AN INTENSIVE STUDY OF THE FISH FAUNA OF THE MUSKEG RIVER WATERSHED OF NORTHEASTERN ALBERTA

by

W.A. BOND and K. MACHNIAK Environment Canada Freshwater Institute Winnipeg, Manitoba

for

ALBERTA OIL SANDS ENVIRONMENTAL RESEARCH PROGRAM Project AF 4.5.1

July 1979 ix

TABLE OF CONTENTS

Page

DECLARATION •••••••..•.••••.•.....•••••..••..•.•.•.•••....••• ii

LETT ER 0 F TRA NS MITT AL ••.•.••..••.•••••••.••.•••...•••.•...•• iii

DES CRIP T I VE SUMMA RY .•••••.••..••.•••.•.•..•••.••..•••..•••.• i v

LIST OF TABLES .••.•••.••.•••...... •..•.•••.••...•...•...••.• xiii

LIST OF FIGURES .•••.•••••.••••...•••.•••.••••••••••••••.•••• xv i i

ABSTRACT ••••••••••••.•••••••.•.•.•.••..•••••.••.••..•••••••. x i x

AC KNOWLEDGEMENTS •••••••.••...••••.••••..•.••••..•..••..•.••• xx i

1 . I NTRODUCT I ON •.•.•..••••••..••..•.•••.••••••••.••..

2. R~SUM€ OF CURRENT STATE OF KNOWLEDGE •• '! •••••••••• 4

3. DESCRIPTION OF STUDY AREA .•.••••.•••••.•••••..•.•• 6

4. MATERIALS AND METHODS •.•••.•••••••••••••.•.• 0 ••••• 17 4. 1 Counting Fence Construction •.••••.••••••••.•..••.• 17 4.2 Counting Fence Operation "0' •••••••••••••••••••••• 17 4.2. 1 Samp 1 i ng Schedu 1e ••.•...•••.•.•••..••••.•..••.. 19 4.2.2 Trap Checks ...... ,...... 19 4.2.3 Tagging ••••..••.•••.•...•.••..••••••..••.•••••. 19 4.2.4 Dead Samples ••.••••.••••.•••••••••••..••.•.•... 21 4 .. 3 Other Fish Collection Techniques ••••.•.•••••••••. 22 4.3. 1 Small Fish Collection Sites •..••.••.••••...•.•. 22 4.4 Laboratory Techniques .••••••.••.•••..•....•..•.••. 23 4.4. 1 Fish Identification and Measurement •••...•••.•• 23 4.4.2 Age Determination •••••••••••.••••.••• , •.•.•••.• 23 4.4.3 Fecundity ...... 25 4.4.4 F00 d Ha bit s ...... 2 5 4.4.5 Data Analysis •.••.•••.•.•...•••.••.•••••••••••• 25 4.5 Aquatic Habitat Analysis ••.•••••••••••.••••...•••• 26 4.5. 1 Reach Definition and Description •••••.••••.•••• 26 4.5.2 Point Samples .•.•..••••••.••••••••.••••••..•••• 26 4.5.3 Mapping Procedures •••..•.•..••••••••..•••.••••. 27 4.6 Limitations of Methods ••••.•..•.••••.••••.••..•••. 27 5. RESULTS AND DISCUSSION •.••..•••..•.•.•..•••••.•••• 30 5. 1 Fish Fauna of the Muskeg River .•.•••••.••••..••.•. 30 5.2 Relative Abundance and Distribution •••.••..••••••• 32 5.3 Tagging Results ••.••.•.••...•..• ~ ••..••.••....•••• 35 5.3. 1 Tag Re I eases and Recaptures ••.••.•••.•.•••. , ••• 35 5.3.2 Movement of Tagged Fish ...... 35 5.3.2.1 Wh i te suckers ...... 37 x TABLE OF CONTENTS (CONTINUED)

Page

5.3.2.2 Longnose Suckers •...... •...... •.•.•.•..•.•. 38 5.3.2.3 .•...... ••.•...•.••..•.•...••.• 39 5.3.2.4 Arctic Grayl ing ....••....•...••.•..•..••....•• 39 5.2 Life Histories of Fish •.....•....••••.•.. 39 5.4. 1 Wh i te S ucke rs •.••.•...•...•.....•.••..•...... 39 5.4.1.1 Seasonal Timing of Upstream Mi9ration ..•....•. 39 5.4.1.2 Diel Timing of Upstream Migration •....••....•. 40 5.4.1.3 Spawning Period .•.•.••..••.•.••.....•...... •.• 40 5.4.1.4 Spawn i n g Area s .•..•..•..••..•..•.•••.•..••.... 46 5.4.1.5 Length of Time Spent in the Muskeg River ..•... 46 5.4.1.6 Seasonal and Diel Timing of Downstream Migration ..••...•••.••.•.••.•.•.•....•...•... 47 5.4.1.7 Spawning Mortality .•..•.•.••.••....•.••.•.•.•. 47 5.4.1.8 Size Composition of Migrant White Suckers •.... 49 5.4.1.9 Age Composition of Migrant White Suckers •.•.•. 53 5.4.1.10 Sex Ratio of Migrant White Suckers .••••.••..•. 53 5.4.1.11 Homing of White Suckers ...... ••.•.•..•••...... 57 5.4.1.12 Fecund i ty .....•....•..•••..••.•..•.....•..•••. 57 5.4.1.13 Age and Growth ..•...•..•.....•.•.•...... •..... 58 5.4.1.14 Sex and Ma t uri t y •...•...•....•....•.•.••.•.... 63 5.4.1.15 Length-Weight Relationship.; ..•...... 65 5.4.1.16 Growth of Young-of-the-Year ...... •...... •.. 65 5.4.1.17 Food Ha bit s ...... •...... •...... 66 5.4.1.18 Rea r i n g Area .•...... •...... •....•...... 66 5.4.1.19 ov e rw i n t e r i n g ...•.•...... •...... •.....• 66 5.4.2 Longnose Suckers ...... •...... •.....•.•..• 69 5.4.2.1 Seasonal Timing of Upstream Migration .....•... 69 5.4.2.2 Diel Timing of Upstream Migration ...... •..•..• 69 5.4.2.3 Spawn i ng Per iod .••.....•..•...... •...... ••..• 72 5.4.2.4 Spawn i ng Areas ...•..•.•....••••.•..••.•...... 72 5.4.2.5 Length of Time Spent in Muskeg River ...... • 73 5.4.2.6 Seasonal and Diel Timing of Downstream Migration ....•...... •...••....•..•..•....•.. 73 5.4.2.7 Spawning Mortality ••...... ••.•.•..•.•.... 73 5.4.2.8 Size Composition of Migrant Longnose Suckers .. 75 5.4.2.9 Age Composition of Migrant Longnose Suckers ..• 75 5.4.2.10 Sex Ratio of Migrant Longnose Suckers , •...••.• 75 5.4.2.11 Homing of Longnose Suckers ...... •.....•.....•• 80 5.4.2. 12 Fecundity •...•.••.....•..•...•.••.•...•••••••• 80 5.4.2.13 Age and Grow t h ..•.•..•....•••...•....••...•.•. 82 5.4.2. 14 Sex and Ma t uri t y .•••...••. ••...•....•••.•.••.• 86 5.4.2.15 Length-Weight Relationship ...... •.•...•..••... 86 5.4.2.16 Growth of Young-of-the-Year ••.•.•••..•...... 88 5.4.2.17 Food Hab i ts ...... •...•..•...••••..•.•.•.•..... 88 5.4.2.18 Rearing Areas ....•....•.••••.••.•...•.•••....• 89 5.4.2.19 Overwintering ••.•....•.•..•.••••••..•.•...•.•. 89 5.4.3 Arctic Gray1 ing ....••.••.••..•••.••••.•.••..• 89 5.4.3.1 Spr i ng Movements .•••..•..•..••••••••.•...•.••• 89 xi i

TABLE OF CONTENTS (CONCLUDED)

Page

5.4.8.6 Food Ha b j t s .•.•••••••.•••••••.••.••• ,...... 133 5.4.9 S 1 i my Sc u 1pin •.•..•••.••••.•.•••...••.•.•.•••• 13 3 5.4.9.1 Distribution and Relative Abundance ...... 133 5.4.9.2 Age and Growth .••..•.•.••.•.••••.•.....••.•••• 133

5.4.9.3 Sex and Ma t uri t y ..••..••..•.•....•.•.••••• 0 • •• 133 5.4.9.4 Length-Weight Relationship '0 •••••••••••••••••• 135 5.4.9.5 S paw n i n g ••.•••.••••.•..•.••..•...••.•.•.••...• 1 35 5.4.9.6 Food Habits ..•.•.••.•..•.••••.••••...... •..... 135 5.4.10 Longnose Dace .•.••••.•••••..•.•..•••.••.•.••.. 136 5.4.10.1 Distribution and Relative Abundance ...... 136 5.4.10.2 Age and Growth •.••...•.•.•.•.•..•.••••.•.•.••. 136

5.4.10.3 Sex and Ma t uri t y .•...... •.•..•.••. 0 • • • • •• 136 5.4.10.4 Length-Weight Relationship •••.••••..•••.•••••• 136 5.4.10.5 Spawn i n g ••••.••••••..•...••••.••...••••.•.•••• 1 36 5.4.10.6 Food Ha bit s •.••.••. ,...... 138 5.4.11 Other .Small Fish Species ••••.•..•.•.•...•.•..• 138 5.4.11.1 Pearl Dace •...••.•.••.•••....•..•••.•..•••.••. 138 5.4.11.2 -Perch .••..•..••••••.•.••....••••....•••. 138 5.4.11.3 Ninespine Stickleback ••..••••••••..•.••....•.• 139

5.4.11.4 Fathead Mi nno\tJ •. 0 ••••••••••••••• 0 •••••••• o •••• 139 5.4.11.5 Spottail Shiner ••••.••....•...... •.. , ••.....•. 140 5.5 Habit8t /'\nalysis ...... 140 5.5. 1 ttuskeg ftiver t1ainstem ..••..••.....•..•.•.....• lifO 5.5.1. 1 Reach 1M •..•....•...••.•.••••..•••••.•.•..•..• 14() 5.5.1.2 Reach 2M •.•....••.••....•••..•.••••••.••.•..•• 149 5.5.1.3 Reach 3M .•••..••.•••••.•••••.•.•.•.•..•.••.•.• 151 5.5.1.4 Reach 4M •.•••• ,...... "153 5.5.1.5 Reach 5M •.•••.••••••.•..•..•.••••....•..•••••• 154 5.5.2 Ha rt 1ey Creek and Kea r 1 Creek ••.•.•..••••••••• 155 5.5.2.1 Rea c h 1 H •.••.••.•.•..•••••..••...•.••••••••.•• 155 5.5.2.2 Reaches 2H, 3H, and 4H ••••••••••.•••....•.•.•. 158 5.5.2.3 Reach 5H ••••••.•.••.••.•••••..••.•••••••.•.••• 159 5.5.2.4 Kea r 1 Cree k ...••.••••....•••.•••••.•.••.•.•.•. 159

6. CONCLUSiONS •..•••••••.•.•••••••...••.••..•.••••.• 161

7. REFERENCES CITED ...... 164

8. APPENDICES 172 8. 1 Maximum and Minimum Daily Water Temperatures Recorded at the Muskeg River Counting Fence, 1977 •••••••••.••••••••.•.•.•..•••.••••••••••••••• 172 8.2 Dates of Tagging and Recapture, Location of Recapture, Distances Travelled, and Elapsed Time Between Release and Recapture for Fish Tagged at the Muskeg River Counting Fence in 1976 and 1977, and Subsequently Recaptured Outside the Muskeg Watershed in 1977 and 1978 ...... 174

9. LIST OF AOSERP RESEARCH REPORTS 178 xi

TABLE OF CONTENTS (CONTINUED)

Page

5.4.3.2 Size of Migrant Grayling ••••••••••••••.••••••• 90

5.4.3.3 Spawning .••.••• 0" 0 •••••••••• 0 •••••••••••••••• 93 5.4.3.4 Summer Residence of Arctic Grayl ing •••.••••••• 94

