i n d e e r . The study concludes that ground-truth and l a r g e - s c a l e CIR 70 mm photo-graphs can play an important and i n t e g r a t i v e r o l e i n the study of Rangifev w i n t e r rangeland. Use of l a r g e - s c a l e 70 mm photographs i s e s p e c i a l l y advo-cated; there are numerous advantages f o r using l a r g e - s c a l e remote sensing s y s -tems to study northern rangelands, i n c l u d i n g p a r t i c u l a r l y a p r o v i s i o n of base-l i n e i n f o r m a t i o n that permits future m o n i t o r i n g . i v TABLE OF CONTENTS Page Chapter ABSTRACT i i LIST OF TABLES v i LIST OF FIGURES v i i i ACKNOWLEDGEMENTS x I INTRODUCTION 1 1. General 1 2. Study O b j e c t i v e s 2 I I LITERATURE REVIEW 3 1. H i s t o r i c a l Background on the Reindeer Herd 4 2. Winter Use of Ground Lichens by Rangifer 5 3. Remote Sensing of Rangifer Rangeland 12 3.1 A e r i a l Observations 12 3.2 LANDSAT to Study Rangifer Rangeland • 13 3.3 Medium-Scale (1:20,000-1:60,000) A i r Photographs . . . 16 3.4 Large-Scale ( 1:20,000) A i r Photographs 18 4. 70 mm Photographs f o r Rangeland Studies 19 I I I STUDY AREA . 23 -1. General L o c a t i o n - 24 2. Geology, Landform and S o i l s 24 3. Climate 30 4. Vegetation 33 4.1 General Vegetation Zonations 33 4.2 B o t a n i c a l I n v e s t i g a t i o n s 33 4.3 E c o l o g i c a l Studies of Ve g e t a t i o n 34 5. Remote Sensing 35 IV METHODS 38 1. Photo A c q u i s i t i o n 39 1.1 The M u l t i s t a g e Program 39 1.2 F l i g h t l i n e S e l e c t i o n 39 1.3 Photographic M i s s i o n 45 2. Ground-Truth 45 2.1 P r e - f i e l d S i t e S e l e c t i o n 45 2.2 A i r Photo Annotation i n the F i e l d 47 2.3 Data C o l l e c t i o n at Ground-Truth S i t e s 49 2.4 Numerical A n a l y s i s of Ground-Truth Data 52 3. I n t e r p r e t a t i o n and A n a l y s i s of Large-Scale A i r Photographs 53 3.1 Summaries by Reindeer Management Zones 53 3.2 M i c r o d e n s i t o m e t r i c Studies of 'Lichen Types' . . . . . . 55 V TABLE OF CONTENTS (Cont'd.) Page V RESULTS 59 1. Ground-Truth . 60 1.1 C l a s s i f i c a t i o n of Four V e g e t a t i o n Groups 60 1.2 C o r r e l a t i o n of Environmental Parameters to the Four Vegetation Groups 74 1.3 Determination of Li c h e n Standing Crop 79 2. I n t e r p r e t a t i o n and A n a l y s i s of Large-Scale A i r Photographs 84 2.1 General 84 2.2 Patterned Ground and T e r r a i n Disturbance by V e h i c l e s . 89 2.3 M i c r o d e n s i t o m e t r i c Measurements 93 2.4 E s t i m a t i o n of Lichen Standing Crop 101 VI DISCUSSION 108 1. Ground-Truth 109 1.1 C l a s s i f i c a t i o n of the Four V e g e t a t i o n Groups 109 1.2 The Use of I n d i c a t o r Species 110 1.3 F l o r a and Species D i v e r s i t y 112 1.4 General Environment and S o i l s Measurements 114 1.5 Lichen Standing Crop 117 2. I n t e r p r e t a t i o n and A n a l y s i s of Large-Scale A i r Photographs 119 2.1 General 119 2.2 Ice-Wedge Polygons 123 2.3 T e r r a i n Disturbance by V e h i c l e s 125 2.4 Lichen Types and Percent L i c h e n Cover 127 2.5 E s t i m a t i o n of C a r r y i n g Capacity 130 2.6 O v e r u t i l i z a t i o n E f f e c t s 135 3. Future Recommendations 139 3.1 Follow-up Studies on the Rangifer Rangeland 139 3.2 Other P o s s i b l e A p p l i c a t i o n s f o r the Large-Scale A i r Photographs 143 VII CONCLUSIONS 146 LITERATURE CITED 152 APPENDIX I 169 VI LIST OF TABLES Selected c l i m a t i c data f o r the Tuktoyaktuk P e n i n s u l a area, N.W.T. General l o c a t i o n and approximate length of 12 t r a n s e c t s i n the study area. Measurements recorded f o r each f l i g h t l i n e photo-frame. Summary data on reindeer management zones and c o r r e l a t i o n w i t h Ecoregion and E c o d i s t r i c t map u n i t s . T e n t a t i v e h i e r a r c h i c a l c l a s s i f i c a t i o n of the v e g e t a t i o n , Tuk-toyaktuk P e n i n s u l a area, Northwest T e r r i t o r i e s . The percent of t o t a l plant taxa encountered i n the ground-truth s t u d i e s , summarized f o r each v e g e t a t i o n group by s i x s t r u c t u r a l c a t e g o r i e s . Slope p o s i t i o n and aspect of ground-truth s i t e s summarized by the four v e g e t a t i o n groups (A, B, C and D). Percent cover by general environment parameters i n 10 m x 10 m p l o t s at 112 s i t e s , summarized by the four v e g e t a t i o n groups. Subsurface s o i l t e x t u r e f o r mineral s o i l s and the occurrence of organic s o i l s at 112 s i t e s , summarized by the four v e g e t a t i o n groups. Subsurface s o i l p h y s i c a l and chemical data summarized by the four v e g e t a t i o n groups. Mean percent ground cover and, i n parentheses, frequency for the four most abundant l i c h e n species i n v e g e t a t i o n groups A, B and C. L i c h e n biomass measurements from d i v o t s , percent l i c h e n cover, and estimates of top and bottom components of l i c h e n standing crop at t h i r t y - s i x s i t e s where t e r r e s t r i a l l i c h e n s were abundant ( i . e . , ground cover _ 2 0 % ) . Summarized t e c h n i c a l data f o r l a r g e - s c a l e 70 mm CIR photographs acquired Aug. 5-8, 1980. Numbers of ground-truth l o c a t i o n s and s i t e s , and l a r g e - s c a l e a i r photographs i n each of the seven reindeer management zones. v i i LIST OF TABLES (Cont'd.) Table P a § e XV The occurrence of ice-wedge polygons, and t e r r a i n d i s t u r b a n c e by v e h i c l e s , summarized f o r l a r g e - s c a l e photographs i n the reindeer management zones. X V I I I Recombination matr i x (percentage values) f o r step 3 of the stepwise LDF a n a l y s i s using m i c r o d e n s i t o m e t r i c readings o f Lic h e n Types. XIX Number of photo-frames, t o t a l surface area, t o t a l land area and percent l i c h e n cover summarized f o r reindeer management zones according to those w i t h no l i c h e n , and those assigned to Lichen Types I , I I and I I I . 92 XVI M i c r o d e n s i t o m e t r i c readings of o p t i c a l d e n s i t y values w i t h w h i t e , r e d, green and blue l i g h t of three L i c h e n Types on l a r g e - s c a l e (1:1,400-1:3,600) CIR photos. 94 XVII Summary of stepwise L i n e a r D i s c r i m i n a n t Function (LDF) a n a l y s i s based on m i c r o d e n s i t o m e t r i c readings of the three Lichen Types. 99 100 102 XX Estimates of bottom and top components of l i c h e n standing crop based on 1) percent cover i n t e r p r e t e d from l a r g e - s c a l e photo-graphs, and 2) f i e l d s i t e estimates of l i c h e n biomass per u n i t area from Table X I I . 103 XXI Estimates of t o t a l l i c h e n standing crop summarized by rei n d e e r management zones. 107 XXII Selected northern ecosystems used f o r comparison of l i c h e n standing crop. 120 X X I I I E s t i m a t i o n of wi n t e r range c a r r y i n g c a p a c i t y f or the Tuktoyak-tuk P e n i n s u l a area, N.W.T. . 136 XXIV Parameters and scores used to c a l c u l a t e an index of the i n -t e n s i t y of recent Rangifev g r a z i n g . 138 V i l l LIST OF FIGURES Figure Page 1. Modern technology i s used to update r e i n d e e r ranching i n the Tuktoyaktuk Pe n i n s u l a area, N.W.T. (a,b) . 6 2. Map of the study area showing l o c a t i o n of 12 t r a n s e c t s alcng which f l i g h t l i n e s were l o c a t e d , and 44 ground-truth l o c a t i o n s at which s i t e s were s t u d i e d . 25 3. Regional zonations of the area east of the Mackenzie R i v e r D e l t a (a, b, c, d) . 27 4. A m u l t i s t a g e sampling program to i n v e s t i g a t e r e i n d e e r rangeland. 40 5. The areas occupied by the Mackenzie D e l t a r e i n d e e r herd, summer 1978 to f a l l 1982 (hatched l i n e s ) , and the l o c a t i o n of 13 t r a n s e c t s . 41 6. A p o r t i o n of the study area southwest of Tuktoyaktuk showing the random . l o c a t i o n of f l i g h t l i n e s f o r l a r g e - s c a l e 70 mm photography along t r a n s e c t s 4 and 5. 44 7. Twin 70 mm cameras mounted on Cessna 180 wing t i p s f o r s t e r e o -photography. 46 8. Cibachrome 2X enlargement of 1:36,000 ( o r i g i n a l s c a l e ) 70 mm CIR photograph showing path of l a r g e - s c a l e f l i g h t l i n e 4-2. 48 9. C o l l e c t i o n of a 20 cm x 20 cm l i c h e n d i v o t to estimate l i c h e n standing crop. 51 10. Map of the seven reindeer management zones defined i n the study area. 57 11. Two-way i n d i c a t o r species a n a l y s i s (TWINSPAN) dendrogram of 112 ground-truth s i t e s based on v e g e t a t i o n cover of 420 s p e c i e s , Tuktoyaktuk P e n i n s u l a area, N.W.T. 61 12. Normal c o l o u r 35 mm photographs of v e g e t a t i o n group A and B examples ( a , b). 65 13. Example of v e g e t a t i o n group C ( a , b ) . 68 14. Example of v e g e t a t i o n group D. 70 IX LIST OF FIGURES (Cont'd.) Figure P a S e 15. Bottom versus top components of l i c h e n biomass f o r v e g e t a t i o n groups A (open t r i a n g l e s ) , B ( c l o s e d c i r c l e s ) and C (open squares). 16. Cibachrome 2.5X enlargement of 1:1,600 ( o r i g i n a l -scale) 70 mm CIR photograph, f l i g h t l i n e 4-2. 85 90 17. L a r g e - s c a l e (1:1,800) CIR photo-pair showing examples of v e g e t a t i o n groups A and B, and low damage l e v e l s to tundra r e s u l t i n g from m u l t i p l e passes of tracked v e h i c l e s . 91 18. Examples of Li c h e n Types on l a r g e - s c a l e CIR s t e r e o - p a i r s (a,b). 95 19. P l o t of means and standard d e v i a t i o n s of three L i c h e n Types i n blue and white l i g h t o p t i c a l d e n s i t y (O.D.) space. 98 20. Histogram showing f o r each of the seven r e i n d e e r management zones, top and bottom standing crops of Li c h e n Type I ( l e f t ) , I I (centre) and I I I ( r i g h t ) . 104 21. A flow-diagram f o r a s s i g n i n g "unknown" f i e l d s i t e s among the four v e g e t a t i o n groups based on TWINSPAN i n d i c a t o r species and d e c i s i o n r u l e s . I l l 22. Comparison of oblique normal c o l o u r and v e r t i c a l CIR photographs of the same area (a, b). 131 X ACKNOWLEDGEMENTS This study was j o i n t l y funded by Canadian Reindeer (1978) Ltd. and the Department of Indian A f f a i r s and Northern Development. A d d i t i o n a l f i e l d sup-port was provided by the Polar Continental Shelf P r o j e c t , Department of Energy, Mines and Resources. For advice and guidance, I am p a r t i c u l a r l y indebted to my thesis super-v i s o r , Dr. Peter Murtha, and members of my committee: Dr. Roy Strang, Dr. Gary B r a d f i e l d , Dr. Robert Woodham and Dr. Tim B a l l a r d . Helpful comments were provided by thesis examiners Dr. F. Bunnell, Dean J.K. Stager and Dr. P. T u e l l e r . The majority of the project was conducted during an educational leave from the Great Lakes Forest Research Centre, Canadian Forestry Service, and I thank establishment d i r e c t o r Mr. Jim Cayford and program d i r e c t o r Dr. Cal S u l l i v a n for allowing me the opportunity to pursue this t o p i c . I am g r a t e f u l to the following for t h e i r valuable c o n t r i b u t i o n s : Mr. G. Abrahamson, Mr. W. Nasogaluak, Dr. D. B i l l i n g s l e y and Dr. G. Godkin for t h e i r i n t e r e s t and encouragement i n the study; Ms. N. Holm for her help i n the lab and with diagrams; Mr. P. Williams of Integrated Resource Photography Ltd. for his r o l e i n obtaining e x c e l l e n t a e r i a l photographs; Ms. D. Weeks f o r word pro-cessing; Mr. B. Wong and Dr. S. Nash who provided answers to my numerous questions on data processing and s t a t i s t i c s ; Dr. R. Ireland, Dr. G. Argus, Dr. J . G i l l e t t , Dr. W. Sc h o f i e l d , Dr. I. Brodo, Mr. P. Wong, Ms. L. Ley and Dr. J . Stein for t h e i r help i n i d e n t i f i c a t i o n and v e r i f i c a t i o n of plant taxa; Mr. J . Ostrick of the Western A r c t i c Resource Centre, Inuvik for l o g i s t i c a l help i n the f i e l d ; and Mr. E. Moussali f o r lab assistance. x i A s i g n i f i c a n t c o n t r i b u t i o n was made by my capable f i e l d and lab a s s i s t -ant, Mr." M. S i l t a n e n . F i n a l l y , I wish to express my a p p r e c i a t i o n to my w i f e , Ruth, and c h i l d r e n Joan and Jesse f o r t h e i r p a t i e n c e , love and encouragement throughout the study. INTRODUCTION - u -1. General Remote sensing has a prominent r o l e to play i n northern resource study and management. Sev e r a l f a c t o r s focus a t t e n t i o n on remote sensing as a way of o b t a i n i n g t i m e l y , c o s t - e f f e c t i v e and accurate i n f o r m a t i o n f o r a r c t i c areas. These i n c l u d e a c c e l e r a t i n g searches f o r minerals and new energy sources, the severe l o g i s t i c s problems and c l i m a t i c c o n d i t i o n s , the vast r e l a t i v e l y unpopulated expanses, and the s c a r c i t y of data on v i r t u a l l y a l l aspects of the environment (McQuillan 1975, Thie 1979). Dramatic advances over the past decade i n a c q u i s i t i o n , processing and i n t e r p r e t a t i o n techniques have made s a t e l l i t e remote sensing data an import-ant t o o l f o r r e g i o n a l study and management of Canadian a r c t i c resources (Thie et al. 1974, McQuillan 1975, Rubec 1982). LANDSAT with i t s s y n o p t i c and r e p e t i t i v e coverage has been of p a r t i c u l a r value f o r recent b r o a d - l e v e l northern t e r r a i n surveys (Wiken et al. 1981). At intermediate l e v e l s c o n s i d -e r a b l e use i s made of r e a d i l y a v a i l a b l e N a t i o n a l A i r Photo L i b r a r y (NAPL) bl a c k and white 1:60,000 a e r i a l photographs, supplemented as r e q u i r e d w i t h c o l o u r photographs or other remote sensing data (Rubec 1982). There has how-ever been l i t t l e c o n s i d e r a t i o n of the r o l e of l a r g e - s c a l e remote sensing sys-tems i n a r c t i c t e r r a i n , although there are o f t e n needs f o r i n f o r m a t i o n at a l o c a l l e v e l that could b e n e f i t from such technology. L a r g e - s c a l e 70 mm c o l o u r and c o l o u r - i n f r a r e d a e r i a l photographs have been advocated g e n e r a l l y f o r rangeland study and management (Carneggie & Reppert 1969, D r i s c o l l 1969, T u e l l e r 1982). However, w i t h i n a r c t i c Canada's over 2.3 m i l l i o n sq km of rangeland ( K l e i n 1970), v i r t u a l l y no attempts t o assess the usefulness of 70 mm a e r i a l photographs have been made. - 2 -2 . Study O b j e c t i v e s The present study deals with a ground-truth and remote sensing program to assess reindeer (Rangifer tarandus tarandus L.) w i n t e r rangeland on the tundra, Tuktoyaktuk P e n i n s u l a area, Northwest T e r r i t o r i e s . A question was posed: can an i n t e g r a t i v e r o l e be c l e a r l y defined for ground-truth and l a r g e - s c a l e 70 mm a i r photographs i n the study of arctic Rangifer rangeland? R e s u l t s of the study were r e q u i r e d at a r e g i o n a l l e v e l f o r e f f e c t i v e general management of a l a r g e commercial r e i n d e e r herding o p e r a t i o n but these r e s u l t s had to evaluate l i c h e n , the p r i n c i p a l w i n t e r forage, which could only be r e l i a b l y detected and q u a n t i f i e d on the ground or at very l a r g e photo-graphic s c a l e s (e.g., 1:2,000). A c c o r d i n g l y , two s u b - o b j e c t i v e s were de-f i n e d : ( i ) to r e l a t e ground-truth of v e g e t a t i o n and other s i t e parameters, i n c l u d i n g l i c h e n cover and abundance measurements, to 70 mm l a r g e - s c a l e photographs, and ( i i ) to i n v e n t o r y and i n t e r p r e t the 70 mm l a r g e - s c a l e photo-graphs according to a r e g i o n a l scheme so that r e s u l t s could help form a b a s i s f o r long-term management of the r e i n d e e r herd and rangeland. The present i n v e s t i g a t i o n was n e i t h e r j u s t a remote sensing study or a v e g e t a t i o n a l ecology study. I t was purposely aimed at "middle-ground" be-tween ^ i e l d - b a s e d plant ecology, as the dominant component of the ground-t r u t h work, and remote s e n s i n g ; i t could not have been conducted without con-t r i b u t i o n s from both s c i e n c e s . This "middle-ground" i s an area i l l - a d d r e s s e d by the m a j o r i t y of past researchers and so has i n part been r e s p o n s i b l e f o r some animosity between s t r o n g l y f i e l d - o r i e n t e d v e g e t a t i o n s c i e n t i s t s and r e -mote sensing s p e c i a l i s t s who o f t e n l a c k s i g n i f i c a n t f i e l d experience. - 3 -I I . L I T E R A T U R E REVIEW - 4 -1. H i s t o r i c a l Background on the Reindeer Herd In 1919 the Canadian government established a Royal Commission to i n -v e s t i g a t e p o s s i b i l i t i e s of reindeer and muskox husbandry i n northern Canada. Envisioned was an opportunity to supplement the w i l d l i f e resources of the north and improve the poor economic conditions faced by native inhabitants (Rutherford et al. 1922). A preliminary survey was conducted between the Alaska/Yukon border and the Coppermine River i n 1927 and 1928, and i t was determined that excellent rangeland existed i n the area east of the Mackenzie Delta ( P o r s i l d 1929). A f t e r an arduous s i x year reindeer drive from Alaska, the Mackenzie Delta Reindeer Grazing Preserve was put into operation i n 1935 with the a r r i v a l of approximately 3,200 animals (Abrahamson 1963, H i l l 1968, Scotter 1982). From 1935 to about 1960 numerous unsuccessful attempts were made to break o f f from the government-owned reindeer herd smaller herds to be passed on to Inuit ownership. It i s now believed that attempts f a i l e d for a combi-nation of reasons in c l u d i n g i n e f f i c i e n t herding p r a c t i c e s , over-harvesting of animals, overgrazing near settlements, and losses through disease, predation, poaching and s t r a y i n g animals (Scotter 1968, Stager & Denike 1972, Treude 1979). The "experiment" nonetheless demonstrated that herded reindeer could survive and multiply i n the Mackenzie Delta area, and although the reindeer project was government-subsidized to a considerable extent during t h i s period, d i s t i n c t economic and s o c i a l b enefits for the regional population had developed ( H i l l 1967). A f t e r about 1960, "open herding" techniques of herd management were adopted. Unlike the previous "closed herding" approach, the reindeer were allowed to run f r e e l y over the range with an annually organized roundup. The revised approach was far more l a b o u r - e f f i c i e n t ( H i l l 1967, F r i e s e n & Nelson - 5 -1978). The herd was managed under c o n t r a c t u a l arrangements to s e v e r a l i n d i -v i d u a l s u n t i l 1968 when the Canadian W i l d l i f e S e r v i c e was given i n t e r i m con-t r o l of the f l o u n d e r i n g o p e r a t i o n c o n s i s t i n g of only 2,700 animals (Stager & Denike 1972, Treude 1979). In 1974 the e n t i r e government re i n d e e r h o l d i n g was s o l d to the former c h i e f herder, who i n turn s o l d i t i n 1978 to Mr. W i l l i a m Nasogaluak. Both men are n a t i v e I n u i t . Employing sound herd manage-ment, the herd s i z e s t e a d i l y increased to 13,000 i n s p r i n g , 1980 (Nasogaluak & B i l l i n g s l e y 1981) and about 16,000 i n f a l l , 1981 ( D i c k i n s o n 1982). With access to new southern markets f o l l o w i n g the recent c o n s t r u c t i o n of the Dempster Highway, and the emergence of a l u c r a t i v e cash crop, namely the s a l e o f deer a n t l e r s to o r i e n t a l markets as a component i n medicines, Canadian Reindeer (1978) L t d . has an o p t o m i s t i c economic outlook (Nasogaluak & B i l l i n g s l e y 1981). Modern technology i s being drawn upon to improve herd management as much as p o s s i b l e ( F i g . 1) and at the present time, the herd i s c o n t i n u i n g to expand i n s i z e (Nasogaluak, pers. comm.^). A comprehensive annotated b i b l i o g r a p h y p e r t a i n i n g to a l l aspects of the reindeer o p e r a t i o n has been r e c e n t l y prepared (Sims & Murtha 1983). 2. Winter Use of Ground Lichens by Rangifer' Lichens c o n s t i t u t e the c h i e f w i n t e r source of n u t r i t i o n f o r most r e i n -deer and c a r i b o u . In the USSR where over 3.4 m i l l i o n head or over t h r e e -quarters of the world's reindeer e x i s t ( K l e i n & Kuzyakin 1982) approximately 80 to 95% of the winter food i n most re i n d e e r d i s t r i c t s c o n s i s t s of l i c h e n ( K u r k e l a 1976). Three s p e c i e s , Cladina svellaris^C. vangifevina and C. iOwner, Canadian Reindeer (1978) L t d . , Tuktoyaktuk, N.W.T. 20 June, 1980. ^nomenclature f o r a l l p l a n t s given i n the text f o l l o w s Appendix I . - 6 -F i g . 1. Modem technology i s used to update reindeer ranching i n the Mackenzie Delta area, N.W.T. (a) Lapp herder on foot, with herd dog, tends reindeer on Richards Island fawn-ing grounds. Public Archives of Canada, Ottawa, Ont., Photo no. PA-121721. 24 A p r i l , 1936. (b) Forty-four years l a t e r , h e l i c o p t e r s a s s i s t a spring round-up for censusing, ear-tagging and dehorning reindeer at Atkinson Point, Tuktoyaktuk Peninsula. 21 June, 1980. - 8 -arbuscula may comprise from 75 to 90 % of a l l lichens eaten by Russian r e i n -deer (Andreev 1954). In Scandinavia, reindeer consume l i t t l e l i c h e n i n summer, accounting for only about 20% of the d i e t , but th i s figure r i s e s i n autumn from 20 to 50%, to i n winter between 85 and 90% (Ahti 1961, Skogland 1975). S i m i l a r figures are reported for some North American Rangife popula-tions (Cringan 1957, Scotter 1967, K e l s a l l 1968, M i l l e r 1976). Although adaptations to non-lichen winter diets are known (Shank et al. 1978, O r i t s -land et al. 1980, K l e i n 1982), the use of li c h e n s , p a r t i c u l a r l y t e r r e s t r i a l l i c h e n s , i s widespread i n northern areas because these plants represent e a s i -ly a c c e s s i b l e carbohydrate nourishment i n otherwise r e l a t i v e l y unproductive, sub-marginal tundra and subarctic landscapes (Llano 1956, P u l l i a i n e n 1971). The abundance of t e r r e s t r i a l lichens can vary considerably from area to area from pure, dense carpets to i s o l a t e d i n d i v i d u a l t h a l l i . R e l i e f and climate are c o n t r o l l i n g features that make a region generally favourable f o r l i c h e n growth, as i s the r o l l i n g topography and subhumid climate of c e n t r a l Labrador (Hustich 1951) or much of north- c e n t r a l USSR (Andreev 1954, K l e i n & Kuzyakin 1982), or unfavourable for growth, as i s the f l a t meadow and marsh tundra of the North Slope, north of Alaska's Brooks Range (Thomson 1979). Recurrent w i l d f i r e i n some areas patterns the landscape; l i c h e n cover may ex i s t i n broad t r a c t s of land that r e f l e c t a h i s t o r y of f i r e s (Johnson & Rowe 1975, Johnson 1980, 1981). Lichens, when dry, burn e a s i l y and so are p a r t i c -u l a r l y susceptible to the e f f e c t s of f i r e . The r e s u l t s of burning are not homogeneous and depend on many f a c t o r s , including the fuel type, the sub-s t r a t e , antecedent and post-event moisture and weather conditions, the i n t e n -- 9 -s i t y and extent of the burning and the time period involved. Often as a re-sult of changing conditions during a f i r e , there w i l l be as much hetero-geneity within a burn as between burns ( F u l l e r & Rouse 1979). Thus the e f f e c t s of f i r e on l i c h e n stands are complex and the time required for recov-ery can vary (Johnson 1980, K l e i n 1982). For example, a wind-driven "crown f i r e " occurring under conditions of r e l a t i v e l y high surface moisture might produce l i t t l e or no damage to ground lichens (Johnson & Rowe 1975). Even l o c a l l y , changes of just a few centimeters in r e l i e f can have pro-nounced e f f e c t s on the occurrence and vegetatiorial composition of l i c h e n stands (Lambert 1972, Williams et al. 1978). A r c t i c lichens are adapted to e x i s t i n g i n h e a t - d e f i c i e n t ecosystems, i n part because they are h i g h l y re-s i s t e n t to f r o s t i n j u r y and can survive long periods of i n a c t i v i t y while i n a frozen state (Llano 1956, Kershaw 1977). However, low and wet areas normally have few lichens associated with them, and many areas, such as rocky or g r a v e l l y s i t e s , do not have su i t a b l e substrates for f r u t i c o s e l i c h e n s . D i s t r i b u t i o n of t e r r e s t r i a l l i c h e n taxa can also vary s i g n i f i c a n t l y , with only a few genera and species c o n s t i t u t i n g s u i t a b l e l i c h e n forage for Rangifer (Andreev 1954, Pegau 1968). Other taxa occur in low amounts, are appressed c l o s e l y to the groundsurface so they remain unavailable, or contain b i t t e r l i c h e n i c acids that render them unpalatable (Llano 1956, K e l s a l l 1968, Skuncke 1969). While about two thousand l i c h e n species have been described from the a r c t i c (Thomson 1979), only a small number are considered mainstays i n Rangifer winter d i e t s . These species include the well-known and circum-polar "reindeer moss" lichens (Cladina stellaris3 C. rangiferina3 C. arbuscu-la3 C. mitis) that grow with branching intertwined podetia often as dense ground mats up to 30 cm deep. Other important forage lichens are members of - 10 -the genera Cetraria, Stereocaulon, Peltigera,. Alectoria and Cladonia (Andreev 1954, Llano 1956, K e l s a l l 1968). It i s believed c e r t a i n species are s e l e c t i v e l y sought a f t e r by Rangifer for d i f f e r e n t reasons i n c l u d i n g : ( i ) higher d i g e s t i b i l i t i e s (e.g. the hollow-stemmed podetia of Cladina give a high surface area to volume r a t i o ) ; ( i i ) higher p a l a t a b i l i t i e s due to lower concentrations of pungent, b i t t e r l i c h e n i c acids; ( i i i ) vitamin contents (e.g. vitamin C i s present i n some Cetraria spp.), and (iv) higher a v a i l a b l e nitrogen contents (e.g. some Peltigera spp. and Stereocaulon spp. have a c t i v e blue-green algae producing nitrogen i n cephalodia) (Andreev 1954, Llano 1956, Skuncke 1969, P u l l i a i n e n 1971, Williams et al. 1978). The mech-anisms by which Rangifer might perceive small d i f f e r e n c e s , for example i n n u t r i t i o n a l q u a l i t y , are not known. A Rangifer winter diet of p r i m a r i l y l i c h e n , supplemented with small amounts of protein and minerals from i n c i d e n t a l l y - i n g e s t e d evergreen plant parts, etc. ( K e l s a l l 1968), i s a "maintenance d i e t " . Because of the low pro-t e i n content of the foodstuff, by the a r r i v a l of spring most animals are pro-t e i n - d e f i c i e n t (Palmer & Rouse 1945). Reindeer overwintering on l i c h e n may even gain weight because of the carbohydrate content (Palmer 1934, Holleman et al. 1979), but i n s u f f i c i e n t p rotein w i l l r e s u l t i n dete r i o r a t e d muscle t i s s u e and often weak and disease-prone animals (Palmer & Rouse 1945, Skuncke 1969). Upper parts of ground l i c h e n are normally selected for foraging; they are generally more n u t r i t i o u s and palatable than lower senescing parts of t h a l l i that contain a build-up of l i c h e n i c acids (Llano 1956, P u l l i a i n e n 1971). - 11 -Lichens are not only important i n w i n t e r . I t has been recognized f o r some time that l i c h e n s are r e q u i r e d as intermediary agents i n d i g e s t i o n d u r ing the t r a n s i t i o n periods of the s p r i n g and autumn between the w i n t e r d i e t of mainly carbohydrate, and the p r o t e i n food of summer (Skuncke 1969) . P o s s i b l y there are summer d i e t requirements f o r l i c h e n to s u s t a i n the rumen m i c r o f l o r a important for d i g e s t i o n of l i c h e n carbohydrates (White & T r u d e l l 1980a). Winter snow cover i n v a r i a b l y r e s t r i c t s access to the l i c h e n fodder and so serves as a secondary "mask" on the a c t u a l d i s t r i b u t i o n of food patches. When snow depths are s l i g h t , Rangifer may feed on exposed v e g e t a t i o n , but w i t h deeper snow, most dig feeding c r a t e r s . The concave shape of the hoof, with i t s sharp fore-edges enables the animals to e a s i l y scrape snow away dur i n g f e e d i n g . When necessary, c r a t e r s may be over 50 cm deep although l e s s e r depths are u s u a l l y sought ( P r u i t t 1959, Henshaw 1968, Skogland 1978). Numbers, o r i e n t a t i o n and shapes of c r a t e r s are dependent on l o c a l snow depth, surface and subsurface hardness and d e n s i t y , snow-crystal t e x t u r e , and on the h a b i t of the i n d i v i d u a l r e i n d e e r or c a r i b o u ( K e l s a l l 1968, M i l l e r 1974, Thing 1977). Considerable areas may be c r a t e r e d . K e l s a l l (1968) noted that i n ten days, an enclosed herd of 32 r e i n d e e r c r a t e r e d over about 18% of the t o t a l a v a i l a b l e s u r f a c e . Moreover, s i n c e c r a t e r i n g d i s t u r b s and compacts snow cover i n the immediate areas of the c r a t e r s , up to ten to twenty times the area of the a c t u a l c r a t e r may be made u n a v a i l a b l e f o r fu t u r e use i n any one wi n t e r ( P r u i t t 1959). Because of trampling and snow aging Rangifer may be able to u t i l i z e s i t e s only twice i n any winter (Bergerud 1974). Snow cover may thus s e v e r e l y r e s t r i c t u t i l i z a t i o n of b e t t e r q u a l i t y l i c h e n patches; c r a t e r i n g i t s e l f causes c o n s i d e r a b l e areas to become u n a v a i l a b l e while other - 12 -areas may remain for much of the winter under snow too deep or too hard for c r a t e r i n g . 3. Remote Sensing of Rangifer Rangeland 3.1 A e r i a l Observations The species composition of reindeer l i c h e n stands may be v i s u a l l y deter-mined from the a i r at low a l t i t u d e s of 100 to 200 m (Andreev 1961). For ex-ample, pure white stands are t y p i c a l l y dominated by Cladina stellaris3 greyish-white stands may be mixed stands of C. mitis3 C. arbusaula and C. rangiferina, with Alectoria ochroleuca when present imparting a bluish-green tinge (Andreev 1940, Ahti & Hepburn 1967). P o t e n t i a l problems i n d i s t i n -guishing forage lichens from c e r t a i n dry ground mosses or rocky outcrops are avoided by employing experienced a e r i a l observers who have an intimate knowl-edge of the ground vegetation i n an area. Studies i n the USSR i n d i c a t e that such i n d i v i d u a l s can estimate the biomass of shrubs and c e r t a i n l i c h e n pas-tures within 10 to 20% (Andreev 1961). A e r i a l - v i s u a l rangeland assessment for Rangifer has been r o u t i n e l y ap-p l i e d i n the USSR and Northern Europe for years. From 1945 to 1960 Soviet workers cover mapped 4.83 m i l l i o n sq km of reindeer rangeland using mainly a e r i a l observations (Andreev 1961, 1970). In Sweden, Skuncke (1969) adapted from Russian researchers a method for a e r i a l a p p r a isal of rangelands. From small a i r c r a f t f l y i n g along p a r a l l e l transects 20 to 30 km apart and at an a l t i t u d e of 150 to 200 m, quantity and q u a l i t y of as many as seventeen d i f f e r e n t vegetation types may be estimated within 1 sq km sample p l o t s . The a i r c r a f t f l i e s at a constant precalculated height and speed, and at regular time i n t e r v a l s two or more observers e s t i -- 13 -mate forage abundance i n plots below the a i r c r a f t . Using t h i s scheme, seasonal ranges can be outlined usually on broad-scale maps (Pegau 1968, Skuncke 1969). With some modifications, the technique has been used to i n -ventory reindeer rangeland i n Sweden (Skuncke 1969, Eriksson 1980), Norway (Lyftingsmo 1965, c i t e d i n Eriksson 1980), and Greenland (Thing 1980). In northern Ontario, systematic observations along 7,700 km of transects were used to produce broad-scale (1:500,000) maps of woodland caribou rangeland (Ahti & Hepburn 1967). Most Soviet a e r i a l mapping i s supplemented with a e r i a l photographs, with r e s u l t s t r a n s f e r r e d on to maps at the 1:100,000 or 1:200,000 scale and reassessed every ten to twelve years for changes due to grazing and f i r e s (Andreev 1970). S p e c i f i e d l e v e l s of p r e c i s i o n are obtained with a s t r a t i f i e d sampling m o d i f i c a t i o n of the Russian a e r i a l - v i s u a l tech-nique (Eriksson 1980). 