5.4.3.5 Age and Growth ..••••••.••••.•.••.•••.• 0 ••••••• 95 5.3.4.6 Length-Weight Relationship •.••••••.••••••.•••• 99 5.3.4.7 Sex and Matur i ty .•••••••••••••••.••••••••••••. 99 5.3.4.8 Fecund i ty ..•.••.•••••...••••.•.••.••••••.••.•• 99 5.3.4.9 Food Hab its ...•.•..••••••.••.••••••.••••••.••• 100 5.4.3.10 Rea r i n g A rea s 102 5.3.4.11 Overwintering ...... 102 5.4.4 Northern Pike 103 5.4.4.1 Movements and Distribution .••••.•.•.•.•••••••. 103 5.4.4.2 Spawn i ng •..•.•..•••.••••..•••••••••••.••••.•.• 105 5.4.4.3 Length-Frequency Distribution ••••••••••••••••• 106 5.4.4.4 Age and Grow t h ••...••...••.•••••.•• •••••.••••• 106 5.4.4.5 Length-Weight Relationship •••••••••..••••••••• 11 1 5.4.4.6 Sex and Maturity .•..•..•.•.••.•.•.••.••••••••• 11 1 5.4.4.7 Fecund i ty , .•••.•.••.•..•.•.•.•.•.••••.•.•.•••• 11 1 5.4.4.8 Food Habits ..•...••..•...... •••.••••••••..•.•.• 113 5.4.4.9 Rearing and Overwintering .•••••••••••••.••.••• 11 3 5.4.5 Moun ta i n \~h i tef ish .•....•.••.••..••.••.••.••.• 115 5.4.5.1 Spring Movements and Distribution .••.•...•...• 115 5.4.5.2 Spav"ning ..•.•.....•.•.•...••.•••••••••••••••.. 116 5.4.5.3 Length-Frequency Distribution ••.•••••••.•.•.•• 116 5.4.5.4 Ag e and Grow t h ..••.•..•..••••. , ••••••.•. 116 5.4.5.5 Sex and Maturity ••••.•...••••.••••••••••..•••• 119 5.4.5.6 Length-Weight Relationship ••.•••....••.••••••. 119

5.4.5.7 Food Habits •.•.•..•..•... , .••••••••.•••.. 0 •••• 119 5.4.6 Other Large Fish Species ••••••.••••.•.••.••.•• 119 5.4.6.1 Lake \~h i tef ish •.••.••.•.•..••.•••.••.••.•...•• 119 5.4.6.2 Bur bo t ...... ••.••••...... •••••..••.••••••••. 120 5.4.6.3 Wa 1 1 eye .•..• ~ •..•••..•••.•.•..•••..•• \ ••••••.. 121

5.4.6.4 Dolly Varden •..•••••••.••••••••.. 0 ••••••••••• 121 5.4.6.5 La ke Cis co ...•.•••...•..•••••.••.••••••••..••• 121

5.4.6.6 Ye 1 1ow Per c h ••. 0 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 121 5.4.7 Brook Stickleback •.••••••••.••••••.• , ••••.•••• 122 5.4.7.1 Distribution and Relative Abundance ...... 122

5.4.7.2 Age and Grow th .• 0 ••••••••••••••••••••••••••••• 122 5.4.7.3 Sex and Maturity ••••••••••••• , .••••••.•••••••• 122 5.4.7.4 Length-Weight Relationship •••••••••••••.. , .••• 127 5.4.7.5 Spawn i ng •..•••••••.• • •••••••••••••••••..••••• 127 5.4.7.6 Food Habits ••••.••••.•.•.••••••••..••••.•••.•• 127 5.4.8 Lake Chub ..•••••••.•••••••••••.••••.•••.•.• •• 130 5.4.8.1 Distribution and Relative Abundance ...... 130 5.4.8.2 Age and Growth .••••.•••••••••••.••••••••••.••• 130 5.4.8.3 Sex and Matur i ty .•••••••••..••••••••••••••.•.• 132 5.4.8.4 Length-Weight Relationship ••••••••• ! •••••••• o. 132 5.4.8.5 Spawn i ng ••.•••••• • •••••.•••.••.••••••••••..••• 132 xi i i

LIST OF TABLES

Page

1. Summary of Physical and Chemical Characteristics of the Muskeg River on Several Dates, 1976 ...... 16

2. Sampl ing Schedule for Muskeg River Counting Fence, 1977 ...... 20

3. Dates of Small Fish Collections at Each Col lection Site, Muskeg River Watershed, 1977 ...... 24

4. List of Fish Species Captured in the Muskeg River Drainage during 1976 and 1977 ...... 31

5. Summary of Fish Recorded at the Muskeg River Counting Fence, 1977 ...... 33

6. Number of Fish Captured by Seine, Minnow Trap, Drift Net, and Dipnet at Each Smal 1 Fish Collecting Site in the Muskeg River Drainage, 1977 ...... 34

7. Summary of Tag Releases and Recaptures by Species for Fish Tagged at Muskeg River Counting Fence, 1976 and 1977, and Recaptured Outside the Muskeg River Watershed ...... 36

8. Summary of Fish Enumerated during the Counting Fence Operation in the Muskeg River, 1977 ...... 41

9. Summary of Diel Timing of the Upstream Migration of White Suckers in the Muskeg River, 1977. Fish that were Counted at Times other than those Indicated were Included in the Next Time Check Period ...... 45

10. Summary of Diel Timing of the Downstream Migration of White Suckers in the Muskeg River, 1977. Fish that were Counted at Times other than those Indicated were Included in the Next Check Period ...... 48

11. Length-Frequency Distribution of White Suckers during Migration, Muskeg River, 1977 ...... 50

12. Sex Ratio for White Suckers during the Upstream Migration, Muskeg River, 1977 ...... 56

13. Fecundity Estimates for White Suckers Sampled during the 1977 Muskeg River Spawning Migration ...••...... • 59 xiv

LIST OF TABLES (CONTINUED)

Page

14. Age-Length Relationship (Derived from Fin Rays and Otol iths) for White Suckers Captured in the Muskeg River Watershed, 1977, Sexes Separate and Combined Sample (Includes Unsexed Fish) ...... •••...•.• , ...•.•• 60

15. Age-Weight Relationship for White Suckers Captured in the Muskeg River Watershed, 1977, Sexes Separate and Combined Sample (Includes Unsexed Fish) ....••..... 61

16. Age-Specific Sex Ratios and Maturity for White Suckers from the Muskeg River Drainage, 1977. Sex Ratios Were Based Only on Fish for which Sex was

De term i ned •..•.....•••.•.....•...••.. 6 • • • • • • • • • • • • • • • • 64

17. Food Habits of Adult Longnose Suckers, White Suckers and Lake Whitefish Captured from the Muskeg River,

1977 to & ...... e * ••••••••• ., ., ••• c. .. e .... " ...... 67

18. Food Habits of Young-of-the-Year and Juveniles of the Larger Species Captured in the Muskeg River, 1977 68

19. Summary of Diel Timing of the Upstream Migration of Longnose Suckers in the Muskeg River, 1977. Fish that were Counted at Times other than those Indicated were Included in the Next Time Check .•....•.•••...... ••.•.. 71

20. Summary of Diel Timing of the Downstream Migration of Longnose Suckers in the Muskeg River, 1977. Fish that were Counted at Times other than those. Indicated were Included in the Next Time Check ..••.••.•..••..•.•...•. 74

21. Length-Frequency Distribution of Longnose Suckers During the Upstream Migration in the Muskeg River,

1977 ...... c ...... e"' ••••••••••• 76

22. Sex Ratio for Longnose Suckers during Upstream Migration, Muskeg River, 1977 ...... •...... •..• 79

23. Fecundity Estimates for Longnose Suckers Sampled During the 1977 Muskeg River Spawning Migration •....•• 81

24. Age-Length Relationship (Derived from Fin Rays and Otoliths) for Longnose Suckers Captured in the Muskeg River Watershed, 1977, Sexes Separate and Combined Sample (Includes Unsexed Fish) •..•.•.••••.••• 83 xv

LIST OF 'TABLES (CONTI NUED)

Page

25. Age-Weight Relationship for Longnose Suckers Captured in the Muskeg River Watershed, 1977, Sexes Separate and Combined Sample (Includes Unsexed Fish) 84

26. Age-Specific Sex Ratios and Maturity for Longnose Suckers From the Muskeg River Drainage, 1977. Sex Ratios were Based Only on Fish for which Sex was Determi ned ...... 0...... 87

27. Length-Frequency Distribution of Arctic Grayling during the Upstream Migration in the Muskeg River,

1977 ...... 0 •••••••••••••••••••••••• •••••••••• 91

28. Age-Length Relationship (Derived from Scales) and Age-Weight Relationship for Mountain Whitefish and Arctic Grayl ing Captured in the Muskeg River, 1977, Sexes Combined (Includes Unsexed Fish) ...... 96

29. Size and Weight Relationships for Young-of-the-Year and Juveniles of Larger Fish Species Collected from the Muskeg Ri ver, 1977 ...... o. ....•• .•.. .••• •..•. • 97

30. Food Habits of Arctic Grayl ing Collected ~rom the

Muskeg River during 1976 and 1977 0 ••• 0 •••••••••••••••• 101

31. Age-Length Relationship (Derived from Scales) for Northern Pike Captured in the Muskeg River, 1977, Sexes Separate and Combined Sample (Includes Unsexed

F ish) .. 0 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 108

32. Age-Weight Relcltionship for Northern Pike Captured i~ the Muskeg River, 1977, Sexes Separate and Combined Sample (Includes Unsexed Fish) ...... 110

33. Age-Specific Sex Ratios and Maturity for Northern Pi~e from the Muskeg River, 1977. Sex Ratios were Based Only on Fish for which Sex was Determined ...... 112

34. Actual Egg Counts of Two Northern Pike Females Sampled during the 1977 Spawning Period, Muskeg

Rive r 0 ••••••• 0 ••••••••••••••••• 0 • • • • • • • • • • • • • • • • • • • • •• 114

35. Age-Length (Derived from Otol iths) and Age-Weight Relationships, Age-Specific Sex Ratios, and Maturity of Small Fishes Captured in the Muskeg River, 1977 .... 124 xvi

LIST OF TABLES (CONCLUDED)

Page

36. Sex and Maturity Ratios, by Size C1ass~ for Brook· Stickleback Captured from the Muskeg River Water­ shed, 1977. Sex Ratios were Based Only on Fish for which Sex was Determined ...... •...•...... •.. 128

37. Food Habits for Small Fishes Collected from the Muskeg River, 1977 ....•.•.•...... •..•...... •. 129

38. Physical Characteristics of the Muskeg River

Mainstem, 22-23 June 1978 .. o •••••••••••••••••••••••••• 143

39. Summary of Physical and Chemical Information Collected at Each Sampling Point in the Muskeg Rive r Wa t e r she d, 22 - 23 June 1978 ...... 144

40. Number of Fish Captured in Small Mesh Seines at Each Sampling Point in the Muskeg River Watershed, 22-23 June 1978 ....•..•....•..•..•.....•...... •...... 145

41. Percentage Composition by Numbers for the Major Benthic Macro-invertebrate Groups Taken at Each Samp1 ing Point in the Muskeg River Watershed, 22-23 June 1978 ...... •...... •...... ••...... •.. 146

42. Documented Distribution of Adult and Young Fish in the Muskeg River Mainstem Based on Catch Data Obtained in 1976-1978, and on Reports by Other I n d i v i d ua 1s •••..••.••••••••••••••••.••.••..••••••.•••• 147

43. Physical Characteristics of Hartley Creek, 22-23 June 1978 ...•...... •.....••..•...•. 156

44. Documented Distribution of Adult and Young Fish in Hartley Creek and Kear1 Creek Based on Catch Data Obtained in 1976-1978, and on Reports by Other I nd i vi d ua 1s ...... •.•...... •.....•...... ••.•...• 157

45. Maximum and Minimum Daily Water Temperatures Recorded at the Muskeg River Counting Fence, 1977 .•... 173

46. Dates of Tagging and Recapture, Location of Recapture, Distances Travelled, and Elapsed Time Between Release and Recapture for Fish Tagged at the Muskeg River Counting Fence in 1976 and 1977, and Subsequently Recaptured Outside the Muskeg River Watershed in 1977 and 1978 ...... 175 xv i i

LIST OF FIGURES

Page

1. Map of the AOSERP Study Area Indicating the Location of the Muskeg River ...... 7

2. Map of the Muskeg River Drainage Basin Indicating the Location of the Counting Fence and Small Fish Collection Sites...... 8

3. The Upper Muskeg River at Small Fish Collection Site 6 .•...... •...... •...... 9

4. The Confluence of the Muskeg River and Kearl Creek at Sma 11 Fish Co 11 ect ion Site 9 ...... 10

5. The Muskeg River Downstream of the Counting Fence .. 0 •• 11

6. The Canyon Section of the Muskeg River ...... •... 13

7. Discharge of the Muskeg River from 1 April to 31 October 1977 ...... •...... 14

8. Discharge of the Muskeg River from 1 August to 30 November 1978 ...... 15

9. The Muskeg River Counting Fence ...... 0 18

10. Seasonal Timing of the Migration in the Muskeg River, 1977 ...... 44

11. Length-Frequency Distribution for White Suckers during the Upstream Migration in the Muskeg River, 1977 ...... 51

12. Seasonal Changes in Length-Frequency Distribution for White Suckers during the Upstream Migration in the Muskeg River, 1977 ...... 52

13. Length-Frequency Distribution for White Suckers during Three Time Periods in the Downstream Migration in the Muskeg River, 1977 '0 ••••••••••••••••• 54

14. Age Composition for White Suckers Sampled during the Counting Fence Operation, Muskeg River, 1977 ...... 55

15. Growth in Fork Length for White Suckers from the Muskeg River and from Several Other Areas ...... • 62

16. Seasonal Timing of the Longnose Sucker Migration in the Muskeg River, 1977 ...... 70 xvii i

LIST OF FIGURES (CONCLUDED)

Page

17. Length-Frequency Distribution for Longnose Suckers during the Upstream Migration in the Muskeg River, 1977 " ...... ,...... 7 7

18. Age Composition for Longnose Suckers Sampled during the Counting Fence Operation, Muskeg River, 1977 ...... 78

19. Growth in Fork Length for Longnose Suckers from the Muskeg River and .from Severa 1 Other Areas ...... 85

20. Length-Frequency Distribution for Arctic Grayling during the Upstream Migration in the Muskeg River, 1977 ..... 0...... 0.. ... 92

21. Growth in Fork Length for Arctic Grayl ing from the Muskeg River and from Several Other Areas...... 98

22. Seasonal Timing of the Northern Pike Migration in the Muskeg River, 1977 .... 00...... 104

23. Length-Frequency Distribution for Northern Pike during the Upstream Migration in the Muskeg River, 1977 ...... •....•...... •...... •. 107

24. Growth in Fork Length for Northern Pike from the Muskeg River and from Several Other Areas ...... 109

25. Length-Frequency Distribution for Mountain Whitefish during the Upstream Migration in the Muskeg River,

1977 ...... 0 •••••••• 117

26. Growth in Fork Length for Mountain Whitefish from the Muskeg River and f rom Severa 1 Other Areas ...... 118

27. Length-Frequency Distribution for Brook Stickleback from the Muskeg River, 1977 ...... 123

28. Length-Frequency Diostribution for Lake Chub from the Muskeg River, 1977 .... 0...... 131 29. Length-Frequency Distribution for Slimy Sculpin from the Muskeg River, 1977 ...... 134

30. Length-Frequency Distribution for Longnose Dace from the Muskeg River, 1977 .... 1...... 137 31. Biophysical Map of the Muskeg River, Hartley Creek, and Kear1 Creek •...•...... •....•.....•.•...•.••••. 141 xix

ABSTRACT The fish fauna of the Muskeg River was studied during the open water period in 1976 and 1977. Additional work in 1978 served to define the aquatic habitat of this watershed in terms of various physical parameters. Fish movements into and out of the Muskeg River were monitored by means of a two-way counting fence. The fence was operated from 28 April to 30 July 1976, and from 28 April to 15 June 1977. Small mesh beach seines were used throughout the watershed to collect ~mall fishes. Tags were applied to 3898 migrant fish in order to determine the length of time spent in the Muskeg River watershed by individual fish, and to define migration patterns within the lower Athabasca River system. The general biology of the various fish species was described in terms of their age and growth patterns, food habits, fecundity, etc. White and longnose suckers were the most abundant fish taken at the upstream trap in both years of the study. These two species occurred in equal numbers in 1976 when they accounted for 92% of the 6153 fish enumerated. Arctic grayling (5%) and northern pike (2%) made up most of the remainder. The total 1977 catch at the upstream trap was 5275 fish, of which 56% were white suckers, 31% were longnose suckers, 8% were northern pike, and 3% were Arctic grayl ing. Upstream runs of white suckers, longnose suckers and Arctic grayling represented spawning movements. Some pike also spawn within the Muskeg River system although most of the pike movement observed appeared to be associated with feeding. Small numbers of mountain whitefish, lake whitefish and also undertook spring feeding movements into the Muskeg River. After spawning in the lower 35 km of the Muskeg River and in the lower reaches of Hartley Creek, suckers of both species began to leave the Muskeg River watershed. Most spawners had probably left the stream by mid-June. Sucker fry hatched and began to migrate out of the Muskeg River watershed in early June. Arctic grayling remained in the Muskeg River throughout the summer to feed. Young-of-the-year grayling may overwinter in the Muskeg xx

River, and join the migrant population in the autumn of their second year. Only 2% of all fish tagged were recaptured outside the Muskeg River watershed. Recaptures suggest that white and longnose suckers that spawn in the Muskeg River are part of the Lake Athabasca population and return to the lake to overwinter. A homing tendency was demonstrated by both species. Northern pike showed little tendency to move around. The resident fish fauna of the Muskeg River consists largely of brook stickleback, lake chub, longnose dace, and sl imy sculpin. xxi

ACKNOWLEDGEMENTS The authors greatly appreciate the direction and encouragement provided by R.R. Wallace and the members of the Aquatic Fauna Technical Research Committee, and by M.R. Falk of the AOSERP head office staff during the early planning stages of this project. We also thank D. Hadler and C. Boyle of the AOSERP Fort McMurray office for many services rendered, especially in the area of field communications and logistics. Technical support, both in the field and in the labora­ torY,was provided by M.R. Orr, B. Corbett, D. Miller and D. Rudy. R. Gavel, J. Hardin, B. Anholt, H. Schneider and D. Loucks served as field assistants during the study. Many other people provided field assistance and useful comments throughout the study and we are grateful to all of them. Thanks are due S. Zettler and his staff of the Graphics Department at the Freshwater Institute who prepared the figures for the report, and J. Allan and W. Thompson who typed the manuscript. To the residents of northeastern Alberta who expressed interest in the study and who assisted us through the return of fish tags, we offer our sincere appreciation. Funding for this project was provided by the Alberta Oil Sands Environmental Research Program, a joint Alberta-Canada research program establ ished to fund, direct and co-ordinate envi­ ronmental research in the Athabasca Oil Sands area of northeastern Alberta. 1. INTRODUCTION The proposed development of the Athabasca Oil Sands may introduce disturbance to some lake and river systems of the lower Athabasca River drainage. Especially susceptible is that section of the surface-mineable area for which the Alberta Energy Resources Conservation Board (AERCB) has granted development approval. Local disruption in the form of land clearing, muskeg drainage and removal, stream diversions, and the construction of access routes may affect the water quality and quantity of streams in addition to the physical alterations produced. Other activi­ ties that could affect water quality include tailings pond seepages and saline minewater discharges. The diversion or blockage of streams may affect fish spawning runs. Traditional fish rearing, feeding, and overwintering areas may be disturbed or lost altogether. In the case of migrant fish populations, such local disruptions could be felt over much wider areas. In order to provide information that could be used to minimize the adverse effects of development on fish populations of the Athabasca River and its tributaries, the Alberta Oi 1 Sands Environmental Research Program (AOSERP) through its Aquatic Fauna Technical Research Committee, initiated an integrated series of projects to assess the baseline state of the fish resources of the area. The work, which began in 1976, involves a broadly based fisheries investigation of the Athabasca River as well as site-intensive study of selected tributaries. Tributaries selected for intensive study are those considered to be most immediately imperilled by future surface mining operations or by increased pressure from a growing human population. The Muskeg River, a medium sized watershed on the east side of the Athabasca River, was the first tributary selected for intensive study. Initially, this tributary was selected because a large portion of its drainage lies within the surface-mineable area and because the AERC.B was involved in considering the construction there of two synthetic crude oil plants. The possibil ity was anticipated that construction of one or both of these plants could involve massive watershed disturbance including 2 the eventual diversion of both the Muskeg River and its major tributary, Hartley Creek. Development plans for this area have recently changed with the proposal by the Alsands Project Group~ a consortium of nine oil companies, to develop a single plant north of the Muskeg River. The new proposal will not require the diversion of either the Muskeg River or Hartley Creek, and total destruction of this watershed will, therefore, be avoided. However, construction and operation of the proposed plant, the construction of a proposed new town, and increased access through extension of roadways are expected to place considerable pressure on the fish populations of the Muskeg River. The present study was conducted over a period of three years with the general objective, as outlined in the terms of reference agreed to by AOSERP and the Department of Fisheries and Environment, of describing the baseline states of the fish resources of the Muskeg River watershed and providing a quanti­ tative estimate of the significance of the watershed to the fisheries of the Athabasca River system. Specific objectives for the study were as fol lows: 1. To enumerate the migrant populations of those fish species utilizing the Muskeg River watershed on a seasonal basis; 2. To describe the timing of the seasonal and daily movements of the various fish populations into and out of the Muskeg River watershed, and to obtain information concerning the age and growth, sex ratio, fecundity, food habits, etc., of these fish; 3. To determine the extent of movement of the various non-resident fish populations within the Muskeg River watershed, and to locate critical spawning and nursery areas; 4. To apply conventional (Floy) tags to migrant fish to permit definition of their migration routes within the Athabasca River system; 3

5. To monitor the downstream migration of fry of various species hatched within the Muskeg River watershed, and to estimate recruitment of these species to the Athabasca River system; and 6. To assess the resident fish species of the Muskeg River watershed in terms of relative abundance, distribution and general biology. During the second year of the study it was also our objective to describe in detail the aquatic habitat of the Muskeg River watershed. This work was postponed until year three because the classification system and key adopted by AOSERP for this purpose was unavailable in 1977. Results of work done in 1976 have already been reported in interim form (Bond and Machniak 1977). This report presents the results of work done in 1977 and attempts to draw together the results from both years. It also presents results of the habitat characterization done in 1978 and attempts to relate habitat to fish util ization. 4

2. RESUME OF CURRENT STATE OF KNOWLEDGE Prior to 1976, information relative to the fish fauna of the Muskeg River was limited to that generated by Griffiths' (1973) preliminary survey and subsequent baseline studies conducted by Lombard-North Group Ltd. (1973) and Renewable Resources Consulting Services Ltd. (1974). The latter two studies were performed as part of an environmental assessment of Shell's lease 13 mining project and a summary of the work is included in the lease 13 environmental impact assessment that was filed with Alberta Environment in 1975 (Shell Canada Ltd. 1975). Since Griffith's work was part of a broad regional study intended to evaluate the sport fishery potential of a large number of streams in the oil sands area, his treatment of anyone stream was, of necessity, cursory. He did, however, document the occurrence of eight fish species in the Muskeg River and identified the presence of a grayl ing popUlation in the lower reaches. He did not examine the upper Muskeg River watershed nor did he sample Hartley Creek. The work by Lombard-North Group Ltd. (1973) and Renew­ able Resources Consulting Services (1974), while extending knowledge of the fish fauna of the Muskeg River, left many questions unanswered. These studies suggested an important role for the Muskeg River in terms of providing spawning areas for longnose suckers and white suckers although they were unable to enumerate the runs. The capture of Arctic grayling, longnose and white suckers, and mountain whitefish in Hartley Creek suggested a greater importance for that tributary than was predicted by Griffiths. The significance of the mouth region for fish popula­ tions from the Athabasca River was implied and an attempt was made to relate fish utilization to habitat type. However, because these studies concentrated on the region within leases 13 and 30, they provided no information on the resident fish populations of the upper watershed or the extent to which this region is utilized by migrant popUlations. Since no attempt was made to capture small fish, the likely presence of several species was not detected, nor were the younger age classes of larger 5

species sampled. Small sample sizes precluded an adequate description of the life history and general biology of several species. Previous studies did not permit an adequate description of the fish resources of the Muskeg River watershed. Quantifica­ tion of the migrant populations that utilize the Muskeg River watershed on a seasonal basis and a clear description of such seasonal utilization patterns were required. Areas within the watershed that may be critical in the life histories of the various species required definition. The composition and distribution of resident fish populations required description. Life history patterns and general biological features of all species required further elucidation. 6

3. DESCRIPTION OF THE STUDY AREA The Muskeg River originates in the Muskeg Mountain uplands and travels approximately 100 km before joining the Athabasca River 58 km downstream from Fort McMurray (Figure 1). The total area drained by the Muskeg River system is 1464 km2 , of which 80% is forest and 20% muskeg (Northwest Hydraulic