3.2 LANDSAT to Study Rangifer Rangeland In 1972 a f e a s i b i l i t y study was undertaken i n northeastern Alaska to de-termine the LANDSAT s a t e l l i t e ' s p o t e n t i a l a p p l i c a t i o n for several s p e c i f i c tasks r e l a t e d to caribou management (Lent & LaPerriere 1974, LaPerriere 1976). A v a r i e t y of v i s u a l and d i g i t a l processing techniques were employed on multidate s a t e l l i t e imagery, and a h e l i c o p t e r reconnaissance program pro-vided ground-truth data. General vegetation maps were subsequently produced for over 19,000 sq km of caribou rangeland. Simple photo i n t e r p r e t a t i o n of images was u s e f u l f or synoptic analyses of regional vegetation such as forest or tundra. These units were usually too general to permit w i l d l i f e habitat evaluations but the mapping of recent w i l d f i r e burns was an exception. D i g i -t a l analyses were s u i t a b l e for rapid, more d e t a i l e d mapping of vegetation associations over large areas (Lent & LaPerriere 1974). - 14 -Using LANDSAT imagery, Lent & L a P e r r i e r e (1974) were unable to detect disturbed snowcover, h e a v i l y used t r a i l systems or aggregations of caribou but could c o r r e l a t e snow-free areas on May imagery to pre-calving migration routes (LaPerriere 1976). They suggested that snow cover data from r e p e t i t i v e s a t e l l i t e imagery might allow predictions of annual v a r i a t i o n s in caribou migration and range-use patterns (Lent & LaPerriere 1974) . Investigating t h i s aspect, Lent (1980) found that the calving areas chosen by caribou during the 1970's have generally been associated with zones of e a r l y m e l t - o f f . In p a r t i c u l a r c e r t a i n drainages and routes were used during pre-calving migrations because they represent snowfree or shallow snowcover avenues (Lent 1980). Lent & L a P e r r i e r e 1 s (1974) f e a s i b i l i t y study concluded that there was considerable p o t e n t i a l a p p l i c a t i o n of LANDSAT to w i l d l i f e habitat inventory and general vegetation mapping. Encouraged by LANDSAT's apparent usefulness for broad-scale h a b i t a t mapping, the Alaska S o i l Conservation Service undertook i n 1976, 1979, and 1980 inventories of 65,000 sq km of wildlands on the Seward Peninsula, west-ern Alaska. The primary objective was to produce vegetation maps at the 1:250,000 scale for use i n reindeer range management (George et al. 1977, George 1981). LANDSAT d i g i t a l data were used as the primary database but were augmented with f i e l d h e l i c o p t e r reconnaissance and occassional ground checks (George et al. 1977, C l i f f o r d 1979). S a t e l l i t e s p e c t r a l data was com-puter c l a s s i f i e d using a c l u s t e r i n g algorithm and the product displayed on a colour t e l e v i s i o n monitor so that f i n a l maps could be e i t h e r hand-drawn or computer plotted (George 1981). For the 1979 and 1980 surveys, recent high-l e v e l CIR photographs of the Seward Peninsula were used to f i r s t subdivide the landscape into physiographic units for sampling purposes, and afterwards - 15 -to check the accuracy of computer-classified maps (George 1981). K l e i n & White (1979) predicted i n the future the eventual refinement of vegetation and range mapping using s a t e l l i t e data so that i n t e r p r e t i v e maps could be prepared of g r a z i n g - s o i l r e l a t i o n s h i p s , growth rates of lichens and other vegetation components, assessment of q u a l i t y , stocking l e v e l s and p o t e n t i a l s , migration routes and c o r r i d o r s , c a l v i n g grounds, wintering areas, and other range c h a r a c t e r i s t i c s . Computer c l u s t e r i n g of LANDSAT d i g i t a l data was ca r r i e d out for a study of reindeer rangeland i n the Adventdalen Valley, Svalbard, northern Norway ( O r i t s l a n d et al. 1980, Odeegard et al. 1981). Eight s p e c t r a l classes were delineated but only two were considered of importance to the approximately 600 reindeer i n the area, one representing sedge and gr a s s - r i c h summer range, and the other a dry heath winter range ( O r i t s l a n d et al. 1980). A 1:50,000 map was prepared and, based on estimates from the l i t e r a t u r e on optimum stocking l e v e l s , the suggestion was made for an immediate reduction of the reindeer population i n the v a l l e y ( O r i t s l a n d et al. 1980). In conjunction with an experimental i n t r o d u c t i o n of reindeer to the Belcher Islands, N.W.T. a preliminary range survey was undertaken that i n c o r -porated v i s u a l i n t e r p r e t a t i o n of LANDSAT imagery (Edmonds 1979). A 1:250,000 scale s a t e l l i t e image was interpreted with the aid of older 1:40,000 and 1:60,000 black and white a e r i a l photographs, so that three seasonal range types (summer, s p r i n g / f a l l , winter) were delineated (Edmonds 1979). Barrenground caribou (Rangifer tarandus groenlandiaus) rangeland i n southern D i s t r i c t of Keewatin, N.W.T. was studied using d i g i t a l and v i s u a l analyses of LANDSAT data ( C i h l a r et al. 1978). An unsupervised c l a s s i f i c a -t i o n algorithm was f i r s t applied to d i g i t a l data to delineate general vegeta-- 16 -t i o n a l patterns i n a 95,500 sq km area. Preliminary maps were c a r r i e d into the f i e l d and used as sampling bases for habitat studies, and as bases for evaluating the r e l a t i v e importance of areas as caribou h a b i t a t . F i e l d work, and a d d i t i o n a l v i s u a l studies of LANDSAT scenes helped i n the i d e n t i f i c a t i o n of nine vegetation mapping units and f i v e vegetation complexes that were mapped at 1:250,000 and 1:1,000,000 r e s p e c t i v e l y for the e n t i r e study area ( C i h l a r et al. 1978). P r i n c i p a l components colour enhancement of d i g i t a l data was used to dev-elop vegetation maps for a 76,500 sq km area of northwestern Manitoba (Dixon 1981, Dixon & Horn 1982). As a preliminary assessment of barrenground caribou h a b i t a t , three categories of burn and twelve major vegetation associations of the tundra and subarctic forest were represented on s i x NTS mapsheets at a scale of 1:250,000 (Horn 1981, Dixon 1981). Vegetation f i e l d data was c o l l e c t e d to provide d e t a i l e d descriptions of the vegetation a s s o c i a t i o n s . Image enhancements gave appropriate r e s u l t s based on f i e l d studies and i n t e r p r e t a t i o n of older (1957) conventional black and white photographs (Horn 1981, Dixon 1981). V i s u a l analyses of LANDSAT image transparencies and d i g i t a l . c l a s s i f i c a t i o n s did not provide useful r e s u l t s for caribou management (Dixon 1981, Dixon & Horn 1982). 3.3 Medium-Scale (1:20,000-1:60,000) A i r Photographs Conventional medium-scale a e r i a l photographs have not received wide-spread use i n the study of Rangifer rangeland. Probably the expense of ob-t a i n i n g h i g h - q u a l i t y photo-coverage for the large land areas t y p i c a l l y i n -volved, and the time required i n i n t e r p r e t i n g and c a l c u l a t i n g areas on large numbers of photos have served as deterrents. Although 1:60,000 black and - 17 -white panchromatic air-photos are widely a v a i l a b l e and cover most northern areas, they may be dated and usually are of v a r i a b l e q u a l i t y for rangeland mapping. More importantly, 1:20,000 or larger scale photographs are i n v a r i -ably required; smaller scales cause i n t e r p r e t e r s to overlook small stretches of water, bedrock and other features that s i g n i f i c a n t l y lower a pasture's q u a l i t y (Skuncke 1969, Andreev 1970). As a p i l o t study of caribou winter range, Beckel (1958) mapped vegeta-t i o n of an 11,432 sq km tra c t i n subarctic northern Manitoba using 1:60,000 NAPL black and white photographs acquired i n 1955. With topography, s o i l types and drainage as main c r i t e r i a for d i v i s i o n , she recognized eight cover types. The percent cover figures for the types were subsequently t a l l i e d by K e l s a l l (1968: Table 5). To provide information on Rangifer rangeland types for a large area, a por t i o n of the medium-scale (e.g. 1:20,000-1:60,000) black and white photo-coverage may be sub-sampled. To e s t a b l i s h reindeer grazing areas f o r family-owned herds i n Finmark, northern Norway, a sampling g r i d was o v e r l a i n on an 18,000 sq km t r a c t and a selected sample of 1:20,000 a i r photos t o t a l l i n g only 5% of the area was interpreted ( E i n e v o l l 1968). Percent cover was e s t i -mated for nine t e r r a i n types i n c l u d i n g two considered as prime winter range-land ( E i n e v o l l 1968). On parts of Somerset Island, N.W.T. selected black and white 1:60,000 photos were used to s e l e c t f i e l d sampling locations and gener-a l l y map major range types for caribou and muskoxen (Russell & Edmonds 1976). Using photographs of a s i m i l a r scale, and i n t e g r a t i n g intensive vege-t a t i o n ground sampling into t h e i r study, Komarkova & Webber (1980) produced two maps (1:21,000 and 1:10,500 scales) of habitat types near Atkasook, northern Alaska. These maps were then used as a framework for d e t a i l e d - 18 -studies of plant p r o d u c t i v i t y and standing crop (Komarkova & Webber 1980), s o i l d i s t r i b u t i o n and v a r i a b i l i t y (Everett 1980), and caribou habitat preferences and forage consumption (White & T r u d e l l 1980b) . Caribou rangelands were investigated on Southamptom Island, N.W.T., a 43,000 sq km i s l a n d i n northwestern Hudson Bay (Parker 1976). Seven classes of moisture regime were f i r s t mapped on 1:60,000 black and white a i r photos, then f i e l d surveys were conducted to r e l a t e the classes to vegetation ground cover, d i s p e r s i o n and above-ground biomass. Problems were encountered i n c o r r e l a t i n g moisture regime classes to range types i d e n t i f i e d i n the f i e l d , p a r t i c u l a r l y on the undulating, and v e g e t a t i o n a l l y varied Precambrian Shield portions of the i s l a n d . It was concluded that the usefulness of the approach was l i m i t e d by the scale of the photographs (Parker 1976). 3.4 Large-Scale (<1:20,000) A i r Photographs In the Southampton Island study of caribou rangeland, the 1:60,000 black and white photographs were compared to l i m i t e d coverage by normal colour 1:12,000 photos acquired the summer of 1970. The l a t t e r proved far superior for i d e n t i f i c a t i o n of vegetation types described during ground-truth studies (Parker 1976). During summer, 1967, colour a e r i a l photography at a scale of 1:15,840 was conducted for a rangeland inventory of the northwestern Manitoba mapsheet (64K, Whiskey Jack Lake) that Beckel (1958) had previously mapped ( M i l l e r & Barnhardt 1973, M i l l e r 1976). A dot-grid overlay was used to provide percent cover figures for eight major habitat types int e r p r e t e d on the colour photos, and annual rates of w i l d f i r e burn were estimated ( M i l l e r 1976, 1980). Based on the comparative abundance of habitat types as determined from the colour - 19 -photos, twenty-five ground s i t e s were a l l o t e d among the four most common types for d e t a i l e d study. F i e l d measurements were made of vegetation composition and cover, standing crops by important forage species, and two-year plant recovery on clipped plots ( M i l l e r 1976). In western Greenland, a hand-held 35 mm camera system was used to acquire large scale (1:8,500) colour i n f r a r e d photographic p r i n t s (20 cm x 25 cm format) for approximately 300 sq km of caribou winter range and 100 sq km of summer range (Holt 1980). Vegetation maps were prepared from photo-over-l a y s , and were checked against ground surveys. E f f e c t s of grazing were docu-mented during the ground surveys, however attempts to c o r r e l a t e these obser-vations with the a i r photos were unsuccessful except for i s o l a t e d cases of severe overgrazing (Holt, pers. comm.*). 4. 70 mm A e r i a l Photographs to Study Rangeland App l i c a t i o n s of large-scale remote sensing systems for range resource study have become widespread i n recent years. Up u n t i l the l a t e 1960's, only standard black and white panchromatic photographs of medium scale (1:15,840 to 1:60,000) were used for mapping broad vegetation types (e.g. grassland vs shrubland vs woodland) and l o c a t i n g c u l t u r a l features such as roads, fences or seeded areas ( D r i s c o l l 1970). It was then r e a l i z e d that large-scale re-mote sensing systems could provide information for many c r i t i c a l aspects of rangeland management, including range inventory and c l a s s i f i c a t i o n , the de-termination of carrying c a p a c i t i e s and s i t e p r o d u c t i v i t y , studies of u t i l i z a -l s . Holt, I n s t i t u t e of Systematic Botany, Univ. of Copenhagen, Denmark. L e t t e r dated 24 Feb., 1981. - 20 -t i o n l e v e l s , range readiness and trend monitoring, and the evaluation of multiple-use p o t e n t i a l s , improvement p o t e n t i a l and w i l d l i f e habitat values (Carneggie & Reppert 1969, T u e l l e r 1977, 1982). Research concentrated on the use of large-scale (e.g. 1:600-1:5,000) 70 mm photographs, and there are now some conclusive r e s u l t s on the information which can be obtained by such re-mote sensors. There are several advantages for choosing 70 mm photographic systems over other remote sensors for range studies. T y p i c a l l y the cameras have fast shutter speeds and rapid f i l m advances that permit low l e v e l f l i g h t s while maintaining stereo-overlap. Equipment costs are r e l a t i v e l y low, the cameras are l i g h t and portable and allow interchangeable lens and f i l t e r systems, t h e i r normal narrow angles of view reduce t i l t e f f e c t s on p a r a l l a x measure-ments, and the f i l m r o l l s are r e a d i l y processed for. d i r e c t viewing, without c u t t i n g , on easily-modified l i g h t table systems (Carneggie & Reppert 1969, D r i s c o l l 1971). Colour or colour i n f r a r e d (CIR) films are normally recom-mended for use, with the l a t t e r preferred for separation and i d e n t i f i c a t i o n of shrub, forb and grass species ( D r i s c o l l 1969, D r i s c o l l et al. 1970, D r i s c o l l & Coleman 1974), the determination of s o i l surface condition ( D r i s c o l l et al. 1970, T u e l l e r 1977), and the study of plant vigour and bio-mass ( D r i s c o l l et al. 1974, E v e r i t t et al. 1980). For the study of large rangeland t r a c t s 70 mm a e r i a l photographs may be-come too expensive and numerous. In such instances multistage sampling schemes can incorporate them along with other remote sensing platforms at smaller scales into comprehensive information-gathering systems (Carneggie & Reppert 1969, D r i s c o l l 1974, T u e l l e r 1982). The future monitoring c a p a b i l i t y provided by large-scale photographic systems is of p a r t i c u l a r value; the - 21 -photographs are important permanent records for future comparison (Carneggie & Reppert 1969, D r i s c o l l 1970). Microdensitometry of f i l m dye layer densi-t i e s has proven to be a useful technique for the q u a n t i t a t i v e evaluation and comparison of wildland plant communities and components viewed on 70 mm large-scale photographs ( D r i s c o l l et al. 1970, 1974, E v e r i t t et al. 1980, 1981). Studies regarding the a p p l i c a t i o n of 70 mm photograpic systems to range resources have been concentrated i n the a r i d rangelands of the southwestern United States and western A u s t r a l i a , but also have covered other areas of the world. There are few studies documented from the a r c t i c or subarctic even though these areas c o n s t i t u t e the largest rangeland expanses i n the world ( K l e i n 1970). Lesser snow goose (Anser oaerulescens) colonies i n the eastern Canadian a r c t i c were examined on 1:10,000 and 1:4,000 scale 70 mm photographs (Kerbes 1975). The photographs provided estimates on goose nesting densi-t i e s , geographical l i m i t s of col o n i e s , and sex and age class d i s t r i b u t i o n s (Kerbes 1975). Landscape change and vegetation succession was monitored f o l -lowing upstream h y d r o - e l e c t r i c dam construction that caused a sudden drop i n water l e v e l i n the Peace-Athabasca Delta, northeastern A l b e r t a . 70 mm photo-coverage at 1:6,000 s c a l e , and ground-truth studies were used i n the prepara-t i o n of large-scale maps that documented changes over a four year period i n the Delta vegetation ( D i r s c h l et al. 1974). In a multistage study of lesser snow goose habitat at Cape Henrietta-Maria, southern Hudson Bay, large-scale 70 mm photographs were used in conjunction with ground-truth and smaller-scaled remote sensing data to estimate the d i s t r i b u t i o n of major vegetation community types (Wickware et al. 1980). - 22 -Undoubtedly 70 mm a e r i a l photographs w i l l be an important source of remotely-sensed data i n the future and w i l l be h e a v i l y r e l i e d on for many a p p l i c a t i o n s . This i s p a r t i c u l a r l y true i n l i g h t of e s c a l a t i n g costs for ground-truth and increasing requirements for l a r g e - s c a l e information on t e r r a i n areas (Carneggie & Reppert 1969, T u e l l e r 1982). - 23 -- 24 -STUDY AREA 1. General Location The study area i s located immediately east of the Mackenzie River Delta, Northwest T e r r i t o r i e s ( F i g . 2), and comprises about 15,000 sq km of land area, approximately the northern one-third of the Mackenzie Delta Reindeer Grazing Preserve ( H i l l 1968). It includes the e n t i r e Tuktoyaktuk Peninsula and adjacent mainland to the southeast, Richards Island, and a s t r i p of tundra and forest-tundra south of Eskimo Lake and Liverpool Bay that extends inland up to 35 km. The area spans almost two degrees of l a t i t u d e from 68°50'N to 70°20'N, and s i x degrees longitude from 129°W to 135°W, and i s included on the 1:500,000 scale Inuvik mapsheet (Map no. 107SW & 107SE, National Topographic S e r i e s ) . The study area was defined based on consultations with the reindeer herd owner, Mr. William Nasogaluak, and h i s business manager, Dr. Douglas B i l l i n g -s l e y . It includes those rangelands where the reindeer have grazed i n recent years, and those areas to where future expansion might conceivably be extend-ed. While good rangeland may exist south of the study area, the herders are apprehensive to move into closed woodlands where the reindeer quickly s c a t t e r and are l o s t . Pursuit of animals i n the deeper snow of wooded areas is d i f -f i c u l t with snowmobiles and impossible on foot. 2. Geology, Landform and S o i l s Sedimentary rocks, predominantly shales and sandstones underlie much of the region and date as far back as the Cambrian period (Young et al. 1976). Estimates for f i n d i n g hydrocarbons i n Paleocene and older sedimentary rocks range from f a i r to excellent i n the Richards Island area, and n i l to good i n - 25 -Figure 2. Map of the study area showing l o c a t i o n of 12 transects along which f l i g h t l i n e s were located, and 44 ground-truth locations at which s i t e s were studied. S o l i d and open c i r c l e s are, r e s p e c t i v e l y , 1980 and 1981 locat i o n s , and the study area boundary i s indi c a t e d by a broken l i n e . Inset map shows study area i n r e l a t i o n to mainland NW Canada. - 26 -the Eskimo Lakes-Tuktoyaktuk P e n i n s u l a area (Young et al. 1976) and there has thus been con s i d e r a b l e e x p l o r a t i o n i n the past throughout the area f o r com-m e r c i a l o i l and gas d e p o s i t s . Bedrock i s o v e r l a i n i n most of the study area by P l e i s t o c e n e f l u v i a l , d e l t a i c and e s t u a r i n e sediments. G l a c i a l t i l l s d a t i n g to the Wisconsin i c e r e t r e a t form t h i n surface mantles, except on the n o r t h e a s t e r n part of the Tuktoyaktuk P e n i n s u l a (Mackay 1963, Rampton 1972). G l a c i a l a c t i o n i s evidenced by e s k e r s , kames, ground, l a t e r a l and end moraines, p i t t e d outwash d e p o s i t s , g l a c i a l l i n e a t i o n s and deformation pat-t e r n s , and g l a c i a l drainage systems (Mackay 1963). The study area i s l a r g e l y w i t h i n the P l e i s t o c e n e Coastland Region (Mac-kay 1963) which i n c l u d e s Richards I s l a n d , the Tuktoyaktuk P e n i n s u l a and the mainland 16 to 32 km south of the Eskimo Lakes ( F i g . 3b). Most of the P l e i s t o c e n e Coastlands l i e s below 60 m e l e v a t i o n with the topography v a r y i n g .from n e a r l y f l a t along the coast to u n d u l a t i n g i n l a n d . L a c u s t r i n e , morainal and f l u v i a l genetic m a t e r i a l s dominate, and lakes cover over 15% of the sur-face (Mackay 1963). Tabular i c e and pingo growth, thermal e r o s i o n and thermokarst i n c l u d i n g p a r t i c u l a r l y thermokarst lake development, water flow and wind a c t i o n have s i g n i f i c a n t l y modifed l o c a l surface morphologies throughout most of the study area. Pingos and t a b u l a r ground i c e are w i d e l y d i s t r i b u t e d except bn the P e n i n s u l a east of Hutchinson Bay where ground i c e i s v i r t u a l l y absent and pingos occur with lower frequency (Mackay 1963, Stager 1956).' Rampton (1972) has mapped major s u r f i c i a l d e posits arid geomorphic f e a t u r e s . The e n t i r e area i s w i t h i n the continuous permafrost zone (Brown 1967) and p e r e n n i a l l y frozen ground i s encountered at most l o c a t i o n s w i t h i n 1 m of the s u r f a c e . Permafrost depths range from under 400 ra i n the Reindeer - 27 -F i g . 3. Regional zonations of the area east of the Mackenzie River Delta. (a) Reindeer seasonal rangelands based on use of the en t i r e Mackenzie Delta Grazing Reserve, and a v a i l a b i l i t y of herbaceous and graminoid vegetation for summer fodder, lichens for winter fodder, winter fuelwood, s u i t a b l e fawning grounds, etc.; as defined by P o r s i l d (1947, broken l i n e s ) and Cody (1963, s o l i d l i n e s ) . (1) summer range, (2) s p r i n g / f a l l range, (3) winter range. (b) Physiographic regions in t e r p r e t e d from a e r i a l photographs (Mackay 1963). (1) Pleistocene Coastlands, (2) Unfluted P l a i n s , (3) Fluted P l a i n s , (4) Caribou H i l l s , (5) Mackenzie Delta, (6) Campbell Lake H i l l s , (7) Anderson River Uplands. (c) Ecoregions defined as broad areas mapped at l e v e l s of about 1:1 m i l l i o n that have d i s t i n c t i v e e c o l o g i c a l responses to climate as expressed by general vegetation, s o i l s , physiographic and hydrologic patterns, etc. (Houseknecht 1981). (1) Mackenzie Coastal P l a i n , (2) Liverpool Coastal P l a i n , (3) Anderson Coastal P l a i n , (4) Wolverine Creek, (5) Mackenzie River. (d) Climatic zones ( s o l i d l i n e s ) based on major c l i m a t i c v a r i a b l e s (mean annual r a i n f a l l , minimum and maximum annual temperatures, e t c . ) , physio-graphy and vegetation (Burns 1973-1974). (1) marine tundra, (2) c o n t i -nental tundra, (3) tai g a . Forest regions (broken l i n e ) based on forest vegetation and general physiography (Rowe 1972) with tundra (north and east of broken l i n e ) , and forest-tundra (south and-west). - 29 -S t a t i o n area to over 600 m on the coast northeast of Tuktoyaktuk (Mackay 1979). At Tuktoyaktuk, mean annual ground temperature i s -8°C, permafrost reaches a depth of 365 m on undisturbed s i t e s and the active layer i s from 8 to s l i g h t l y more than 100 cm in depth (Rampton & Bouchard 1975). Permafrost ice occurs as coatings, grains, wedges and massive beds and i s incorporated into many landform features common to the area: massive i c y beds, pingos, r a i s e d - and low-centred ice-wedge polygons, s o l i f l u c t i o n s t r i p e s and lobes, and cryoturbated earth hummocks (Stager 1956, Mackay 1963, 1979, Rampton & Mackay 1971, Tarnocai & Z o l t a i 1978, Z o l t a i & Tarnocai 1981). Tundra d i s -turbances can a l t e r or disrupt surface vegetation which subsequently leads to thermokarst subsidence and t e r r a i n damage that may p e r s i s t for decades (Mac-kay 1970, Kerfoot 1972, Strang 1973, Rampton & Bouchard 1975). S o i l s developed on these p e r e n n i a l l y frozen materials are c l a s s i f i e d as Cryosolic (Canada S o i l Survey Committee, Subcommittee on S o i l C l a s s i f i c a t i o n 1978). Within the study area, s o i l s developed on medium and fine-textured materials are cryoturbated, have a high ice content and are r e f e r r e d to as Turbic Cryosols, while i n areas of coarse-grained deposits S t a t i c Cryosols are found ( Z o l t a i & Tarnocai 1975). Organic Cryosols occur i n poorly drained areas, e s p e c i a l l y on ice-wedge polygons and l a c u s t r i n e f l a t s (Rampton & Bouchard 1975, Z o l t a i & Tarnocai 1975) Wetlands including peatlands (cf Z o l t a i et al. 1974) are common, cover-ing over 25% of the study area north of the t r e e l i n e and 5 to 25% south of the t r e e l i n e (Wetlands Working Group, Canada Committee on E c o l o g i c a l Land C l a s s i f i c a t i o n 1981). Most of the area belongs to the Low A r c t i c Wetland^ Region, where c h a r a c t e r i s t i c wetlands are lowland polygons, both r a i s e d -- 30 -and low-centred v a r i e t i e s , and, i n h y d r o l o g i c a l l y suitable areas, f l o o d p l a i n fens, t i d a l marshes and other wetland types (Wetlands Working Group, Canada Committee on E c o l o g i c a l Land C l a s s i f i c a t i o n 1981). E c o l o g i c a l land survey (ELS) i s an integrated, h o l i s t i c approach to landscape survey and mapping by which areas of land, as ecosystems, are c l a s s i f i e d according to t h e i r e c o l o g i c a l unity. At the broadest l e v e l s of the ELS h i e r a r c h i c scheme, ecoregion and e c o d i s t r i c t map units.are defined to aid regional planning and management (Wiken 1980). A preliminary 1:500,000 ecoregion and e c o d i s t r i c t map prepared for the Tuktoyaktuk Peninsula area (Houseknecht 1981) extends in context and concept ecoregion and e c o d i s t r i c t information on the northern Yukon (Wiken et al. 1981). The study area f a l l s l a r g e l y within two ecoregions, the Mackenzie Coastal P l a i n , and L i v e r p o o l Coastal P l a i n ( F i g . 3c). 3. Climate The climate of the study area i s the Cl Subhumid Cold Microthermal type described by Sanderson (1948). The southern h a l f has a continental climate but the northern portion including the Tuktoyaktuk Peninsula experiences a maritime influence from the cold Beaufort Sea ( F i g . 3d; Burns 1973-1974). Winters are long, very cold and lack s i g n i f i c a n t daylight .periods. Summers are short, cool to warm, and have extended daylengths, p a r t i c u l a r l y i n June and J u l y , that can b r i e f l y r e s u l t i n amounts of a v a i l a b l e s o l a r energy equiv-alent to those at temperate l a t i t u d e s (Burns 1973-1974). Ranges of c l i m a t i c parameters across the study area are represented by selected data from two weather stations i n the study area, Inuvik and Tuktoyaktuk, and from an - 31 -a d d i t i o n a l s t a t i o n immediately east of the study area at Nicholson Point that is representative of conditions near the t i p of the Tuktoyaktuk Peninsula (Table I ) . Inuvik has a continental climate with the three summer months having mean d a i l y temperatures at or above 10.1°C (Atmos. Environ. Serv. 1982a). The two coastal stations of Tuktoyaktuk and Nicholson Point r e f l e c t maritime influences i n warmer winter and much cooler summer temperatures (Table I, Atmos. Environ. Serv. 1982a). The growing season averages 106 days, and the mean dates for l a s t and f i r s t f r o s t are i n mid-June and lat e August although f r o s t s may occur i n any month (Burns 1973-1974). Mean annual p r e c i p i t a t i o n and the proportion f a l l i n g as snow decrease northward i n the study area (Table I ) . Total p r e c i p i t a t i o n however fluctuates considerably from year to year with the c o e f f i c i e n t of v a r i a t i o n ranging from 15-20% in the south to near 50% on the Beaufort Sea coast (Burns .1973-1974). A c q u i s i t i o n of high q u a l i t y remote sensing data normally depends upon cl e a r weather conditions which are infrequent i n the study area during summer months. Fog in. p a r t i c u l a r reduces v i s i b i l i t y along c o a s t a l areas, and when advected inland by winds often becomes low-level blanket cloud. In a d d i t i o n , there i s extensive stratus and stratocumulus cloud cover i n summer and f a l l due to a p r e v a i l i n g thermal i n v e r s i o n structure and the "watery s t a t e " of the underlying ground surface (Burns 1973-1974). The mean proportion of sky ob-scured by clouds i n July for most of the study area i s seven to eight tenths, although there i s tremendous v a r i a b i l i t y from year to year i n t h i s figure (Hare & Thomas 1974, Burns 1973-1974). - 32 -Table I. Selected c l i m a t i c data for the Tuktoyaktuk Peninsula area, N.W.T. Station Inuvik Tuktoyaktuk Nicholson Point E l e v a t i o n (m a s l ) l 60 18 98 Mean Temperature, 1951-1980C"C) 1 - Annual - January - July - 9.8 -29.6 13.6 -10.9 -28.4 10.6 -12.0 -29.1 8.0 Average no. of f r o s t - f r e e days, 1941-1970 2 45 55 25 Degree-days over 5°C, 1941-1970 2 530 250 — Average P r e c i p i t a t i o n , 1951-1980 1 - Annual (mm) - June-August (mm) - Snowfall (cm) 266.1 100.7 176.6 137.6 60.6 65.2 109.4 57.7 40.6 ^•Atmos. Environ. Serv. 1982a,b 2Burns 1973-1974 - 33 -4. Vegetation 4.1 General Vegetation Zonations Most of the study area i s i n the treeless low a r c t i c where complex pat-terns of drainage and near-surface permafrost strongly c o n t r o l vegetation pattern and d i v e r s i t y (Lambert 1972). D r i e r s i t e s support various types of low shrub, moss and l i c h e n heaths while tussocky sedges (Eriophorum spp., Carex spp.), scrub willow (Salix spp.) and ground b i r c h (Betula glandulosa) t y p i c a l l y dominate poorly drained lower slopes and f l a t s . In the southern portions just i n s i d e the Forest Tundra Zone ( F i g . 3d, Rowe 1972), widely-spaced and stunted white spruce (Piaea glauoa) with a moss, l i c h e n and heath ground cover e x i s t i n sheltered l o c a t i o n s . C e r t a i n assemblages of plants are associated with s p e c i f i c landscape p o s i t i o n s , such as the slopes of pingos, surfaces of ice-wedge polygons, thermokarst lake s h o r e l i n e s , and brackish depressions along the seacoast (Lambert 1972, Corns 1974, Ito 1978). Mackay (1963) noted that a general sequence from north to south i s : tundra; tundra with scrub willow and ground b i r c h ; scrub willow and ground b i r c h ; woodland and tundra with much scrub willow and ground b i r c h , and open woodland. 