Consultants Ltd. 1974). Only 2% of the total wat~rshed area is lakes, the largest of which, Kear1 Lake (Figure 2), is only 5.4 km 2 in surface area and 2 m in maximum depth. Hartley Creek (Figure 2), the major tributary of the Muskeg River, drains 325 km 2 to the south of the main stream and enters the Muskeg River about 33 km upstream from its confluence with the Athabasca River. The water of the Muskeg River and Hartley Creek is stained brown as a result of the presence of humic and fu1vic acids. The climate of the study area is continental, charac­ terized by cold winters, short, cool summers, and wide temperature fluctuations (Intercontinental Engineering of Alberta Ltd. 1973). Precipitation records for the Muskeg Mountains show the annual precipitation to be 49.8 cm, of which 33.6 cm falls between May and September (NHCL 1974). The upper portion of the Muskeg River (Figure 3) is well drained and vegetated by mixed spruce and areas of treed muskeg. Surficial deposits consist of relatively thick drift composed mainly of ti 11 (NHCL 1974) while the bedrock material is largely Cretaceous shales and sandstones. The large central area of the watershed is flat, poorly drained and covered with marshland and treed muskeg (Figure 4). A thin surficial layer of outwash sand is underlain in this area by the McMurray Oil Sands Formation. The Muskeg River leaves the flat, central portion of its watershed in the lower 16 km of its course and begins to cut through the McMurray Oil Sands and Waterways limestone (NHCL 1974). The lower reaches of the river valley are stream cut and the channel is frequently confined by limestone outcroppings. The stream channel in this area is fairly stable, the substrate, consisting of large areas of gravel (Figure 5) with occasional areas of boulders and 7

sru~~~_

Alherta Birch Mountains Edmonton •

Km 10 0 10 20 30 laQiff Mi 10 0 20 30 40

Figure 1. Map of the AOSERP study area indicating the location of the Muskeg River. 8

LIMIT OF MUSKEG /' RIVER WATERSHED

13

AOSERP STUDY AREA

MILES 5 0 5 10 I .,~, I I JI1 I I I I 5 0 5 10 15 20 KILOMETRES

Figure 2. Map of the Muskeg River drainage basin indicating the location of the counting fence and small fish co 1 1e c t ion sit e s . 9

Figure 3. The upper Muskeg River at small fish collection Site 6. 10

Figure 4. The confluence of the Muskeg River and Kearl Creek at small fish collection Site 9. 11

Figure 5. The Muskeg River downstream of the counting fence. 12 bedrock (Figure 6). The Muskeg River generally freezes over in late October and remains ice-covered unti 1 late Apri 1. Ice left the stream on 15 Apri 1 in 1976 and on 22 Apri 1 in 1977. Under ice cover, water temperatures remain near OOC but the stream can warm quickly in the spring and reach high temperatures in mid-summer. A maximum water temperature of 25°C was recorded in 1976 and daily temper­ ature fluctuations of up to SoC were observed (Bond and Machniak 1977). Discharge records for the Muskeg River (Water Survey of Canada 1975) showed a mean dai ly discharge in 1977 of 2.3 m3/s (range 0.2 to 13.5 m3/s). After the spring flood, water levels generally decl ine through the summer although considerable fluctuation may occur as a result of heavy precipitation (Figures 7andS). A brief description of the physical and chemical characteristics of Muskeg River water is given in Table 1. More complete physical and chemical data for this stream are presented by Seidner (in prep.). 13

Figure 6. The canyon section of the Muskeg River. 14

12 -cj Q.) ~ rc") IO~ I\ Last Days of E JflJ Ice Conditions ----W (!) 8 0:: \~ . _.. _- ......

2

15 30 15 31 15 30 15 31 15 31 15 30 15 30 APRIL MAY JUNE JULY AUGUST SEPTEMBER OCTOBER Figure 7. Discharge of the Muskeg River from 1 Apri 1 to 31 October 1977. 30 -o Q) (/) ...... I'() ~ 20

FREEZE-UP V; «~ I ~ .. I ~ 10 o

15 31 15 30 15 31 15 30 AUGUST SEPTEMBER OCTOBER NOVEMBER Figure 8. Discharge of the Muskeg River from 1 August to 30 November 1~78. 16

Table 1. Summary of physical and chemical characteristics of the Muskeg River on severa1 dates, 1976a .

Date Parameterb 11 Feb 14 May 27 July 7 Sept

Discharge (m 3!s) 0.4 2.6 0.9 2,9

pH (pH units) ].7 8. 1 1 8 7.8

Specific conductance 367 259 380 270 (llmhos!cm @ 25C)

Turbidity (JTU) 6.,3 2.8 1·">, I . 14,6

Apparent colour 65 70 35 80 (Relative units)

Total alkal inity 119 136 228 148

Total hardness 139 137 196 137

Humic acid 8 4 9

Fulvic acid 10 20 9

Filterable residue NO 181 276 162

----,--_. -- a Data provided by Mr. C. ~ Froel ich, Alberta Oi 1 Sands Environmental Research Program. b Except as indicated, data are expressed as mg/l. 17

4. MATERIALS AND METHODS The fish fauna of the Muskeg River was studied during the open water period of 1976 and 1977. During spring and summer of both years various methods were employed to collect fish through­ out the watershed although the major emphasis was placed on the construction and operation of a two-way fish counting fence to monitor spring movements of fish into and out of the Muskeg River. The fence was established approximately 1 km upstream of the mouth of the tributary, thus permitting enumeration of a large proportion of the fish moving from the Athabasca River into the Muskeg River watershed. The counting fence was operated from 28 April to 30 July in 1976 and from 28 April to 15 June in 1977. The absence of a fall fence operation in 1976 and 1977 was seen as a serious omission in the study and it was planned to conduct such an operation in 1978. However, extremely high water in September and October (Figure 8) made this impossible. During June 1978, a biophysical inventory of the Muskeg River was conducted in order to describe the aquatic habitats of the watershed and relate habitat types to fish uti lization.

4.1 COUNTING FENCE CONSTRUCTION The counting fence (Figure 9) was constructed of 2.5 cm by 2.5 cm welded wire fabric and was installed in such a way as to form a complete temporary barrier to fish. Fish travell ing upstream or downstream encountered the fence at some point and were led into one of the holding boxes where they could be worked with. Complete detai ls of construction and installation are given in Bond and Machniak (1977).

4.2 COUNTING FENCE OPERATION The operation of a counting fence of this type is highly labour intensive, especially during the high water period generally encountered in the spring. Debris carried by the river tends to clog the openings in the wire mesh, placing great pressure on the structure. Frequent cleaning is required to remove such debris and prevent the fence being washed out. 18

Figure 9. The Muskeg River counting fence. 19

4.2. 1 Sampling Schedule The 1977 counting fence was operated from 28 April to 15 June. The downstream trap, however, was kept closed until 12 May to prevent upstream migrants from drifting into it. The upstream trap was checked six or more times daily as required during the early part of May when fish movement was intense. After 12 May, the traps were usually checked three or four times daily (Table 2).

4.2.2 Trap Checks Each trap check was performed by two persons, one _ working inside the trap and the other serving as recorder. The number of fish of each species was recorded and as many fish as possible were measured and sexed. The development of pearl organs by male white and longnose suckers often made it possible to distinguish between the sexes for these species without sacrificing the fish. Smaller fish, that were either females or immature males, could not be sexed by this method, and where doubt existed, no sex was recorded. Handling of fish was minimized by using a scoop constructed of PVC pipe and rochelle netting, and fish were passed through the fence in the direction in which they were moving. Relative water level was recorded at each trap check using a metre stick anchored in the stream. A continuous record of stream temperatures was provided by a Ryan Model D15 recording thermometer. Temperature data are summarized in Appendix 8.1. The fence was cleaned as required and examined regularly for holes.

4.2.3 Tagging Numbered Floy anchor tags (Type FD-68S) were applied to as many fish (mainly suckers and pike) as was practicable. Tags were inserted into the left side of the fish near the base of the dorsal fin. The risk of infection was minimized by rinsing the tagging gun in disinfectant and in fresh water before each insertion. Suckers, retained in a holding pen for up to 15 minutes, rarely showed any ill effects. However, in 1976, 20

Table 2. Samp 1 i ng schedule for Muskeg River counting fence, 1977.

Time of Fence Checka Date 0300 0900 '1200 1500 1800 2100 2400

28 Apri 1 + + 29 + 1300 1600 + 30 + + + 1 May + + + 2 + + + + + + 3 + + + + + + 4 + + + + + + 5 + 1300 + 2200 + 6 + + + + + + 7 + 0500 + + + + + 8 + 0600 + + + + + 9 + 0500 + + + + + 10 + + + + + + 11 + + + + + + 12 + + + + + 13 + + + + 14 + + + + 15 + + + + 16 + + + 17 + + + 18 + + + 19 + + + + 20 + + + 21 + + + 22 + + + + 23 + + + 24 + + + + 25 + + + 26 + + 27 + + + 28 + + + 29 + 30 + + + 31 + + + 1 June + + + + 2 + + 3 + + + 4 + + + 5 + + + 6 + + + 7 + + + 8 + + + 9 + + + + 10 + + + + + 11 + + + 12 + + + + 13 + + + + + 14 + + + 15 + Operations terminated

a Actual check time indicated where different from scheduled check time. 21 grayling did not appear to cope well with the stress imposed by the application of Floy tags. Therefore, metallic clip tags were utilized for this species in 1977. These tags were affixed to the left operculum and no mortality was observed. Depending on the species, either fork or total length (±1.0 mm) was recorded for each fish tagged and the sex was noted if possible. Tagged fish were not weighed and no body structures were retained for age determination. Tagging was done only during the day in 1976 but in 1977. floodlights, operated from a portable generator, enabled the fence crew to tag fish during the late evening and at night. Care was taken at all times not to impede the progress of the fish any more than necessary. When fish were observed to be backing up in front of the trap. tagging was curtailed and the remaining fish were simply passed through and enumerated. The tagging program was well publicized by posters and press releases and a two dollar reward was offered for returned tags. Tag returns were made by sport fishermen along the Athabasca River, by domestic fishermen on the Athabasca River and Lake Athabasca, and by commercial fishermen on Lake Athabasca. Personnel of LGL Ltd., Environmental Research Associates, Edmonton, and Aquatic Environments Ltd., Calgary, also returned tags, while others were recovered by fishery crews working on the Athabasca River (Bond and Berry in prep.b), the Steepbank River (Machniak and Bond in prep.), and the MacKay River (Machniak et a 1. in prep.).

4.2.4 Dead Samples Small numbers of fish were sacrificed each day for biological analysis. Fork or total length (±l.O mm) and weight (±20 g) were recorded for each fish. Weights of some small fish were determined on a triple beam balance (±O.l g). Sex and stage of maturity were determined by examination of the gonads. A fish was considered to be mature if it appeared that it would spawn or had already spawned in the year of capture. A ripe fish was a mature fish whose gonads were close to spawning condition and 22 from which sexual products could be expressed by application of light pressure to the abdomen. A spent or spawned out fish was a mature fish which had obviously spawned shortly prior to its capture. Ovaries for fecundity work were removed from a number of longnose suckers, white suckers, and Arctic grayling and weighed fresh on a triple beam balance (±0.1 g). These ovaries were then preserved in Gilson's fluid. Stomach contents were noted and a small number of stomachs were preserved in 10% formalin for a more detailed assessment of food habits. Scales were removed from the appropriate body location (Hatfield et a1. 1972) for ageing of grayl ing, mountain whitefish, pike, walleye, and lake whitefish. Otoliths (ear bones) were taken from , and for suckers, the left pectoral fin was retained for age determination.

4.3 OTHER FISH COLLECTION TECHNIQUES Apart from the counting fence, fish were collected by various methods including small mesh beach seines (3.2 mm oval mesh), commercial minnow traps, gill nets, electrofishing, drift nets, dip nets, and angl ing. Large fish captured by these methods were either dead sampled or measured and tagged. Small fish were initial ly preserved in 10% formalin and later transferred to 40% isopropyl alcohol. An attempt was made to monitor the downstream fry migrations in 1977 using a bomb drift sampler (Burton and

Flannagan 1976). However, the 202 ~m Nitex uti1 ized in the construc­ tion of the sampler quickly became clogged with debris, rendering the sampler ineffective. Drift samplers, as a consequence, were useful only in identifying the starting time of the fry migration.

4.3.1 Small Fish Collection Sites Small fish were collected from 10 general areas of the Muskeg River watershed. The sampling sites utilized in 1977 were essentially the same as we sampled in 1976 (Bond and Machniak 1977). Sampling Sites 1, 2, and 3 were located downstream of Shell lease 13, Sites 5, 6, 8, 9, and 10 were upstream of the lease, while Sites 4 and 7 were situated within the lease boundaries 23

(Figure 2). Each site consisted of from 10 m to 3 km of stream channel that was sampled in such a way as to obtain a representa­ tive sample of the fish population of the area. No standard unit of effort was used. The dates on which each location was sampled in 1977 are shown in Table 3. Numerous col lections were made at Sites 1, 2 and 3 on dates other than those indicated in Table 3 and these fish were included in the results.