4.2 Botanical Investigations The h i s t o r y of botanical c o l l e c t i n g i n the study area i s summarized by P o r s i l d (1947), Cody (1965) and P o r s i l d & Cody (1980). Vascular plant records also are given by these authors, and the l a t t e r reference includes recent d i s t r i b u t i o n a l and habitat information, and a h i e r a r c h i c i d e n t i f i c a -t i o n key based on diagnostic plant features. The vascular f l o r a of the en-t i r e Mackenzie Delta Reindeer Grazing Preserve i s r i c h , t o t a l l i n g over 420 - 34 -taxa (Cody 1965). Vascular species that have r e s t r i c t e d ranges and might be considered rare are given i n Cody's (1979) l i s t f o r the continental Northwest T e r r i t o r i e s . L i s t s of non-vascular plants i n the Reindeer Preserve are com-p i l e d for lichens (Ahti et al. 1973), mosses (Steere 1958, Holmen & Scotter 1971) and hepatics (Steere 1958, Scotter 1968). 4.3 E c o l o g i c a l Studies of Vegetation A comprehensive vegetation c l a s s i f i c a t i o n i n the study area has not been made, i n s p i t e of v i s i t s by numerous plant e c o l o g i s t s . On a regional l e v e l , Mackay (1963) b r i e f l y discussed vegetation units i n terms of h i s de-fined physiographic regions of the Mackenzie Delta area. Lambert (1972) further elaborated on the vegetation associations within the framework of Mackay's (1963) regions but contended that while extremes of wet and dry have unique vegetation assemblages, intergradations over short distances are so common that a vegetation c l a s s i f i c a t i o n on anything more than a broad scale is not p l a u s i b l e (Lambert 1972). Useful f i e l d d e s c r i p t i o n s of plant communi-t i e s are provided by P o r s i l d (1929, 1947), and Cody (1963), but neither of these authors employed numerical techniques. Vegetation communities were q u a n t i t a t i v e l y described at f i v e locations i n the open tundra east of the Mackenzie Delta, and on the basis of physio-gnomy and f l o r i s t i c composition, f i v e major community types ( t a l l shrub-herb; medium shrub-herb; low shrub-heath; herb-low shrub-heath; and herb) and eleven subgroups were p r o v i s i o n a l l y defined (Corns 1974). Ito (1978) tabu-lated percent cover data for vegetation quadrat studies on pingo slopes and raised-centred polygons near Tuktoyaktuk. Further plant cover data were sum-marized from locations at Urquhart Lake, Atkinson Point and near Tuktoyaktuk - 35 -( R i t c h i e 1974), and, i n conjunction with an environmental impact study around o i l d r i l l i n g s i t e s , from several locations on Richards Island and at the north end of Parsons Lake (Anon 1974). For a study of reindeer winter use of l i c h e n - r i c h woodlands west of S i t i d g i Lake, I n g l i s (1975a,b) used or d i n a t i o n analyses and tabular summaries to c l a s s i f y the vegetation types i n which he was working. Seven community types were recognized by I n g l i s (1975a), i n -cluding four open woodland (white s p r u c e - t a l l shrub-moss; heath-spruce-l i c h e n ; spruce-shrub-heath-lichen-moss; spruce-lichen-heath-sphagnum) and three non-wooded types (lichen-heath-shrub; lichen-heath; shrub-sedge-moss). Vegetation types of disturbed s i t e s have also been q u a n t i t a t i v e l y described at various locations i n the study area (Hernandez 1973, Younkin 1973, Strang 1973). Summer droughts occur only o c c a s i o n a l l y (Burns 1973-1974) but conditions have o c c a s i o n a l l y been s u f f i c i e n t l y dry to permit serious w i l d f i r e s . One f i r e extended over 2,000 sq km (Cody 1963, 1965, Wein 1976). F i r e e f f e c t s and p o s t - f i r e recovery sequences have been described for tundra (Wein and B l i s s 1973, Wein 1975, Haag 1974) and woodland (Black & B l i s s 1978) communi-t i e s within the study area. 5. Remote Sensing For a l l of the Canadian a r c t i c and subarctic there ex i s t s National A i r Photo L i b r a r y (NAPL) black and white panchromatic 22.9 x 22.9 cm format photo-coverage at about the 1:60,000 scale- and with 60% forward overlap. Photo-missions date from several d i f f e r e n t years and for most locations there i s multi-year coverage. Aside from t h e i r use i n the production of National Topographic Series base-maps, NAPL photographs have provided the basis f or - 36 -many studies i n the Mackenzie Delta region i n c l u d i n g the inventory and map-ping of pingos (Stager 1956, Mackay 1963), ground-ice slumps, g l a c i a l land-form features, percent cover by lakes and physiographic regions (Mackay 1963). Multidate NAPL photos have been used to determine rates of shoreline, erosion at Tuktoyaktuk (Rampton & Bouchard 1975) and to monitor pingo growth and the drainage and subsequent changes of near-coast thermokarst lakebeds (Mackay 1979). Vegetation mapping at 1:125,000 scale for the western h a l f of the study area was conducted s o l e l y using 1:60,000 NAPL Photos (Forest Man-agement I n s t i t u t e 1974, 1975). Medium-scale vegetation maps for Garry Island (Kerfoot 1969; 1:49,000), an a c t i v e portion of the Mackenzie Delta near Rein-deer Sta t i o n ( G i l l 1971; 1:16,000) and the v i c i n i t i e s of exploratory o i l d r i l l i n g r i g s on Richards Island and north of Parsons Lake (Anon 1974; v a r i -able scales, mostly about 1:54,000) were produced using NAPL photographs. Black and white photos were used to extend a p r o v i s i o n a l c l a s s i f i c a t i o n of tundra vegetation types developed on the ground to provide preliminary e s t i -mates of percent cover (Corns 1974). LANDSAT t e r r a i n studies of only a preliminary nature have been conducted in the Mackenzie Delta region. Using photographic transparencies of s a t e l -l i t e scenes, Sayn-Wittgenstein (1973) was able to recognize major physio-graphic zones, the lo c a t i o n of the t r e e l i n e and a number of w i l d f i r e , scars, i n c l u d i n g a large 1967 f i r e near Inuvik. Burned and unburned t e r r a i n near S i t i d g i Lake has been separated using density s l i c i n g techniques (Tarnocai & Thie 1974). Automatic c l a s s i f i c a t i o n of d i g i t a l LANDSAT data was used to define broad classes of vegetated land, unvegetated land and water bodies on Richards Island (Tarnocai & K r i s t o f 1976). - 37 -Offshore, considerable use i s presently being made of s a t e l l i t e and airborne remote sensing systems, inc l u d i n g radar, to study ice dynamics around Beaufort Sea o i l d r i l l i n g platforms (Dey et al. 1979, Dey 1980). - 38 -IV METHODS - 39 -1. Photo A c q u i s i t i o n 1.1 The Multistage Program The study reported here constitutes only a portion of a larger m u l t i -stage sampling program ( F i g . 4) i n the Tuktoyaktuk Peninsula area undertaken by the author and Dr. P.A. Murthal for the Department of Indian A f f a i r s and Northern Development and Canadian Reindeer Ltd. That program i s presently underway and w i l l terminate i n mid-1983 with the" production of a f i n a l re-port. The multistage work ( F i g . 4) involved analysis of remote sensing data acquired within a nested subsampling scheme using m u l t i s p e c t r a l scanner (MSS) images and photographs (black & white, normal colour and CIR films) at several d i f f e r e n t s c a l e s . Considerably more information about the earth resources of an area can be obtained on multistage images or photos than on those obtained at only one scale (Wiken 1980, Rubec 1982). The present study reports on findings based on 'Stage I I I ' and 'Ground-Truth' phases of the o v e r a l l program ( F i g . 4). 1.2 F l i g h t l i n e S e l e c t i o n At a broad scale of 1:500,000, twelve transects were chosen within the study area as being representative of general vegetation and t e r r a i n condi-tions ( F i g . 5, Table I I ) . Selection of transects was based on a number of c r i t e r i a : ( i ) a e r i a l observations, and study of medium- and large-scale ob-liq u e 35 mm photographs from June, 1980 o v e r f l i g h t s in a l i g h t a i r c r a f t on the northern Tuktoyaktuk Peninsula and i n the Tuktoyaktuk-Inuvik region; ( i i ) i F a c u l t y of Forestry, Univ. B r i t i s h Columbia. MULTISTAGE SAMPLING PROGRAM STAGE I 1 : 1 , 0 0 0 , 0 0 0 t o 1 : 5 0 0 , 0 0 0 -L A N D S A T s c e n e s ; v a r i o u s s u m m e r d a t e s , 1 9 7 2 - 1 9 8 0 . 6 0 , 0 0 0 - N A P L b l a c k a n d w h i t e s L e r e o - p h o t o g r a p l i s ; 1 9 7 0 a n d 1 9 7 3 5 0 , 0 0 0 - n o r m a l c o l o u r a n d C I R s t e r e o - p h o t o g r a p h s , a n d a i r b o r n e m u l t i s p e c t r a l s c a n n e r d a t a ; A u g u s t , 1 9 8 2 . 1 : 3 4 , 0 0 0 - C I R 70 mm s t e r e o - p h o t o g r a p h s ; ' A u g u s t , 1 9 8 0 . 1 : 1 , 4 0 0 t o 1 : 3 , 4 0 0 C I R 7 0 mm s t e r e o - p h o t o g r a p h A u g u s t , 1 9 8 0 . GROUND-TRUTH v e g e t a t l o n / s o i l s & p e r m a f r o s t , e n v i r o n m e n t a l f e a t u r e s ) Figure 4. A multistage sampling program to i n v e s t i g a t e r e i n d e e r rangeland. The present study deals w i t h r e s u l t s at 'Stage I I I ' and 'Ground-Truth' l e v e l s . 0 20 40 I I 1 km Figure 5. The areas occupied by the Mackenzie Delta reindeer herd, summer 1978 to f a l l 1982 (hatched l i n e s ) , and the l o c a t i o n of 13 transects. The herd summers generally east of - t r a n s e c t 7 throughout the t i p of the Peninsula, uses the c e n t r a l Peninsula as s p r i n g / f a l l range, and winters south and southeast of Tuktoyaktuk i n the hatched areas around transects 4 and 5. Transect 10 f a l l s j u s t outside the study area boundary and i s excluded from r e s u l t s reported on here. - 42 -Table I I . General location and approximate length of 12 transects in the study area. Transect no 1 Location 1 Aklavik mapsheec2 - W.of S i t i d g i Lake Co N t i p of unnamed lake - approx. 8 km. 2 Aklavik mapsheet - from Reindeer Depot NE past Scissor Lakes, West Round Lake - approx. 11 km. 3 Mackenzie Delta mapsheet - across narrowest portion of Richards Island, NW to SE from mouth of Burnt Creek to just short of large i s l a n d i n Mackenzie R. East Channel mouth - approx. 19 km. 4 Mackenzie Delta mapsheet - E from East Channel, where Holmes Creek meets the channel (cabin located at t h i s point a l s o ) , to Inuvik-Tuk main winter road - approx. 44 km. 5 Mackenzie Delta mapsheet - from Peninsula Point area, Beaufort Sea coast SSE to main peninsula on Eskimo Lakes, bi s e c t i n g winter road - approx. 41 km. 6 Mackenzie Delta/Stanton mapsheecs - SW to NE transect, endpoints at edges of 2 major lakes, centre l i n e of f l i g h t l i n e passes between 2 other major lakes ( i . e . , I t k r i l e k L., Other unnamed lake) - approx. 35 km. 7 Stanton mapsheet - SSE from Atkinson Pt W of McKinley Bay, bisec t i n g 2 major lakes and passing to the south shore of the Tuk Peninsula j u s t W of a major peninsula -approx. 40 km. 8 Cape Dalhousie/Stanton mapsheets - passing through narrowest part of Tuk Peninsula NW Co SE from Russell Inlet b i s e c t i n g major lakes, to S side bordering Liverpool Bay - approx. 15 km. 9 Stanton mapsheet - passing from S end of Rufus Lake due W to Liverpool Bay, b i s e c t i n g larger lake and Sanders Creek - approx. 22 km. 11 Stanton mapsheet -' NW to SE l i n e between two mid-sized lakes just S of major "arms" of Eskimo Lakes; just W of Kugalik River muth - approx. 10 km. 12 Mackenzie Delta/Aklavik mapsheets - passing from S shore of Eskimo L. SE between the large Urquhart L. and Old Man Lake, terminating at edge of other larger lake, after passing through tree l i n e - approx. 20 km. 13 Aklavik mapsheet - SW from Old Man L. to Stanley Point -approx. 21 km. ( t o t a l Line length approx. 292 km) las shown on F i g . 5. 2l:250,00O scale National Topographic Series mapsheets 107B ( A k l a v i k ) , C (Mackenzie Delta), D (Stanton), E (Cape Dalhousie). - 43 -using a v a i l a b l e equipment at the U n i v e r s i t y of B r i t i s h Columbia, preliminary o p t i c a l and d i g i t a l analyses and i n t e r p r e t a t i o n s of summer LANDSAT scenes covering the e n t i r e study area; ( i i i ) p r e - f i e l d observations and mapping on selected 1:60,000 NAPL black and white panchromatic photos, and ( i v ) d i s -cussions with the reindeer herd owner, William Nasogaluak, on recent past and proposed future seasonal movements of the herd. In e a r l y phases of the study, the herd owner expressed p a r t i c u l a r con-cern over the winter rangelands generally southeast, south and southwest of the town of Tuktoyaktuk as far as the Eskimo Lakes. This area had received heavy use by the reindeer i n recent winters ( F i g . 5) and being i n the v i c i n -i t y of town, i t has .been convenient and thus preferred by the herders. The area is bisected by the Tuktoyaktuk-Inuvik overland winter road which pro-vides general access and many sui t a b l e locations for l a t e winter round-ups and slaughters using portable abbatoir f a c i l i t i e s on sledges.. During s e l e c t i o n of f l i g h t l i n e s for large-scale photography, i n accordance with the herd owner's requests, no less than one-third of the line-km coverage was located along transects 4, 5 and 6 ( F i g . 5) i n t h i s area of i n t e r e s t , so that i t c onstituted an area of more intensive i n v e s t i g a t i o n . Along the twelve transects, midpoints of f o r t y - f o u r large-scale photo f l i g h t l i n e s were randomly located. An appropriate weighting was applied to locate over 33% of the line-km coverage within the more intensive study area southeast, south and southwest of Tuktoyaktuk ( F i g . 6). For ease of naviga-t i o n by the a e r i a l photography crew, ends of f l i g h t l i n e s were oriented at or adjacent to r e a d i l y i d e n t i f i a b l e features on the landscape such as a lake margin or i n l e t , a stream system or a prominent pingo. Transects and f l i g h t -l i n e s were mapped onto two i d e n t i c a l sets of 1:60,000 black and white NAPL 44 -F i g . 6. A p o r t i o n of the study area l o c a t i o n of f l i g h t l i n e s f o r 4 and 5. southwest of Tuktoyaktuk showing the random la r g e - s c a l e 70 mm photography along transects - 45 -photos and 1:250,000 National Topographic Series colour base-maps so that one set could be used by the a e r i a l photography crew while the second set was being c a r r i e d into the f i e l d by the ground-truth team. 1.3 Photographic Mission Transects and f l i g h t l i n e s were photographed from a Cessna 180 fixed-wing a i r c r a f t during August 5-8, 1980 under c l e a r sky conditions by P.G. Williams and an a s s i s t a n t , Integrated Resource Photography Ltd., Vancouver, B.C. Con-t r a c t u a l arrangements c a l l e d for standard s p e c i f i c a t i o n s (Interdepartmental Committee on A i r Surveys 1973, 1979) and the use of vertically-mounted wing-t i p cameras ( F i g . 7, Williams 1978) to obtain colour i n f r a r e d (CIR) stereo photo-coverage of transects and f l i g h t l i n e s at approximate scales of 1:34,000 and 1:2,000 r e s p e c t i v e l y . Kodak Aerochrome Type 2443 f i l m was exposed using two 76.2 mm f o c a l length Vinten 492 S/N cameras (1/1000 sec, / 2.0) f i r e d i n tandem by an intervalometer. Both cameras were f i t t e d with a yellow Wratten 12 f i l t e r normally required for CIR photography as well as a magenta colour compensating (CC20M) f i l t e r recommended p a r t i c u l a r l y for low a l t i t u d e CIR ap-p l i c a t i o n s (Interdepartmental Committee on A i r Surveys 1980). Together these f i l t e r s attenuate v i s i b l e r a d i a t i o n while allowing almost unimpeded trans-mission of near-infrared r a d i a t i o n thus enhancing the i n f r a r e d s e n s i t i v i t y of the f i l m (Fleming 1980). Film processing was subsequently carried, out by NAPL, Ottawa, Ontario. 2. Ground-Truth 2.1 P r e - f i e l d S i t e S e l e c t i o n P r i o r to the f i r s t f i e l d program of J u l y to mid-August, 1980, ground-truth s i t e s were chosen during i n t e r p r e t a t i o n of 1:60,000 NAPL photographs - 46 -F i g . 7. Twin 70 mm cameras are mounted on Cessna 180 wingtips for stereo-photography. 3 August, 1980. - 47 -of the transects. Sites were selected to be representative of major landscape and vegetation conditions occurring i n the photos. I d e n t i f i e d along f l i g h t -l i n e s , s i t e s were marked onto mylar-overlaid airphotos which were then c a r r i e d into the f i e l d . Before the second f i e l d program of July to ea r l y August, 1981, the 1980-acquired CIR photographs, superior i n q u a l i t y and r e s o l u t i o n to the 1:60,000 photos, were used for s i t e s e l e c t i o n . As before, s i t e s were chosen to represent p r e v a i l i n g landscape and vegetation conditions but i n the second year an e f f o r t was made to include any conditions not en-countered during the 1980 f i e l d program. For both programs attempts were made to locate s i t e s d i r e c t l y within f l i g h t l i n e paths. For the 1981 program, Cibachrcme 4X enlargement p r i n t s of 1:34,000 CIR photo-frames were obtained for portions of 24 of the 44 f l i g h t l i n e s . F l i g h t l i n e paths were pl o t t e d frame by frame onto the enlargements, and s i t e s were then chosen d i r e c t l y within areas of coverage (e.g., F i g . 8). The Cibachrome p r i n t s were f i t t e d i nto protective acetate sleeves that could be written on with black ink pens, and were c a r r i e d i n t o the f i e l d with other photo and map materials as used i n the f i r s t summer's program. 2.2 A i r Photo Annotation i n the F i e l d During the two f i e l d programs, every f l i g h t l i n e was traversed at a low a l t i t u d e (60 to 120 m agl) by h e l i c o p t e r or flo a t p l a n e . D e s c r i p t i v e notes on vegetation, topography, general physiognomy, etc... were noted with an i n -d e l i b l e ink pen d i r e c t l y onto the 1:60,000 or 1:34,000 a i r photos. Supple-mentary 35 mm oblique normal colour and CIR photos were acquired, with ap-proximate positions and angles annotated on the photos. - 48 -F i g . 8. Cibachrome 2X enlargement of 1:36,000 ( o r i g i n a l scale) 70 mm CIR photograph showing path of large-scale f l i g h t l i n e 4-2. Mixed shrub tundra communities impart a red-brown colour, ice-wedge polygons with high lichen-cover are grey-toned, and large sedge-rich wetlands f i l l i n g most of drained lake basin are pink to red. The lake was undrained i n 1956 1:60,000 NAPL a e r i a l photographs but drained sometime p r i o r to summer 1971 (Dr. J.R. Mackay, Dep. Geography, Univ. B r i t i s h Columbia, pers. comm.). (69°05,N, 133°54,W). 7 August, 1980. - 49 -2.3 Data C o l l e c t i o n at Ground-Truth Sites Using h e l i c o p t e r and floatplane support, 44 ground-truth locations ( F i g . 2) were v i s i t e d during summer, 1981 and 1982. At each l o c a t i o n , one to four s i t e s were examined. A two-person (R. Sims and M. Siltanen) ground-truth team c o l l e c t e d data on vegetation (Sims), general environment (Sims, Siltanen) and s o i l s (Siltanen) at 65 s i t e s i n 1980 and 47 s i t e s i n 1981. C r i t e r i a for s e l e c t i o n of a s i t e on the ground were: ( i ) that i t be located as accurately as possible within a f l i g h t l i n e path; ( i i ) that i t be repre-sentative of vegetation and landscape conditions i n the broader, general v i c i n i t y of the f l i g h t l i n e ; ( i i i ) that i t be a minimum of 0.1 ha and the veg-e t a t i o n composition be v i s u a l l y homogeneous, and (iv) that areas disturbed by man (e.g., cabin s i t e s , t e r r a i n v e h i c l e paths, etc.) be avoided. S i t e s were c l a s s i f i e d i n the f i e l d according to a h i e r a r c h i c a l vegeta-t i o n c l a s s i f i c a t i o n o r i g i n a l l y proposed for Alaska (Viereck and Dyrness 1980). The scheme has f i v e l e v e l s of r e s o l u t i o n that range from broad forma-tions ( I ; f o r e s t , tundra, shrubland, herbaceous vegetation, aquatic vegeta-tion) through intermediate l e v e l s (II to IV) based on general physiognomy. At the f i n e s t l e v e l (V) d i s c r e t e plant communites are named based on dominant vegetation i n p r i n c i p a l layers ( i . e . , tree, t a l l shrub, low shrub, herb, ground). Vegetation cover at each s i t e was estimated o c u l a r l y in nine to fourteen 1 m x 1 m quadrats by means of a twelve c l a s s scale: p < 1%, + = 1-5%, 1 = 6-15%, 2 = 16-25%, 3 = 26-35%, 4 = 36-45% 9 = 86-95%, 10 = 96-100%. Other vegetation i n the immediate v i c i n i t y was recorded as present. Voucher plant specimens were c o l l e c t e d and l a t e r i d e n t i f i e d and deposited i n herbaria - 50 -at the U n i v e r s i t y of B r i t i s h Columbia (UBC 1),'the National Museum of Natural Science (CAN, CANM, CANL), and the Great Lakes Forest Research Centre (SSMF). At 36 s i t e s where ±_ 20% cover by t e r r e s t r i a l lichens occurred, biomass c o l l e c t i o n s were made from fourteen to seventeen 20 cm x 20 cm divots within l i c h e n patches ( F i g . 9). C o l l e c t e d l i c h e n material was a i r - d r i e d , and l a t e r separated from l i t t e r and into l i v i n g and decadent portions using the c r i t e r i a of Pegau (1968) and P u l l i a i n e n (1971). Lower moribund portions of li c h e n t h a l l i were separated, using s c i s s o r s , from l i v i n g , actively-growing l i c h e n at the point where mottled darkenings and/or general d i s c o l o u r a t i o n indicated the onset of decomposition. Samples were oven-dried at 90°C to constant weight, and weighed on an e l e c t r i c a l pan balance. Lichen biomass, in kg.ha -!, was then estimated, employing the 1 m x 1 m quadrat measurements of percent cover t o t a l l e d for a l l l i c h e n species. General environment was described at s i t e s and included l o c a t i o n , aspect and slope p o s i t i o n data. In one representative 10 m x 10 m plot at each s i t e percent cover was estimated, using the twelve class scale, f o r : (1) woody standing dead; (2) graminoid^ standing dead; (3) bare mineral ground; (4) bare organic ground; (5) t o t a l shrub; (6) t o t a l graminoid; (7) t o t a l l i c h e n vegetation, and (8) t o t a l open water. Ground photographs, using both normal colour and CIR 35 mm f i l m , were obtained at a l l s i t e s . Depth to permafrost was measured at ten locations at each s i t e and the s o i l was t e n t a t i v e l y c l a s s i f i e d (Canada S o i l Survey Committee, Subcommittee ^abbreviations for herbaria follow Woodland (1980). 2'graminoid' includes grass and g r a s s - l i k e vascular plant taxa belonging to Gvamineae, Cypevaceae and Juncaaeaee. - 51 -F i g . 9. C o l l e c t i o n of a 20 cm x 20 cm l i c h e n d i v o t to estimate l i c h e n standing crop at s e l e c t e d ground-truth s i t e s . 21 J u l y , 1981. - 52 -on S o i l C l a s s i f i c a t i o n 1978) based only on act i v e layer s o i l p r o f i l e s from one representative s o i l p i t dug at each s i t e . S o i l was sampled from the top 10 cm (minus l i t t e r layers) and, when mineral s o i l was encountered at the base of s o i l p i t s , from the bottom 10 cm. S o i l samples were frozen within 20 h of c o l l e c t i o n u n t i l mixed and a i r - d r i e d for analyses. 'Top' samples were mechanically analysed: gravel f r a c t i o n s were seived, and sand, s i l t and c l a y f r a c t i o n s were hydrometrically separated using the Bouyoucos method (Anon 1979). Standard a n a l y t i c a l methods (Anon 1979) were followed to determine d e t a i l e d chemistry of 'top' samples: s o i l pH, ca t i o n exchange capacity, organic and inorganic carbon, t o t a l K j e l d a h l nitrogen, potassium, magnesium, sodium and pyrophosphate-extractable i r o n and manganese. 'Bottom' samples were only analyzed for organic and inorganic carbon. 2.4 Numerical Analysis of Ground-Truth Data Vegetation, general environment and s o i l ground-truth data were placed on computer f i l e s and analyzed at the U n i v e r s i t y of B r i t i s h Columbia's Com-puting Centre. A l l o r i g i n a l datasets used i n the study are stored on stand-ard 9-track computer tapes, and copies along with documentation have been lodged at two l o c a t i o n s : (1) Computing Services, Faculty of Forestry, Uni-v e r s i t y of B r i t i s h Columbia, Vancouver, B.C., and (2) Computing Services, Great Lakes Forest Research Centre, Canadian Forestry Service, Sault Ste. Marie, Ont. A p o l y t h e t i c d e v i s i v e c l a s s i f i c a t i o n technique named two-way i n d i c a t o r species analysis or TWINSPAN ( H i l l 1979, Gauch & Whittaker 1981) was used to p a r t i t i o n s i t e s , based on mean vegetation cover at s i t e s , among a few major groupings. TWINSPAN ordinates data by a r e c i p r o c a l averaging algorithm ( H i l l 1973), which emphasizes species on the ordination's f i r s t axis to p o l a r i z e - 53 -s i t e s , then i n t e r a t i v e l y divides s i t e c l u s t e r s , r e p o l a r i z e s species and re-c l a s s i f i e s s i t e c l u s t e r s u n t i l each has no more than a chosen minimum number of members ( H i l l 1979, Gauch 1982). A corresponding species c l a s s i f i c a t i o n i s produced and the sample and species h i e r a r c h i c a l c l a s s i f i c a t i o n s are then used together to produce an arranged data matrix. TWINSPAN i s a FORTRAN pro-gram i n the p u b l i c l y - a v a i l a b l e Cornell Ecology Program serie s (Gauch 1982). Although the program permits the user to choose c e r t a i n options, default values supplied by TWINSPAN were used i n present analyses. TWINSPAN's computer output includes a l i s t i n g of the successive dichoto-mizations that r e s u l t from the program's i t e r a t i v e r e c i p r o c a l averaging steps. For each dichotomization, species c o n t r i b u t i n g most to the d i v i s i o n (termed " i n d i c a t o r " and " p r e f e r e n t i a l " species)•and those s i t e s assigned to each branch of the d i v i s i o n are tabulated ( H i l l 1979). This computer output can then be used to plot by hand the dichotomizations as branches of a dendrogram, a method which e f f i c i e n t l y summarizes and displays the c l a s s i f i -cation r e s u l t s . Summary s t a t i s t i c s were obtained on general environment and s o i l ground-truth data using UBC ISP, an i n t e r a c t i v e s t a t i s t i c s package (Kita & T e n i s c i 1978). 3. I n t e r p r e t a t i o n and Analysis of Large-Scale A i r Photographs 3.1 Summaries by Reindeer Management Zones Fi l m r o l l s were mounted as 'rig h t ' and ' l e f t ' pairs on a h o r i z o n t a l l i g h t t able, annotated by f l i g h t l i n e and frame number using the method des-cribed by Goba et al. (1982), and viewed frame by frame with a 2X pocket stereoscope. For each stereo photo-pair a number of measurements (Table III) - 54 -Table I I I . Measurements recorded for each f l i g h t l i n e photo-frame. 1. Technical Information: 1. transect, f l i g h t l i n e & photo-frame number 2. percent forward o v e r l a p 1 3. percent crab (deviation from the mid-line of forward d i r e c t i o n ) ^ 4. photo scale (estimated from a i r c r a f t ' s barometric altimeter) 2. General Cover Features: 1. open water (e.g. lakes, streams, p o o l s ) 2 2. ice-wedge polygons (patterned ground) 2 3. number of ice-wedge polygons 4. t e r r a i n disturbance by vehicles ( i ) cover 2 ( i i ) l e v e l of damage-^ 5. l i c h e n ( i ) c over 2 ( i i ) t y p e 4 : I. patchy white l i c h e n cover and bare sandy patches on uplands and upper slopes; mixed with Dvyas} Salix, Betula, low herbaceous vegetation which impart pink/red component to colour; I I . patchy white l i c h e n cover on lower slopes and f l a t s ; mixed with brown mosses, including Sphagnum spp., bare organic surfaces, which impart dark component to colour; also, Salix, Betula, low herbaceous vegetation which impart pink/red component to colour; I I I . patchy to continuous white l i c h e n cover on f i n e l y hummocky, cryoturbated, mainly f l a t surfaces, e s p e c i a l l y raised-centred ice-wedge polygon systems. Ameasured to nearest 1 mm with a r u l e r , and converted to percent values. 2percent cover measured within 80% non-forward-overlap portion of photo-frame using a 100-cell gridded acetate overlay. Prefer to Table XV for s c a l e . 4 o n l y recorded for the dominant l i c h e n type i n the photo-frame. - 55 -were t a l l i e d d i r e c t l y onto FORTRAN coding forms. Measurements of 'general cover features' (Table III) were made i n the 80% non-forward-overlap region of photo-frames. Side-overlap by the two cameras ranged from 95% to 100% but measurements from within the small non-side-overlap portion of the 80% non-forward-overlap region were standardly acquired from ' r i g h t ' f i l m r o l l s . Record manipulation and s t a t i s t i c a l summaries were subsequently c a r r i e d out using MIDAS, the Michigan I n t e r a c t i v e Data Analysis System (Fox & Guire 1976) at the U n i v e r s i t y of B r i t i s h Columbia's Computing Centre. The c o n f i g -uration of transects and f l i g h t l i n e s allowed preliminary data summary on a regional basis within 'reindeer management zones'. The zones were adapted from Houseknecht (1981) and represent E c o d i s t r i c t s , or geographical sub-d i v i s i o n s of E c o d i s t r i c t s (Table IV, F i g . 10). Two of these zones^C and D, c o n s t i t u t e d the area of more intensive study, where a minimum of one-third of the line-km coverage was located. 3.2 Microdensitometric Studies of 'Lichen Types' 'Lichen Types' were interpreted (Table III) from the CIR photographs. The dominant Lichen Type i n a photo-frame was the condition that accounted for greater than 50% of the land area. . Lichen Types that were not dominant were ignored. They often covered small land areas only or occurred near photo-edges where v i g n e t t i n g and r a d i a l d i s t o r t i o n made assignment among Lichen Types more d i f f i c u l t . In attempting to c o r r e l a t e dominant Lichen Types to ground-truth studies, microdensitometric readings were made of l i c h e n patches on the o r i g -i n a l ' r i g h t ' f i l m r o l l s . The photo-frames sub-sampled for microdensitometric study had to have an appreciable l i c h e n cover, at least 5%. As w e l l , the - 56 -T a b l e IV. Summary d a t a on r e i n d e e r management zones and c o r r e l a t i o n w i t h E c o r e g i o n and E c o d i s t r i c t map u n i t s . R e i n d e e r management zone^ E c o r e g i o n ^ E c o d i s t r i c t ^ E c o d i s t r i c t s u b - a r e a Approx. t o t * l a n d a r e a ( s q i l km)3 P e r c e n t w a t e r ^ O r i g i n a l t r a n s e c t no.4 A Mackenzie C o a s t a l P l a i n T u k t o y a k t u k P e n i n s u l a (203) — 1,943 28 7,8 B L i v e r p o o l C o a s t a l P l a i n R i c h a r d s I s l a n d (301) — 1,214 25 3 C rf Eskimo Lakes (302) w. T u k t o y a k t u k P e n i n s u l a 2,208 26 5,6 D l l i t P a r s o n s Lake 1,631 24 4 E l l i i S. Eskimo Lake9 5,531 18 9, 11-13 F Anderson C o a s t a l P l a i n C a r i b o u H i l l s (403) — 1,366 9 2 G W o l v e r i n e R i v e r N o r r i s Creek (501) — 512 10 1 P r e f e r t o map, F i g . 10. ^ E c o r e g i o n s , mapped at about 1:1 m i l l i o n are c h a r a c t e r i z e d by d i s t i n c t i v e e c o l o g i c a l r e s p o n s e t o c l i m a t e as e x p r e s s e d by v e g e t a t i o n , s o i l s , w a t e r , f a u n a , e t c . E c o d i s t r i c t s , mapped at 1:500,000 t o 1:125,000 are p a r t s o f e c o r e g i o n s c h a r a c t e r i z e d by d i s t i n c t i v e p a t t e r n s o f r e l i e f , g e o l o g y , geomorphology, v e g e t a t i o n , s o i l s , water and fauna ( d e f i n i t i o n s a f t e r Wiken 1980). For E c o d i s t r i c t s , numbers i n parentheses' are Houseknecht's (1981) map u n i t numbers. -'estimated from dot g r i d (2.48 d o t s / s q cm) o v e r l a y s on N a t i o n a l T o p o g r a p h i c S e r i e s 1:250,000 mapsheets. ^ r e f e r t o map, F i g . 