4.4 LABORATORY TECHNIQUES

4.4. 1 urement Preserved fish specimens were identified using taxonomic keys and descriptions given by Paetz and Nelson (1970) and McPhail and Lindsey (1970). While most fish could be iden­ tified to species, larval catostomids could often be identified only to genus. Small, preserved fish specimens were measured to the nearest 1.0 mm (0.5 mm for some larval fishes) and weighed to the nearest 0.1 g on a triple beam balance.

4.4.2 Age Determination Ages were determined by the scale method for Arctic grayling, mountain whitefish, lake whitefish, walleye, and northern pike. Several scales from each fish were cleaned and mounted between two glass slides and the annuli were interpreted from the image produced by an Eberback microprojector. Longnose and white suckers were aged from cross sections of pectoral fin rays as described by Beamish and Harvey (1969) and Beamish (1973). After embedding the dried fin rays in epoxy, thin sections (0.5 to 1.0 mm) were cut by hand using a jeweller's saw with No.6 or r~o. 7 blades. The sections were then mounted in Permount on glass slides and read under a compound microscope. Ages for all other fish were determined from otol iths. Otoliths were stored in a 1:1 glycerine and water mixture and read whole under a dissecting microscope using reflected light. Where required, the otol ith was ground by hand on a carborundum. 24

Table 3. Dates of sma 11 fish collections at each co 11 ect i on site, Muskeg River watershed, 1977a .

Date of Co 11 ect ion Site No. 20 19 18 30 16 15 13 May June July July Aug. Sept. Oct.

+ + + +

2 + + + + +

3 + + + + + +

4 + + + + +

5 + +

6 + + + + +

7 + + +

8 + + + + +

9 + + +

10 + + + + + al n addition to the above, samples were collected from Site 1 on 5 June and 6 August; from Site 2 on 10 other dates; from Site 3 on five other dates, and from Site 7 on 3 June 1977. 25

Independent age determinations were made by three people in all cases. Where discrepancies existed among the three results, the readers conferred until a consensus was achieved.

4.4.3 Fecundi ty Fecundity was determined for longnose and white suckers using the gravimetric method of estimation described by Healey and Nicol (1975). The ovarian tissue was removed from the sample and the separated eggs dried to constant weight. The weight of a subsample of eggs was determined and the total number of ova then derived by extrapolation. The accuracy of the estimates was assessed by performing total counts on several ovaries.

4.4.4 Food Habits The stomach contents of preserved fish were removed and the food items identified to the lowest possible taxon using keys and descriptions from Pennak (1953). Results were expressed as percentage frequency of occurrence, percentage of total number, and, in some cases, percentage of total volume.

4.4.5 Data Analysis Biological data were analyzed for graphic and tabular presentation using a Hewlett-Packard Model 9810-A programmable ca lculator. Length-weight relationships were described by the power equat i on: a + b (10910L); sb where: W weight in grams L fork or total length in mill imetres a = y-intercept b slope of the regression 1 ine, and sb standard deviation of b. Data summaries and raw data are presently on file at the Freshwater Institute in Winnipeg. 26

4.5 AQUATIC HABITAT ANALYSIS An effort was made to characterize the aquatic habitat of the Muskeg River util izing the procedures described by Brown et ale (1978). In this system, streams are divided into reaches which differ from each other in their physical characteristics. A hel icopter survey is used to produce average values for various parameters over each entire reach and site-specific information is gathered from sample points within each reach.

4.5.1 Reach Definition and Descri ion A reach is a section of stream whose physical properties (habitat characteristics) are relatively homogeneous throughout its length. According to Brown et al. (1978), IIreach boundaries are located in regions where the topography changes drastically, or significant changes in water quality, channel form and/or flow character" occur. Tentative reach boundaries for the Muskeg River and Hartley Creek were assigned by reference to National Topographical Series maps (1 :50,000) and available gradient information (RRCS 1975). These were later verified in the field. Aerial photo interpretation, a recommended method for assigning tentative reach boundaries, was not used in he present study, General descriptions of each reach were acquired during an aerial survey of the Muskeg River. At that time observations were recorded on various aspects of the aquatic habitat. These characteristics, which include velocity, substrate, pools, riffles, riparian vegetation etc., are presented as averages of these parameters over the length of the reach.

4.5.2 Point Samples Site-specific information on biological and physical parameters was collected on 22 and 23 June 1978. The sites sampled included small fish collection sites uti 1 ized in 1976 and 1977 plus several additional locations. At each site, stream width was measured and the depth was taken at three locations across the channel. A rough estimate 27

of stream velocity was obtained by floating a small chip a distance of 5 m and timing it. This was also done at three locations across the channel. The substrate composition at the site was estimated in terms of fines « 2 mm), gravel (2 to 64 mm), larges (> 64 mm) and bedrock. Riparian vegetation and aquatic vegetation were noted and water temperature was recorded using a pocket thermometer. At every second site, dissolved oxygen was determined using a Hach field kit (Model AL-36-B) and pH was estimated by means of colour comparator. Specific conductance was measured using a Beckman RB-3 conductivity meter. From five to seven seine hauls (3.2 mm oval mesh) were made at each location. Fish captured were preserved in 10% formalin in the field and were later identified to species, measured, and weighed. Benthic macroinvertebrates were collected

using the kick method and a net fitted with 202 ~m Nitex. These were also preserved in 10% formalin. Kick samples were later divided into major groups (Chironomidae, Ephemeroptera, Plecoptera, Trichoptera, Simuliidae, 01 igochaeta and "others"). No attempt was made to identify these samples further as extensive inverte­ brate data from the Muskeg River watershed have already been presented by Barton and Wallace (in prep.).

4.5.3 Mapping Procedures A map of that portion of the Muskeg River watershed surveyed was prepared at a scale of 1:50 000 to summarize the biophysical data gathered during this study. Fish data col lected in 1978 were supplemented by those gathered during 1976 and 1977.

4.6 LIMITATIONS OF METHODS The primary objective of the present study was to enumerate and describe the migrant fish populations that utilize the Muskeg River on a seasonal rather than a year-round basis. The best possible means of achieving such an objective is undoubt­ edlya counting fence. However, this apparatus, like any other, has certain limitations. The 2.5 x 2.5 cm wire mesh used in the construction of 28 the counting fence is believed to have been highly effective in catching fish longer than 150 mm in fork length. Smaller fish, although sometimes taken in the trap, were able to pass through the apertures. Seasonal movements of small fish (such as trout­ perch) could not, therefore, be monitored. The Muskeg River counting fence could be operated effectively only at discharge rates of less than 7 m3 /s. Thus, the fence could not be installed unti 1 the spring flood had begun to subside, and fish movements occurring during the peak of the flood could not be monitored. Once the fence was installed, no problems were encountered in either year of the study as water levels remained low during the period of operation. Because of the highly compacted nature of the substrate at the fence site, 1 ittle problem was encountered with holes developing under the structure. Although small numbers of fish may have avoided the traps through such holes, we bel ieve the number to be small relative to the total number counted. We believe that our catch data are highly representa­ tive of the nature and timing of the upstream migrations. Suckers moving upstream quickly located the entrance to the trap and showed no hesitation in entering it. At times of heavy upstream movement, suckers backed below the fence but continued actively to seek a way through. Their progress was delayed as 1 ittle as possible by continuous trap work at such times. Downstream data are considered to be less representative. The downstream trap, especially in 1976 (Bond and Machniak 1977), was inefficient in terms of holding fish. This problem was considerably reduced in 1977 by a modifi­ cation of the trap entrance, whereby the entrance was from the long axis of the trap. The second problem encountered in mon i tor,i ng the downs t ream run was the apparent re 1 uctance of suckers to enter the downstream trap. Fish moving downstream would often stop just ahead of the entrance to the trap and hold there. Many times they would refuse to enter and would move back upstream. Thus, the situation in the Muskeg River was similar to that described by Kendel (1 ) where post-spawning, downstream movements of longnose suckers were delayed by the presence of a 29 counting fence. The counting fence was operated from 28 April to 30 July 1976, and from 28 April to 15 June 1977. Thus, spring and early summer fish movements in the Muskeg River have been fairly accurately described. The exception appears to be Arctic grayl ing, many of which undoubtedly passed upstream prior to fence installa­ tion. The absence of a fall fence operation leaves a serious gap in the study. No firm plans for a fall operation were included in the 1976 and 1977 studies. High water thwarted a planned fall operation in 1978 and prevented a complete enumeration of Arctic grayling during their downstream migration which is suspected to occur just prior to freeze-up. The small mesh seines (3.2 mm) used in the present study are considered to have been highly effective in capturing small fish in the Muskeg River watershed o However, in deep water, and in areas with an uneven bottom (rocks, logs etc.) their value was limited. Such areas might have been more efficiently sampled by an electrofisher or toxicant. Because no winter sampling was conducted, the present study produced no information on fish util ization of the Muskeg River at that time of the year. 30

5. RESULTS AND DISCUSSION

5.1 FISH FAUNA OF THE MUSKEG RIVER Nineteen fish species representing nine famil ies (Table 4) were captured in the Muskeg Rive watershed during 1977. All species taken in 1976 (Bond and Machniak 1977) were represented in the 1977 catch with the exception of spottail shiner. Species captured in 1977 that represent additions to the 1976 results include Dolly Varden, fathead minnow, yellow perch, ninespine stickleback, and lake cisco. The fish fauna of the Muskeg River can be divided into three categories on the basis of the extent to which this watershed forms part of the home range of the various populations. The first category includes a number of species that appear to be more typical of the Athabasca River or other areas outside the Muskeg watershed. It includes lake whitefish, walleye, yellow perch, burbot, Dolly Varden, lake cisco, fathead minnow, ninespine stickleback, and spottail shiner. These species are seldom encountered upstream of the fence site and are most 1 ikely to be taken in the vicinity of the river mouth, an area that may be of considerable importance in te~ms of providing resting areas during migrations within the Athabasca River or nursery areas for young­ of-the-year. The second category includes five species that appear to have establ ished resident populations within the Muskeg River watershed and whose home range is more or less restricted to that watershed. These are lake chub, brook stickleback,longnose dace, slimy sculpin, and pearl dace. For these species the Muskeg River satisfies all requirements of all life stages on a year round basis. The third category includes a number of species to which the Muskeg River represents a small but important portion of their home range. These species, whi le inhabiting areas outside of, and, in some cases, great distances from the Muskeg River for part or most of the year, re urn to the tributary periodically to spawn and/or feed. The Muskeg River watershed may also provide 31

Table 4. List of fish species captured in the Muskeg River drainage during 1976 and 1977.

Family and Species Names Common Names

Family Coregonidae Coregonus cZupeafoNnis (Mitchill) Lake whitefish Prosopium wiZZiamsoni (Girard) Mountain whitefish Coregonus artedii Lesueura Lake cisco ThymaZZus arcticus (Pallas) Arctic grayling SaZveZinus maZma (Walbaum)a Dolly Varden

Fami ly Esocidae Esox Zucius Linnaeus Northern pike

Family Cyprinidae SemotiZus margarita nachtriebi (Cox) Northern pearl dace Couesius pZumbeus (Agassiz) Lake chub Rhinichthys cataractae (Valenciennes) Longnose dace Notropis hudsonius (Clinton)b Spottail shiner PimephaZes promeZas Rafinesquea Fathead minnow

Family commersoni (Lacepede) White sucker Catostomus catostomus (Forster) Longnose sucker

Family Percopsidae Percopsis omiscomaycus (Walbaum) Trout-perch

Fam i 1y Gad j dae Lota Zota (Linnaeus) Burbot

Family Gasterosteidae CuZaea inconstans (Kirtland) Brook stickleback Pungitius pungitius (Linnaeus)a Ninespine stickleback

Fami ly Cottidae Cottus cognatus Richardson Slimy sculpin

Fami 1y Perc i dae Stizostedion vitreum vitreum (Mitchill) Wa 11 eye Perca fZavescens (Mitchill)a Yellow perch a Captured in 1977 but not in 1976. b Captured in 1976 but not in 1977. 32

rearing and overwintering areas for juvenile members of some of these populations. Species included in this group are white sucker, longnose sucker, Arctic grayl ing, mountain whitefish, northern pike, and, perhaps, trout-perch.

5.2 RELATIVE ABUNDANCE AND DISTRIBUTION A total of 5275 fish (10 species) were passed through the upstream trap between 28 April and 15 June 1977 (Table 5). As in 1976 (Bond and Machniak 1977), white suckers (56.3%) and long­ nose suckers (31.1%) dominated the catch. Northern pike (8.2%), Arctic gray1 ing (3.1%), and mountain whitefish (1.1%) accounted for most of the remainder. By 15 June, 2487 fish had been counted at the downstream trap (Table 5). Remaining in the watershed beyond 15 June were 1505 white suckers (53.4% of the total number of white suckers enumerated at the upstream trap), 637 longnose suckers (38.8%), 150 Arctic grayling (93.2%), 374 northern pike (86.4%), and small numbers of several other species. These numbers are certainly con­ servative, especially in the case of Arctic grayl ing which probably began to move into the Muskeg River several days prior to fence installation. Collections throughout the Muskeg River watershed during the summer produced 261 11 fish (Table 6). Suckers accounted for 75.5% of this total, the majority (>96%) being young-of-the­ year. Excluding suckers, brook stickleback was the most abundant small fish in the samples accounting for 47.9% of the total catch. Also occurring commonly were lake chub (25.7%), slimy sculpin (6.2%), and longnose dace (4.4%). Pearl dace, which dominated the resident fish population in the Steepbank River (Machniak and Bond in prep.), comprised only 1.7% of the sma1 1 fish sample in the Muskeg River in 1977 and only 0.4% in 1976 (Bond and Machniak 1977). Brook stickleback were captured at eight of the 10 sampl ing sites in 1977, but, as in 1976 (Bond and Machniak 1977), they were most abundant in the more tranquil water upstream of Site 3. Lake chub were also taken at eight locations and were found in association with brook stickleback at Site 6. However, this 33

Table 5. Summary of fish recorded at the Muskeg River counting fence, 1977.

Number of Fish Species Upstream Trap Downstream Trap

White sucker 2970 1 385

Longnose sucker 1641 1004

Arctic grayl ing 161 11

Northern pike 433 59 a Mountain whitefish 57 17

Lake whitefish 0 6

Wa 11 eye 8 5

Burbot 0

Lake cisco 0

Do 1 1y Va r den 3 0

Total 5275 2487 a Includes a small number of lake whitefish which were misidentified prior to 11 May 1977. Table 6. Number of fish captured by seine, minnow trap) drift net and dipnet at each small fish collection site in the Muskeg River Drainage, 1977.

Muskeg River Hartley Creek Kear 1 Creek Total Area Area 2 Area 3 Area 4 Area 5 Area 6 Other Area 7 Area 8 Area 9 Area 10 N % N % N % N % N N % N % N % N % N % N % N %

Arctic grayling 0.2 5 0.5 13 4.0 1.8 20 0,8 Pear 1 dace 10 1.0 1.4 11 D.4 Lake chub 6 0.9 37 3.7 54 16.7 2.3 42 35.3 6 lD.7 12 48.0 2.6 165 6.3 Longnose dace 6 0.9 19 1.9 1 0.3 1 5.3 4.D 28 1.1 Sucker spp. 610 92.1 857 86.8 174 53.8 275 91.1 1916 73.2 Wh i te sucker 5 0.5 18 5.6 2 0.7 2 1.7 21 37.5 1 4.0 4 10.3 53 2.0 w .,J:- Longnose sucker 0.1 4 1.3 2 3.6 7 0.3 Trout-perch 6 0.9 0.1 7 0.3 Burbot 3 0.3 3 D.l Brook stickleback 56 17.3 13 4.3 15 100.0 75 63.0 14 73.7 25 44.6 5 20.0 34 87.2 71 98.6 308 11.8 51 imy scul pin 6 0.9 18 1.8 5 1.5 4 21.1 1.8 6 24.0 40 1.5 Northern pike 6 0.6 0.3 7 0.3 Fathead minnow 1 0.1 1 < D. 1 Lake whitefish 14 1.4 2 0.6 16 0.6 Yellow perch 27 4.1 6 0.6 33 1.3 Ninespine stickleback 0.1 1 < 0.1 Mountain whitefish 3 0.3 3 D.l Totals 662 987 323 302 15 119 19 56 25 39 72 2619 35 species seems to be most abundant at Sites 2 and 3. In 1976, chub were taken in large numbers at Site 7 of Hartley Creek (Bond and Machniak 1977). Longnose dace were captured as far upstream as Site 3 and one was reported from Site 7 of Hartley Creek in 1976 (Bond and Machniak 1977). However, this species appears to be most abundant in the lower reaches of the watershed (Sites and 2) as is also the sl imy sculpin (Sites 1, 2, and 3).

5.3 TAGGING RESULTS

5.3. 1 Tag Releases and Recaptures F10y tags were appl ied to 1629 fish during 1977, bringing to 3898 the total number of fish tagged over two years (Table 7). The majority of fish tagged were longnose suckers (51.9%), white suckers (42.8%), and northern pike (4.9%). Fish were tagged during both the upstream and downstream runs. Recaptures at the downstream trap in 1976 provided an indication of the length of time spent by some individual fish in the Muskeg River watershed (Bond and Machniak 1977). Recaptures of 1976 tags at the fence site in 1977 demonstrated a homing tendency in both white and longnose suckers. Considering only fish that were tagged at the fence site and recaptured outside the Muskeg River watershed, 77 recaptures have been reported for a tag return rate to date of 2.0% (Table 7). The highest recapture rates obtained outside the watershed were for northern pike (14.1%). White and longnose suckers had recapture rates of 1.9 and 0.8% respectively. In addition to the Floy tags mentioned above, metal cl ip tags were applied to 40 Arctic grayling in 1977, of which one has been recaptured.

5.3.2 Movement of Tagged Fish The recapture of tagged fish can provide useful infor­ mation concerning the extent and timing of fish movements. However, a degree of caution usually must be exercised in the interpretation of the results. In the first place, one can never 36

Table 7. Summary of tag releases and recaptures by species for fish tagged at Muskeg River counting fence, 1976 and 1977, and recaptured outside the Muskeg River watershed.

Number Tagged Percent of Number Percent Species Total Number Recaptured Recaptured 1976 1977 Tagged

White sucker 876 793 42.8 32 1.9 Longnose sucker 1267 757 51.9 17 0.8 Northern pike 119 73 4.9 27 14. 1 Arctic grayl ing 3 2 O. 1 20.0 Walleye 4 4 0.2 o 0.0

Total 2269 1629 100.0 77 2.0 37

be absolutely certain that the movement exhibited by an individual fish is representative of all fish in the population. Secondly, since it is obvious that no tags will be recovered from areas where no fishing effort occurs, it can be argued that recaptures serve merely to identify fishing areas. There is no question that, in the AOSERP study area, considerably more fishing effort is expended downstream from Fort McMurray than upstream. As well, in some cases, low recovery rates make it impossible to form firm conclusions as to general movement trends. Nevertheless, results from the present and several other recent studies (Machniak and Bond in prep.; Bond and Berry in prep.a, in prep.b; Machniak et a1. in prep.; Jones et a1. 1978; Kristensen and Pidge 1977) are beginning to identify patterns of fish movements within the AOSERP study area.

5.3.2.1 White suckers. Floy tags were applied to 1669 white suckers in the Muskeg River during 1976 and 1977, of which 32 have been recaptured outside the Muskeg River watershed (Appendix 8.2) (Bond and Machniak 1977). Of this number, 12 fish were recaptured in the lower Athabasca River or Lake Athabasca. Only three white suckers were recaptured upstream of the Muskeg River, none of which was taken upstream of the Steepbank River. One fish, tagged 20 May 1976 as it left the Muskeg River, was recaptured at the Muskeg River upstream trap on 8 May 1977 and subsequently in the Athabasca delta on 21 June 1977. Another was recaptured at the Muskeg River downstream trap on 27 May 1977, 16 days after it had entered the tributary. This fish was recaptured again on 15 May 1978 at the upstream fence of the MacKay River (Machniak et ale in prep.), and in June 1978, it was recaptured in Lake Athabasca at the mouth of the Athabasca River. Four other white suckers, tagged in the Muskeg River in 1977, were also recaptured in the MacKay River upstream trap in 1978. Another fish, tagged in May 1976 in the Muskeg River had been at large for 724 days when it was recaptured at the MacKay River trap on 14 May 1978. One Muskeg River fish was recaptured in the Steepbank River. This fish, tagged 16 July 1976, was recap­ tured moving upstream in the Steepbank River on 4 May 1977. A 38 total of 176 white suckers, tagged in 1976, were recaptured in the Muskeg river during 1977. Tag return evidence from this and other studies (Bond and Berry in prep.b; Machniak and Bond in prep~; Shell Canada Ltd, 1975: Machniak et al, in prep.) suggests that white suckers that spawn in the Muskeg River and other tributaries of the AOSERP study area belong to the Lake Athabasca population and return to the lake during summer or fall to overwinter. There is also an indication of a strong homing tendency on the part of this species although some individuals apparently enter other tributaries.

5.3.2.2 Longnose suckers. A total of 2204 longnose suckers were tagged in the Muskeg River during 1976 and 1977, of which 17 have been recaptured outside the Muskeg River watershed (Appendix 8.2) (Bond and Machniak 1977). Three fish, tagged in May 1977, were recaptured in Lake Athabasca between 31 May and 22 June, indicating a rapid downstream movement of from 264 to 296 km. One sucker, tagged 13 June 1976, was recaptured at the Muskeg River fence on 9 June 1977 and was recaptured again at the mouth of Clark Creek (km 11) in the Athabasca River on 23 June. Another, tagged 29 May 1976, was observed spawning in the lower reaches of Beaver Creek on 14 May 1977 (D. Tripp, Fishery Biologist, Aquatic Environments Ltd. verbal communication with W. A. Bond, June 1977). Nine 10ngnose suckers, tagged in the Muskeg River in 1976, were recap­ tured in May 1977 at the Steepbank River counting fence while one fish, tagged 18 May 1977, was recaptured on 1 May 1978 at the

Mac~y River upstream trap. A total of 260 10ngnose suckers, tagged in 1976, were recaptured in the Muskeg River during 1977. Tag return evidence from this and other studies (Bond and Berry in prep.b; Machniak and Bond in prep.; Machniak et al. in prep.) suggests that longnose suckers that spawn in the Muskeg River and other tributaries of the AOSERP study area belong to the Lake Athabasca population and return to the lake during summer or fall to overwinter. There is also an indication of a strong homing tendency in this species although some individuals apparently enter other tributaries. 39

5.3.2.3 Northern ike. Fourteen percent of all pike tagged in the Muskeg River in 1976 and 1977 have been recaptured outside the Muskeg River watershed or near the tributary mouth (Appendix 8.2) ~ond and Machniak 1977). Although Bond and Machniak 0977) noted that one pike moved 72 km between tagging and recapture. pike in general demonstrated little tendency to move around and most were recap­ tured within 15 to 20 km of the tagging site. Bond and Berry (in . prep.a, in prep.b) and Machniak and Bond (in prep.) reported similar results. Pike in the AOSERP study area appear to concentrate in the lower reaches of tributary streams during the summer and to move up and down the tributaries to some extent. They probably leave the tributaries in late summer or fal 1 to overwinter in the Athabasca Rive r.

5.3.2.4 Arctic gray] ing. Floy tags were appled to only five Arctic grayling, one of which has been recaptured (Appendix 8.2). This fish, tagged in the Muskeg River on 30 April 1976, was recap­ tured 10 October 1977 moving downstream in the Steepbank River. It had been at large for 528 days at the time of its recapture. One grayling, tagged with a metal cl ip at the upstream trap in May 1977, was recaptured near smal 1 fish collection site 4 during August 1977.

5.4 LIFE HISTORIES OF FISH SPECIES

5.4. 1 White Suckers

5.4.1 . 1 Seasonal timing of upstream migration. White sucker spawning migrations appear to be initiated by increasing water temperatures fol lowing spring break-up, and often begin when the daily maximum water temperature in the spawning stream approaches 10°C (Geen et al. 1966; Bond 1972). Whi te suckers were present in small numbers and moving upstream in the Muskeg River on 28 April 1977, on which date the maximum daily water temperature was 9°C. Water temperature decreased to 5°C during the next two 40 days and few fish entered the trap. The number of migrant suckers then increased daily from 1 to 6 Mayas the water temperature rose steadily (Table 8, Figure 10). The main part of the migration occurred between 7 and 9 May, the first three days on which the maximum daily water temperature exceeded 10°C. On these days, 45.9% of the total migration passed through the upstream trap. The upstream run was essentially complete by 12 May although small numbers of fish continued to move up after this date. The 1977 white sucker run into the Muskeg River fol lowed by several days the migration into the adjacent Steepbank River, but the pattern was similar in both cases. The Steepbank River first reached 10°C on 1 May and the peak of the upstream white sucker run occurred between 2 and 4 May (Machniak and Bond in prep.).