5. F i g . 10. Map of the 7 r e i n d e e r management zones d e f i n e d i n the study area. - 58 -Lichen Type had to be represented at or near the photo-centre, to avoid e f f e c t s of lens f a l l o f f that occur at the periphery ( E v e r i t t et al. 1980). Using a Macbeth TR524 transmission/reflectance densitometer i n transmission mode, yellow, magenta, cyan and t o t a l dye layer d e n s i t i e s on the CIR f i l m were measured, r e s p e c t i v e l y , by blue (Wratten #94), green (Wratten #93), red (Wratten #92) and white (Wratten #106) f i l t e r s . A 1 mm c i r c u l a r aperture with an approximate ground r e s o l u t i o n on 1:2,000 photographs of 3.1 sq m was used for spot readings over l i c h e n patches. The r e s u l t s were placed on computer f i l e s and analyzed at the U n i v e r s i t y of B r i t i s h Columbia's Computing Centre. To help determine i f Lichen Types could be r e a d i l y separated, standard analysis of variance and Duncan's new multiple range test (Sokal & Rohlf 1969) were employed to test for s i g n i f i -cant d i f f e r e n c e s among the microdensitometric spot readings of l i c h e n patches. A stepwise Linear Discriminant Function (LDF) analysis (Jennrich & Sampson 1977) was c a r r i e d out (1) to rank the dye layer density v a r i a b l e s that best described the three a priori groups of Lichen Types i n descending order of importance, and (2) to generate p r o b a b i l i t y groups of Lichen Types with the same structure as the o r i g i n a l input. - 59 -V RESULTS - 60 -1. Ground Truth 1.1 C l a s s i f i c a t i o n of Four Vegetation Groups A dendrogram was produced from the TWINSPAN analysis of vegetation cover data at 112 ground-truth s i t e s ( F i g . 11). At the f i r s t r e c i p r o c a l averaging d i v i s i o n (cut l e v e l I) two groups are formed, one proceeding to further div-i s i o n s on the l e f t side of the dendrogram, the other, l a b e l l e d as 'D' at the bottom r i g h t of F i g . 11 and c o n s i s t i n g of fourteen s i t e s , proceeding to d i v -isio n s on the r i g h t . Other major c l u s t e r s r e s u l t i n g from l e f t side d i v i s i o n s at cut l e v e l s II and III are l a b e l l e d A, B and C and c o n s i s t , r e s p e c t i v e l y , of 32, 25 and 41 s i t e s . Based on vegetation cover, the four major c l u s t e r s i n d i c a t e d i n F i g . 11 represent aggregates of broadly s i m i l a r vegetation types and are here r e f e r r e d to as "vegetation groups". Although not s p e c i f i c a l l y dealt with here, plant assemblages may be de-fined at f i n e r levels of the dendrogram. At cut l e v e l V, for example, the eighteen c l u s t e r s ( F i g . 11) correspond to, with only a few exceptions, vege-t a t i o n communities that can be d i s t i n c t l y separated based on cover of the dominant vegetation species (Table V). Most of the vegetation communities given i n Table V have been previously l i s t e d by Viereck & Dyrness (1980) as occurring from mainly t r e e l e s s tundra areas in nearby Alaska. The four vegetation groups (A, B, C and D) correspond c l o s e l y to i n t e r -mediate l e v e l s of the h i e r a r c h i c a l vegetation c l a s s i f i c a t i o n applied i n the f i e l d (Table V). Group A includes dry upland plant communities (e.g. F i g 12a): Dryas open, and closed mat and cushion tundra, mixed shrub tundra and an open, low alder-grass community t r a n s i t i o n a l to group B. Group B includes mesic s i t e s of intermediate and lower slope positions (e.g. F i g . 12 b) : open, low alder tundra, b i r c h and ericaceous shrub-sphagnum tundra, c o n i f e r CUT LEVEL Cladina stellaris Arctostaphylos rubra Ledum decumbens Cetraria cucullata Salix alaxensis Lupinus arcticus Dryas integrifolia Carex aquatilis Eriophorum angustifolium 0/1 Cladina stellaris C. rangiferina Ledum decumbens Rubus chamaemorus Carex atrofusca IV _ _ V _ F i g . 11. 0/1 -1/0 Dryas integrifolia Salix reticulata Vaccinium uliginosum Carex lugensj Pyrola grandiflora Tomenthypnum nitens] Aulacomnium palustre Alnus crispa Pohlia nutans Salix alaxensis Ledum decumbens Cladonia pleurota Sphagnum fuscum -1/0 n= X2) Ct) ® © (§> (4) (10) (15) (1) J Bryoria nitidula Cinclidium arcticum Scorpidium scorpioides -1/0 Aulacomnium lurgidum Carex lugens Cladonia cornuta -1/0 A (n=32) (6) © ® ( § ) (1) (7) (5) (12) V J Cladina rangiferina C. stellaris Bryoria nitidula Vaccinium uliginosum Hierochloe pauciflora 1/2 B (n=25) <3) © (4) (20) (9) (8) V , J Arctophila fulva Q/1 -1/0 -1/0 C (n-41) (1) <§) % ® (3) (3) (4) D (n= 14) ® (3) Two-way i n d i c a t o r species a n a l y s i s (TWINSPAN) dendrogram of 112 ground-truth s i t e s based on vegeta t i o n cover of 420 s p e c i e s , Tuktoyaktuk P e n i n s u l a area, N.W.T. Four major c l u s t e r s (A,B,C,D) are d e l i n e a t e d , and f o r cut l e v e l s I , I I and I I I , i n d i c a t o r s p e c i e s and t h e i r d e c i s i o n r u l e s are given. At c u t - l e v e l V 18 v e g e t a t i o n a s s o c i a t i o n s are separated. Tflble v- ^^:art^l ^ " ' [ ^ ^ ^ T - • N O R T H W E S T T E R R I T O R I E A < A D A » - « L e v e l I 1. F o r e s t 2. Tundra L e v e l I I L e v e l I I I L e v e l IV L e v e l V 1 A. C o n i f e r F o r e s t TWINSPAN TWINSPAN v e g e t a t i o n c ut l e v e l group V membership^ membership 2 ( 3 ) C o n i f e r Woodland A. Sedge-grass ( 1 ) Wet sedge-grass Tundra C. Tussock Tundra D. Shrub Tundra ( 3 ) Sedge-shrub (2) Sedge t u s s o c k -shrub (2) B i r c h and e r i c a c e o u s shrubs White spruce Hoea glauca/Cladina spp. Wet sedge meadow Carex aquatilia/Seorpidium eaorpioidea Carex aquatilia - Eriophorum anguatifolium Carex aquatilia - C. rotundata C. chordorhizza Carex rariflora Wet sedge-herb meadow Carex aquatilia - Potentilla paluetria ' S e d g e - w i l l o w Carex aquatilia - Salix spp. Eriophorum vaginatum - Ledum deoumbena - Vaocinium vitia-idaea Eriophorum vaginatum - Betula glanduloaa - ledum decumbena B i r c h and e r i c a c e o u s Betula glanduloaa - Ledum deoumbena shrubs - sphagnum - Vacoinium spp.- Rubue chamaemorua/'Sphagnum spp. Sedge t u s s o c k -e r i c a c e o u s shrub Sedge t u s s o c k - m i x e d shrub U n d i f f e r e n t i a t e d u n d e r s t o r y Betula glanduloaa - Ledum deoumbena - Vaooinium spp./ Cladina spp. - Cetraria spp. Betula glanduloaa - Ledum decumbena 8 . 15 16 17 \l> 18 11 10 11 10 ON ro L e v e l I L e v e l I I L e v e l I I I L e v e l IV , / 2. Tundra (3) Mixed shrub U n d i f f e r e n t i a t e d ( c o n t ' d ) u n d e r s t o r y (4) Open low a l d e r Low a l d e r g r a s s Low alder-sphagnum D. Shrub (A) Qpen low a l d e r U n d i f f e r e n t i a t e d E. Mat and Cushion Tundra (1) Open mat and Dryas - l i c h e n cush i on (2) and c u s h i o n Dryas L i c h e n TWINSPAN v e g e t a t i o n • group L e v e l V J membership* Salix glauaa - S. alaxensis -Betula glanduloaa - Bnpetrum nigrum ssp hermaphroditumj'Cetraria spp. Salix alaxensis - Betula glanduloaa Salix pulchra - Betula glanduloaa -Pyrola grandiflora/Cetraria spp. Salix glauaa - Betula glanduloaa -Briophorum vaginatum/Sphagnum spp. Alnus orispa - Salix alaxensis/ Calamagrostie neglecta Alnua crispa - Salix spp./ Sphagnum spp. Alnua crispa - Betula glanduloaa -Salix spp./ Ledum decumbens/ Sphagnum spp. Alnua crispa — Salix glauaa/ Carex lugena/Tomenthypnum niteno Dryas integrifolia/Cetraria cucullata - o t h e r l i c h e n s Dryas integrifolia - Arctoataphyloa rubra Dryas integrifolia - Salix reticulata Cladina spp. - Cetraria spp./ Eriophorum vaginatum L e v e l I 2. Tundra ( c o n t ' d . ) 5. A q u a t i c L e v e l I I L e v e l I I I A. Freshwater (1) Ponds and la k e s L e v e l IV Emergent v e g e t a t i o n L e v e l V 1 TWINSPAN TWINSPAN v e g e t a t i o n c u t l e v e l group V membership^ membership^ Cladina spp./ Ledum decumbena-Betula glandulosa - o t h e r l i c h e n s Cetraria spp./ Ledum decumbens -Betula glandulosa Cladina spp. - Cetraria spp. - o t h e r l i c h e n s Arctophila fulva - Eriophorwn anguetifolium 12 12 13 18 ' S p e c i e s i n community names s e p a r a t e d by hyphens are i n the.same l a y e r ; a s l a s h between s p e c i e s i n d i c a t e s a change i n l a y e r ( t r e e l a y e r t o shrub l a y e r , shrub l a y e r t o herb l a y e r , e t c . ) ^as g i v e n i n F i g . 11. 3 n o f i e l d s i t e s were s t u d i e d ; i d e n t i f i e d from l a r g e - s c a l e CIR a i r photographs, and g e n e r a l ground o b s e r v a t i o n s . - 65 -F i g . 12. Normal colour 35 mm photographs of vegetation group A and B examples. (a) An example of vegetation group A: A mixed shrub tundra community on an upper slope p o s i t i o n ( s i t e 80-43, f l i g h t l i n e 6-1; 69°26 IN, 132°23'W). Vegetation dominants, with cover-class i n parentheses as given i n Appendix I, are: Salix alaxensis (5), Empetrum nigrum spp. hermaphroditum (5), Betula glandulosa (4), Salix arbusculoides (4), Lupinus arcticus (3), Pohlia nutans (3), Pyvola grandiflora (3). T o t a l l i c h e n cover 6.5%. 31 J u l y , 1980. (b) An example of vegetation group B: An open white spruce woodland community on a mid-slope p o s i t i o n ( s i t e 81-71, f l i g h t l i n e 13-2; 68°49'N, 132°46'W). Vegetation dominants are: Picea glauca (4), Tomenthypnum nitens (4), Alnus cvispa (3), Arctostaphylos rubra (3), Cladina stellaris (3), Ledum decumbens (3), Vaccinium uliginosum (3). T o t a l l i c h e n cover 15.2%. Range pole i s marked i n 10 cm i n t e r v a l s . 18 J u l y , 1981. - 67 -woodland, and mixed shrub tundra with u n d i f f e r e n t i a t e d understory. Group C in-cludes dry to raesic s i t e s c h a r a c t e r i s t i c of f l a t l a n d s , e s p e c i a l l y pat-terned ground (e.g. F i g . 13 a and b), and includes: sedge tussock-shrub tundra, bi r c h and ericaceous shrub with u n d i f f e r e n t i a t e d understory, and l i c h e n closed mat and cushion tundra. Group D consists of open fen and sha l -low marsh wetlands t y p i c a l l y associated with standing water or with water tables near the surface (e.g. F i g . 14): wet sedge-grass tundra, and emergent vegetation along ponds and lakes. On th i s basis alone, a preliminary en-vironmental i n t e r p r e t a t i o n of the TWINSPAN ordering across the four vegeta-t i o n groups A to D can be hypothesized: a dry-to-wet s o i l moisture gradient. C a l c u l a t i o n of 'i n d i c a t o r species' and 'decision rules' for t h e i r use are unique features of TWINSPAN ( H i l l 1979). L i s t e d i n general order of e f f e c t i v e n e s s , i n d i c a t o r species are given i n F i g . 11 just above the dendro-gram branch to which they p e r t a i n . As given i n the present study, i n d i c a t o r species are those that best d i f f e r e n t i a t e d i v i s i o n s based just on presence or absence c r i t e r i a , and they provide a technique for assigning future 'unknown' s i t e s among the four vegetation groups. For example, at cut l e v e l I, the presence at a s i t e of Carex aquatilis and Eriophorum angustifolium3 two plants ubiquitous i n study area wetlands, indicates that the c l a s s i f i e r move to the right side of the dendrogram. In a s i m i l a r manner, at cut l e v e l I the occurrence of Cladina stellaris3 Arctostaphylos rubra. Ledum decumbens and Cetraria cucullata moves the c l a s s i f i e r to the l e f t side. Not a l l the i n d i -cator species may be present, or i n d i c a t o r species from l e f t side and r i g h t side l i s t s at a d i v i s i o n may both be found at an 'unknown' s i t e . Such s i t u -ations are solved by applying the d e c i s i o n r u l e , found just below d i v i s i o n points of the dendrogram ( F i g . 11). For the '0/1' of cut l e v e l I, summing +1 - 68 -Fig. 13. Example of vegetation group C. (a) Normal colour 35 mm ground photograph of lichen closed mat and cushion tundra on raised-centred ice-wedge polygons (site 81-110, f l i g h t l i n e 4-6; 69°07'N, 133°23'W). Vegetation dominants are: Cladina rangiferina (5), C. s t e l l a r i s (5), Betula glandulosa (4), Bryoria nitidula (4), Cetraria aucullata (4), Ledum decumbens (4), Alectoria ochroleuca (3), Cladina mitis (3), Rubus chamaemorus (3). Total lichen cover 71.3%. 2 August, 1981. (b) CIR 35 mm ground photograph of the same si t e . Colour infrared film en-hances the lichen cover because of the particularly high near-infrared re-flectance by the te r r e s t r i a l lichens. 2 August, 1981. - 69 -- 70 -F i g . 14. Example of ve g e t a t i o n group D. Normal colour ground photograph of wet sedge meadow tundra ( s i t e 81-105, f l i g h t l i n e 12-3; 69°01,N, 132°12,W). Vegetation dominants are: Carex aquatilis ( 5 ) , Scorpidium scorpioides ( 5 ) , Bryion pseudotriquetrum ( 3 ) , E. russeolum var. albzdum (3). Open standing water covers about 5% and ground l i c h e n s are absent. Note the orange a l g a l mat i n the background c o n t a i n i n g mainly blue-green genera: Nostoa, Osailla-toria, Anabaena, Merismopedia and Coelosphaeri-um ( i d e n t i f i e d by Dr. J . S t e i n , Dep. Botany, Univ. B r i t i s h Columbia). 30 J u l y , 1981. - 71 -values for each r i g h t side i n d i c a t o r species present, and -1 for l e f t side species present, a score i s obtained that i s e i t h e r 0 or less (the c l a s s i f i e r moves to the l e f t ) or 1 or greater (the c l a s s i f i e r moves to the r i g h t ) . Note that the de c i s i o n rule has changed to '-1/0' for the three d i v i s i o n s at cut l e v e l III ( F i g . 11). The i n d i c a t o r species are t a l l i e d i n an i d e n t i c a l manner, but a score of -1 or less moves the c l a s s i f i e r to the l e f t , and a score of 0 or greater moves the c l a s s i f i e r to the r i g h t . In the present case, a c l a s s i f i e r using the d e c i s i o n rules i n the f i e l d can assign without d i f f i c u l t y 'unknown' s i t e s among the four vegetation groups based on a knowl-edge of the presence or absence of only a small number of i n d i c a t o r plant species. P r e f e r e n t i a l species, those e x h i b i t i n g c l e a r e c o l o g i c a l preferences, and i n d i c a t o r species are used by TWINSPAN to construct ordered two-way tables ( H i l l 1979, Gauch 1982). The tabular arrangements c l o s e l y resemble the hand-sorted Braun-Blanquet releve tables as described by Mueller-Dombois & Ellenburg (1974, chapt. 9), except the l a t t e r approach, while i t concen-trates matrix values i n blocks for strongly p r e f e r e n t i a l species, places weakly p r e f e r e n t i a l species i n a separate, f i n a l l i s t with no i n d i c a t i o n of t h e i r place i n the species sequence. In the tabular ordering here (Appendix I) the TWINSPAN species sequence i s provided, however s i t e data has been sum-marized according to the four vegetation groups. Appreciation of the re-l a t i o n s h i p of some species sequences is lo s t i n t h i s summary method but general trends can be seen. For example, the f i r s t species "block", from the beginning of the l i s t to the f i r s t broken h o r i z o n t a l l i n e , consists of taxa with obvious a f f i n i t i e s for vegetation group B. Subsequent species blocks can s i m i l a r l y be c o r r e l a t e d with other groups, or combinations of groups. - 72 -Species near one another and not separated by a d i v i s i o n i n the TWINSPAN table can be expected to be found i n the f i e l d at the same or s i m i l a r s i t e s . Conversely, i t i s less l i k e l y that plant taxa further apart i n Appendix I would be found growing at the same or s i m i l a r s i t e s . Of the 420 plant taxa encountered i n the f i e l d program (Appendix I ) , 48.6% are vascular plants, while bryophytes and lichens represent 26.7% and 24.8% r e s p e c t i v e l y . Of the vascular taxa, approximately h a l f are broad-leaved herbs, while trees and woody shrubs, and graminoids each account for one-quarter (Table V I ) . Number of taxa i s highest i n vegetation group A (275 taxa), followed by groups C (235), B (225) and D (148). For a l l four groups, percent of taxa among the s i x s t r u c t u r a l categories mirrors that for the t o t a l , with two main exceptions: (1) broad-leaved herbs form a higher per-centage, and mosses a lower percentage of the t o t a l taxa for group A; and s i m i l a r l y (2) mosses comprise a greater, and lichens a lesser proportion of the t o t a l taxa for group D (Table V I ) . Only 70 taxa, or 16.7% of the t o t a l f l o r a , are common to a l l four groups. Three vascular and t h i r t y - t h r e e non-vascular taxa are newly reported for the Mackenzie Delta Reindeer Preserve and they are indicated i n Appendix I. The three vascular taxa - Carex saxatilis var. rhomalea, Oxytropis various and Salix planifolid var. planifoila - are extensions to t h e i r present ranges as given by P o r s i l d & Cody (1980). Barbilophozia attenuata, Cladopodiella fluitanSj Lophozia alpestris, and Saapania irrigua are additions to Scotter's (1968) l i s t of hepatics. Fourteen moss taxa named here were not given by Holmen & Scotter (1971), with four species - Dioranum brevifoliwn, D. leio-neurorij Drepanoaladus oapillifolius and Rypnum plicatulum - new to the North-- 73 -Table VI. The percent of t o t a l plant taxa encountered i n the ground-truth studies, summarized for each vegetation group by s i x s t r u c t u r a l c a t egories. Vegetation Group S t r u c t u r a l category-*- A B C D t o t a l 1. trees and woody shrubs 11.3 13.8 11.1 12.2 9.8 2. broad-leaved herbs 27.6 2*1.3 20.4 20.3 26.4 3. graminoids 14.9 10.2 15.3 18.2 12.4 ( t o t a l vascular taxa) (53.8) (45.3) (46.8) (50.7) (48.6) 4. hepatics 3.3 4.4 6.8 6.8 6.7 5. mosses 14.5 21.8 17.0 30.4 20.0 6. lichens 28.4 28.4 29.4 12.2 24.8 ( t o t a l non-vascular taxa) (46.2) (54.6) (53.2) (49.4) (51.5) t o t a l no. of taxa 275 225 235 148 420 ^•corresponds to categories i n Appendix I. - 74 -west T e r r i t o r i e s , according to the l i s t of Ireland et al. (1980). F i f t e e n l i c h e n species were not previously reported by Ahti et al. (1973). 1.2 C o r r e l a t i o n of Environmental Parameters to the Four Vegetation Groups Slope p o s i t i o n measurements at the 112 s i t e s support the notion of a dry-to-wet s o i l moisture gradient operating across the four vegetation groups (Table V I I ) . The majority (68.7%) of vegetation group A s i t e s occupied better-drained locations or h i l l c r e s t s , or upper and middle slopes. Three-quarters (72.0%) of group B s i t e s were on slopes, and a sim-i l a r proportion (73.1%) of group C s i t e s were on lower or toe slopes, or on l e v e l ground. The open wetlands, group D, were a l l associated with low or f l a t t e r r a i n . General aspect showed no c l e a r c o r r e l a t i o n s other than the a f f i n i t y of group C and D s i t e s for l e v e l locations (Table V I I ) . Although general environment measurements i n 10 m x 10 m plots were of only broad physiognomic features, some notable comparisons among the four vegetation groups can be made (Table V I I I ) . Standing dead mat e r i a l , both woody and graminoid, covers about three times the surface area i n group D s i t e s as i n others. While the greater proportion of standing dead i n groups A, B and C was accounted for by woody ma t e r i a l , i n group D i t was p r i m a r i l y graminoid. Unvegetated ground covered a small area at most s i t e s , with bare organic surfaces more prevalent than bare mineral ground, p a r t i c u l a r l y for those group C and D s i t e s on low landscape p o s i t i o n s . T o t a l tree and woody shrub cover, with an o v e r a l l mean of 26.8%, was gener-a l l y high, but as indicated by the large standard deviations, v a r i a b l e (Table V I I I ) . Graminoid cover was high i n group D but l i c h e n cover was ab-sent i n the same group. Standard deviations for l i c h e n cover i n d i c a t e con-- 75 -Table VII. Slope p o s i t i o n and aspect of ground-truth s i t e s summarized by the four vegetation groups (A, B, C and D). Percent of s i t e s A B C D (n = 32) (n = 25) (n = 41) (n = 14) SLOPE POSITION 1 cres t 25.0 8.0 4.9 — upper slope 28.1 12.0 7.3 — middle slope 15.6 28.0 17.1 lower slope 12.5 20.0 7.3 toe slope 3.1 12.0 12.2 7.1 depression - - — 14.3 l e v e l 15.6 20.0 53.6 78.6 ASPECT 7.1 north 9.4 24.0 4.9 east 18.8 16.0 12.2 7.1 south 12.5 12.0 7.3 — west 31.3 20.0 22.0 7.1 l e v e l 28.1 28.0 53.6 78.6 ^Classes as defined by Walmsley et al. (1980). \ - 76 -Table VIII. Percent cover by general environment parameters i n 10 m x 10 m plots at 112 s i t e s , summarized by the four vegetation groups. Vegetation group A (n = 32) B (n = 24) C (n = 41) D (n = 14) Parameter X (+ SD) X (+_ SD) X (+ SD) X (+ SD) woody standing dead 4.8 ( + 2.7) 4.0 (+ 2.2) 3.5 (+_ 3.3) 2.0 (+ 2.7) graminoid standing dead 2.9 ( + 4.2) 1.8 (+ 2.4) 3.3 (+ 5.5) 16.6 (+ 11.9) bare mineral ground .9 (+ 3.6) .1 (+ .3) .3 (+ .9) .1 (+ .3) bare organic ground 3.8 (+3.7) 2.6 (+ 1.7) . 4.1 (+_ 6.7) 4.3 (+_ 2.5) t o t a l tree and woody shrubl 35.0 (+ 15.3) 25.6 (+_ 10.1) 32.4 (+17.7) 16.1 (+12.3) t o t a l graminoid^ 8.2 (+ 7.1) 5.9 (+5.7) 7.4 (+8.2) 44.6 (+11.3) t o t a l lichens- 12.9 (+_ 18.3) 18.4 ( + 13.3) 40.9 (+ 19.6) .2 (+ .2) t o t a l open water — — .1 (+ .2) 8.1 (+_ 9.0) I t o t a l tree and woody shrub, t o t a l graminoid and t o t a l l i c h e n include a l l species belonging to s t r u c t u r a l categories 1, 3, and 6, r e s p e c t i v e l y , i n Appendix I. - 77 -si d e r a b l e v a r i a b i l i t y from s i t e to s i t e , however group C had a decidedly higher mean cover. Open water attained appreciable cover only at group D s i t e s (Table V I I I ) . Mineral s o i l was encountered within the active layer at about two-thirds (61.6%) of the s i t e s . S o i l texture classes were c o r r e l a t e d with the four vegetation groups because s o i l texture can provide an accurate, a l b e i t i n -d i r e c t measure of s o i l moisture. Of group A s i t e s , 62.1% of the mineral s o i l s are sandy (Table IX) and a s i m i l a r percentage (63.0%) of group B miner-a l s o i l s are s i l t y and clay s o i l s . These percent f i g u r e s , however, do not c l e a r l y show the trends. For example, there i s an overlap between group A and B s o i l texture c l a s s e s : a number of group A s i t e s are s i l t y - l o a m s , while some group B s i t e s are loams (Table IX). In a d d i t i o n , the broad 'vegetation group 1 designation masks a s o i l texture gradient e x i s t i n g at f i n e r l e v e l s of the TWINSPAN c l a s s i f i c a t i o n . At cut l e v e l V of the dendrogram ( F i g . 11), there are f i v e c l u s t e r s belonging to vegetation group A and c a l c u l a t i o n of mean sand f r a c t i o n s of these s o i l s y i e l d s for c l u s t e r s 1, 2, 3, 4 and 5, r e -s p e c t i v e l y , 88.0, 70.0, 54.6, 32.8 and 33.5%. While 63.4% of group C s i t e s have organic s o i l s within the a c t i v e layer, with the exception of two s i t e s , s o i l s at a l l other group C s i t e s are h i g h l y cryoturbated and have only small amounts of mineral s o i l banded at or near the bottom of the p i t . Organic s o i l s are found at about h a l f (57.1%) of group D s i t e s . Patterned t e r r a i n of both low-centred and raised-centred ice-wedge poly-gons i s associated mainly with group C. Of the 29 s i t e s studied that had patterned ground, 26 were group C s i t e s . Groups B and D had no s i t e s on polygonal ground, and group A had just 3. Of those 26 s i t e s i n group C, most - 78 -Table IX. Subsurface s o i l texture for mineral s o i l s and the occurrence of organic s o i l s at 112 s i t e s , summarized by the four vegetation groups. Vegetation group S o i l type C D (numbers of s i t e s ) Totals 1) mineral s o i l s (a) sandy s o i l texture c l a s s e s 1 sand 3 loamy sand 1 1 sandy loam 8 3 sandy clay loam 2 loam 5 3 (b) s i l t and clay s o i l texture c l a s s e s 1 s i l t y loam 6 2 s i l t y clay loam - 3 s i l t y clay clay loam 3 7 c l a y 1 2) organic s o i l s t o t a l s 32 25 1 4 26 41 14 4 4 13 2 11 11 3 3 17 1 43 112 1 follows Canada S o i l Survey Committee, Subcommittee on S o i l C l a s s i f i c a t i o n (1978). - 79 -(87.5%) were on organic s o i l s . However, two seemingly anomalous group C s i t e s that had p a r t i c u l a r l y l a r ge r a i s e d - c e n t r e d ice-wedge polygons, were s i t u a t e d on loamy sand s o i l s . Although they have l a r g e standard d e v i a t i o n s , the s o i l p h y s i c a l and chemical data (Table X) show some general r e l a t i o n s h i p s among the four groups. G e n e r a l l y , the s o i l s are a c i d , have high c a t i o n exchange c a p a c i t i e s w i t h an o v e r a l l mean of 89.1 meq.lOOg -!, and low amounts of a v a i l a b l e macro-n u t r i e n t s (Mg, K, t o t a l N). The organic contents of the sampled upper hor-izons are h i g h . Even the measurements from the bottom of the a c t i v e l a y e r y i e l d a mean value of 15.4% organic carbon. S o i l carbon: n i t r o g e n (C:N) r a t i o s are high i n d i c a t i n g slow rates of organic matter decomposition and probably r e f l e c t i n g s i m i l a r . C:N r a t i o s i n the plant t i s s u e s . Comparing s o i l p h y s i c a l and chemical data among the v e g e t a t i o n groups (Table X) general increases are observed from A to C i n percent organic carbon, of both top and bottom samples, i n o r g a n i c carbon (bottom) and C:N r a t i o s , w h i l e general decreases occur i n s o i l pH, i n o r g a n i c C ( t o p ) , and Mg. Depth to permafrost was g e n e r a l l y deeper i n groups A and D, probably i n the former r e f l e c t i n g the b e t t e r - d r a i n e d , c o a r s e - t e x t u r e d s o i l c o n d i t i o n s of h i l l c r e s t s and upper s l o p e s , and i n the l a t t e r because of the higher thermal con-d u c t i v i t y of standing water t y p i c a l l y present at group D s i t e s . 1.3 Determination of L i c h e n Standing Crop As already noted i n Table V I I I , l i c h e n cover was a p p r e c i a b l e only i n v e g e t a t i o n groups A, B and C. Cover at i n d i v i d u a l s i t e s i n these groups ranges w i d e l y from trace amounts (1.9%) to n e a r l y continuous ground carpets (89.3%), w i t h the most ext e n s i v e l i c h e n cover occuring at group C s i t e s . The - 80 -T a b l e X. S u b s u r f a c e s o i l p h y s i c a l and c h e m i c a l d a t a summarized by the f o u r v e g e t a t i o n groups. V e g e t a t i o n Croup Parameter A B C D depth to permafrost (cm) 41.7 (+ 18.5) 31.6 (+ 7.2) 31.7 (+ 12.3) 44.5 (+ 26.3) n = 32 n = 25 n - 40 n - 14 s o i l pH" 6.20 (+ 1.34) 4.94 (• .85) 4.17 ( + .69) 5.43 (+ 1.20) n = 32 n - 23 n = 41 n = 11 s o i l CEC 77.8 (+ 41.3) 100.2 (+ 34.2) 97.6 (+ 36.1) 69.7 (+ 46.7) (meq.100 g" 1) n = 32 n = 23 n = 41 n • 11 o r g a n i c C 63.2 (+ 29.3) 78.6 (+ 21.6) 85.6 ( + 21.3) 73.9 ( + 30.7) (%) n = 32 n - 23 n » 40 n » 12 c o t a l N .838 ( + .433) .809 ( + .409) .912 (+ .477) .282 (+ .134) (%) n = 21 n - 7 n - 18 n - 3 C:N r a t i o 58.5 (+ 23.9):1 97.3 (+ 87.0):1 98.6 i [+ 59.0):1 93.8 (• 59.1):: n = 21 n - 7 n •"»18 n = 3 i n o r g a n i c C 2.06 (+ 2.08) 1.03 (+ .90) .79 (• .44) 2.43 (+ 1.61) (Z) n = 21 n - 7 n » 18 n - 3 K* .032 (t .030) .039 (+ .017) .039 (+ .017) .013 (+ .003) « ) n = 21 n - 7 n - 18 n = 3 Mg + .215 (<• .109) .175 (+ .055) .111 (+ .077) .066 (+ .012) U ) n « 21 n - 7 n - 18 n » 3 N a + .015 (+ .009) .011 (+ .009) .016 (+ .029) .013 (+ 0.13) (Z) n =» 21 n - 7 n » 18 n = 3 Fe, pyrophosphate .176 (+ .124) .236 (+ .157) .205 (+ .222) .380 e x t r a c t a b L e (X) n » 14 n - 5 n - 15 n - 1 Mn , pyrophosphate .025 (+ .039) .031 (+ .039) .007 (+ .003) .006 e x t r a c t a b l e (X) n. = 14 n - 5 n - 15 n = 1 o r g a n i c C* 13.0 (+ 12.7) 15.5 (+ 18.9) 22.1 (+ 19.8) 9.73 (+ 4.08) (X) n - 29 n = 18 n - 15 n - 6 i n o r g a n i c C* .467 (• .450) .670 (• 1.995) .786 (+ .443) .727 (+ .786) (X) n = 29 n - 18 n = 15 n » 6 'measurements from 'bottom' samples i n s o i l p i t s . - 81 -Table XI. Mean percent ground cover and, i n parentheses, frequency for the four most abundant l i c h e n species in vegetation groups A, B and C. Vegetation group A B C Species (n = 32) (n = 25) (n = 41) Cladina vangiferina 0. .6 (43, .8) 6. .9 (88. .0) 20, .8 (92, .6) C. stellaris 0, .5 (46, .9) 3. .8 (92, .0) 10, .6 (90. .2) Cetraria aucullata 5, .3 (90 .6) 4, .0 (80, .0) 6, .8 (90. .2) C. ericetorum 2, .9 (81 .3) 1, .0 (56, .0) 1, .6 (82. .9) t o t a l cover 9. .3 15, .7 39, .8 - 82 -three vegetation groups have the same four dominant lichen species although in d i f f e r e n t proportions (Table XI). Group A had highest mean cover, i n de-creasing order of importance, by Cetraria cucullata, C. ericetorum, Cladina rangiferina and C. stellaris. For group B s i t e s the order was: Cladina rangiferina, Cetraria cucullata, Cladina stellaris and Cetraria ericetorum. Group C s i t e s had highest mean cover by Cladina rangiferina followed by C. stellaris, Cetraria cucullata, and C. ericetorum. A l l four are recognized as important Rangifer l i c h e n forage species (Andreev 1954, 1961, Pegau 1968, Parker 1976). Differences i n r e l a t i v e cover between the two l i c h e n genera can be elucidated by means of a simple Cladina:Cetraria r a t i o based on the data summarized by vegetation groups i n Table XI. The r a t i o was small (0.13:1) for group A s i t e s , however the two Cladina spp. covered about twice (2.14:1) as much ground surface, on average, as do the two Cetraria spp. at group B s i t e s . At group C s i t e s , the r a t i o was 3.74:1. Values for t o t a l cover by the four most important l i c h e n species (Table XI) c l o s e l y correspond to t o t a l l i c h e n cover estimates made i n the 10 m x 10 m plots at s i t e s (Table V I I I ) . Thus the four taxa c o n s t i t u t e a major propor-t i o n of the l i c h e n ground cover i n the Tuktoyaktuk Peninsula area. In a d d i t i o n to the four taxa ind i c a t e d i n Table XI, other l i c h e n species with frequencies over 50% i n the three vegetation groups were (from Appendix I ) : f o r group A, Cladonia amaurocraea (59), C. chlorophaea (63), C. pocil-lum (63), C. sulphurina (59) and Ochrolechia gyalecta (66); for group B, Cladonia amaurocraea (72), Ochrolechia gyalecta (64) and Thamnolia subuli-formis (64); and, for group C, Bryoria nitidula (76), Cetraria nivalis (59), Cladonia amaurocraea (66), C. chlorophaea (54), C. sulphurina (54) and Ochro-lechia gyalecta (68). - 83 -TabLe XII. Lichen biomass measurements from divots,.percenc lichen cover, and estimates of top and bottom components of Lichen standing crop at t h i r t y - s i x s i t e s where t e r r e s t r i a l . Lichens were abundant ( i . e . , ground cover ^ 207.), lichen biomass from divocs' escimaces of lichen scanding crop (kg.ha" 1) (kg.ha-i) (1) (2) (3) (4) (5) (6) boccom cop mean percenc boccom Cop cocal lichen L l c L Q sice no. x (• SD) * ( + SD) (1) X (3) (2) x (3) (4) + (5) VEGETATION GROUP A 80-13 1017. 6 (• 229.6) 709.4 (• 283. 5) 20. 9 212. 6 148. 2 360.3 80-14 542. 2 (+ 195.2) 411.0 (+ 196. 8) 20. 4 110. 6 83. 3 194.4 30-47 809. 8 (+ 308.2) 1247.4 (+ 446. 0) 35. 3 285. 8 440. 4 726.2 80-54 1045. 2 (* 288.8) 1067.2 (+ 133. 0) 28.2 294. 8 301. 0 595.8 80-55 775. 0 (+ 202.4) 1092.4 (• 415. 0) 58. 2 451. 0 635. 8 1086.3 80-62 1699. 6 (+ 356.4) 994.0 (+ 616. 4) 29. 1 494. 6 289. 2 783.8 80-63 906. 6 (+ 220.0) 976.4 (+ 353. 0) 39. 7 360. 4 387. 6 743.0 81-77 998. .4 (+_ 184.0) 585.0 (+_ 120. 4) 51. 4 513. 2 300. 6 313 .3 X 974. 4 885.4 35. 4 344. 9 313.4 658.3 (.* SD) (+, 336.2) '(+ 286. 2) (+ 13 .7) VEGETATION GROUP B 80-1 2307. ,2 (+ 625.6) 1963.0 (+ 695. 0) 25, .6 613. ,6 522. 2 1135.8 80-19 2209. ,8 (• 268.2) 2675.4 (+ 477. ,4) 52. ,7 1164. 5 1410.0 2574.6 30-20 1940. .0 (~ 360.0) 2019.6 (+ 409. ,6) 22. ,5 436. 6 454. 4 891.0 80-23 1876. ,6 (+ 587.2) 1639.8 (+ 390. 0) 60. .8 1141. 0 997. 0 2138.0 80-31 2401, .0 (+ 305.2) 2689.3 (+ 493. ,8) 22. ,0 528. ,2 491. 8 1120.0 80-32 1655. ,2 (+ 136.8) 1690.6 (* 467. ,6) 42. ,9 710. 0 725. 2 1435.2 80-33 2215, .0 (+ 171.6) 2056.0 (+ 348. ,0) 31. .6 700. ,0 649. ,6 1349.5 31-66 904. ,2 (• 87.8) 1270.2 (+ 194.6) 36. ,5 330. ,0 463. 6 793.5 81-74 2651, .8 (+ 320.2) 2439.8 (+ 500.4) 45, .3 1201. ,2 1105. ,2 2306.4 81-90 1449, .0 (+ 349.4) 1288.4 ( + 371. ,2) 41. ,5 601. ,2 534.6 1135.8 81-95 2189 .0 (+ 334.3) 1541.2 (• 178, .8) 29. .0 634. ,8 447. 0 1081.8 81-99 2298, .2 (+ 351.2) 1588.8 (+ 549, .6) 30. .9 710. ,2 491. 0 1201.0 81-101 1650 .6 432.3) 974.2 (+ 255. .4) 25. .3 417, .6 246. .4 664.0 X 1980 .6 1833.6 40. .0 792, ,2 933. .4 1525.6 (+ SD) (+ 470.2) (+, 537. .4) (+ 12 .0) VEGETATION GROUP C 80-3 2299 .5 (• 286.0) 3315.2 (+ 154, .4) 33 .1 761. .2 1097, .2 1358.4 80-4 2552 .4 (* 82.0) 2733.2 (+ 223 .4) 29 .6 755 .6 809, .0 1570.2 80-5 2230, .6 (+ 258.4) 3213.4 (+ 504 .3) 25, .1 559, .8 806, .6 1366.4 30-16 2950 .8 (* 386.8) 5299.6 '(+ 623 .8) 65 .7 1938 .6 3481, .8 5420.4 80-17 2009 .0 (+ 315.6) 2929.4 ( + 445, .0) 69 .2 1390 .2 2027, .0 3417.2 80-18 2303 .6 (+ 291.6) 2080.4 (+ 403 .4) 77 . 7 1789 .8 1616 .6 3406.4 80-21 2765 .2 (+ 102.1) 4376.6 (+ 323 .8) 89 .3 2469 .4 3908 .2 6377.6 80-22 2293 .2 (+ 751.6) 1493.0 (* 647 .6) 45 .0 1032 .0 671 .8 1703.8 80-25 2816 .2 (+ 575.6) 3600.2 (+ 187 .4) 87 .9 2475 .4 3164, .6 5640.0 80-28 238 7 .4 (+ 326.0) 2687.2 (+ 779 .8) 66 .3 1582 .8 1781 .6 3364.4 80-50 2152 .4 (+ 275.8) 1787.0 (+ 507 .2) 55 .2 118, .0 986, .6 2174.6 80-53 1704 .0 (+ 202.8) 2282.6 (+ 379 .0) 77 .3 1317 .2 1764 .4 3081.6 80-65 2347 .6 (+ 634.0) 3052.2 (• 1106 .2) 36 .9 2040 .0 2652, .4 4692.4 81-81 2573 .4 (• 166.4) 2097.6 T+ 366 .0) 42 .0 1080 .8 881 .0 1961.8 31-110 2011 .6 (+ 457.4fo 1598.0 (+ 134 .8) 71 .3 1434, .4 1139, .4 2573.8 X 2359 .8 2836.4 61 .4 1448 .9 1741, .5 3190.4 (+ SD) (+ 332.3) 1047 .0) (.* 21 .8) 'sixteen 20 x 20 cm divocs per sice excepc for sices 80-33, 80-53, 80-55 and 81-110 in which numbers were 14, 17, 15 and 14 respectively. •mean vaLues from quadrat measurements of percent veget at ion cove r. - 84 -Estimates of l i c h e n standing crop ranged from 194.4 to 6,377.6 kg.ha - 1 for 36 s i t e s where l i c h e n cover ranged from 20.4% to 89.3% (Table X I I ) . A l -though a small number of s i t e s represent each of the three vegetation groups and consequently r e s u l t i n large standard d e v i a t i o n s , the mean t o t a l standing crops of the group B and group C s i t e s are, r e s p e c t i v e l y , about two and fi v e times that of group A (Table X I I ) . While there are exceptions, higher l i c h e n cover at s i t e s may be general-l y c o r r e l a t e d with the larger estimates of l i c h e n biomass from divots (Table X I I ) . Both top and bottom l i c h e n biomass measurements from divots are s i g n i -f i c a n t l y d i f f e r e n t (p = .05) for the three vegetation groups, according to unpaired T-tests (Sokal & Rohlf 1969). -P l o t t i n g mean li c h e n biomass for top and bottom components — the values i n .columns (1) and (2) of Table XII — reveals a c u r v i l i n e a r r e l a t i o n s h i p ( F i g . 15). Vegetation group C s i t e s , with the generally larger biomass values, have a larger proportion of top to bottom. These proportions are re-versed for the group A s i t e s , which have also the lower biomass values. Measurements at group B s i t e s i n d i c a t e approximately equal amounts of top and bottom components. These top to bottom r e l a t i o n s h i p s are r e f l e c t e d i n r e l a -t i v e abundances of the mean t o t a l top and mean t o t a l bottom l i c h e n standing crops, given i n columns (4) and (5) of Table XII. 2. I n t e r p r e t a t i o n and Analysis of Large-Scale A i r Photographs 2.1 General A t o t a l of 1,469 colour i n f r a r e d (CIR) photo-pairs were acquired at a mean scale of 1:1,960 and with a mean forward overlap of 19.7% (Table X I I I ) , thus conforming c l o s e l y to the o r i g i n a l s p e c i f i c a t i o n s for the photo-- 85 -5.5 - 4.5 o i CO JZ 2 3.5 C/5 (fi < CQ Q. o r-1.5 0.5 I I"—" • I • SITE MEASUREMENTS OF - LICHEN BIOMASS • / / / / -• / r 2 = .73 • * • / _ • s * • - • • •s A • ^ « A A A ^ ^ i i 0.5 1.0 1.5 2.0 2.5 3.0 BOTTOM BIOMASS (kg-ha " 1x 1 0 3 ) F i g . 15. Top versus bottom components of l i c h e n biomass f o r vegetation groups A (open t r i a n g l e s ) , B (closed c i r c l e s ) and C (open squares). Vegetation group D had no appreciable l i c h e n biomass. C u r v i -l i n e a r regression i s of the form log y - 2.54 + (.00036) x. - 86 -Table XIII. Summarized t e c h n i c a l data for large-scale 70 mm CIR photographs acquired Aug. 5-8, 1980. No. of Range photo-frames (n) x (+_ SD) min. max. f l y i n g height (m agl) 1,469 150 (+_ 37) 106 274 percent forward overlap l',423 19.7 (+_ 9.1) 0 65.0 percent crab 564 1 8.5 (+_ 6.3) 0 36.2 o r i g i n a l photo-scale 1,469 1:1,963 (+ 490) 1:1,400 1:3,600 non-overlap surface • area of each photo-frame (ha) 1,469 1.09 .(+_ .73) .44 4.21 non-overlap ground area of each photo-frame (ha) 1,469 0.97 (+ .72) 0 4.21 ^measurements from a random sample of 564 photo-frames. - 87 -mission. The average area of coverage by each photo-frame, ca l c u l a t e d from f l y i n g height and subtracting the mean percent forward overlap, i s 1.09 ha. Subtracting percent water cover for each photo-frame, the mean land area per photo-frame was 0.97 ha. The t o t a l land area examined was 1,497.3 ha. Rela-t i v e to the e n t i r e study area (Table IV), only 0.1% of the land area was sampled by the large-scale a e r i a l photographs. This percentage f i g u r e i s low, and allows only tentative observations on the d i s t r i b u t i o n of features among the seven reindeer management zones. However, in defense of making ob-servations based on such l i m i t e d ground coverage of the study area, two points are made: ( i ) Good geographical coverage was attained with the random s e l e c t i o n of f l i g h t l i n e s — photographs were acquired along f l i g h t l i n e s located throughout the study area, and hence should be representative, and ( i i ) systematic observations on small-scale (1:34,000 CIR, 1:60,000 black and white NAPL) photographs and LANDSAT scenes i n d i c a t e the recurrence, through-out the area, of s i m i l a r features. Indeed the broad s i m i l a r i t i e s of land-scape ( r e l i e f , vegetation, d i s t r i b u t i o n of patterned ground, hydrologic features, etc....) are r e f l e c t e d by the lack of conformity among regional boundaries drawn by past researchers ( F i g . 3a-d). Reindeer management zones C and D ( F i g . 10), of p a r t i c u l a r i n t e r e s t to the reindeer herd owner, were sampled more i n t e n s i v e l y ; 54.1% of photo-frames, 38.6% of ground-truth l o c a t i o n s , and 53.6% of s i t e s were located within these two zones (Table XIV). Management zone E, representing the l a r g e s t land area (Table IV), had 19.6% of the photo-frames, and 25.0% of both ground-truth locations and s i t e s . Other zones (A, B, F, and G) were represented by comparatively small sample sizes (Table XIV). - 88 -Table XIV. Numbers of ground-truth locations and s i t e s , and large-scale a i r photos i n each of the seven reindeer management zones. Reindeer No. of large-management No. of ground- No. of ground- scale photo-zone truth l o c a t i o n s 1 t r u t h s i t e s frames A 7 10 88 B 2 5 87 C 14 40 414 D 6 20 380 E 11 28 389 F 2 4 125 G 2 5 86 t o t a l s 44 112 1469 i n d i c a t e d on F i g . 2. At each ground-truth l o c a t i o n , one to four s i t e s were studied. - 89 -2.2 Patterned Ground and T e r r a i n Disturbance by V e h i c l e s Patterned ground i s common throughout the Tuktoyaktuk P e n i n s u l a area i n the form of low-centred and r a i s e d - c e n t r e d ice-wedge polygons that are v i s i -b l e from the a i r ( F i g . 16, 17). With the exception of zone G where they were of low occurrence and cover, ice-wedge polygons were observed i n about h a l f of the photo-frames and, f o r d i f f e r e n t zones, cover ranged from 16.3% to 43.4% of the land surface (Table XV). I t must be emphasized that ice-wedge polygons with v i s i b l e surface e x p r e s s i o n were measured here, however i c e -wedges probably u n d e r l i e most i f not a l l of the non-patterned t e r r a i n as w e l l (J.R. Mackay, pers. comm.1). Ice wedges with no surface expression commonly become v i s i b l e i n the tundra f o l l o w i n g severe t e r r a i n damage along v e h i c l e paths and seismic l i n e s as d i f f e r e n t i a l m e l t i n g and thermokarst subsidence p a t t e r n s (Mackay 1970). Ice-wedge polygons occurred at a mean d e n s i t y of 119.9 per ha (Table XV). Although pingos are widely s c a t t e r e d through the study area (Stager 1956), only four were t r a v e r s e d by f l i g h t l i n e s r e p r e s e n t i n g a low occurrence, i n 14 photo-frames of 0.9%. Because of t h e i r conspicuous appearance and l a r g e s i z e , up to 49.5 m high and 600 m diameter (Mackay 1979), these perma-f r o s t features are more r e a d i l y i n v e n t o r i e d on s m a l l - s c a l e d photographs (e.g. 1:60,000 black and white NAPL photos). Measurements of t e r r a i n disturbance by v e h i c l e s ( F i g . 17) showed gr e a t -est frequency and cover values i n management zones C and F, probably i n the former due to the p r o x i m i t y to the Tuktoyaktuk townsite and i t s w i n t e r road l e a d i n g south, and i n the l a t t e r , to c o n s i d e r a b l e past human a c t i v i t i e s i n the v i c i n i t y of Reindeer S t a t i o n (Table XV). Severe damage l e v e l s were i j . R . Mackay, Dep. Geography, Univ. B r i t i s h Columbia. 6 May, 1982. - 90 -F i g . 1 6 . Cibachrome 2 . 5 X enlargement of 1 : 1 , 6 0 0 ( o r i g i n a l s c a l e) 7 0 mm CIR photograph, f l i g h t l i n e 4 - 2 . Note the high cover by, 'Type I I I ' b r i g h t white ground l i c h e n on cryoturbated surfaces of r a i s e d - c e n t r e d polygons, and i n the ice-wedge troughs, the occurrence of standing water ( b l a c k ) , aquatic mosses (dark brown) and wet sphagnum, sedge and low shrub cover (orange to red) . Located immediately south of area covered i n F i g . 8 at 6 9 ° 0 5'N, 133°42'W. 5 August, 1 9 8 0 . - 91 -Fi g . 17. Large-scale (1:1,800) CIR photo-pair showing examples of vege-t a t i o n groups A (raised-centred and low-centred polygons, bottom r i g h t and centre) and B (lower h i l l slope, top and l e f t ) , and low damage l e v e l s to tundra r e s u l t i n g from multiple passes of tracked v e h i c l e s . (frame 27, f l i g h t l i n e 5-4; 69°15'N, 133° 05'W). 5 August, 1980. Table XV. Frequency and percent cover of ice-wedge polygons, and t e r r a i n disturbance by v e h i c l e s , summarized for large-scale photographs i n the reindeer management zones. T e r r a i n disturbance by vehicles Ice-wedge polygons frequency^ cover Reindeer management zone frequency cover ha area % of density of land polygons 1 area (no. polygons.ha" ~ l) low mod. severe ha area % of land area A 68. .2 29 .9 43, .4 178. .4 4. .5 - - 0. .1 0. .1 B 40. .2 16 .8 29 .0 132. .4 17. .2 - - 0. .3 0. .4 C 40. .6 57 .4 19 .1 108. .4 24. ,6 9.4 5.6 6. .1 2, .0 D 41. .8 58 .9 22, .4 78. .2 10. .5 0.3 - 0. .9 0. .3 E . 32. .9 43 .4 16, .3 163. .0 3. .1 - - 0. .3 0, .1 F 52. .1 110 .6 33, .8 150. .8 34. .4 - - 1. .8 0, .6 G 2. .4 0 .7 4, .6 76, .9 3. .5 - - 0. .1 0, .1 ^expressed i n terms of area in which polygons were present. 2 l ow - v i s i b l e tracked depressions but no permafrost degradation or vegetation change; mod. - minor changes to permafrost and/or vegetation cover, water pools i n low areas; severe - permafrost degrada-t i o n , d i s t i n c t vegetation changes, bare s o i l patches, water pools, and slumping of surface material. - 93 -observed only i n photo-frames from zone C, and occurred mainly along f l i g h t -l i n e 5-3, located only a few km from Tuktoyaktuk ( F i g . 6). For other zones, frequency of low damage le v e l s varied from 3.1 to 17.2 percent, and cover was low (0.1 to 0.9%). Vehicle t e r r a i n disturbance was observed i n 279 (19.0%) of the 1,469 photo-frames. 2.3 Microdensitometric Measurements Lichen patches on a t o t a l of 296 larg e - s c a l e CIR photographs were studied using microdensitometry. Based on the c r i t e r i a for the determination of 'Lichen Types' presented i n Table I I I , 103 of the photographs were assigned to Type I, 110 to Type II and 83 to Type I I I . O p t i c a l density of the developed f i l m dye-layers as measured with blue l i g h t was elevated i n Lichen Types I and II because the l i c h e n cover occurs i n close a s s o c i a t i o n with herbaceous and shrub vegetation (Table XVI, F i g . , 18a). Blue l i g h t microdensitometry measures the yellow dye layer density on CIR f i l m and pro-vides an index of s p e c t r a l r e f l e c t a n c e i n the green portion (0.5 to 0.6 Jim) of the v i s i b l e spectrum ( F r i t z 1967). Probably because the l i c h e n cover occurs i n as s o c i a t i o n with bare organic patches and brown mosses as well as green vegetation i n Lichen Type I I , o p t i c a l density measurements with green and red l i g h t were higher as well (Table XVI). Lowest mean measurements for o p t i c a l density were obtained using white, red, green and blue l i g h t s on photographs assigned to Lichen Type III (Table XVI). High r e f l e c t a n c e i n green, red, and near-infrared s p e c t r a l regions (0.5 to 0.9 ym) on these photographs res u l t e d i n less dye layer development than i n Lichen Types I or I I ; o p t i c a l density counts were therefore low ( F i g . 18b). A l l three Lichen Types had s i g n i f i c a n t l y d i f f e r e n t (p = .05) mean o p t i -c a l d e n s i t i e s using white and blue l i g h t , while red and green l i g h t provided - 94 -Table XVI. Microdensitometric readings of o p t i c a l densi-ty values with white, red, green and blue l i g h t of three Lichen Types on large-scale (1:1,400 to 1:3,600) CIR photos. Means followed by a common l e t t e r are not s i g n i f i c a n t l y d i f f e r e n t at the 5% p r o b a b i l i t y l e v e l , according to Duncan's new multiple range t e s t . White Red Green Blue Lichen Typel n Light Light Light Light Type I 103 0.61 b 0.58 b 0.69 & 0.78 a Type II 110 0.70 a 0.76 b 0.73 b 0.73 b Type III 83 0.49 a 0.54 a 0.51 a 0.53 a las defined i n Table I I I . F i g . 18. Examples of Lichen Types on large-scale CIR st e r e o - p a i r s . (a) Large-scale (1:1,800) CIR photo-pair showing Lichen Type I on arm of cres c e n t i c sand dune ( r i g h t ) and on the rims of some ice-wedge polygons. Elsewhere i n the photographs, the l i g h t c o l o u r a t i o n i s due to graminoid standing dead i n t h i s sedge-rich tundra heath near the t i p of the Tuktoyak-tuk Peninsula, used mainly as summer rangeland by the reindeer herd (frame .17, f l i g h t l i n e 8-2; 70°02'N, 129°52'W). 6 August, 1980. (b) Bright white l i c h e n (Type III) cover and scattered Salix spp. and Alnus crispa shrubs on patterned ground south of Eskimo Lake near the t r e e l i n e . Photo-scale i s 1:2,000 (frame 11, l i n e 13-1; 68°55'N, 132°27'W). 6 August, 1980. - 96 -- 97 -only p a r t i a l d i f f e r e n t i a t i o n (Table XVI). A plot of o p t i c a l density means and standard deviations using white and blue l i g h t showed a region of overlap between Lichen Type I and II where an unknown sample c l e a r l y could not be assigned to one or the other ( F i g . 19). An LDF analysis of the microdensitometric data found, i n decreasing order of e f f e c t i v e n e s s , that white, blue and red l i g h t discriminated among the three Lichen Types (Table XVII). As a measure of the s e p a r a b i l i t y of the Lichen Types, the LDF program r e c l a s s i f i e d the density data among three groups with the same structure as the a pri.OV% groups. A j a c k n i f e d LDF ( J e n n r i c h & Sampson 1977) re s u l t e d i n an o v e r a l l 81.1% correct r e c l a s s i f i c a -t i o n of the 296 sets of density measures (Table XVIII). Photo-frames that had l i c h e n patches o r i g i n a l l y c l a s s i f i e d as Lichen Type II were the most suc-c e s s f u l l y reassigned (83.6%) by the LDF analysis (Table XVIII). Using j u s t white and blue l i g h t microdensitometric data would l i k e l y have shown mainly m i s c l a s s i f i c a t i o n s between Type I and II (e.g. F i g . 19) but the use a d d i t i o n -a l l y of red l i g h t data i n the LDF analysis' recombination matrix appears to have improved that r e l a t i o n s h i p ; p o t e n t i a l l y m i s c l a s s i f i e d i n d i v i d u a l s were assigned about equally among the three Lichen Types (Table XVIII). Quantitative analyses of the microdensitometric data on l i c h e n patches suggest that the Lichen Types can be r e a d i l y separated on the large-scale CIR photographs. The three Lichen Types I, II and III were generally equivalent to the vegetation groups A, B and C recognized from ground-truth studies. The c r i t e r i a for assigning Lichen Types used r e l a t i o n s h i p s broadly ascribed to vegetation groups e a r l i e r i n the r e s u l t s . For example slope p o s i t i o n deter-mined from stereo-pairs of large-scale CIR photographs was used to help de-- 98 -1.0 .9 .8 3 -TO-TYPE I .7 TYPE n o .6 .5 T Y P E H .4 .3 4 .5 .6 O.D. (white light) .7 .8 F i g . 19. P l o t of means and standard d e v i a t i o n s of three Lichen Types i n blue and white l i g h t o p t i c a l d e n s i t y (O.D.) space; based on 296 micro-d e n s i t o m e t r i c spot readings from l a r g e - s c a l e CIR photographs. - 99 -Table XVII. Summary of stepwise Linear Discriminant Function (LDF) analysis based on microdensitometric readings of the three Lichen Types. Step Variable Entered Removed No. of Variables included U - S t a t i s t i c Approx. F s t a t i s t i c 1 White Light 1 .556 116.71 2 Blue Light 2 .294 123.43 3 Red Light 3 .283 85.36 - 100 -Table XVIII. Recombination matrix (percentage values) for step 3 of the stepwise (LDF analysis using microdensitometric readings of Lichen Types. About 81% were c o r r e c t l y c l a s s i f i e d using the BMD:P7M computer program 1 s ^ j a c k -n i f e d c l a s s i f i c a t i o n . Predicted Type I Type II Type III Type I 76.7 5.8 17.5 Observed Type II 12.7 83.6 3.6 Type II I 7.2 9.6 83.1 - 101 -termine Lichen Types, the c r i t e r i a being those a t t r i b u t e s described for vege-t a t i o n groups. S i m i l a r l y , Lichen Type III was c o r r e l a t e d with patterned ground, previously r e l a t e d to vegetation group C s i t e s . A d d i t i o n a l checks of the general r e l a t i o n s h i p of Lichen Types and vegetation groups were conducted s y s t e m a t i c a l l y during f i e l d studies where ground-truth s i t e s were f i r s t assigned to a vegetation group, then l a t e r annotated t e n t a t i v e l y as a Lichen Type from a e r i a l observations. These were further checked on the large-scale a i r photos in the lab, and c o r r e l a t e d c l o s e l y . 2.4 Estimation of Lichen Standing Crop Computer-assisted summaries of the large-scale photo data i n d i c a t e d 332 photo-frames had no l i c h e n cover, while 252, 593 and 292 r e s p e c t i v e l y were assigned to Lichen Types I, II and III (Table XIX). For each of these four groups of photo-frames, t o t a l surface and land area were ca l c u l a t e d with cor-rections made for v a r i a t i o n s i n photo-scales. The greatest land area (637.5 ha) was represented by Lichen Type II photographs, followed by Type I I I , no l i c h e n , and Type I photographs (Table XIX). Lichen cover, determined as a percent of land area, for i n d i v i d u a l photo-frames ranged up to 97.0%, a l -though the o v e r a l l mean cover by l i c h e n i n photo-frames, in c l u d i n g those i n which no l i c h e n occurred, was 6.8%. Mean percent l i c h e n cover i n the photo-frames was t a l l i e d for the Lichen Types according to the reindeer management zones (Table XIX). Lichen percent cover values, given as the percentages of land area covered by those photo-frames i n which each Lichen Type was dominant, are re-expressed i n Table XX i n terms of the t o t a l land area sampled i n each r e i n -deer management zone. These l a t t e r figures were used, along with mean l i c h e n Table XIX. Number of photo-frames, total surface area, total land area and percent lichen cover summarized for reindeer management zones according to those wi th no 1 ichen and those ass igned to Lichen Types I, II and III. No Lichen Lichen Types II III Reindeer sur face land surface land 1 ichen surface 1 and I ichen surface land 1 ichen management area area n' area area cover n' area area cover area area cover zone n' (ha) (ha) (ha) (ha) (X of land area) (ha) (ha) (% of land area) n< (ha) (ha) (I of Land area) A 27 23.7 17.0 23 18.9 16.5 4.4 37 37.3 34.5 3.4 1 1.0 .9 3.6 B 37 34.8 15. 7 23 17.5 16.7 3.0 27 28.8 25.5 1.4 - - - -C 124 101.1 87.5 153 120. 7 110.5 4.2 96 73. 7 68.9 6.6 41 35.1 33.5 3.6 D 66 49.8 39.8 IB 12.9 12.7 20.0 200 148.4 136.2 19.0 96 75.8 73. 7 12.2 E 34 39.3 19.5 24 29.9 29.2 6.0 138 145.7 140.3 7.6 93 83.4 77.8 14.6 F 38 78.9 63.8 11 39.0 37.5 3.4 63 189.1 175.7 3.8 13 51.5 50.0 4.8 G 6 12.3 7. 7 - - - - 32 57.7 56.4 8.2 48 91.5 88.4 15.0 Totals 332 339.9 251.0 252 238.9 223.1 593 680. 7 637.5 292 338.3 324.3 no. of photo-frames in which Lichen Type was dominant. - 103 -Table XX. Estimates of bottom and top components of lichen standing crop based on 1) percent cover i n t e r -preted from large-scale photographs, and 2) f i e l d s i t e estimates of lichen biomass per unit area from Table XII. Summarized for the three Lichen Types ( l , I I , III) according to seven reindeer management zones. Iichen cover standing crop (percent of (kg.ha~*) tot a l land Reindeer area) bottom top t o t a l management • - - • • • • •- — • • . • • — zone I II III I II III I II III I II l i t A 1.06 1. ,70 ,04 10.4 33. ,6 1. 2 9. ,2 31. 2 1. 2 19.6 64. .8 2. .4 B .86 ,62 - 8.4 12. ,4 - 7. .6 11. 2 16.0 23. ,6 C 1.54 1. .52 .40 15.2 20. ,0 9. .6 13. ,6 27. ,6 11. ,6 28.8 57, .6 21, .2 D .96 9, ,86 3. .42 9.6 95. , 2 80. ,8 8. .4 180. ,8 97. ,2 18.0 376, ,0 178. .0 E .66 4. .00 4. .26 6.4 79, .2 100. .4 6, ,0 73. ,2 120. .8 12.4 152, .4 221, .2 F .40 2 .04 .74 4.0 40, .4 17, .2 3, .6 37. ,6 20, .8 7.6 78, .0 38, .0 G — 3. .04 8, .70 — 60, .0 205, .2 - 55, .6 246. ,8 — 115 .6 452 .0 - 104 -F i g . 20. Histogram showing for each of the seven reindeer management zones, top and bottom standing crops of Lichen Type I ( l e f t ) , I I ( c e n t r e ) and I I ( r i g h t ) . Based on data given i n Table XX. - 105 -biomass estimates for vegetation groups A (applied to Type I ) , B (Type II) and C (Type III) from columns (1) and (2) of Table XII, to estimate l i c h e n standing crop as a weight per unit area measure. Standing crops of l i c h e n were c a l c u l a t e d for top and bottom components, and t o t a l s , according to the Lichen Types (Table XX, F i g . 20). Lichen Type I t o t a l standing crops are low, and ranged from 0 to 28.8 kg.ha -! for the reindeer management zones, while Lichen Type II (23.6 to 376.0 kg.ha"!) and Type III (0 to 452.0 kg.ha -!) ranged considerably higher (Table XX, F i g . 20). Table XXI summarizes the standing crop estimates of Table XX. Geograph-i c a l l y the three more northerly reindeer management zones (A, B and C) had lower t o t a l l i c h e n standing crops, ranging from 39.6 to 107.2 kg.ha -!. The four more southerly zones (D, E, F and G) by comparison had standing crops of 123.2 to 572.0 kg.ha -!. As one would expect, estimates i n Table XXI are con-si d e r a b l y lower than l i c h e n standing crop estimates for ground-truth s i t e s where l i c h e n cover equal to or greater than 20% occurred (Table X I I ) . Even for management zone D where the largest estimate of 572.0 kg.ha -! w a s ob-tained, the standing crop was below the mean standing crop of 658.3 kg.ha -! obtained for eight vegetation group A s i t e s (Table X I I ) . The 'top' component of l i c h e n standing crop, of p a r t i c u l a r importance as reindeer fodder, accounted for almost exactly h a l f of the t o t a l standing crop estimates within a l l zones. Even the extremes are represented by r e l a t i v e l y small percentage v a r i a t i o n : i n reindeer management zone B, one of the more no r t h e r l y zones, li c h e n 'top' standing crop accounted for 47.5% of the t o t a l , and i n the most southerly zone, G, the comparable figure was only s l i g h t l y elevated (53.3%). - 106 -The t o t a l l i c h e n standing crop for the study area can be expressed, using the data i n Table XXI, by (1) m u l t i p l y i n g by the land area represented by each reindeer management zone (Table IV), and (2) d i v i d i n g by the t o t a l land area i n the study area (14,410 sq km, Table IV). A mean value of 276.2 kg.ha - 1 i s obtained for t o t a l l i c h e n standing crop i n the study area. The mean l i c h e n top standing crop s i m i l a r l y i s 141.1 kg.ha - 1 or 51.1% of the t o t a l . - 107 -Table XXI. Estimates of t o t a l l i c h e n standing ; crop summarized by reindeer management zones Reindeer standing crop (kg.ha" 1) management zone bottom top t o t a l A 45.2 41.6 86.8 B 20.8 18.8 39.6 C 54.4 52.8 107.2 D 285.6 286.4 572.0 E 186.0 200.0 286.0 F 61.6 61.6 123.2 G 265.2 302.4 567.6 - 108 -VI DISCUSSION - 109 -1. Ground-Truth 1.1 C l a s s i f i c a t i o n of the Four Vegetation Groups Based on quadrat cover data obtained during the ground-truth program, the vegetation of the study area could be c l a s s i f i e d among eighteen plant community-types (Fig. 11, Table V), and a number of these could undoubtedly be separated into f i n e r l e v e l s such as dominance-types. I n t e r p r e t a t i o n of the data here however has dealt with the d e l i n e a t i o n and d e s c r i p t i o n of just four major vegetation groups. E f f o r t s were concentrated at the group l e v e l i n deference to a d e t a i l e d treatment of community-types even though the l a t t e r i s c e r t a i n l y of academic i n t e r e s t and within the c a p a b i l i t i e s of the present data-base. One of the o r i g i n a l objectives of the study was to provide a simple but workable scheme for i n t e r r e l a t i n g ground-truth and lar g e - s c a l e a i r photos. This objective presupposes ground-truth r e s u l t s that are as l u c i d and s i m p l i f i e d - as p o s s i b l e . The generation of only four ground-truth vegetation units f a c i l i t a t e s a photo-interpretation system that should be easy-to-use and therefore i s more l i k e l y to be accepted and adopted by the reindeer herd owner for reindeer rangeland management in the Tuktoyaktuk Peninsula area. Other researchers examining regional vegetation i n the Mackenzie Delta area have described plant community-landform r e l a t i o n s h i p s that can only be r e l a t e d only roughly to the four vegetation groups defined here. In the Pleistocene Coastlands physiographic region ( F i g . 3b), Lambert (1972) described "heath tundra" occupying h i l l c r e sts and upper slopes, "Eriophovum tussock tundra" on gentle lower slopes and "ice-wedge polygons" on f l a t s and depressions. Corns (1974) i l l u s t r a t e d the r e l a t i o n of topographic p o s i t i o n to f i v e major plant community types defined on the basis of physiognomy and f l o r i s t i c composition. R i t c h i e (1974), i n d i s c u s s i n g tundra vegetation east - no -o f the Mackenzie River Delta noted the following vegetation-topographic r e l a -t i o n : herb communities on steep tundra slopes and scarps (30°-50°) and x e r i c s i t e s (sands and gr a v e l s ) ; Betula glandulosa - e r i c o i d heaths with or without Alnus on moderate slopes ( l O ' ^ O 0 ) ; Eriophorum tussock tundra on gentle slopes (2°-5°), and Eriophorum, Vaacinium - Rubus communities on r a i s e d -centred polygons occupying f l a t surfaces and depressions. While descriptions of plant communities, dominant vegetation taxa, and slope cl a s s d e s c r i p t i o n s vary among the preceding authors, they a l l state that d i s t i n c t vegetation-topographic r e l a t i o n s are evident within the study area. Of the 112 ground-truth s i t e s , 98 were non-wetlands belonging to vegeta-t i o n groups A, B and C ( F i g . 11). The greatest number of s i t e s (41) were assigned to group C. It should be noted however that p o t e n t i a l ground-truth s i t e s were chosen s u b j e c t i v e l y on a i r photographs before going into the f i e l d . While this was considered an appropriate method for ensuring that major vegetation and landscape conditions along a l l f l i g h t l i n e s were sampled, the proportions of s i t e s among the four vegetation groups do not r e f l e c t r e l a t i v e abundances of these groups i n the study area. 1.2 The Use of Indicator Species Using the de c i s i o n rules generated by TWINSPAN, the seventeen i n d i c a t o r species can be used to organize a simple three-step flow diagram for assign-ing "unknown" s i t e s i n the f i e l d among the four vegetation groups ( F i g . 21). Assuming the i n d i c a t o r species and d e c i s i o n rules work s u f f i c i e n t l y well to assign the majority of s i t e s c o r r e c t l y , and only a d d i t i o n a l f i e l d t e s t i n g w i l l a s c e r t a i n t h i s , the flow-diagram could be a most useful t o o l for f i e l d workers. The group to which a s i t e i s assigned immediately indicates i n f o r -- I l l -— + Cladina stellaris Carex aquatilis Arctostaphylos rubra Eriophorum Ledum decumbens angustifol ium Cetraria cucullata 0/! 0 2k Sal ix alaxensis Lupinus arcticus Dryas integrifolia > 1 G R O U P D Cladina stel lar is C. rangi fer ina Ledum decumbens Rubus chamaemorus 0 / 1 G R O U P A 0 > > 1 Carex lugens Pyrola grandiflora Tomenthypnum nitens Aulacomnium palustre; A lnus cr ispal -1/0 Bryor ia ni t idula G R O U P B -1 > > 0 G R O U P C F i g . 21, A flow-diagram f o r a s s i g n i n g "unknown" f i e l d s i t e s among the four v e g e t a t i o n groups based on TWINSPAN-determined i n d i c a t o r species and d e c i s i o n r u l e s . - 112 -mation regarding other s i t e features reported on here, such as the general slope c l a s s , the p r o b a b i l i t y of occurrence of patterned ground or organic s o i l , approximate ranges to be expected of c e r t a i n s o i l p hysical or chemical parameters or s o i l t e x t u r a l c l a s s e s , and i f percent l i c h e n cover i s measured, the estimated l i c h e n standing crop. This l a t t e r measure can be a r r i v e d at using the mean li c h e n biomass values for vegetation groups A, B and C (Table XI I ) . While most appropriately obtained for s i t e s where l i c h e n cover i s equal to or greater than 20%, as the l i c h e n biomass measurements were acquired at such s i t e s , estimates of l i c h e n standing crop can be extrapolated to a l l s i t e s with caution. The i n d i c a t o r species and d e c i s i o n rule technique used here requires that a f i e l d worker know by sight the seventeen species ( F i g . 