5.4.1.2 Diel timing of upstream migration. Bond and Machniak (1977) reported that, in 1976, most white suckers migrated into the Muskeg River between noon and midnight. They observed that maximum movement occurred in the late afternoon and early evening, just following the time of highest daily water temperature. Geen et al. (1966) and Machniak and Bond (in prep.) reported similar results. A different pattern, however, was observed in the Muskeg River during 1977 as most fish moved upstream at night when the water temperature had dropped considerably below the da~ly maximum. The majority of fish (78%) moved upstream between 2100 and 1200 h (Table 9). Thus it is evident that the diel timing of white sucker migrations can vary considerably from year to year.

5.4.1.3 Spawning period. As wil 1 be discussed later, the majority of white suckers mov1ng upstream in the Muskeg River between 28 April and 5 May were immature fish. The main upstream migration of spawners commenced approximately 6 May. Most mature females observed at the fence site were not fully ripe (freely running eggs) until about 6 to 8 May. White suckers were observed spawning downstream of the counting fence during the second week of May. Eggs were collected in drift nets as early as 9 May 1977 and, while not confirmed, Table 8. Summary of fish enumerated during the counting fence operation in the Muskeg River, 1977.

Upstream Trap Downstream Trap

..c: ..!: (j) 0) C C r.r: Q Cil C C (/) (/) C L f.!'; I- U ._, o ~- u .-- (j) (jJ'*- o :~ '- Q) ro4- e C) 0; rv ..:-> 0,) Da i 1y e Q) (1) (J) .'...1 Q) Da i ly O'l ~ .;....: -~ -\..J >- -1-' (ll c: ~ en~ -1-' _"- -I-' >- 4-J (J) e .j...J Date e 'J .- U U n:; L~ :J .- Totals c u U ~\ ro \...~ :J .- Totals o ::J ..!: ::::; ~ L o ._- o..!: o :J .c :J \... L 0'- O..!: -.JVl :3 1/')

28 Apri 1 7 3 5 0 0 15 29 8 27 10 9 9 63 30 0 16 5 21 0 42 1 May 13 58 13 19 0 103 a 2 92 83 7 33 4 220 .j::- 3 64 170 9 53 3 299 4 241 184 15 39 3 482 258 221 10 24 1 5 514 Trap Closed 6 116 211 8 22 0 357 7 71 520 6 24 0 621 8 102 562 4 7 0 675 9 63 282 2 10 1 358 10 42 110 1 30 3 188a 11 49 187 1 29 4 271a 12 22 81 8 20 3 134 3 83 1 4 1 93 b 13 148 44 5 4 6 208a 8 68 0 1 0 77 14 42 30 5 9 2 88 24 46 1 5 0 77 b 15 33 32 4 3 0 72 67 87 2 1 0 158 b 16 9 1 0 0 0 9 10 3 0 2 3 18 17 3 8 2 2 1 16 35 10 2 2 1 51b 18 5 3 2 3 0 13 67 26 1 0 1 95 continued Table 8. Continued.

Upstream Trap Downstream Trap

..c ..c (l.) 0) c C I/) (l.) 0) C C I/) I/) C !... I/) C !... o !... !... u·- (l.) rol+- Da i 1y o !... "- U .- (l.) rol+- C (l.) (l.) (l.) ..c .j..J (l.) C (l.) (l.) (l.) ..c +-J (l.) Da il y Date O)~ +-J ~ +-J >- +-J (l.) C +-J Totals O)~ +-J ~ +-J >- +-J (l.) C +-J Totals C u .- u u ro !...~ ::J .- C U U u ro !...~ ::J .- o ::J ..c ::J !... !... 0·- o..c o ::J ..c ::J !... !... o .- o..c ....JI/) :3 I/)

19 May 17 12 4 0 1 34 69 26 0 a 2 2 99 20 28 13 0 3 .1 46 73 58 0 1 1 133 21 16 14 4 0 1 35 6 13 0 1 0 21b 22 17 8 6 4 3 39a 114 91 0 2 1 207 23 15 13 3 3 0 34 11 0 29 2 0 42 ..t:"" 24 22 7 0 3 0 32 41 62 0 2 0 105 N 25 13 4 3 3 1 24 45 66 0 2 0 115 b 26 20 5 2 2 0 29 24 23 0 3 0 50 27 8 8 3 4 0 23 30 15 0 1 0 47b 28 2 9 2 0 2 15 28 29 1 1 1 60 29 0 3 0 1 0 4 20 6 1 0 0 27 30 13 7 4 1 0 25 26 66 0 0 0 92 31 2 6 0 6 1 15 11 76 1 3 0 91 1 June 27 9 0 3 2 41 54 51 0 1 0 106 2 11 0 3 9 0 23 7 11 0 4 0 23 b a 3 5 4 0 3 0 13 13 20 0 3 0 37 b 4 8 2 3 5 1 20a 22 20 0 2 0 45 b a 5 13 2 1 5 1 23 14 20 0 0 0 34 a 6 1 0 1 2 1 6 42 31 1 0 1 75 7 5 1 0 3 0 lOa 0 12 0 0 1 13 8 1 2 0 3 0 6 23 38 0 4 2 67 cont i nued Table 8. Concluded.

Ups t ream Trap Downstream Trap

u ..c ..c - .j...J - .j...J

9 June 3 0 0 1 1 5 11 45 0 2 1 59 10 1 2 0 2 0 6a 14 70 0 2 0 86 11 0 2 0 0 0 2 20 58 0 1 1 80 12 1 0 0 3 0 4 9 63 0 1 0 73 13 1 2 0 1 0 4 14 29 0 3 0 46 .::- 14 2 0 0 2 0 4 17 28 0 1 1 47 w 15 1 2 0 0 1 4 14 24 0 0 0 38 Total 1641 2970 161 433 57 5275 1004 1385 11 59 1 7 2487 % 31.1 56.3 3. 1 8.2 1 . 1 40.4 55.7 0.4 2.4 0.7 a Other species counted through upstream trap: one lake cisco, 2 May; one burbot, 13 May; eight walleye, 10 May (two fish), 11 May, 20 May, 22 May, 3 June, 4 June and 10 June; three Dolly Varden, 5 June~ 6 June and 7 June.

b Other species counted through downstream trap: five wa 11 eye t 12 May, 17 May, 21 May, 3 June and 4 June; and six lake whitefish, 14 May, 15 May, 25 May (two fish), 27 May and 2 June. c Numbers shown for mountain whitefish between 28 April and 11 May probably include a few lake whitefish that were erroneously identified. WHITE SUCKER

500

400

10- ~ 300 ~u JO ~~ :J: Upstream -0 - CJ) 200 n=2970 LL °E D Males S~ LL ~ 0 EZI Females ,J::- 100 ,J::- a: \ i n~n [JAil w 5 m :E 0 10- :::> G) z 55 0E ~Q 45 ~- 100 ==Qj 35 oQ)0 > .-1 25 ~ ~ 28301 15 311 15 APR. MAY JUNE

Figure 10. Seasonal timing of the white sucker migration in the Muskeg River, 1977. 45

Table 9. Summary of diel timing of the upstream migration of white suckers in the Muskeg River, 1977. Fish that were counted at times other than those indicated were included in the next check period.

Number of Fish Counted at Each Check 0900 1500 1800 2100 2400 Oate (0300)a Total

28 Apri 1 NO NO NO 0 3 NO 3 29 16 NO 8 2 1 NO 27 30 NO 14 NO 1 1 NO 16 1 May NO 5 NO NO 10 43 58 2 21 0 3 3 19 37 83 3 98 0 7 1 0 64 170 4a 75 41 7 11 7 43 184 5a 125 NO 57 6 NO 33 221 6a 79 25 1 18 6 82 211 7a 152 126 37 51 37 117 520 8a 128 187 87 57 42 61 562 9a 106 85 11 10 42 28 282 lOa 27 20 1 8 13 41 110 11 a 51 18 25 10 8 75 187 12a 50 9 NO 20 2 trap closed 81 13 NO 21 NO 7 NO 16 44 14 NO 23 NO 1 1 5 30 15 NO 27 NO 0 0 5 32 16 NO 1 NO 0 NO 0 1 17 NO 6 NO NO 1 1 8 18 NO 2 NO NO 0 1 3 19 NO 2 NO 0 1 9 12 20 NO 0 NO NO 3 10 13 21 NO 4 NO NO 0 10 14 22 NO 0 NO 0 2 6 8 23 NO 9 NO 2 NO 2 13 24 NO 1 NO 0 1 5 7 25 NO 0 NO 0 NO 4 4 26 NO 2 NO NO NO 3 5 27 NO 3 NO NO 2 3 8 28 NO 2 NO 1 NO 6 9 29 NO 3 NO NO NO NO 3 30 1 NO 2 NO NO 4 7 31 NO 0 NO NO 4 2 6 1 Junea 3 2 NO 1 NO 3 9 2 NO 0 NO NO NO 0 0 3 NO 0 NO 2 NO 2 4 4 NO 0 NO 1 NO 1 2 5 NO 1 NO 1 NO 0 2 6 NO 0 NO 0 NO 0 0 7 NO 0 NO 0 NO 1 1 8 NO 0 NO 0 NO 2 2 9 NO 0 NO 0 0 0 0 10 NO 0 0 0 0 2 2 11 NO 0 NO 0 NO 2 2 12 NO 0 0 NO 0 0 0 13 NO 1 0 0 0 1 2 14 NO 0 NO NO 0 0 0 15 NO 2 ooerations terminated 2

Totals 932 642 246 214 206 730 2970 % Grand Total 31.4 21.6 8.3 7.2 6.9 24.6

a Checks were made at 0300 h rather than 0900 h during the peak of the runs. 46 these were thought to be white sucker eggs. The first spent white suckers were taken at the downstream trap on 12 May and by 18 May virtually all fish were spawned out. The white sucker spawning period in the Muskeg River in 1977 was almost identical to that observed in 1976 (Bond and Machniak 1977). However, as the initiation of white sucker spawning migrations appears to be closely related to stream temperature (Geen et al. 1966; Tremblay 1962), the precise timing of this event can be expected to vary considerably from year to year.

Spawning areas. White suckers have been reported to spawn in a variety of habitats, including lake margins and quiet reaches in the mouths of streams (Scott and Crossman 1973). However, optimal conditions probably involve shallow water running over a gravel substrate (Geen 1958). Bond (1972) suggested that the presence of deep pools adjacent to the spawning sites may also be an important factor. Within the lower 35 km of the Muskeg River system there are many areas that appear to satisfy these conditions. As in 1976 (Bond and Machniak 1977), white suckers were observed spawning below the fence site during the second week of May 1977e Although spawning was not seen upstream of the fence, young-of-the-year suckers were abundant at Sites 3 and 4 (Figure 2) by mid-June. Few sucker fry were captured in Hartley Creek either in 1976 or 1977 although some spawning probably occurs in that tributary downstream of Site 8. Only one young-of-the-year sucker was captured from the Muskeg River upstream of Site 4 during the two years of this study. This fish was taken at Site 6 (Figure 2) on 16 August 1977.

5.4.105 Length of time spent in the Muskeg River. By 15 June 1977, when trap operations ceased, only 46.6% of the white suckers counted through the upstream trap had returned downstream. The downstream migration observed clearly represented the departure of spawners from the tributary and began approximately one week 47 following the passage of this group through the upstream trap (see Section 5.4.1.8). On the other hand, fewer than 10% of the immature migrants «350 mm) had returned downstream as of 15 June. No analysis of the 1977 tag data was performed to indicate the length of time spent in the tributary by individual fish. However, the 1976 data indicated that this time varied considerably (from three to 84 days) for fish that had left the Muskeg River by 30 July. On that date, 40.9% of the white suckers enumerated at the upstream trap sti 11 remained in the tributary (Bond and Machniak 1977). Many immature white suckers may remain in the Muskeg River until freeze-up as was the case in the Steepbank River (Machniak and Bond in prep.).