21) so he can quickly determine t h e i r presence or absence at a s i t e . Fortunately a l l seventeen have d i s t i n c t i v e taxonomic features that would help minimize con-fusion with other s i m i l a r taxa that might be encountered i n the f i e l d . While Figure 11 only shows the d e c i s i o n rules and i n d i c a t o r species for cut l e v e l s I, II and I I I , TWINSPAN provides them for a l l f i n e r d i v i s i o n - p o i n t s of c l a s s -i f i c a t i o n s ( H i l l 1979). Within a minute or two, an experienced f i e l d user of the technique could assign a s i t e to a vegetation group and continue on to further assign the s i t e to Level V community-types as indicated i n Table V. 1.3 F l o r a and Species D i v e r s i t y The t o t a l number of vascular taxa named here i s only about h a l f the number reported for the Reindeer Preserve and Mackenzie River Delta by Cody (1965). In his broader study region, Cody (1965) examined many unique r i v e r -- 113 -ine and other habitats that were not encountered i n the present study. Numerous a d d i t i o n a l vascular plant records within the present study area how-ever are given by P o r s i l d & Cody (1980) so the t o t a l vascular f l o r a can be expected to be about 350 species. With the exception of I n g l i s ' (1975b) i n v e s t i g a t i o n of reindeer range-land near S i t i d g i Lake, e c o l o g i c a l studies in the Tuktoyaktuk Peninsula area have f a i l e d to include non-vascular plants to an adequate degree. The present work provides evidence that t h i s i s unacceptable; non-vascular plants represented h a l f of the taxa encountered and i n terms of ground cover, l i c h e n cover was i n excess of 80% at some vegetation group C s i t e s (Table XII) . Non-vascular plants were good i n d i c a t o r s of c e r t a i n vegetation types; for example, 6 of the 17 i n d i c a t o r species used to discriminate among the vegeta-t i o n groups were non-vascular ( F i g . 21). The number of new plant records reported for the study area i n d i c a t e s that, in s p i t e of extensive past botanical c o l l e c t i n g i n the general region, the f l o r a is s t i l l incompletely known. This is e s p e c i a l l y true for non-vas-c u l a r taxa which accounted for most (33 of 36) of the f i n d s , but only repre-sented about h a l f of the t o t a l taxa recorded during ground-truth i n v e s t i g a -t i o n s . A h t i et al. (1973) had e a r l i e r i n d i c a t e d that the l i c h e n f l o r a of the Reindeer Grazing Preserve i n p a r t i c u l a r was only incompletely known. Vegetation group A s i t e s , generally the d r i e r upland s i t e s on coarse-textured s o i l s , supported a greater number of taxa than the other groups (Table VI). Species richness, or number of taxa, may be considered as a simple measure of habitat d i v e r s i t y (whittaker 1972) and i n group A the high-er species richness i s probably due to the d i v e r s i t y of habitats represented, from dry g r a v e l l y ridges to protected mid-slope positions where cryoturbation - 114 -and s o l i f l u c t i o n processes have exposed organic material to create humid peaty microsites (cf Tarnocai & Z o l t a i 1978). The plant communities ranged considerably i n general physiognomy and species composition from Dryas in-tegrifolia - Salix reticulata closed mat and cushion tundra to Alnus crispa -Salix alaxensis / Calamagrostis neglecta open low alder shrub tundra (Table V). Conversely, group D's reduced species richness (Table VI) suggests a narrower range of conditions for establishment and growth of plant taxa. Wetland plant species must be generally adapted to high standing water l e v e l s and perhaps seasonal floodings. In group D, while lichens account for 18 taxa or 12.2% of the t o t a l (Table VI), the cover attained by them was n e g l i -g i b l e (Table V I I I ) . 1.4 General Environment and S o i l s Measurements The 10 m x 10 m plots examined at s i t e s were considered valuable for re-l a t i n g l arge-scale photographs more d i r e c t l y to ground-truth r e s u l t s . The pl o t s covered areas of land and water that were r e a d i l y resolved on the a e r i a l photos and, for example, represented 5 mm x 5 mm c e l l s on 1:2,000 scale photographs. The plots were not p h y s i c a l l y flagged on the ground so they were not marked out on the a e r i a l photos. However, from f i e l d notes and s i t e ground and a e r i a l oblique 35 mm photos at s i t e s , approximate locations could be r e c a l l e d s u f f i c i e n t l y to determine that the data they provided was c l e a r l y representative of ground conditions over the broader area of the s i t e . This observation i s s i g n i f i c a n t since during f i e l d studies, data-gathering from plots constituted a small f r a c t i o n of the time and e f f o r t re-quired for d e t a i l e d quadrat studies. Future s i m i l a r studies might benefit from the "streamlining" of ground-truth data-gathering that use of these - 115 -plots seems to allow. D r i s c o l l et al. (1970) found that 6 m x 6 m plots and 6 m l i n e - t r a n s e c t s , ground-marked within several Colorado and C a l i f o r n i a rangelands, could be used to estimate f o l i a r cover and plant density within acceptable error l i m i t s on 1:600 to 1:4,200 CIR 70 mm air-photos. One of the c r i t e r i a used to define Lichen Type I, equivalent to vegeta-t i o n group A, was the occurrence of bare sand cover (Table I I I ) . That bare s o i l cover, as measured i n the 10 m x 10 m p l o t s , was greatest i n group A (Table VIII) lends support to t h i s c r i t e r i o n . Certain other parameters measured i n the plots y i e l d mean cover values and standard deviations (Table VIII) that would be useful i n the production of a photo-interpretation key to the Lichen Types/vegetation groups f o r smaller-scaled remote sensing data. For example, graminoid standing dead and t o t a l graminoid cover were both lowest i n vegetation group B. The former feature imparted a white yellow colouration on the CIR photos (see F i g . 18a) while the l a t t e r , a bright red c o l o u r a t i o n . Both features could be recog-nized r e a d i l y by an experienced photo-interpreter so might prove, with t e s t -ing, to be valuable parameters for such a key. Lichen cover measurements in the 10 m x 10 m plots (Table VIII) corre-sponded with quadrat measurements for the four most important li c h e n species (Table XI). In vegetation group D, the t o t a l l i c h e n cover was small, only 0.2 (+_ .02)% (Table VIII) and thus, as mentioned elsewhere, the group con-t r i b u t e d very l i t t l e to the l i c h e n standing crop of the area. Tundra s o i l s of the Tuktoyaktuk Peninsula area appear to have a general-ly low nutrient status, and those chemical c h a r a c t e r i s t i c s tabled here (Table X) are comparable to those i n l i c h e n woodland s o i l s of Northern Quebec (Moore 1980) and tundra s o i l s i n Western Alaska (Everett 1980). The measurements - 116 -demonstrate large random v a r i a b i l i t i e s which may be found as inherent among most s o i l s . They are however no doubt enhanced i n cold-region s o i l s subject to freeze-thaw processes, e s p e c i a l l y cryoturbation (Everett 1980). The h i l l crest and upper slope p o s i t i o n s o i l s of vegetation group A have lower amounts of subsurface organic matter (Table X). Consequently, as Janz (1973) reported i n a comparison of s o i l c h a r a c t e r i s t i c s from h i l l top and de-pression areas near Tuktoyaktuk, s o i l moisture-holding c a p a c i t i e s are gener-a l l y lower, the i n s u l a t i n g e f f e c t s are reduced, and the active layer i s deep-er. Conversely, i n lower areas, represented by the group C s o i l s (Table X), the poorly-drained mainly organic s o i l s are w e l l - i n s u l a t e d , and hence temper-atures remain low throughout the season, retarding decomposition by micro-organisms. Where standing water occurs for at least part of the season as i n group C ice-wedge polygon troughs, low-centred polygon centres, or most group D s i t e s , organic matter accumulation may be perpetuated under anaerobic con-d i t i o n s . Because of buildups of humic and other acids r e s u l t i n g from incom-plete decomposition of o r g a n i c - r i c h materials, lowland s i t e s (e.g. those of group C) are t y p i c a l l y more a c i d i c than better-drained l o c a t i o n s . In the present study, the mean group C pH (4.17) i s more than two pH units lower than the mean group A pH (6.20, Table X). Low s o i l temperatures of tundra s o i l s also l i m i t m i c r o b i a l transformation of organic nitrogen and values r e -ported here for t o t a l N are low (Table X). Other studies i n the Tuktoyaktuk area have shown av a i l a b l e ammonium- or n i t r a t e - n i t r o g e n to be extremely low and, along with phosphorus, l i m i t i n g to plant production (Janz 1973, Haag 1974). In part, the development of l i c h e n mats p a r t i c u l a r l y i n vegetation group C s i t e s can be r e l a t e d to the apparently low nutrient s t a t u s . Northern - 117 -lichens have a low net p r o d u c t i v i t y and growth, normally grow well on s l i g h t -ly a c i d i c s o i l surfaces, probably obtain few nutrients from the s o i l and con-t a i n low concentrations of major nutrients ( P u l l i a i n e n 1971, Kershaw 1977, Williams et al. 1978). 1.5 Lichen Standing Crop A s i g n i f i c a n t proportion of the study area's l i c h e n standing crop was undoubtedly accounted for by the four most abundant l i c h e n species (Table XI). These taxa a l l occurred, with the exception of two vegetation group A s i t e s , at s i t e s where l i c h e n biomass measurements were recorded. The taxa e x h i b i t e d a combined mean ground cover, i n vegetation groups A, B, and C re s p e c t i v e l y , of 9.3, 15.7 and 29.8% (Table XI), but t h e i r proportions ranged higher at the 36 s i t e s where biomass c o l l e c t i o n s were made, as evidenced by the t o t a l percent l i c h e n covers at these s i t e s (column 3, Table X I I ) . Other researchers have found these species ubiquitous as w e l l . Among the most common lichens on Russian and Alaskan reindeer ranges are Cladina stellaris and C. rangiferina (Andreev 1954, Pegau' 1968, Holleman et al. 1979). Studies of caribou rangeland on Southampton Island, N.W.T., i n d i -cated Cetraria, i n c l u d i n g Cetraria aucullata, was the dominant l i c h e n genus (Parker 1976). Near S i t i d g i Lake, i n the southwestern portion of the study area, I n g l i s (1975b) found Cladina rangiferina, C. mitis and Cetraria niv-alis composed over 75% of the l i c h e n ground cover i n reindeer feeding c r a t e r s . While c e r t a i n rare l i c h e n species appear strongly selected for or against, the most common on Rangifer winter ranges are often selected i n r e l a t i o n to t h e i r r e l a t i v e abundances (Courtright 1959, Pegau 1968, Skuncke - 118 -1969). In an Alaskan tundra range of raised-centred ice-wedge polygons dominated by Cetraria aucullata and Cladina spp., reindeer were observed to s e l e c t these taxa i n r e l a t i o n to t h e i r r e l a t i v e standing crops (White & T r u d e l l 1980b). Future studies of feeding preferences on the Tuktoyaktuk Peninsula reindeer w i l l l i k e l y confirm that these four species are the winter d i e t mainstays. Differences i n top to bottom l i c h e n biomass components ( F i g . 15) r e f l e c t a composite of two major r e l a t i o n s h i p s i n the lichens growing among the three vegetation groups A, B anc C. F i r s t , there i s a s h i f t i n the r e l a t i v e amounts of the two l i c h e n genera that represent most of the biomass. Both have d i f f e r e n t growth forms. The Cladina: Cetraria r a t i o s show i n vegetation group A the dominant taxa are Cetraria spp. that have f l a t t e n e d , strap-shaped and leathery t h a l l i whereas i n groups B and C, Cladina spp. with t h e i r up-r i g h t , branched and intertwined podetia, are dominant. Secondly, growth con-d i t i o n s are appreciably d i f f e r e n t for lichens within the three vegetation groups. In group A, h i l l crest and upper slope locations and coarser-tex-tured s o i l s i n d i c a t e dry s i t e conditions, reduced humidity at m i c r o s i t e s , and g e n e r a l l y open-growth conditions subject to wind-dessication and cold winter ground-surface condi t i o n s . In groups B and C, lower posi t i o n s are protected from wind-scouring, deeper winter snow accumulations protect against f r o s t i n j u r y , and the greater occurrence of organic s o i l s and generally higher organic G contents in mineral s o i l s (Table X) provide more humid m i c r o s i t e s . Cryoturbation creates an i r r e g u l a r hummocky surface that provides many d i f -ferent microsites for l i c h e n s . The c u r v i l i n e a r r e l a t i o n s h i p of top to bottom l i c h e n biomass may also r e f l e c t a small error factor from f i e l d c o l l e c t i o n s and measurements. Gener-- 119 -a l l y l i c h e n at group A s i t e s tended to grow more interspersed with low, de-cumbent vegetation such as Arctostaphylos rubra, Dryas integrifolia, Pyrola grandiflora and Vaccinium vitis-idaea and c o l l e c t i o n s had, as a r e s u l t , greater amounts of extraneous material to be separated out. Although i n the laboratory the same c r i t e r i a were used to separate top and bottom components, the rate of t h a l l u s decomposition and natural senescence r e l a t i v e to the rate ofgrowth may vary, p a r t i c u l a r l y among the more abundant species. Lichen standing crop measurements at other North American subarctic and a r c t i c locations are comparable to estimates obtained in the present study (Table XXII). Results by other researchers range broadly from 24 kg.ha - 1 for a low-centred ice-wedge polygon trough that had a 2% l i c h e n cover (Williams et al. 1978) to a Picea mariana l i c h e n woodland that had a l i c h e n standing crop of 9,392 kg.ha - 1 with a 97% l i c h e n cover (Rencz & A u c l a i r 1978). Be-cause the standing crop values i n Table XXII are from a range of ecosystem types that have a range of l i c h e n ground cover values, there i s no c o r r e l a -t i o n of decreasing standing crop with increasing l a t i t u d e as might otherwise be expected. Lichen woodland ecosystems occurring south of the t r e e l i n e have l i c h e n standing crops that range higher than those of other ecosystems i n d i -cated i n Table XXII. 2. I n t e r p r e t a t i o n and Analysis of Large-Scale A i r Photographs 2.1 General The present i n v e s t i g a t i o n attempted to define a role for large-scale 70 mm photographic systems in a r c t i c rangeland studies. Numerous LANDSAT and medium-scale (e.g., 1:20,000-1:60,000) photo-interpretation studies have been undertaken of Rangifer winter rangeland (Chapt. I I , Sections 3.2 and 3.3) Table XXII. Selected northern ecosystems used for comparison of li c h e n standing crop. (1) and (3) Parker (1976), (2) Sims & Stewart (1981), (4) Moser et al. (1979), (5) Williams et al. (1*978), (6) present study, (7) M i l l e r (1976), (8) R e n c z & A u c l a l r (1978). Ecosystem Location Percent l i c h e n cover Lichen standing crop (kg.ha-1) (1) Raised l i c h e n - Dryas sedge heaths (2) Subarctic Picea mariana bog on ra i s e d peat plateau (3) Lichen-covered a l l u v i a l shingles along drainageways (4) Alpine l i c h e n heaths grazed by caribou Southampton Island, NWT. Gillam, N. Manitoba Southampton Island, NWT. Anaktuvuk Pass, Alaska (5) Ice-wedge polygons (centres, margins, Barrow, Alaska troughs) and sedge tussock meadows (6) Various ecosystems (low shrub heath, ericaceous heath, lic h e n heath, etc. (7) Upland Picea mariana and P. glauca l i c h e n woodlands (8) Picea mariana lichen woodland Tuktoyaktuk Peninsula area, NWT. Whiskey Jack Lake area, N. Manitoba S c h e f f e r v i l l e , N. Quebec 6-57 10 50 (approx.) 2-32 20-89 18-92 97 63-529 780 767-1,624 1,491-2,238 24-5,841 194-6,378 560-6,777 9,392 - 121 -but the r e s u l t s of these studies gave l i t t l e i n d i c a t i o n of l i c h e n cover and abundance at l o c a l l e v e l s . The broad vegetational typings that are normally described by these studies are not d e f i n i t i v e enough to be p a r t i c u l a r l y use-f u l for rigorous rangeland management. Soviet and northern European r e -searchers i n p a r t i c u l a r have preferred a e r i a l - v i s u a l assessments of rangeland from low f l y i n g a i r c r a f t (Chapt. I I , s e c t i o n 3.1). This technique permits l i c h e n , as the animals' main winter foodstuff, to be properly q u a n t i f i e d . However, only l a r g e - s c a l e photographs allow r e l i a b l e estimates of range q u a l i t y while also providing a basis for long term monitoring. Medium-scale photographs c o n s i s t i n g of a v a i l a b l e NAPL black and white 1:60,000 photos and acquired CIR 70 mm 1:34,000 photos were employed only to a l i m i t e d extent i n the present i n v e s t i g a t i o n , p r i m a r i l y i n l o c a t i n g ground-truth s i t e s and f l i g h t l i n e s . Past work in the study area which used medium-scale remote sensing data e x c l u s i v e l y , provided l i m i t e d information on the rangeland. Vegetation types, for part of the study area were mapped using 1:60,000 NAPL photos (Forest Management I n s t i t u t e 1974, 1975) but r e s u l t i n g map units did not allow i n t e r p r e t a t i o n s of percent l i c h e n cover, mainly be-cause of i n s u f f i c i e n t levels of r e s o l u t i o n , and i n a b i l i t y to d i s c r i m i n a t e l i c h e n from other features having s i m i l a r high reflectances on the black and white f i l m . Over 90% of the study area was mapped (Forest Management I n s t i -tute 1975) within two vegetation-type aggregates: (1) high and low shrubs on upland s i t e s with good drainage, and (2) low shrub, moss, l i c h e n and grass on v a r i a b l e to f l a t t e r r a i n , with poor to f a i r drainage. At a few selected locations within the study area, Corns (1974) c a r r i e d out preliminary mapping based on ground-truth of vegetation. His exercise was l i m i t e d by the scale of the 1:60,000 NAPL photographs used, and at three areas, 'Eskimo Lake', - 122 -'Caribou H i l l s ' and 'Tuktoyaktuk', 88.6%, 92.0% and 75.5% of the land area r e s p e c t i v e l y was interpreted as complexes of low shrub-heath, b i r c h - a l d e r -heath, and raised-centred polygons (Corns 1974, Hernandez 1974). C l e a r l y such broad and complex units provide no basis for determining l i c h e n forage type or abundance on Rangifer winter rangeland. Large-scale photo measurements (Table III) were conducted i n a standard 80% non-forward-overlap area of each photo-frame. Variations i n percent f o r -ward overlap were not accounted for although these were generally minor sources of error (Table X I I I ) . Because of the wingtip c o n f i g u r a t i o n for photography, l e f t and r i g h t cameras photographed n e a r l y - i d e n t i c a l ground areas ( i . e . , 95-100% side-over-lap) so that a f u l l stereographic view was p o s s i b l e . V i g n e t t i n g e f f e c t s par-t i c u l a r l y common with CIR films and shorter f o c a l length lenses (Goba et al. 1982) were not s i g n i f i c a n t on most large-scale photographs but become an ob-vious "edge-effect" on the 1:34,000 scale photographs (e.g., see F i g . 8). Radial d i s t o r t i o n as well was not an appreciable problem with the large-scale photographs although i t l i k e l y contributed a small error factor to percentage cover c a l c u l a t i o n s . Radial d i s t o r t i o n created the i l l u s i o n of minor slopes on photo-edges p a r t i c u l a r l y evident when viewing f l a t t e r r a i n . Adopting a regional zonation scheme for the study area was made d i f f i -c u l t by the lack of consensus among past researchers as to where boundaries should l i e ( F i g . 3a-d). Preliminary d i g i t a l and optical, studies of recent LANDSAT scenes also did not c l a r i f y matters. Ultimately, the ecoregion/eco-d i s t r i c t d i v i s i o n s (Houseknecht 1981) with some a d d i t i o n a l subdivisions to one e c o d i s t r i c t , were adopted as they are i n t e g r a t i v e units based on several - 123 -e c o l o g i c a l factors including climate, t e r r a i n , hydrology and vegetational cover. Being e s s e n t i a l l y e c o l o g i c a l units they were f e l t more appropriate for a study that b a s i c a l l y used an e c o l o g i c a l approach. Houseknecht 1s (1981) e c o d i s t r i c t units can be generally c o r r e l a t e d to the more prominent physio-graphic d i v i s i o n s given by Mackay (1963). For the purposes of the present study, the r e s u l t i n g four e c o l o g i c a l land c l a s s i f i c a t i o n units employed along with three a d d i t i o n a l subdivisions have been re f e r r e d to as reindeer manage-ment zones. Land areas for the photo-frames (Table XIII) were c a l c u l a t e d by sub-t r a c t i n g the areas occupied by water i n each photo-frame from the t o t a l non-forward-overlap surface areas. Figures on the areas occupied by water how-ever do not permit a meaningful estimate of percent water cover i n the study area since cameras were turned o f f to conserve f i l m when the f l i g h t l i n e s passed over water bodies larger than a few hectares. Estimates for percent water cover i n the reindeer management zones are given i n Table IV from dot-g r i d overlays on 1:250,000 mapsheets and compare favourably to estimates given by Mackay (1963). Across the region he found, from systematic samp-li n g s of 1:60,000 NAPL photographs, that percent water ranged from 0-5% around Reindeer Station, to 30-50% for most of the Tuktoyaktuk Peninsula, to over 50% near Cape Dalhousie at the NE t i p of the Peninsula (Mackay 1963). 2.2 Ice-wedge Polygons The abundance and density of ice-wedge polygons i n the Mackenzie Delta area have not been estimated before. Mackay (1963) noted that i n the region east of the Mackenzie Delta polygons were the most conspicuous type of pat-terned ground and were widespread i n poorly drained areas, p a r t i c u l a r l y - 124 -former lake bottoms and sedgy depressions. In many locations i n the study area, p a r t i c u l a r l y low areas among h i l l s , they formed small polygon f i e l d s often less than a hectare i n s i z e . Consequently, numbers and areas of cover-age were not e a s i l y t a l l i e d on small-scaled imagery such as 1:60,000 black and white air-photos. Sager (1951) stated that to resolve ice-wedge polygons on a e r i a l photos, a scale of 1:50,000 or larger was required. Only near the t i p of the Tuktoyaktuk Peninsula, i n the Point Atkinson area and eastward are ice-wedge polygons, p a r t i c u l a r l y the low-centred forms, extremely abundant, extending through the f l a t s unbroken for many kilometres (Mackay 1963). The present studies ind i c a t e that percent of land cover estimates for polygons ranged from 4.6% in reindeer management zone C, to 43.4% in zone A (Table V I I I ) . Using these estimates and t o t a l land area estimates of zones (Table IV) a t o t a l of 3,371 sq km of the 14,410 sq km study area, or 23.4% i s c a l c u -lated t e n t a t i v e l y as covered by ice-wedge polygons. As noted i n the ground-truth r e s u l t s , most ice-wedge polygons were i n d i -c ative of vegetation group C which o v e r a l l supported the most luxuriant l i c h e n growth. As a g e n e r a l i z a t i o n then, ice-wedge polygon rims and crests c o n s t i t u t e some of the best reindeer rangeland i n the Tuktoyaktuk Peninsula area. However, since vegetation group C was not comprised s o l e l y of i c e -wedge polygon s i t e s there is an understandable lack of correspondence between the a i r photo-interpreted percent of land area estimates for polygons (Table XV) and t o t a l l i c h e n standing crop estimates (Table XXI) for the same r e i n -deer management zones. It should be recognized that neither si z e nor density of ice-wedge poly-gons are r e l i a b l e i n d i c a t o r s of the depth and width of subsurface ice-wedges or the t o t a l amount of i c e i n the ground and thus for t r a f f i c a b i l i t y or s i m i -- 125 -l a r a p p l i c a t i o n s the r e s u l t s must be used ca u t i o u s l y . As w e l l , ice-wedges without surface expression are known to ex i s t throughout the Tuktoyaktuk Peninsula area (J.R. Mackay pers. comm.-'-). Mackay (1970) notes that ice-wedges i n the Mackenzie Delta region may be spaced as c l o s e l y as 15 m apart, and that one square km of t e r r a i n may have over 65 l i n e a r km of ice-wedges. Based on the present counts of numbers of ice-wedge polygons per ha on the large-scale CIR photographs (Table XV), i c e -wedges are estimated to be more c l o s e l y spaced, ranging from 11.4 to 7.5 m for d i f f e r e n t management zones. Such closer spacing y i e l d s i n one square km of ice-wedge polygon-covered t e r r a i n , from about 88 to over 133 l i n e a r km of ice-wedges. Mackay (1963) and Kerfoot (1969, 1972) c l e a r l y defined the surface ex-pression of the two main extremes, i . e . , raised-centred and low-centred poly-gons. Gradations ex i s t between them that r e s u l t from many factors i n c l u d i n g p r i m a r i l y age, r a p i d i t y of thaw and growth of ice-wedges, surface erosion by water, wind and mass wasting, vegetation, type of material and water con-tent. The large-scale photos were useful for d i s c r i m i n a t i n g among several intermediate forms and for c a l c u l a t i n g t h e i r r e l a t i v e abundances and densi-t i e s , however those r e s u l t s are not reported here. 2.3 T e r r a i n Disturbance by Vehicles E s c a l a t i n g northern a c t i v i t i e s , i n c l u d i n g resource exploration a c t i v i -t i e s , within and near the Tuktoyaktuk Peninsula area have prompted considera-J.R. Mackay, Dep. Geography, Univ. B r i t i s h Columbia. 6 May, 1982. - 126 -ble public concern over the p o s s i b i l i t i e s of widespread and i r r e p a r a b l e dam-age to the natural environment. The public are apprehensive because the tun-dra i s f r a g i l e , and man-induced stresses at the ground surface may destroy the thermal e q u i l i b r i u m between ground cover and the underlying permafrost, leading to thermokarst subsidence (Mackay 1970, Kerfoot 1972). From examina-t i o n of the larg e - s c a l e photos, most t e r r a i n damage by veh i c l e s i n the study area could be classed as low (Table XV). Severe damage l e v e l s , p o s s i b l y the re s u l t of seismic l i n e construction i n summer, were only encountered i n 1.6% of the photos, and occurred i n reindeer management zone C, i n the v i c i n i t y of the Tuktoyaktuk townsite. The area of coverage was low, ranging from 0.1% i n reindeer management zones A, E and G to 2.0% i n zone C. However, the high frequency of occurrence by disturbed t e r r a i n was notable; v e h i c l e tracks were observed i n 19.0% of the large-scale photographs. T e r r a i n damage was ob-served i n a l l zones (Table XV). Using percent cover estimates (Table XV) and t o t a l land area estimates of zones (Table IV), a t o t a l of 70.1 sq km or.0.5% of the study area is covered by p r i m a r i l y low-level t e r r a i n damage from v e h i c l e s . B l i s s & Wein (1972) have reviewed the early h i s t o r y of o i l e x p l o r a t i o n and technology i n the Mackenzie Delta region, from i n i t i a l b u l ldozing for seismic operations during the summer months of 1965 to, with the r e a l i z a t i o n of the extent of damage th i s caused, winter shear-blading operations. They estimated that while seismic l i n e s and winter roads may form conspicuous landscape features, they occupied less than 0.5% of the Mackenzie Delta area landscape i n 1971 ( B l i s s & Wein 1972). Hernandez (1973) located, aged and sampled various disturbed s i t e s i n the study area with the aid of maps provided by Imperial O i l Limited. Calcu-l a t i o n s showed that 0.56% of the land area of an intensively-surveyed 2,100 - 127 -sq km area on the Tuktoyaktuk Peninsula had been disturbed by seismic opera-tions from J u l y , 1965 to A p r i l , 1970 (Hernandez 1973). The present figures (Table XV) i n d i c a t e that the percent of land area a f f e c t e d by winter roads, seismic l i n e s and v e h i c l e tracks remains a small proportion of the t o t a l land area. However, while the mean cover by damaged t e r r a i n i s only 0.5% for the whole of the study area, i t i s somewhat larger i n c e r t a i n zones. In zone C, which includes a major portion of the area i n t e n s i v e l y surveyed by Hernandez (1973), 2.0% of the land area has been disturbed by v e h i c l e s (Table XV). This figure i s larger than Hernandez's (1973) and includes low-damage-level ve h i c l e tracks that were previously not resolveable on small-scaled remote sensing data. However some increase i n t e r r a i n damage during recent years i s also i n d i c a t e d . The large-scale photographs now w i l l serve as an important baseline in the Tuktoyaktuk Peninsula area for monitoring: Ci) the recovery of those areas where v e h i c l e t e r r a i n damage has occurred, and ( i i ) the future occurr-ence of a d d i t i o n a l disturbances, e s p e c i a l l y in l i g h t of increasing use of snow machines and other tracked vehicles by northern inhabitants, and recently-renewed searches for northern energy resources within the study area. The present areas lost as productive Rangifev winter rangeland because of v e h i c l e t e r r a i n disturbance are small and not s i g n i f i c a n t . 2.4 Lichen Types and Percent Lichen Cover Lichen Types are the air-photo i n t e r p r e t e d equivalents of the vegetation groups A, B and C determined from ground-truth studies. Only the dominant Lichen Type was used to characterize each lar g e - s c a l e photo-frame; on the ground, however, the same land area might be assigned among several vegeta-t i o n groups. - 128 -Quantitative analysis of microdensitometric data on li c h e n patches i n d i -cate that Lichen Types could be "readily separated. Dye layer density measures on the large-scale photos gave mean values (Table XVI) that r e f l e c t , i n the approximate 3.1 sq m r e s o l u t i o n area of the microdensitometer's aper-ture, not only the l i c h e n patch but other vegetation and bare ground cover that i s t y p i c a l l y found i n close a s s o c i a t i o n with i t . Because of t h i s obser-vation, i t i s f e l t that the r e s u l t s reported here are highly dependent on photo-scale and the microdensitometer's aperture s i z e . Rangeland shrub and other vegetation species can be accurately i d e n t i -f i e d on l a r g e - s c a l e 70 mm CIR photographs using microdensitometry ( D r i s c o l l & Coleman 1974, D r i s c o l l et al. 1974). I n d i v i d u a l plant taxa were not system-a t i c a l l y i d e n t i f i e d on the l a r g e - s c a l e photographs in the present work, a l -though microdensitometric studies could have been e f f e c t i v e i n separating c e r t a i n species, p a r t i c u l a r l y low shrubs. Separation of l i c h e n species would probably have met with l i t t l e success, however, as ( i ) they r a r e l y grow i n pure carpets of a s i n g l e l i c h e n taxa, without "contamination" by other li c h e n s , mosses or low vascular species; and ( i i ) a l l l i c h e n species encount-ered with s i g n i f i c a n t ground cover for microdensitometric spot readings were characterized by high r e f l e c t a n c e s and appeared white to yellow-white i n colour. Percent lichen cover was estimated i n the 80% non-forward-overlap area of each photo-frame using a c e l l - g r i d overlay. Lichen cover, because of i t s high r e f l e c t a n c e i n the near-infrared portion of the spectrum ( F u l l e r & Rouse 1979) i s r e a d i l y observed on the large-scale CIR photos ( F i g . 13b). Lichen cover may have been underestimated during measurements on the l a r g e - s c a l e photo-frames due to several reasons. F i r s t , low shrub, p a r t i c u l a r l y - 129 -Betula, Salix spp. and Alnus, or Eriophorum vaginatum tussock cover may mask an unknown p o r t i o n of the l i c h e n cover. Normally t h i s was a small e r r o r source, but systematic comparisons of ground-truth measurements of percent l i c h e n cover with a i r - p h o t o estimates i n d i c a t e i n some community-types the e r r o r may be 10% or more. Second, Eriophorum and Carex tussocks and small c r y o t u r b a t e d hummmocks create numerous small shadowed areas on some l a r g e -s c a l e photos. At these northern l a t i t u d e s , even at midday sun angles are low i n e a r l y August. T h i r d , around photo-edges, lens f a l l - o f f e f f e c t s may pro-duce lower r e f l e c t a n c e l e v e l s by small l i c h e n patches, causing the l i c h e n to be overlooked i n percent cover estimates. Conversely, a few sources of e r r o r p o s s i b l y caused o v e r e s t i m a t i o n of l i c h e n cover i n the photos. F i r s t , bare sandy s o i l patches because of t h e i r s i m i l a r high ground r e f l e c t a n c e s can be o c c a s i o n a l l y m i s i n t e r p r e t e d as l i c h e n . Bare sand occurred commonly on h i l l top or upper slope l o c a t i o n s that were Lichen Type I . Second, graminoid standing dead which sometimes appears b r i g h t white i n the l a r g e - s c a l e photos (e.g. F i g . 18a) was s i m i l a r l y m i s i n t e r p r e t e d . Such m i s i n t e r p r e t a t i o n s however were minimized as photo-i n t e r p r e t a t i o n s were c a r r i e d out by the same personnel who conducted the ground-truth programs. The percent l i c h e n cover estimate i n c l u d e s a number of l i c h e n taxa that have no value as Rangifer forage. These taxa however appear to account f o r a s m a l l p r o p o r t i o n of t o t a l l i c h e n cover i n the study area. Two non-forage l i c h e n taxa w i t h high percent occurrences at s i t e s are Ochrolechia gyalecta and Bryoria nitidula and both have very low cover (Appendix I ) . Nonetheless, i n subsequent c a l c u l a t i o n s of l i c h e n standing crops and c a r r y i n g c a p a c i t y , the percent l i c h e n cover by non-forage species can be considered' an a d d i t i o n -a l source of e r r o r . - 130 -It is f e l t that the vertical format afforded by the large-scale 70 mm photos provided an improved view of the lichen cover over an oblique format (Fig. 22). Both formats were particularly affected by shrub and herb strata that obstruct views of the ground surfaces, and result in underestimations of ter r e s t r i a l lichen, cover. Based on comparisons of the large-scale 70 mm photos and oblique 35 mm photographs, the vertical format is preferred. As Kiichler (1967) noted regarding vegetation cover measurements and mapping, only vertical photographs permit a correct grasp of the geographical d i s t r i -bution of the different vegetation types in the landscape, and this would appear true as well when speaking of particular strata of vegetation. It was stated in the introduction that lichen, as the main winter food-stuff for reindeer, could only be properly quantified at a large-scale but was required for regional management. An indication that lichen occurrence or abundance is not well manifested on a small-scale remote sensing data is reflected in the considerable differences of lichen percent cover and stand-ing crop among reindeer management zones C, D and E (Tables XX, XXI) which according to Houseknecht (1981) belong to the same ecodistrict. Such d i f f e r -ences are apparently not being reflected at the ecodistrict l e v e l . 2.5 Estimation of Carrying Capacity The winter rangeland carrying capacity, the number of animals the study area could support on a long-term basis without appreciable overutilization can be tentatively estimated. Summer rangeland is not considered limiting in the study area although summer rangeland quality largely determines the physical stature of the animals (Palmer 1934, Skuncke 1969, Klein 1970, 1982). Porsild (1929) suggested the summer rangeland of the Tuktoyaktuk Pen-- 131 -F i g . 22. Comparison of oblique normal colour and v e r t i c a l CIR photographs of the same area: (a) Oblique 35 mm normal colour photograph of l i c h e n closed mat and cushion tundra on patterned ground (lower l e f t of photo; s i t e 81-104, with l i c h e n cover of 47.5 percent), and wet sedge meadow (upper r i g h t ; s i t e 81-105) as shown i n F i g . 14 ( f l i g h t l i n e 12-3; 69°01*N, 132°12'W). 30 J u l y , 1981. (b) V e r t i c a l 70 mm CIR (1:1,800 scale) photo-pair of the same area showing s i t e s 81-104 (top centre) and 81-105 (bottom centre by open water). Note lime-green c o l o u r a t i o n of a l g a l mats, the same shown i n F i g . 14. View angles of both F i g . 14 and 22a are from the 10 o'clock p o s i t i o n of F i g . 22b. Systematic comparisons of ground and a e r i a l oblique photographs, both normal colour and CIR, with the v e r t i c a l CIR photo-pairs i n d i c a t e the oblique angle of view c o n s i s t e n t l y leads to lower impressions of ground l i c h e n cover than does the v e r t i c a l view (frame 8, f l i g h t l i n e 12-3). 6 August, 1980. - 133 -i n s u l a could support 25 to 28 animals per sq km. On this basis, reindeer management zone A, which i s presently used e x c l u s i v e l y as summer rangeland f o r the reindeer (e.g., see F i g . 5), could alone support over 48,000 animals. While P o r s i l d ' s (1929) f i g u r e i s not based on any q u a n t i t a t i v e s t u d i e s , i t suggests that summer rangeland i s u n l i k e l y to be l i m i t i n g . In the Tuktoyaktuk Peninsula area i t i s the a v a i l a b i l i t y of l i c h e n f o r -age during the winter which l i m i t s c a r r y i n g capacity. Past estimates of carry i n g capacity have been made, but based only on casual observations and comparisons with stocking l e v e l s i n the U.S.S.R., Northern Europe and Alaska. In his i n i t i a l surveys of the area east of the Mackenzie Delta, P o r s i l d (1929, 1947) f e l t the e n t i r e Grazing Preserve could support about 85,000 reindeer. Reviewing more recent l i t e r a t u r e , H i l l (1967) and Scotter (1968) agreed on a much lower, estimate of 25,000 to 30,00.0 reindeer, with the l a t t e r author s t r e s s i n g that the estimate assumed r o t a t i o n a l use by the herded reindeer of the e n t i r e 46,360 sq km area of the Grazing Preserve (Scotter 1970). Igoshina and Florovskaya (1939, c i t e d i n Parker 1976) i n d i c a t e that a s i n g l e adult reindeer eats 5 to 6 kg of l i c h e n per day. DesMeules and Hey-land (1969) found woodland caribou i n Laurentide Park, Quebec consumed 2.7 to 2.9 kg of l i c h e n per day when confined and provided with an u n r e s t r i c t e d fodder supply. Using f a l l o u t radiocesium t r a c e r s , a method unique i n that i t does not a f f e c t the normal a c t i v i t y of the animals, Holleman et ai. (1975, 1979) measured l i c h e n intakes of 4.9 to 5.0 kg per day for free-ranging adult reindeer and caribou. In present c a l c u l a t i o n s of carr y i n g capacity a d a i l y l i c h e n consumption of 5.0 kg was assumed. - 134 -Stocking rates for Rangifer winter rangeland were estimated using the technique of Pegau (1968) as modified by Parker (1976). Both authors employ a "top cropping" parameter in estimations to restr i c t grazing to the actively-growing portions of lichen t h a l l i only. According to Andreev (1954) this constitutes 45% of the lichen weight and about the top one-third of height. At approximately this level of grazing a lichen rangeland can f u l l y recover in three to five years (Andreev 1954, Skuncke 1969). This practice allows a far greater number of reindeer per unit of productive range than intensive grazing of lichen stands where f u l l recovery may require 30 to 50 years or more (Andreev 1954, Scotter 1970). It was f e l t that in present c a l -culations the 'top' estimates of standing crop derived from actual f i e l d and laboratory measures (Table XXI) would be more acceptable. The provision for a four-year rotation (af. Parker 1976) was retained. Reindeer management zones A, B and C have potential for use as summer rangeland (e.g. see Fig. 3a). Although zone A is presently used exclusively, the Richards Island area, zone B, was highly favoured by early herders (Porsild 1947, Cody 1963, Abrahamson 1963). On summer rangeland, lichens are consumed as a small portion of Rangifer diets (Pegau 1968, Skuncke 1969) and growing t h a l l i , especially Cladina podetia are subject to trampling effects (Pegau 1970, Moser et al. 1979). The three zones available as summer range-land but having sufficient lichen cover to support limited winter use were considered u t i l i z e d 365 days a year in present calculations, while other zones (D, E, F, and G) were considered as lichen range only for winter months, approximately October 1 to May 31. Of the study area's total estimated carrying capacity of 20,373 animals, 80.5% or 16,398 reindeer could be supported in management zones D and E which - 135 -represent 49.7% of the land area (Table XXIII). This estimate suggests a mean animal stocking density of 1.4 reindeer per sq km, a figure that falls, between earlier estimates of 0.5 to 0.7 reindeer ( H i l l 1967, Scotter 1968, 1970) and 1.8 reindeer per sq km (Porsild 1929, 1947). Estimates by other authors are for the entire Grazing Preserve, and include consideration of the extensive lichen woodlands in areas south of the study area where presumably lichen standing crop is high. The greatest animal density estimated here is 3.2 in zone G, while the lowest (0.1) is in zone B (Table XXIII). The three northerly zones, A, B and C would support a combined total population of only about 1,491 overwintering animals. Given the reindeer herd's present size of about 16,000 animals (Dickin-son 1982) and the carrying capacity estimate arrived at here that is not much larger, an updated rotational grazing plan involving winter use particularly of management zones D and E is recommended. A less desirable, alternative arrangement is to maintain the herd within the rangelands of management zones A, C and D (as in Fig. 5). This arrangement would have the herd reduced to about 6,200 animals using the figures of Table XXIII, and the imposition of a four-year rotational grazing plan for the 5,787 sq km area. For this latter scheme, l i t t l e use would be made of zones A and C for winter rangeland, as is presently the case (Fig. 5). 2.6 Overutilization Effects It is d i f f i c u l t to assess the extent of Rangifer overutilization, i n -cluding both overgrazing and trampling effects, in the study area. No rigor-ous past baselines such as permanent ground quadrats or large-scale photo-graphic records exist within the study area, except for a few exclosures con-structed near Si t i d g i Lake (Inglis 1975a,b). While no quantitative data Table X X I I I . E s t i m a t i o n of winter range c a r r y i n g c a p a c i t y f or the Tuktoyaktuk Pe n i n s u l a area, N.W.T. (method adapted from Parker 1976). •top' range a v a i l a b l e p r o v i s i o n f o r a re i n d e e r t o t a l Reindeer management zone l i c h e n standing crop (kg.ha" 1) under snow cover 1 (x .5) 4-year r o t a t i o n ' 6 (x .25) g r a z i n g days-* ( d a y s . h a - 1 ) land area (sq km) rein d e e r per sq knA t o t a l r e i n d e e r A 41.6 20.8 5.2 1.0 1,948 0.27 526 B 18.8 9.4 2.4 .5 1,214 .14 170 C 52.8 26.4 6.6 1.3 2,208 .36 795 D 286.4 143.2 35.8 7.2 1,631 3.00 4,893 E 200.0 100.0 25.0 5.0 5,531 2.08 11,505 F 61.6 30.8 7.7 1.5 1,366 .63 861 G 302.4 151.2 37.8 7.6 512 3.17 1,623 20,373 i e s t i m a t e d 50% of l i c h e n forage i s u n a v a i l a b l e due to snow cover each year. ^4 yr g r a z i n g r o t a t i o n allows for continuous use at one-quarter animal d e n s i t y . ^based on d a i l y l i c h e n forage requirements of 5 kg f o r adult reindeer (Igoshina & Florovskaya 1939 [ c i t e d i n Parker 1976], DesMeules & Heyland 1969, Holleman et al. 1975, 1979) ^ f o r management zones A, B and C based on year-round (365 day) use; f o r others based only on Oct. 1 to May 31 (240 day) winter use. - 137 -e x i s t s , o v e r / u t i l i z a t i o n has been described by P o r s i l d (1947), Cody (1963) and Scotter (1968) for the v i c i n i t y of Reindeer S t a t i o n , on Richards Island and on a small i s l a n d adjacent to i t , and at a few a d d i t i o n a l l o c a t i o n s . During ground-truth for the present study what were believed to be o v e r u t i l i z a t i o n e f f e c t s were o c c a s i o n a l l y encountered mostly in reindeer management zone C. P a r t i c u l a r l y at vegetation group C s i t e s i n t h i s zone, the l i c h e n had the appearance of being trampled, with shattered l i c h e n t h a l l i scattered about, clumps of li c h e n dislodged from the substrate and numerous patches of recently-exposed organic surfaces present. At such s i t e s , reindeer droppings were t y p i c a l l y found i n abundance. During the 1981. ground-truth program preliminary use was made of an i n -dex of recent Rangifer grazing (Table XXIV). The index has been s u c c e s s f u l l y employed i n ground studies of caribou habitat preference studies i n northern Alaska (White & T r u d e l l 1980b). Results here, a v a i l a b l e for only a few s i t e s and therefore not reported i n d e t a i l , i n d i c a t e the index i s an appropriate t o o l for c h a r a c t e r i z i n g reindeer use on any rangelands suspected as over-grazed. With magnification equipment some parameters i n Table XXIV can be in t e r p r e t e d on the l a r g e - s c a l e 70 mm photos. I f further studies s u c c e s s f u l l y adapt the index for use with large-scale remote sensing data, further ground-trut h i n g may be. g r e a t l y reduced or eliminated. Some o v e r u t i l i z a t i o n may have already occurred in reindeer management zone C. It has a comparatively low l i c h e n standing crop (Table XXI, F i g . 20) and i t s carrying capacity i s estimated at only 795 animals (Table XXIII) or about one-twentieth the number that traversed parts of the zone in both spring and f a l l of past years ( F i g . 5). Trampling e f f e c t s may be p a r t i c u l a r -l y c r i t i c a l ; studies have shown that a s i n g l e pass over l i c h e n - r i c h tundra by a small reindeer herd may destroy 15% or more of the lichens (Skuncke 1969, - 138 -Table XXIV. Parameters and scores used to c a l c u l a t e an index of the i n t e n s i t y of recent Rangifer grazing (from White & T r u d e l l 1980b). The i n -dex i s c a l c u l a t e d as the sum of the scores f o r each parameter. A. Trampling e f f e c t s 0 No sign of Rangifer hoofprints 1 Occasional hoofprint v i s i b l e within 3 m 2 Broken plant parts which could be a t t r i b u t a b l e to trampling 3 Surface moss or tussocks broken by digging a c t i o n (pieces of moss and tussocks may be l y i n g on the surface) and/or the occurrence of "game" t r a i l s showing recent use B. Fecal abundance 0 No "new" feces w i t h i n 3 m of s i t e 1 1 to 2 groups of p e l l e t s within 3-m radius 2 3 to 5 groups of p e l l e t s w i t h i n 3-m radius 3 more than 5 groups of p e l l e t s within 3-m radius C. Damage to lic h e n s 0 Lichen beds i n t a c t ; Cladina, Cetraria, and Stereocaulon undisturbed i n the moss layer 1 Some lichens removed from the moss bedding 2 Many lichens l y i n g loose on the surface 3 Most lichens loose on the surface; obvious disturbance to the moss l a y -er D. Vascular plant c l i p p i n g 0 No leaves c l i p p e d 1 Signs of a small amount of leaf c l i p p i n g i n the sward and tussock 2 C l i p p i n g of sward and tussock obvious; signs of l i t t e r accumulation ( r e j e c t a ) which could be a t t r i b u t a b l e to c l i p p i n g 3 Most of sward or tussock clipped o f f ; l i t t e r r e j e c t a very obvious E. Browsing signs 0 No signs of clipped twigs 1 A few terminal twigs broken (0-3 mm diameter) 2 Many terminal twigs broken, some twigs of 5 mm diameter broken o ff 3 Twigs with greater than 5 mm diameter taken - 139 -Pegau 1970). For the l a s t two years, Canadian Reindeer (1978) Ltd. have taken steps to reduce p o t e n t i a l o v e r u t i l i z a t i o n by moving the animals across Eskimo Lake i n late February and grazing them i n management zone E u n t i l crossing back on to the Tuktoyaktuk Peninsula near Campbell Island i n l a t e May (D. B i l l i n g s l e y , pers. comm. ).! In zone C and elsewhere throughout the study area, the large-scale 70 mm photos w i l l now serve as permanent records against which any future o v e r u t i l i z a t i o n can be compared and properly quanti-f i e d . 3. Future Recommendations 3.1 Follow-up Studies on the Rangifer Rangeland (1) A c q u i s i t i o n , analysis and i n t e r p r e t a t i o n of s u i t a b l e intermediate-and small-scale remote sensing data for the study area w i l l help to place the large-scale studies with a better regional context by allowing ( i ) improved d e f i n i t i o n s of reindeer management zones, and ( i i ) the development of i n t e r -mediate l e v e l mapping units that are e s s e n t i a l l y "aggregates" of vegetation groups or Lichen Types. Such mapping units could r e f l e c t varying amounts of Lichen Types within known proportions so that estimates of li c h e n cover and other parameters obtained from large-scale studies are not l o s t . The Viereck & Dyrness (1980) vegetation c l a s s i f i c a t i o n scheme adapted here (Table V) lends i t s e l f to remote sensing studies although i t was not o r i g i n a l l y developed for that purpose. Intermediate l e v e l s of the h i e r a r c h -i c a l physiognomically-based scheme may be r e a d i l y related to d i f f e r e n t scales of remote sensing data. P o s s i b l y the system may be used for future m u l t i -stage remote sensing studies i n the Tyktoyaktuk Peninsula study area. ID. B i l l i n g s l e y , Business Advisor, Canadian Reindeer (1978) Ltd., 28 March, 1983. - 140 -The l a r g e - s c a l e photographs represent an actual sampling of only 0.1% of the study area. While this proportion probably would be i n s u f f i c i e n t f o r any area with high landscape d i v e r s i t y , the Tuktoyaktuk Peninsula area, along with many other a r c t i c areas, exhibits r e l a t i v e l y l i t t l e landscape d i v e r s i t y over broad expanses. Although preliminary analyses based, on the l a r g e - s c a l e photos appear appropriate, multistage information ( F i g . 4) should help to v e r i f y i f the l a r g e - s c a l e studies adequately represent the reindeer manage-ment zones. To this end, experimental simulated LANDSAT-4 imagery and approximately 1:50,000 s c a l e CIR photographs of the transects ( F i g . 5) were acquired on August 26, 1982 by the Airborne Operations D i v i s i o n , Canada Centre f o r Remote Sensing, Ottawa, Ontario, under s p e c i f i c a t i o n s provided by the author and Dr. P.A. Murtha 1. Evaluations of these data have r e c e n t l y been i n i t i a t e d and attempts are being made to c o r r e l a t e them to the l a r g e -scale photo and ground-truth s t u d i e s . Results w i l l be reported at a future date. (2) Supplementary a e r i a l photography at large-scales (1:1,400-1:3,600) would be h e l p f u l to b u i l d up the o r i g i n a l data-base with a d d i t i o n a l area coverage and provide multi-year coverage of the 44 f l i g h t l i n e s already photo-graphed. Rephotography of the f l i g h t l i n e s at about '5 yr i n t e r v a l s i s f e l t i n i t i a l l y d e s i r a b l e . Normal colour or CIR films should be used instead of black and white films f o r rephotographys as they ( i ) provide considerably more information on many tundra rangeland features, p a r t i c u l a r l y minor vege-t a t i o n , and ( i i ) represent n e g l i g i b l e increases i n the o v e r a l l cost of photo-missions conducted i n northern areas ( M i l l e r and Barnhardt 1973). While the ^Faculty of Forestry, Univ. B r i t i s h Columbia. - 141 -wingtip 70 mm camera system using CIR f i l m i s advocated here, a 35 mm motor-driven camera which can obtain forward-overlap stereophotographs might be an acceptable and less c o s t l y a l t e r n a t i v e . Other rangeland studies have found such a system useful (Heintz et al. 1979) and the photography could be con-ducted from the Canadian Reindeer Ltd. observation plane without any modi f i -cations to the a i r c r a f t . Goba et al. (1982) have discussed the r e l a t i v e advantages of the two camera systems. A useful addition to future large-scale photo studies i n a r c t i c rangeland would be a radar altimeter which f o r each photo-frame gives a highly accurate measure of f l y i n g height (Kirby and H a l l 1980). Future photo-missions should be conducted as c l o s e l y as pos s i b l e to the phenological stage captured by the Aug. 5-8 timing of the present study. Phenological developments of plants i n a r c t i c areas proceed r a p i d l y through the short summer months. Later f l y i n g dates might provide photos i n which senescence of graminoid vegetation has proceeded to a point where separation of the standing dead from l i c h e n cover i s d i f f i c u l t . Photography of e a r l i e r phenological stages could be affected by other problems. For example, the ubiquitous tundra cottongrass Eriophorum vaginatum has a promi-nent w h i t e - b r i s t l e d i nflorescence i n f r u i t that could from the a i r be mis-taken as l i c h e n . On open tundra by early August the f r u i t i n g heads have usu a l l y thinned and l i k e l i h o o d of confusion with li c h e n cover has decreased. (3) Future ground-truth programs of the rangeland merely for baseline are probably unnecessary, however i f a monitoring mode i s established with the a c q u i s i t i o n of multi-year, la r g e - s c a l e photographs i t may be d e s i r a b l e to f i e l d check a proportion of the s i g n i f i c a n t changes perceived on such photos. During an i n i t i a l probationary period, the flow diagram for assigning s i t e s to vegetation groups ( F i g . 21) could be assessed. I f found to perform - 142 -adequately then a 'streamlined' ground-truth approach at s i t e s could involve: ( i ) assignment to vegetation groups using i n d i c a t o r species and d e c i s i o n rules ( F i g . 11, 21); ( i i ) measurements i n 10 m x 10 m plots of major cover features as given i n Table VIII; ( i i i ) additonal notes that include general observations, slope p o s i t i o n measurements, and ground cover by dominant l i c h e n taxa, and ( i v ) estimation on the ground of recent Rangifer use (Table XXIV). At l e a s t portions of the c r i t e r i a l i s t e d i n Table XXIV could be remotely-sensed on the l a r g e - s c a l e (1:1,400-1:3,400) CIR photo-pairs, p a r t i c -u l a r l y i f stereo-magnification i s used. Future monitoring should incorporate aspects of the index of i n t e n s i t y of recent Rangifer grazing into the p h o t o - i n t e r p r e t a t i o n work. Present observations i n d i c a t e that damaged l i c h e n beds, trampling e f f e c t s , and f e c a l abundance ( p a r t i c u l a r l y black feces on white l i c h e n mats) are d i s c e r n i b l e under magnification. Development of a r e l i a b l e i n t e r p r e t a t i o n scheme f o r the index on remote sensing data could obviate the need f o r f u r t h e r c o s t l y ground-truth. (4) It i s d e s i r a b l e that an i n i t i a l r o t a t i o n a l grazing plan f o r the Tuktoyaktuk Peninsula reindeer herd be developed now. P r o v i s i o n s , however, should be made to modify and redevelop the plan as more information becomes a v a i l a b l e from several sources. Large-scale photo coverage of a d d i t i o n a l areas may be otained. The computer-based, photo-frame data obtained i n the present study e a s i l y accept a d d i t i o n a l photo-frame data as they become a v a i l a b l e . Results of multistage studies ( F i g . 4) may lead to more informed d i v i s i o n s of management zones into subareas for reindeer r o t a t i o n a l use. Better approaches for estimating carrying capacity, or improved estimates f o r - 143 -some parameters such as the proportion annually unavailable to the reindeer because of snow cover may become a v a i l a b l e . Of p a r t i c u l a r value i n this r e -gard would be d e t a i l e d forage preference and ingestion-rate studies on the Tuktoyaktuk Peninsula area's reindeer population. Results from long term monitoring studies may be used to revise the grazing plan. In this regard range that i s l o s t over time due to o v e r u t i l i z a t i o n , w i l d f i r e s , t e r r a i n dam-age or other causes must be removed from the grazing plan, and c a r r y i n g cap-a c i t y estimates adjusted accordingly. 3.2 Other P o s s i b l e Applications f o r the Large-Scale A i r Photographs Numerous other p o t e n t i a l a p p l i c a t i o n s e x i s t f o r the l a r g e - s c a l e 70 mm photographs already acquired i n the course of this study. Results reported here on the frequency and cover by ice-wedge polygons and v e h i c l e t e r r a i n disturbance are two examples of a p p l i c a t i o n s not s o l e l y of i n t e r e s t to Rangifer rangeland management. As a d d i t i o n a l examples, three f u r t h e r a p p l i -cations are suggested f o r the photographs. (1) A p h o t o - i n t e r p r e t a t i o n study of s i l t a t i o n patterns and r i p p l e s on the sublacustrine benches of shallow lakes could elucidate the e f f e c t s of wind dynamics on lake shapes. Mackay (1963) e a r l i e r discussed the r o l e of wind and wave ac t i o n i n developing the c h a r a c t e r i s t i c lemniscate to oval lakes found i n p a r t i c u l a r on the northeast Tuktoyaktuk Peninsula. Numerous lake margins i n this area are covered by the l a r g e - s c a l e 70 mm photographs. (2) The photographs could be a valuable component i n c e r t a i n a u t e c o l o g i -c a l vegetation studies. For example, sedge tussock d e n s i t i e s i n various habitats or at various slope p o s i t i o n s could be measured. For a few taxa such as Eriophorum vaginatum a highly accurate t a l l y of flowering on f r u i t i n g - 144 -heads per unit area could be r e a d i l y obtained from the photographs. When coupled with seed v i a b i l i t y studies such as those conducted by B l i s s & Wein (1973) f o r E. vaginatum i n the Inuvik area, v i a b l e seed production p o t e n t i a l s on a per area basis could be developed. (3) A r c t i c ground s q u i r r e l s (Spermophilus parvyii Richards.) are common throughout the study area along riverbanks, rims of lake basins, and on sandy slopes. Entrances to resident burrows i n s q u i r r e l colonies are p l a i n l y v i s i -b l e , as are most runways among burrows, on many of the l a r g e - s c a l e 70 mm photographs. B a t z l i & Sobaski (1980) have noted that resident burrows are u s u a l l y occupied by one or more ground s q u i r r e l s and that most burrows have s i x or more entrances i n the Atkasook area of Alaska. E x t r a p o l a t i o n of these or s i m i l a r values along with i n t e r p r e t a t i o n of burrow entrance d e n s i t i e s on the photos could lead to i n i t i a l estimates of the ground s q u i r r e l population s i z e i n the Tuktoyaktuk Peninsula area. It i s worthwhile to note that a l l of the preceding studies and numerous others would benefit from the temporal aspect afforded by rephotography of the f l i g h t l i n e s at future dates. A more general point can be made regarding the future usefulness of l a r g e - s c a l e a i r photographs. Within a few years new s a t e l l i t e systems such as LANDSAT-D and SPOT w i l l be providing comprehensive, r e p e t i t i v e coverage of earth resources at g r e a t l y improved resolutions (Audet & Thomson 1982). Medium-scale 1:60,000 photographs, presently so widely-used p a r t i c u l a r l y f o r northern studies (Rubec 1982), may soon become v i r t u a l l y obsolete as r e -searchers instead make use of up-to-date s a t e l l i t e imagery at comparable or s l i g h t l y smaller s c a l e s . For i n t e r n a t i o n a l s e c u r i t y and many tech n o l o g i c a l reasons, l a r g e - s c a l e photographic systems w i l l not be replaced i n the f o r s e e -able future, at l e a s t f o r widespread p u b l i c use, by e a r t h - o r b i t i n g s a t e l l i t e - 145 -remote sensors. The impending a v a i l a b i l i t y however, of improved-resolution s a t e l l i t e coverage w i l l undoubtedly only enhance future requirements f o r a i r -borne l a r g e - s c a l e remote sensing data and data analysis techniques. Large-scale a i r photographs w i l l be valuable aids f o r i n t e r p r e t a t i o n of higher-r e s o l u t i o n s a t e l l i t e scenes. - 146 -V I I CONCLUSIONS - 147 -(1) Based on cover by 420 plant taxa at 112 s i t e s , vegetation of the study area can be c l a s s i f i e d among at l e a s t eighteen community-types by a two-way i n d i c a t o r species analysis (TWINSPAN). Four 'vegetation groups' (A,B,C, and D) can be defined at broad l e v e l s of the c l a s s i f i c a t i o n . Using d e c i s i o n r u l e s , the TWINSPAN analysis indicates that s i t e s can be r a p i d l y assigned during ground-truth to one of the four groups based on presence or absence of only seventeen plant species. The two-way tabular arrangement of plant taxa and vegetation groups show d i s t i n c t preferences by most taxa f o r one or another of the groups; only 70 taxa are common to a l l four groups. Species richness varies among the groups. F i e l d observations at s i t e s i n d i -cate the groups represent a s o i l moisture gradient. Group A consists of dry upland plant communities, B are mesic communities of intermediate and lower slopes, C are poorly-drained f l a t l a n d s and include ice-wedge polygons covered by low moss, l i c h e n and shrub heaths, and D are open fen and shallow marsh wetlands. (2) Examination and summary of a d d i t i o n a l f i e l d data according to the four vegetation groups indi c a t e d the groups generally could be characterized and separated by a range of s i t e parameters, i n c l u d i n g slope p o s i t i o n c l a s s e s , general cover features as measured i n 10 m x 10 m p l o t s , mineral s o i l texture c l a s s e s , the occurrence of organic s o i l s and ice-wedge polygons, c e r t a i n s o i l p h y s i c a l and chemical parameters and, of p a r t i c u l a r importance i n the present study, dominant l i c h e n taxa, and l i c h e n ground cover, biomass and standing crop estimates. Thus, the four groups are not ju s t vegetation-a l , they are ecosystemic u n i t s . (3) Lichens are of p a r t i c u l a r i n t e r e s t because they are a mainstay i n reindeer winter d i e t s . Lichens constituted one-quarter of the t o t a l plant - 148 -taxa, and l i c h e n cover varied up to 89.3% at ground-truth s i t e s . Standing crop estimates ranged from 194.4 to 6,377.6 kg.ha"! based on measurements made at 36 s i t e s where lich e n cover was greater than or equal to 20%. The majority of the l i c h e n standing crop at a l l s i t e s consisted of l i c h e n species known to be important as Rangifer forage. Top and bottom components of l i c h e n biomass, when plotted against one another, y i e l d a c u r v i l i n e a r r e -l a t i o n s h i p that r e f l e c t s , across vegetation groups A, B and C, s h i f t s i n dominant lic h e n species and general conditions for growth or decomposition of l i c h e n t h a l l i . Largest values for l i c h e n percent cover, biomass and standing crop were measured at group C s i t e s . Group D wetlands have n e g l i g i b l e amounts of l i c h e n . (4) Ice wedge polygons, both raised-centred and low-centred v a r i e t i e s , are common t e r r a i n features i n poorly-drained locations and are estimated to cover 23.4% of the study area, according to estimates "made on large-scale photographs. Among various management zones ( F i g . 10), percent of land cover by ice-wedge polygons ranges from 4.6% to 43.4%. Based on d e n s i t i e s of poly-gons as measured on the large-scale photos, one square km of polygon-covered t e r r a i n may have over 133 l i n e a r km of ice wedges. (5) T e r r a i n disturbance by vehicles accounted for low ground cover (0.5%) over the whole of the study area, but ranged upwards to 2.0% i n one management zone. Frequency was considerably higher, with disturbance noted i n 19.0% of the 1,469 photo-frames. Compared to previously published i n f o r -mation, the r e s u l t s here may ind i c a t e increased t e r r a i n disturbance i n the l a s t decade. The photos serve now as permanent records against which ocur-rences of future disturbances can be compared. - 149 -(6) Lichen Types ( I , I I , III) are the air-photo i n t e r p r e t e d equivalents of vegetation groups (A, B, C). They are assigned on the l a r g e - s c a l e photos according to i n t e r p r e t a t i o n s of vegetation cover, general slope p o s i t i o n , the occurrence of patterned ground, and a few a d d i t i o n a l parameters (Table I I I ) . (7) To determine i f Lichen Types could be r e a d i l y separated on the l a r g e - s c a l e CIR photos, microdensitometric measurements were made of l i c h e n patches. Quantitative analyses showed white and blue l i g h t measurements of dye layer d e n s i t i e s provided s i g n i f i c a n t (p = .05) separations among Lichen Types. Linear Discriminant Function (LDF) analysis of the data provided an 81.1% correct reassignment of 296 sets of density measures among the three Lichen Types. (8) Lichen Types are not mappable units at other than large scales (e.g., 1:1,400-1:3,400) even though within three broad categories they repre-sent: ( i ) the complete range of non-wetland types occurring i n the study area, and ( i i ) d i s t i n c t d i f f e r e n c e s i n a number of s i t e parameters, i n p a r t i c u l a r percent l i c h e n cover ranges, top to bottom l i c h e n biomass r a t i o s and standing crop ranges. The dilemma i s that some of the r e s u l t s are required at a regional l e v e l to be u s e f u l f o r general rangeland management but can only be r e l i a b l y obtained at large s c a l e s . Percent l i c h e n cover f o r example can only be measured e f f e c t i v e l y on l a r g e - s c a l e photos but i s necessary as a v a r i a b l e i n determining regional rangeland c a r r y i n g capacity. (9) Percent l i c h e n cover and l i c h e n standing crop are summarized accord-ing to the reindeer management zones. Percent l i c h e n cover ranged from 1.48% i n zone B to 14.24% i n zone D (Table XX). Standing crop was c a l c u l a t e d f o r both top and bottom components, using l i c h e n biomass measurements from ground-truth s t u d i e s . T o t a l standing crop estimates ranged from 39.6 kg.ha -! - 150 -i n zone B to 572.0 kg.ha - 1 i n zone D. On a more general l e v e l , the four more southerly zones have l i c h e n standing crops l a r g e r than the three n o r t h e r l y zones. (10) Using top estimates of l i c h e n standing crop, winter c a r r y i n g cap-a c i t y i s c a l c u l a t e d f o r the study area to be 20,373 reindeer. Over three-quarters of these animals can be accommodated i n two zones (D and E) which represent half of the study area's t o t a l land area. I t i s recommended that these ca r r y i n g capacity estimates be used to develop an improved r o t a t i o n a l grazing plan f o r the present population of about 16,000 animals. Carrying capacity estimates here should only be considered as p r o v i s i o n a l ; severe winter snow conditions, w i l d f i r e s and numerous other factors can a l t e r the a v a i l a b l e l i c h e n i n any given time period. (11) In the i n t r o d u c t i o n i t was pointed out that the study would deal with the "middle-ground" between plant ecology and remote sensing, and would aim to c l a r i f y the l i n k between the two. The outcome of the study i n d i c a t e s that by no means are the two sciences mutually exclusive and, i n f a c t , the l a r g e - s c a l e remote sensing work d i r e c t l y and l o g i c a l l y builds on the ground-t r u t h work. (12) The l a r g e - s c a l e photographs remain a v a i l a b l e f or numerous other po s s i b l e a p p l i c a t i o n s , i n c l u d i n g use with multistage schemes as a d d i t i o n a l remote sensing data becomes a v a i l a b l e . (13) The present i n v e s t i g a t i o n demonstrates that l a r g e - s c a l e 70 mm CIR photographs can play an important and i n t e g r a t i v e r o l e i n the study of a r c t i c Rangifer rangelands when coupled with ground-truth studies. Some primary advantages f o r using l a r g e - s c a l e remote sensing systems i n northern range studies can be l i s t e d : ( i ) the p o t e n t i a l e x i s t s f o r measurement of im-- 151 -p o r t a n t parameters (e.g., l i c h e n cover) that cannot be r e a d i l y i n t e r p r e t e d using s m a l l - s c a l e remote sensing data; ( i i ) w i t h i n c r e a s i n g costs of f i e l d l o g i s t i c s f o r ground-truth i n the no r t h , l a r g e - s c a l e remote sensing data-a c q u i s i t i o n provides an a l t e r n a t i v e that w i l l i n the f u t u r e only become more c o s t - e f f e c t i v e , and ( i i i ) a permanent record i s provided so that b a s e l i n e data can be obtained w h i l e p r o v i d i n g f u t u r e monitoring c a p a b i l i t i e s f o r many important range c h a r a c t e r i s t i c s (e.g., range trend, c o n d i t i o n , u t i l i z a t i o n e f f e c t s , m u l t i p l e - u s e e f f e c t s , e t c . ) . - 152 -LITERATURE CITED - 153 -Abrahamson, G. 1963. 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Mean percent cover class and, in parentheses, frequency are summarized for the four vegetation groups. Main dichotomies for the 420 plant taxa ( i . e . , cut levels I, 11 and 111 for the species ordinations) are indicated in the table by broken horizontal lines. Percent cover class: i » 0-1Z; 2 = 2-42; 3 - 5-92; 4 - 10-19%; 5 = 20-1003!. Frequency is occurrence of the taxa at sites expressed as a percentage of the to t a l number of sites in each vegetation group. S t r u c t u r a l 1 A B C 0 category Spec i es^ (n - 32) (n - 25) (n - 41) (n = 14) 2 RubuB ohamaemoruB L. 1 (16) 2 (72) 3 (83) 1 (39) *5 Sphagnum rubellum Wils. - 2 (24) 1 (17) 1 (7) 5 S. fuBOum (Schimp.) Klinggr. 1 (9) 2 (44) 2 (39) 1 (7) 6 Cetraria ialandiaa (L.) Ach. ssp. ialandica 1 (19) 1 (28) 2 (29) 1 (7) 1 Picea mariana ( M i l l . ) B.S.P. - 1 (4) -2 Melandrium taimyrense To lin. - 1 (4) - -2 Arabia divaricarpa A. Nies. - 1 (4) - -2 Hedysarwn alpinum L. var. americanum Michx. - 1 (4) - -1 Arat08taphyloa alpina (L.) Spreng. - 1 (4) - -2 Pedicularia lapponica L. - 1 (12) 1 (2) -*5 Plagiothecium denticulatum (Hedw.) B.S.G. - 1 (8) - -6 Rhizocarpon geographicum (L.) DC. - 1 (4) - _ 6 R. grande (Flbrke ex Flot.) Arn. - 1 (4) - -5 Surhynchium pulchellum (Hedw.) Jenn. - 1 (4) - -I Aratostaphyloa urva-urai (L.) Spreng. - 1 (4) - -6 Peltigera canina (L.) Willd. - 1 (4) - -6 Ochrolechia androgyna (Hoffm.) Arn. - 1 (4) - -*6 A s p i c i l i a nikrapenaie Darb. - 1 (4) - -6 Candelariella v i t e l l i n a (Ehrh.) Mull. Arg. - 1 (4) - -6 Caloplaca sp. - 1 (4) - -6 Pfiy8cia caesia (Hoffm.) Hampe - 1 (4) - -5 Myurella julacea (Schwaegr.) B.S.G. - 1 (4) - -5 Ditrichum flexicaule (Schwaegr.) Hampe - 1 (4) - -5 Drepanocladus lycopodioides var. brevifoliua (Lindb.) Monk. - 1 (4) - -2 Potentilla norvegica L. - 1 (4) - -2 S t e l l a r i a laeta Richards. - 1 (4) - -1 Ledum groenlandicwn Oeder - 1 (4) - -2 S t e l l a r i a calycantha (Ledeb.) Bong. - 1 (4) - -2 Pi'aba cinerea Adams - 1 (4) - -5 Pohlia cruda (Hedw.) Lindb. - 1 (8) - -5 Isopterygium pulchellum (Hedw.) Jaeg 4 Sauerb. - 1 (4) - -*4 Lophozia alpeatria (Schleich ex Web.) Evans - 1 (4) - -5 Encalypta rhaptocarpa Schwaegr. - 1 (4) - -*5 Bryum lisae var. cuspidatum (B.S.G.) Marg. 1 1 (4) - -5 Plagiomnium medium (B.S.G.) Kop. - 1 (4) - -5 Plagiothecium laetum B.S.G. - 1 (4) - -4 Mylia anomala (Hook.) S.F. Gray - 1 (4) - -*5 Leptodictyum riparium (Hedw.) Warnst. - 1 (4) - -*5 Drepanocladua c a p i l l i f o l i u s (Warnst.) Warnst. - 1 (4) - -1 Picea glauca (Moench) Voss - 1 (40) 1 (7) -1 Rosa acicularie L i n d l . 1 (3) 1 (28) 1 (2) -1 OxycoccuB microcarpuB Turcz. - 1 (20) 1 (2) -6 Cladonia gracilis (L.) Willd. ssp. turbinata - 1 (12) - -*6 C. rei Schaer. - 1 (12) - -3 ElymuB arenarius L. ssp. mollis (Trin.) lin 1 ten _ _ 1 (2) -3 Luzula parviflora (Ehrh.) Desv. - - 1 (2) -2 Polygonum bietorta L. ssp. plumoaum (Small) llutt. - - 1 (2) -S t r u c t u r a l 1 A B C U category Species^ (n = 32) (n = 25) (n - 41) (n - 14) 2 Chrysanthemum arcticum L. - - 1 (2) 6 Alectoria nigricans (Ach.) Nyl. 1 (3) 1 (4) 1 (27) -6 Cladonia fimbriata (L.) Fr. - 1 (4) 1 (7) -6 C. gracilis (L.) Willd. ssp. nigripes - - 1 (2) -6 Pachyospora verrucosa (Ach.) Mass. - - 1 (2) -6 Peltigera malacea (Ach.) Funck. - - 1 (2) -6 Psoroma hypnorum (Vahl.) S. Gray - 1 (2) -6 Stereocaulon glareosum (Sav.) Magn. - - 1 (5) -2 Wilhelmsia physodes (Fisch.) McNeill - - 1 (2) -*3 Carex saxatilis L. var. rhomalea Fern. - - 1 (2) -2 Rorippa islandica (Oeder) Borbas - - 1 (2) -2 Draba longipes Raup - - 1 (2) -2 Ranunculus nivalis L. - - 1 (2) -2 Mertensia maritime (L.) S.F. Gray - - 1 (2) -1 Salix polaris Wahlenb. ssp. peeudopolaris (Flod.) Hult. - - 1 (2) -2 Ranunculus cymbalaria Pursh (2) -3 Puccinellia phryganodes (Trin.) Scribn. 4 Merr. - - 1 (2) -2 Hippurie tetraphylla L. - - 1 (2) -2 Myriophyllum exalbescens Fern. - - 1 (2) -2 Potamogeton vaginatua Turcz. - - 1 (5) -*6 Agyrophora rigida (DuReitz) Llano - - 1 (2) -6 Xanthoria candelaria (L.) Th.Fr. - - 1 (2) -6 X. elegana (Link.) Th.Fr. - - 1 (2) -6 Cladonia thomaonii Ahti - - 1 (2) -*4 Cladopodiella fluitana (Nees) Joerg. - - 1 (2) -5 Sphagnum fimbriatum Wils. ex J. Hook. - - 1 (2) -5 Dicranum groenlandicum Brid. - - 1 (2) -5 Calliergon giganteum (Schimp.) Kinds. - - 1 (2) -2 Potentilla norvegica L. - - 1 (2) -4 Barbilophozia kunzeana (Hub.) Gams. - - 1 (2) -4 Calypogeia sp. - - 1 (5) -4 Cephalozia lunulifolia (Dum.) Dum. - - 1 (2) -4 Gymnocolea inflata Huds.) Dum. - - 1 (5) -4 Lophozia incisa (Schrad.) Dum. - - 1 (7) -4 Odontoschism macounii (Aust.) Underw. - - 1 (5) -4 Ptilidium ciliare (L.) Hampe - 1 (4) 1 (10) -*4 Scapania irrigua Schust. - - 1 (5) -4 Tritomaria quinquedentata var. turgida (Lindb.) Weim. - - 1 (2) -3 Carex livida Willd. var. grayana (Dew.) Fern. - 1 (4) 1 (2) -2 Corallorhiza trifida Chat. - I (4) 1 (2) -2 Melandrium offine J. Vahl - 1 (4) I (2) -2 Petasites arcticua P o r s i l d - 1 (8) 1 (10) -5 Sphagnum anguatifolium (C. .lens, ex Russ.) C. Jens. - 1 (8) 1 (2) -6 Cladina stellaris (Opiz) Brodo 1 (47) 3 (92) 4 (90) 1 (7) 6 Cetraria nivalis (L.) Ach. I (28) 1 (24) 2 (59) -6 Cladina arbuecula (Wallr.) Hale & W. Culb. 1 (6) 1 (40) 2 (32) -6 C. rangiferina (L.) Harm. 1 (44) 3 (88) 4 (93) 1 (7) 6 Pertusaria panyrga (Ach.) Mass. - 1 (4) 1 (7) -6 Sphaerophorua globosus (Huds.) Vain - 1 (8) 1 (2) -6 Stereocaulon paschale (L.) Hoffm. - 1 (4) 1 (7) -5 Polytrichum juniperinum Hedw. - 1 (12) i (2) -2 Boschniakia rossiaa (Cham & Schlecht.) Fedtsch. - 1 (4) I (2) -category Spec iea 2 Lycopodium annotinum L. 3 Calamagroatia inexpanaa A. Gray 3 Eriophorum vaginatum L. sep. vaginatum 2 Tofieldia puailla Michx.) Pers. 1 Ledum deoumbena ( A i t . ) Lodd. *5 Dioranum brevifolium (Lindb.) Lindb. 5 0. epadiaeum Zett. *5 llynum plicatulum (Lindb. ) Jaeg. & Sauerb. 6 Alectoria oahroleuca (Hoffra.) Mass. 6 Bryoria nitidula 6 Cladina mitie (Sandst.) Hale & W. Culb. 6 Cladonia squamosa (Scop.) Hoffm. 6 Coeloncaulon divergens (Ach.) R.H. Howe 6 Peltigera scabrosa Th. Fr. 4 Anastrophylum minutum (Schreb.) Schust. 4 Scapania paludicola Loeske i K. H i i l l . 3 Calamagrostia neglecta (Ehrh.) Gaertn., Mey & Schreb. 3 Carex rupeatria A l l . 3 Luzula confusa Lindebl. 1 Spiraea beauverdiana Schneid. 5 Hylocomium aplendena (Hedw.) B.S.G. 6 Cladonia deformia (L.) Hoffm. 6 C. gracilis (L.) Willd. ssp. gracilis 6 C. macrophylla (Schaer.) Stenham. 6 C. sp. 6 Dactylina aratiaa (Hook.) Nyl. 6 Ochrolechia frigida (Sw.) Lynge *6 Peltigera praetextata (Flbrke ex Somm.) Vain 6 P. epuria (Ach.) DC. 6 Stereocaulon alpinum Laur. 4 Barbilophozia binsteadii (Kaal.) Loeske 2 Rumex arcticus Trautr. 5 Meesia uliginosa Hedw. *5 Oicranum leioneuron Kindh. 5 Sphagnum n&noreum Scop. *5 S. ruasowii Wamst. 2 Pedicularis labradorica Wirsing 5 Sphagnum girgensohnii Russ. 5 S. lenenae H. Lindb. ex Pohle 6 Ichmadophilia ericetorum (L.) Zahlbr. 4 Blepharostoma trichophylla (L.) Dum. 4 Cephalozia plenicepa (Aust.) Lindb. 4 lophozia wenzellii (Nees) Steph. 3 Carex vaginata Tausch 3 Arctagrostie latifolia (R. Br.) Griseb. ssp. latifolia 3 Carex conaimilis Holm 3 Luzula nivalis (Laest.) Beurl. 1 Betula glandulosa Michx. 5 Ceratodon purpureua (Hedw.) Brid. 6 Cladonia acuminata (Ach.) Norrl. A (n - 32) B (n = 25) C I) (n = 41) (n = 14) (3) 1 (8) -(3) 1 (12) 1 (5) -(31) 3 (56) 3 (73) I (7) (13) 1 (28) 1 (21) -(75) 4 (96) 5 (100) 1 (29) (6) 1 (28) 1 (15) -(22) 1 (28) 2 (29) -- 1 (4) 1 (2) -(16) 1 (12) 2 (44) -(44) 1 (24) 2 (76) -(9) 1 (32) 1 (37) -(9) 1 (16) 1 (15) -(9) 1 (12) 1 (24) -(3) 1 (4) 1 (7) _ (16) 1 (60) 1 (37) -(3) 1 (4) 1 (7) -(13) 1 (12) 1 (12) -(6) - 1 (7) -(13) - 1 (12) -(3) 1 (4) 1 (5) _ (3) 1 (4) 1 (5) -(3) - 1 (5) -(13) - 1 (12) -(13) 1 (8) 1 (10) -(9) 1 (16) 1 (12) -(28) - 1 (44) (6) 1 (4) 1 (12) -(3) 1 (8) 1 (2) -(3) - 1 (5) -(3) - 1 (5) -(6) 1 (4) 1 (12) -(3) 1 (12) 1 (2) 1 (7) - 1 (4) - 1 (7) (3) 1 (12) 1 (7) 1 (7) - 1 (8) 1 (5) 1 (14) (3) 1 (20) 1 (20) 1 (21) (28) 1 (36) 1 (39) 1 (21) (3) 1 (4) 1 (2) 1 (7) - 1 (8) 1 (12) 1 (21) - - 1 (7) 1 (7) (9) 1 (20) 1 (20) 1 (21) - 1 (4) 1 (2) 1 (7) (3) 1 (4) 1 (7) 1 (7) (6) 1 (20) 1 (10) 1 (7) (75) 1 (64) 1 (54) 1 (29) (19) 1 (32) 1 (29) 1 (14) (22) 1 (4) 1 (24) 1 (14) (88) 4 (92) 3 (93) 2 (64) (13) 1 (4) 2 (7) 1 (7) (13) 1 (12) 1 (15) 1 (7) St ructural* category Spec Les^ A (n = 32) B (n = 25) C (n = 41) D (n = 14) 6 C. cornuta (L.) Hoffm. 1 (28) I (44) 1 (34) 1 (14) 6 C pyxidata (L.) Hoffm. 1 (16) - 1 (17) 1 (7) 1 /Mnu8 criapa ( A i t . ) Pursh 2 (9) 3 (68) 1 (44) 1 (21) 1 Bmpetrum nigrum L. ssp. hermaphroditum (Lge.) Bocher 3 (78) 3 (84) 2 (88) 1 (36) 1 Vacctnium vitis—idaea L. var. minus Lodd. 3 (78) 2 (92) 3 (100) 1 (29) 2 Petasites aagittatua (Banks) A. Gray 1 (41) 2 (56) 1 (12) 1 (7) 5 Aulacomnium tuvgidum (Wahlenb.) Schwaegr. 2 (44) 2 (56) 2 (54) 1 (14) 5 Dicranum elongatum Schleich ex Schwaegr. 2 (78) 2 (80) 2 (88) 1 (21) 5 Polytrichum atrictum Brid. 1 (66) 2 (84) 2 (78) 1 (21) 5 Tomenthypnum nitena (Hedw.) Loeske 2 (44) 3 (60) 1 (7) 1 (7) 6 Cladonia cenotea (Ach.) Schaer. 1 (19) 1 (36) 1 (41) 1 (7) 6 C. coccifera (L.) Willd. 1 (44) 1 (48) 1 (46) 1 (7) 6 C. pleuvota (Flbrke) Schaer. 1 (31) 1 (28) 1 (37) 1 (7) *6 C. Bulphurina (Michx.) Fries 1 (59) 1 (40) 1 (54) 1 (7) 6 Peltigera aphthoaa (L.) Willd. 2 (53) 2 (68) 1 (44) 1 (7) 6 Thamnolia subuliformia (Ehrh.) W. Culb. 1 (38) 1 (64) 1 (41) 1 (7) 1 Arotoataphyloe rubra (Rehd. & Wils.) Fern. 3 (81) 3 (80) 3 (83) 1 (7) 5 Pohlia nutans (Hedw.) Lindb. 2 (66) 2 (68) 3 (68) 1 (7) 6 Cladonia amaurocraea (Flbrke) Schaer. 1 (59) 1 (72) 2 (66) 1 (7) 6 Ochrolechia gyalectina (Nyl.) Zahlbr. 2 (66) 2 (64) 2 (68) 3 Hierochloe alpina (Sw.) R. & S. 1 (31) 1 (12) 1 (27) 1 (7) 6 Cetraria cucullata ( B e l l ) Ach. 4 (91) 3 (80) 3 (90) 1 (7) 6 C. ericetorum Opiz 3 (81) 2 (56) 2 (83) 1 (7) 6 Cladonia chlorophaea (Flbrke ex Somm.) Spreng. 2 (63) 1 (44) 1 (54) 3 Carex lugens Holm 3 (50) 3 (72) 2 (44) I (14) 6 Cladonia pocillum (Ach.) 0. Rich. 1 (63) 1 (44) 1 (37) 1 (7) 2 Cardamine digitata Richards 1 (16) 1 (16) 1 (10) 2 Epilobium anguatifolium L. 1 (3) 1 (4) 1 (2) -1 Rhododendron lapponicum (I..) Wahlenb. 1 (9) 1 (20) 1 (2) -1 Vaccinium uliginosum L. 3 (44) 2 (68) 2 (41) 1 (7) 2 Androsace aeptentrionalis L. 1 (3) 1 (8) -2 Sauaaurea anguatifolia (Willd.) DC. 1 (28) 1 (48) 1 (12) -6 Cetraria laevigata Rass. 1 (25) 1 (40) 1 (12) -6 C. pinastri (Scop.) S. Gray I (19) 1 (32) 1 (5) -6 C. aepinicola (Ehrh.) Ach. I (13) 1 (28) 1 (2) -6 Cladonia cyanipea (Somm.) Nyl. 1 (13) 1 (12) 1 (5) -6 C. phyllophora Hoffm. 1 (3) 1 (8) _ -*6 C. acabriuacula (Del. ex Duby) Nyl. 1 (13) 1 (32) - -6 Hypogymnia aueterodee (Nyl.) Ras. 1 (6) 1 (8) 1 (2) -6 Parmelia eeptentrionali8 (Lynge) Ahti 1 (19) 1 (32) 1 (2) _ 6 Parmeliopaia ambigua (Wulf.) Nyl. 1 (16) 1 (28) 1 (5) _ 6 Solorina biepora Nyl. 1 (3) 1 (4) - _ 3 Festuca brochyphylla Schultes 2 (16) 1 (8) 1 (20) _ 2 Stellaria longipes Coldie 1 (44) 1 (24) 1 (27) -*6 Ochrolechia inaequatula (Nyl.) Zahlbr. 1 (6) 1 (8) 1 (2) -3 Poa prateneia L. 2 (28) 1 (28) 1 (17) -3 P. arctica R.Br. ssp. arctica 2 (9) 1 (4) 1 (10) -3 Kobreaia myoauroidea ( V i l l . ) F i o r i & Paol. 1 (3) - 1 (2) 3 Carex maritima Gunn. I (3) - 1 (2) -S t r u c t u r a l 1 category Species 2 3 C. gynocratea Wormskj 3 C. scirpoidea Midi*. 3 C. atrofusca Schk. 2 Habenaria obtusata (Pursh) Richards. 1 Salix niphoelada Rydb. 1 S. alaxensis (Anderss.) Cov. 2 Cevaatium beeringianum Cham. & Schlecht. 2 Minuartia rubella (Wahlenb.) lliern. 2 Chrysosplenium tentandrum (Lund) Fries 1 Potentilla frutioosa L. 2 Lupinus arcticus Wats. 2 Astragalus alpinus L. 2 ConioBelenum cnidiifolium (Turcz. ) P o r s i l d 1 Pyrola secunda L. var. secunda 1 P. grandiflora Radius 1 Cassiope tetragona (L.) D. Don ssp. tetragona 2 Androsace chamaejasme Host. var. arctica Knuth 2 Polemonium acutiflorum Willd. 2 Pedicularis capitata Adams 2 Petaaites frigidus (L.) Fries 2 Arnica alpina (I..) O l i n 2 Senecio lugens Richards 2 S. atropurpureus (Ledeb.) Fedtsch. 2 Taraxacum dumetorum Greene 5 Thuidium abietinum (Hedw.) B.S.G. 5 Rhytidium rugosum (Hedw.) Kindb. 5 Diatichium capillaceium (Hedw.) B.S.G. 5 Hypnum hamulosum B.S.G. 6 Bacidia sphaeroides (Dicks.) Zahlbr. 6 Cladonia bacillaria (Ach.) Nyl. *6 C. carneola (Fr.) Fr. 6 C. lepidota Nyl. 6 Hypogymnia physodea (L.) W. Wats. 6 Lecanora epibryon (Ach.) Ach. 6 '•opadium pezizoideum (Ach). Kb'rb. 6 Parmelia eulcata Tayl. 6 Peltigera leucophlebia (Nyl.) Gyeln . *6 P. neckeri Muell. Arg. 6 Phyaconia muBcigena (Ach.) Poelt. 6 Rinodina turfacea (Wahlenb.) Korb 6 Solorina spongiosa (Sm.) Anzi. 1 Ribes triste P a l l . 4 Scapania sp. 1 Salix glauca L. 2 Stellaria eduardsii R. Br. 3 Bromua pumpellianus Scribn. var. pumpellianus 3 Trisetum spicatum (L.) Richt. 3 Alopecurus alpinus J.E. Smith 3 Carex miaandra R. Br. 1 Salix reticulata L. 1 S. phlebophylla Anderss. 1 S. chami830nis Anderss. A B C D (n = 32) (n - 25) (n = 41) (n = 14) I (22) 1 (8) 1 (12) _ 1 (19) 1 (4) 1 (10) -I (6) - 1 (2) 1 (3) 1 (4) - -1 (3) - - -4 (59) 2 (44) 1 (17) 1 (14) 1 (6) 1 (4) - -1 (3) 1 (4) - -1 (3) 1 (4) - -1 (3) 1 (4) - -2 (69) 1 (28) 1 (10) -1 (19) 1 (8) 1 (5) -1 (6) 1 (4) - -1 (59) 1 (48) 1 (10) -2 (78) 1 (68) 1 (7) -2 (25) 1 (16) 2 (5) -1 (3) 1 (4) - -1 (3) - 1 (2) -1 (53) 1 (24) 1 (5) -1 (3) - 1 (2) 1 (7) 1 (13) 1 (8) 1 (5) -1 (53) 1 (40) 1 (20) -1 (6) 1 (4) - -1 (3) 1 (4) - -2 (53) 2 (28) 1 (22) -2 (63) 2 (48) 1 (32) -1 (34) 1 (16) 1 (17) -1 (6) - 1 (2) -1 (6) 1 (8) - -1 (3) - - -1 (13) 1 (4) 1 (5) - ' 1 (3) 1 (4) - -1 (34) 1 (40) 1 (7) -1 (16) - 1 (7) -1 (3) - 1 (5) -1 (22) 1 (12) 1 (5) -1 (34) 1 (12) 1 (7) -1 (6) - 1 (5) -1 (25) 1 (4) 1 (10) -1 (22) I (4) 1 (10) -1 (3) 1 (4) - -1 (6) 1 (4) - -1 (3) 1 (4) - -3 (66) 2 (36) 1 (20) 2 (29) I (25) 1 (4) 1 (5) 1 (7) 1 (3) - - -1 (3) - - -1 (9) - 1 (2) 1 (3) - - -2 (34) - - -1 (6) - - -1 (6) - - -Structural* category Species 2 Anemone parviflora Michx. 2 RanunouluB pedatifiduB Sm. var. leiocarpua (Trautv.) Fern. 2 Draba alpina L. 2 D. nivalie L i l j e b l . 2 D. glabella Pursh 2 Veaaurainia aophioidea (Fisch.) O.E. Schulz 2 Soxifraga folioloaa R. Br. 2 5. hiroulua L. var propinqua (R.Br.) Siram. 2 S. triouapidata Rottb. 2 Parnaasia kotzebuei Cham. & Schlect. 2 Potentilla nivea L. 1 Vryaa integrifolia M . Vahl 2 0xytropi8 deflexa ( P a l l . ) DC. var. folioloaa (Hook.) Barneby 2 0. maydelliana Trautv. 5 Ditrichum flexioaule (Schwaegr.) Hampe *2 Oxytropia varians (Rydb.) K. Schum. 2 Armeria maritima ( M i l l . ) W i l l d . 2 Caatilleja elegans (Ostenf.) Malte 1 Linnaea borealis I., var. amerioana (Forbes) Rehd. 2 Achillea lanuloaa Nutt. 2 Artemeaia tileaii Ledeb. *6 Bacidia epixanthoides (Nyl.) Lett. 6 Caloplaca atillicidiorum (Vahl.) Lynge 6 Cladonia cryptochlorophaea As ah. *6 Gyalecta foveolaris (Ach.) Schaer. 6 Ochrolechia upaaliensia (L.) Mass. 6 Peltigera rufescens (Weis.) Humb. *6 Rinodina roscida (Somm.) Arn. 6 Solorina saccata (L.) Ach. 6 Toninia lobulata (Sonn.) Lynge 2 Astragalus eucosmus Robins. 2 Potentilla pulchella R. Br. 2 Arenana capillaris Poir var. nardifolia (Ledeb.) Regel 2 Rhodiola integrifolia Raf. 2 Gentiana propinqua Richards. 2 Matricaria ambigua (Ledeb.) K r y l . 6 Buellia papillata (Somm.) Tuck. *6 Collema bachmanianum (Fink) Degel. var baokmanianum 6 C. sp. 6 Leptogium tenuissimum (Dicks.) Fr. *6 Rinodina septentrionalis Walme 6 Buellia zahlbruckneri J. Stein. 6 Lecanora coilocarpa (Ach.) Nyl. 6 Phyacia aipolia (Ehrh.) Hampe 5 Brachythecium turgidum (C.J. Hartm.) Kindb. 1 Salix farriae B a l l 4 Lophozia heterocolpoe (Thed.) M.A. Howe 2 Taraxacum lacerum Greene 5 Deamatodon heimii var. arctica (Lindb.) Crum 5 D. cernuus (Hub.) B.S.G. 5 Tortella fragilia (Drumm.) Limpr. 3 Carex holoetoma Drej. A B C D (n = 32) (n = 25) (n = 41) (n = 14) (9) (9) (3) (3) (6) (6) (3) (6) (9) - 1 (2) (13) - - -(6) -(72) 1 (32) 1 (20) (6) -(22) - 1 (2) (3) - 1 (7) -(3) - 1 (2) (6) -(22) 1 (4) (3) -(3) - - " I (3) -(3) - - - S (6) - - - U i (3) -(3) ' (16) 1 (4) (16) - 1 (2) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) (3) 1 (4) (3) (3) (3) (3) St ructuraI 1 category Species 2 A (n = 32) (n = 25) C (n - 41) (n = 14) 2 2 2 2 1 2 2 3 3 2 2 2 2 6 5 5 *6 2 5 5 3 5 5 *5 *5 4 1 2 2 2 5 1 1 5 3 2 2 2 2 2 3 5 2 3 1 2 5 3 1 Chrysanthemum integrifolium Richards I (3) Erigeron humilis Crah. 1 (3) Antennaria angustata Greene 1 (6) Stellaria subvestita Greene 1 (6) Dryas arenulata J«z. 1 (6) Potentilla rubricaulis Lehra. 1 (3) Achillea nigresaens (E. Mey.) Rydb. 1 (3) Kobresia hyperborea P o r s i l d 1 (3) Agropyron sericeum llitchc. 1 ( 3 ) Artemesia frigida Willd. 1 (3) Solidago multiradiata A i t . 1 (3). Honckenya peploides (L.) Ehrh. var. diffusa (Hornera.) Mattf. I (3) Draba corymbosa R. Br. 1 (3) Lecanora eymmicta (Acli.) Ach. 1 (3) Leptobryum pyriforme (Hedw.) Wils. I (3) Brachythecium salebrosum (Web. & Mohr) B.S.G. 1 (3) Cladonia coniocraea (Flbrke) Spreng. 1 (3) Polygonum alaskanum (Small) Wight 1 (3) Oncophorus uahlenbergii Brid. 1 (3) Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. 1 (6) Carex glareosa Wahlenb. var. amphigena Fern. 1 (3) Timmia austriaca Hedw. 1 (3) Brachythecium sp. 1 (3) fl. velutinum (Hedw.) B.S.G. 1 (3) Amblystegium serpens (Hedw.) B.S.G. 1 (3) Preissia quadrata (Scop.) Nees Salix arbusculoides Anderss. Equisetum scirpoides Michx. Pedicularis lanata Cham & Schlecht. P. arctica R. Br. Drepanocladus vernicosus (Lindb. ex C. Salix arctica P a l l . S. arctophila Cockerel 1 Aulacomnium palustre (Hedw.) Schwaegr. Poa alpigena (Fr.) Lindm. Pulsatilla ludoviciana (Nutt.) Heller Anemone richardsonii Hook. Ranunculus lapponicus L. Oxytropis glutinosa P o r s i l d Festuca rubra L. Poa glauaa M. Vahl. Drepanocladus sp. Hartm.) Warnst. Equisetum arvense L. Carex biaolor A l l . Salix pulchra Cham. Rubus acaulis Michx. Drepanocladus uncinatuts (Hedw.) Warnst. Carex capillaris L. Salix lanata L. ssp. richardsonii (Hook.) Skvortsov 1 (6) - - -3 (44) I (12) 1 (12) 1 (14) 2 (25) ! (28) 1 (20) 1 (14) 1 (31) 1 (44) 1 (17) 1 (14) 1 (31) 1 (28) 1 (17) 1 (14) 1 (22) 1 (12) 1 (2) 1 (7) 1 (19) 1 (32) 1 (27) 1 (36) 2 (22) 1 (28) 1 (17) 1 (14) 2 (66) (80) 1 (44) 2 (50) 1 (9) 1 (4) 1 (5) 1 (7) 1 (3) 1 (4) - 1 (7) 1 (6) - - 1 (7) 1 (6) 1 (4) 1 (5) 1 (7) 1 (6) - - 1 (7) I (6) - - 1 (7) 1 (6) - - 1 (7) 1 (3) - - 1 (7) 1 (13) 1 (16) 1 (7) 1 (36) 1 (3) - 1 (2) 1 (7) 3 (38) 2 (32) 1 (20) 1 (36) 1 (3) 1 (4) - 1 (7) 1 (16) 1 (8) - 1 (14) I (9) 1 (12) 1 (2) 1 (21) 1 (6) 1 (8) 1 (2) 1 (14) Structural' category 3 2 5 5 3 2 2 3 5 4 3 3 5 *5 3 2 *5 6 2 5 3 2 5 2 3 3 3 3 3 3 3 1 2 2 2 2 5 5 5 5 *5 2 3 2 *1 5 5 Species^ Deachampaia oaespitosa (L.) Beauv. Pinguicula villoaa L. Drepanooladua badiua (C.J. Hartm.) Roth Paludella aquarrosa (Hedw.) Brid. luzula uahlenbergii Rupr. Polygonum viviparum L. Pediaularia audetioa W i l l d . Calamagroatia oanadenaia (Michx.) Beauv. flryum paeudotriquetrum (Hedw.) Gaerto., Meyer & Scherb. Marchantia jiolymorpha L. Dupontia fisheri R. Br. ssp. p8iloaantha (Rupr.) Ilulten Carex membranacea Hook. Andromeda polifolia L. Bryum sp. Calliergon megalophyllum Mik. Carex rariflora (Wahlenb.) Sm. Senecio oongeatus (R. Br.) DC. Drepanooladua aduncua (Hedw.) Warnst. Nephroma arcticum (L.) Torss. Galium trifidum L. Sphagnum obtusum Warnst. Carex rotundata Wahlenb. Ranunculus gmelenii DC. Polytrichum commune Hedw. Equisetum variegatum Schleich. Arctophila fulva (Trin.) Rupr. Hierochloe pauciflora R. Br. Eriophorum ruaseolum Fr. var. albidum Nyl. E. brachyantherum Trautv. E. angustifolium Honck. Carex chordorrhiza L. C aquatilia Wahlenb. Salix fusaeacena Anderss. Ranunculus hyperboreus Rottb. Potentilla paluatria (L.) Scop. Epilobium palustre L. Rippuria vulgaris L. Calliergon richardsonii (Mitt.) Kindb. ex Warnst. Plagiomnium ellipticum (Brid.) Kop. Catosoopium nigritum (Hedw.) Brid. Cinclidium subrotundum Lindb. Calliergon sarmentoaum (Wahlenb.) Kindb. Cochlearia officionalie L. asp. arctica (Schlecht.) Hult. Carex physocarpa P r e s l . Montia lamproaperma Cham. Salix planifolia Pursh ssp. planifolia Hypnum pratenae Koch ex Brid. Rhizomnium gracile Kop. (n = 32) (n = 25) (n = 41) (n = 14) 1 (6) 1 (8) - 1 (14) - 1 (4) - 1 (7) 1 (6) 1 (4) 1 (2) 1 (7) - 1 (4) - 1 (7) 1 (3) - - 1 (7) 1 (13) - 1 (2) 1 (14) 1 (13) - 1 (5) 1 (14) 1 (3) - - - 1 (14) 1 (13) 1 (8) 1 (5) 2 (29) 1 (6) - - 1 (14) 1 (6) - ' 1 (2) 1 (21) 2 (16) 1 (4) I (5) 3 (21) - 1 (20) 1 (20) 1 (79) - - 1 (2) 1 (7) - - 1 (5) 1 (7) 1 (3) 1 (8) 1 (47) 4 (64) - - 1 (2) 1 (7) - - 1 (5) 1 (7) - 1 (2) 1 (7) - - - 1 (7) - - - 2 (14) - 1 (8) 1 (7) 3 (36) - - 1 (2) 1 (21) - - 1 (5) 2 (21) - - 1 (7) - - 2 (21) - - - 1 (7) 1 (3) - 1 (7) 3 (64) - - 1 (2) 1 (29) 1 (3) 1 (4) 1 (17) 3 (93) - - - 3 (50) 2 (13) 1 (8) 1 (20) 4 (93) 1 (3) - - 2 (7) - - - 1 (7) - 1 (4) 1 (2) 2 (43) - - 1 (2) 1 (43) - - - 1 (14) - - - 2 (7) - - 1 (14) - - - 1 (7) - - - 1 (7) - - - 1 (7) - - - 1 (7) - - - 2 (14) - - - 1 (7) - - - 1 (7) - - - 1 (7) - 1 (4) - 1 (7) -~1 St ru c t u r a l ' A B C D category Species^ (n = 32) (n - 25) (n = 41) (n = 14) 5 Cinalidium stygium Sw. - _ _ 1 (7) 5 Calliergon atramineum (Brid.) Kindb. - - - 2 (29) 5 Campylium atellatwn (Hedu.) C. Jens. - 1 (4) - 2 (29) 5 Cinalidium arotioum (B.S.G.) Schimp. 1 (6) 1 (4) - 2 (57) 5 Drepanooladua exannulatus (B.S.G.) Warnst. 1 (6) - 1 (13) 3 (36) 5 D. fluitana (Hedw.) Warnst. - - - 2 (29) 5 Sphagnum aongstroemii C. Hartin. - - - 1 (21) 5 Scorpidium scorpioidea (Hedw.) Limpr. 1 (6) - 1 (5) 4 (50) 2 Cardomine pratanaia L. 1 (3) - - 1 (29) 2 Caltha paluatria L. var. paluatria - - - 1 (21) 4 Anuura pinguia (L.) Dun. - - - 1 (14) 4 Calypogeia muelleriana (Schiffn.) K. Mi i l l . - _ - 1 (7) 4 Cephalozia ap. - - - 1 (7) *4 Barbilophozia attenuata (Mart.) Loeske - - - I (7) 4 Lophozia rutheana (Limpr.) M.A. Howe - - - I (7) 4 I. ventriooaa (Ricks.) Dum. - - _ 1 (7) 5 Sphagnum oompaotum OC. ex Lam. & DC - 1 (4) 1 (5) 2 (21) 5 S. aquarroaum Crome - 1 (8) 1 (17) 3 (50) 5 S. teres (Scliiinp.) Angstr. ex C. Hartm. 1 (3) 1 (4) 1 (5) (14) 1 Chamaedaphne oalyaulata (L.) Moench - 1 (12) 1 (2) I (29) 5 S. majua (Russ.) C. Jens. - 1 (8) - 1 (7) 'structural category: 1. trees and woody shrubs; 2. broad-leaved herbs; 3. graminoids ( i . e . grass and g r a s s - l i k e plants of the families Gramineae, Cyperaoeae and Junoaoeae) ; 4. hepatics; 5. mosses; 6. lichens. 2 nomenclature follows P o r s i l d 4 Cody (1980) for vascular plants, S t o t l e r & Crandal 1-Stot l e r (1977) for hepatics, Ireland et al. (1980) for mosses, and Hale 4 Culberson (1970) for lichens, excepting Cladina atellaris (Opiz) Brodo (Brodo 1976) and Cladonia thomaonii Ahti (Ahti 1978). *new record or range extension for taxa v e r i f i e d by G.W. Argus (Salix), J.M. G i l l e t t (other vascular taxa), L.M. Ley (hepatics), R.R. Ireland (mosses), I.M. Brodo and P.Y. Wong (li c h e n s ) . Publications Sims, R.A. and P.A. Murtha. (In press.) Reindeer at Mackenzie: a selected annotated bibliography. Dep. Indian A f f a i r s & North. Develop., Ottawa, Ont. Environ. Stud. Rep. 115 pp. Cowell, D.W., A.N. Boissonneau, J.K. Jeglum, G.M. Wickware and R.A. Sims. (In press.) Hudson Bay Lowland peatland inventory. _In Proc. Peatland Inventory Worksh., 9-10 Mar., 1982, Ottawa, Ont. Spons. by NRC's Peat Forum and the Can. Wetland Working Group, CCELC. Sims, R.A., D.W. Cowell and G.M. Wickware. 1982. C l a s s i f i c a t i o n of fens near southern James Bay, Ontario using vegetational physiognomy. Can. J . Bot. 60:2608-2623. Sims, R.A., D.W. Cowell and G.M. Wickware. 1982. The use of vegetational physiognomy i n c l a s s i f y i n g treed peatlands near southern James Bay, Ontario. Nat, can. (Rev. E c o l . Syst.) 109:611-619. Cowell D.W., R.A. Sims and G.M. Wickware. 1982. Frozen beach ridge s o i l s i n the Hudson Bay Lowland, Ontario. Can. J . S o i l S c i . 62:421-425. Sims, R.A. and J.M. Stewart. 1981. A e r i a l biomass d i s t r i b u t i o n i n an undisturbed and disturbed subarctic bog. Can. J . Bot. 59:782-786. Sims, R.A. and P.A. Murtha. 1981. A multistage remote sensing program to assess reindeer rangeland. Proc. Ecolog. Data Process. & Interp. Worksh., Terr. Stud. Br., Min. of Environ., V i c t o r i a , B.C.:345-352. Wickware, G.M., D.W. Cowell and R.A. Sims. 1981. Peat resources of the Hudson Bay Lowland Coastal Zone. Proc. VI Internat. Peat Congr., Duluth, Minn., Aug. 18-21, 1980:138-153. Glooschenko, W.A., R. Sims, M. Gregory and T. Mayer. 1981. Chapt. 18: Use of bog vegetation as a monitor of atmospheric input of metals. In Atmospheric Input of Pollutants to Natural Waters. Ann Arbor Science Publ., Amer. Chem. Soc.:389-399. Wickware, G.M., R.A. Sims, K. Ross, and D.W. Cowell. 1981. The a p p l i c a t i o n of remote sensing and techniques for an e c o l o g i c a l land survey of the Snow Goose colony at Cape Henrietta Maria, Hudson Bay. Proc. 6th Can. Symp. Rem. Sens.:387-395. 2 Publications Page 2 Wickware, G.M., D. Cowell, K. Ross and R.A. Sims. 1980. U t i l i z a t i o n of e c o l o g i c a l land c l a s s i f i c a t i o n data for the study and management of waterfowl resource i n the Hudson Bay Lowland. In Land/Wildlife Integration, Lands D i r e c t . , Ottawa. E c o l . Land C l a s s i f . Rep. No. 11:45-50. Barclay-Estrup, P. and R.A. Sims. 1979. Epiphytes on Ulmus americana L. near Thunder Bay, Ontario. Can. Field-Nat. 93(2):139-143. Cowell, D.W., G.M. Wickware and R.A. Sims. 1979. E c o l o g i c a l land c l a s s i f i c a t i o n of the Hudson Bay Lowland coastal zone, Ontario. Proc. Second Can. Comm. E c o l . Land Class., V i c t o r i a , A p r i l 4-7, 1978. E c o l . Land C l a s s i f . Rep. No. 7:165-175. Sims, R.A., J.L. R i l e y and J.K. Jeglum. 1979. Vegetation, f l o r a and vegetational ecology of the Hudson Bay Lowland - a l i t e r a t u r e review and annotated bibliography. Can. For. Serv., Sault Ste. Marie, Ont., Inf. Rep. O-X-297. 177 p. Sims, R.A. 1978. The use of 'muskeg caps' to deter f r o s t penetration under transmission l i n e tower bases. Proc. 17th Muskeg Res. Conf., Nat. Res. Council Tech. Memo. No. 122:116-131. Haworth, S.E., D.W. Cowell and R.A. Sims. 1978. Bibliography of published and unpublished l i t e r a t u r e on the Hudson Bay Lowland. Can. For. Serv., Sault Ste. Marie, Ont., Inf. Rep. O-X-273. 270 p. 

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