5.4.1 .6 Seasonal and diel timing of downstream migration. The first spent fish were observed upstream of the counting fence on 12 May 1977, on which date the downstream trap was opened. White suckers moved downstream from 12 May through 15 June, the final day of trap operations (Table 8 and Figure 10). While the number of fish passing through the downstream trap varied each day, the downstream run was not characterized by a discrete peak. Bond and Machniak (1977) reported that, after a definite peak between 15 and 20 May, white suckers continued to pass downstream through 30 July. The majority of downstream migrants were captured at night as 71.2% were taken between 2100 and 1200 h (Table 10). A similar timing of downstream movement was observed in the Steepbank River (Machniak and Bond in prep.). The maximum movement of white suckers occurred each day following the period of highest water temperature. Bond (1972) noted that the down­ stream migration usually occurred when stream temperatures were decreasing.

5.4.1.7 Spawning mortal ity. Only a few white suckers were found dead prior to the termination of fence operations on 15 June 1977. Results in 1976, however, indicated that the number of mortalities increased and the general condition of the fish 48

Table 10 0 Summary of diel timing of the downstream migration of white suckers in the Muskeg River, 1977. Fish that were counted at times other than those indicated were included in the next check period.

Number of Fish Counted at Each Trap Check Oate Total 1200 1800 2100 2400

12 May trap opened 18 4 61 83 13 3 31 NO 34 68 14 7 22 0 17 46 15 4 43 10 30 87 16 2 0 NO 1 3 17 0 NO 3 7 10 18 7 NO 2 17 26 19 5 6 0 15 26 20 6 NO 6 46 58 21 1 NO 3 9 13 22 2 41 18 30 91 23 1 2 NO 8 11 24 0 2 7 53 62 25 49 17 NO trap closed 66 26 18 NO NO 5 23 27 5 NO 7 3 15 28 29 0 NO 0 29 29 6 ND NO NO 6 30 2 15 NO 49 66 31 44 NO 11 21 76 1 June 43 It NO 4 51 2 9 NO NO 2 11 3 5 2 ND 13 20 4 17 0 NO 3 20 5 4 3 NO 13 20 6 15 9 NO 7 31 7 7 1 NO 4 12 8 12 11 NO 15 38 9 14 11 8 12 45 10 33 19 5 13 70 1 1 24 22 NO 12 58 12 34 7 9 13 63 13 9 12 2 6 29 14 11 ND 7 10 28 15 24 opera t ions term ina ted 24 Total 452 298 102 533 1385 % Grand 32.7 7.4 Total 21.5 38.5 49 decreased between 18 June and 30 July (Bond and Machniak 1977). Fish taken at that time were often blind in one or both eyes, displayed signs of physical deterioration, and were heavi ly infested with the parasitic copepod Argulus sp. A mortality rate of 16 to 20% was observed by Geen et al ~ (1966) for spawning white suckers in Frye Creek, British Columbia.

5.4.1.8 Size ition of mi rant white suckers Fork lengths were determined for 155] white suckers during the upstream migration in 1977 (Table 11, Figure 11). Migrant suckers ranged in fork length from 157 to 599 mm, but the length-frequency distribution varied considerably as the migration proceeded. The early stages of the upstream migration (28 April to 3 May) were dominated by fish in the 180 to 280 mm fork length range (Figure 12). Fish of this size remained abundant on 4 and 5 May, but at that time a second group of migrants appeared whose fork lengths ranged from about 300 to 400 mm. The large group of immature fish comprising the smaller mode either did not occur in the Muskeg River in 1976 or it had already passed upstream by the time that fence operation began. The middle mode in the length frequency distribution (Figure 11) consists largely of maturing fish. A certain proportion of these fish probably spawned for the first time in 1977, although most were likely non-spawners. Fish in this size range dominated the Muskeg River white sucker run in 1976 (Bond and Machniak 1977) and comprised the vast majority of the 1977 run in the Steepbank River (Machnlak and Bond in prep.). In both streams, these immature fish were proceeding upstream while maximum daily water temperatures ranged from 5 to 9°C. Between 6 and 10 May 1977, the migration was dominated by large fish ranging in fork length from about 400 to 600 mm (Figure 12). This segment is believed to have comprised the main spawning group of white suckers in the Muskeg River in 1977 and was also well represented in the 1976 run. Within this mode, but in neither of the other two, females were clearly larger than males (Figure 11). Interestingly, this large mode did not appear in the Steepbank run in 1977 (Machniak and Bond in prep.). Table 11. Length frequency distribution of white suckers during the upstream migration in the Muskeg River, 1977.

Fork Length Male Female Unknown Total Fork Length Male Female Unknown Total (10 mm intervals) (10 mm intervals)

150 - 159 1 0 0 1 390 - 399 11 8 8 27 160 - 169 0 0 0 0 400 - 409 12 10 2 24 170 - 179 1 1 7 9 410 - 419 9 15 6 30 180 - 189 2 1 9 12 420 - 429 7 14 2 23 190 - 199 7 4 23 34 430 - 439 22 15 4 41 200 - 209 8 2 28 38 440 - 449 24 16 3 43 210 - 219 15 4 41 450 - 459 19 5 2 26 220 - 229 16 4 53 73 460 - 469 30 9 1 40 230 - 239 17 49 69 470 - 479 31 21 1 53 3 V'1 240 - 249 11 1 43 55 480 - 489 25 18 0 43 0 250 - 259 12 2 52 66 490 - 499 20 20 1 41 260 - 269 14 2 45 61 500 - 509 16 20 0 36 270 - 279 4 0 25 29 510 - 519 4 23 0 27 280 - 289 7 0 19 26 520 - 529 2 32 0 34 290 - 299 1 7 0 17 34 530 - 539 1 22 0 23 300 - 309 5 2 29 36 540 - 549 0 11 0 11 310 - 319 8 5 26 39 550 - 559 0 12 0 12 320 - 329 5 4 37 46 560 - 569 0 10 0 10 330 - 339 19 11 36 66 570 - 579 0 1 0 1 340 - 349 14 4 42 60 580 - 589 0 0 1 350 - 359 15 15 36 66 590 - 599 0 0 1 360 - 369 17 11 19 47 370 379 17 9 21 47 Totals 474 380 697 1551 380 - 389 9 1 1 10 30 150l WHITE SUCKERS ... All Fish n 1551 L::,. Males n= 474 125l / \ (\ o Females n=380 V5 100 LL LL 0 75 I \ r j \ V1 a:: W £D ~ 50 ::::> Z

25

200 300 400 500 600 FORK LENGTH (mm)

Figure 11. Length-frequency distribution for white suckers during the upstream migration in the Muskeg River, 1977. 52

20 28 Apr-3 May n=307 15 10 5

15 11-15 May n= 279 10 5

200 300 400 500 600 FORK LENGTH (mm)

Figure 12. Seasonal changes in length-frequency distribution for white suckers during the upstream migration in the Muskeg River, 1977. 53

Fork lengths were obtained from 1050 white suckers during the downstream migration, of which approximately 95% were longer than 350 mm. Most of the downstream fish not measured (n 335) had been tagged previously, and most of these fish also exceeded 350 mm fork length. The length-frequency distribution of downstream migrants remained constant from 12 May to 15 June (Figure 13). The virtual absence of small white suckers from our downstream counts suggests that immature fish tend to remain in the tributary longer than spawners. This is supported by evidence from the adjacent Steepbank River. In that study, spawners also left the stream first, whi le fish that remained in the tributary through the summer tended to be small individuals. Eighty-five percent of the white suckers captured during the fall fence operation in September and October were less than 350 mm in length (Machniak and Bond in prep.).

5.4.1 .9 Age composition of migrant white suckers. Because our age sample was not drawn randomly, it may not reflect accurately the age composition of the white sucker migration. However, the data do ill ustrate the age range of mi grant suckers and our knowledge of the age and growth characteristics of this population (presented in a later section), combined with the length-frequency data (Table 11, Figure 11) permit a fairly accurate description of the age composition of these fish. White suckers in the run ranged in age from three to 16 years (Figure 14). The early part of the migration was dominated by young fish (age 3 and 4) but the age composition shifted toward older age groups as the migration progressed. The main spawning group (>400 mm fork length) consisted largely of fish age 7 and older with most spawners belonging to age groups 8 to 12 inclusive.

5.4.1.10 Sex was deter- mined for 1850 white suckers during the upstream migration, of which 1014 (54.8%) were males (Table 12). This represents a significant deviation from the usual 1:1 ratio (X2 =17.2; P

15 12May-22May n =413 10

5

23May-3June I-- 15 Z n=336 w u 10 0:: w a.. 5

15 4-15 June n=301 10

5

200 300 400 500 600 FORK LENGTH (mm)

Figure 13. Length-frequency distribution for white suckers during three time periods in the downstream migration in the Muskeg River, 1977. 20 I I I I i , WHITE SUCKERS I 10 Males n=123 I en I D Females n = 148 I LL LL V'l ,] V'l 0 Jl a::: w £D ~ :::> z 5

po 'I3 4 5 6 7 8 "I-P--h15 16 FIN RAY AGE (YEARS)

Figure 14. Age composition for white suckers sampled during the counting fence operation, Muskeg River, 1977. 56

Table 12. Sex ratio for white suckers during the upstream migration, Muskeg River, 1977.

Number of Fish Percent Date Malesa Males Females Unknown Total

28 Apri 1 0 0 3 3 0 29 1 3 23 27 25 30 9 1 6 16 90 1 May 14 12 32 58 54 2 18 13 52 83 58 3 41 22 107 170 65 4 50 30 104 184 63 5 60 40 121 221 60 6 95 64 52 11 60 7 251 185 84 520 58 8 269 201 92 562 57 9 44 183 55 282 19 10 27 42 41 110 39 11 53 26 108 187 67 12 25 9 47 81 74 13 9 2 33 44 82 14 6 0 24 30 100 15 7 1 24 32 88 16 0 0 1 1 0 17 2 0 6 8 100 18 0 0 3 3 0 19 1 0 11 12 100 20 6 0 7 13 100 21 3 0 11 14 100 22 2 0 6 8 100 23 3 0 10 13 100 24 0 0 7 7 0 25 0 0 4 4 0 26 0 0 5 5 0 27 4 0 4 8 100 28 5 0 4 9 100 29 2 0 1 100 30 1 0 6 7 100 31 0 0 6 6 0 1 June 2 2 5 9 50 2 0 0 0 0 0 3 1 0 3 4 100 4 2 0 0 2 100 5 0 0 2 2 0 6 0 0 0 0 0 7 0 0 1 1 0 8 0 0 2 2 0 9 0 0 0 0 0 10 0 0 2 2 0 11 1 0 1 2 100 12 0 0 0 0 0 13 0 0 2 2 0 14 0 0 0 0 0 15 0 0 2 2 0

Totals 1014 836 1120 2970 % 55 45

a Based on fish of known sex. 57

Male white suckers usually precede the females onto the spawning grounds (Geen et al. 1966; Bond 1972; Bond and Machniak 1977). However, because of the large number of immature fish in the 1977 Muskeg River migration this trend is not immediately obvious. Among the fish captured between 28 April and 4 May (rrost1'y immatures), males outnumbered females every day. During the main spawning run, however, which occurred from 6 to 10 May, males outnumbered females on the first three days while females were dominant on 9 and 10 May. The ratio of males to females in the downstream run showed no clear pattern.

5.4.1.11 Homing of white suckers. Tagging studies by several authors (Olsen and Scidmore 1963; Geen et al. 1966) have indicated a tendency on the part of white suckers to return to the same spawning stream each year in preference to other streams that might be available. During 1977, clear evidence was produced to indicate that white suckers in the AOSERP study area behave in a similar manner. If, as we suspect, Muskeg River white suckers are part of the Lake Athabasca population, these fish are performing in excess of a 500 km round trip to return to this stream. During the 1976 study (Bond and Machniak 1977), Floy tags were appl ied to 876 white suckers in the Muskeg River. Twenty­ one tagged fish are known to have been dead prior to the 1977 migration, but of the remainder, 20.6% were recaptured in the Muskeg River during the 1977 study. White suckers demonstrated considerable fidelity to the Muskeg River. The counting fence operation on the Steepbank River, for instance, recovered only one tagged white sucker from the 1976 study. McCart et al. (in prep.) in a 1977 study of the MacKay River, did not recover any tagged white suckers from the Muskeg River although five were recorded in a counting fence operation on this tributary in 1978 (Appendix 8.2).

5.4.1.12 Fecundity. Fecundity was estimated gravimetrically for 10 female white suckers from the Muskeg River. The data in 58

Table 13 represent additions to those given by Bond and Machniak (1977). Considering the fecundity data for both years, the esti- mated total number of eggs per female (fork length 397 to 525 mm) ranged from 21 402 to 64 175 with a mear of 42 729 ova per female. Actual counts on five ovaries revealed errors of from +3.2% to -0.5% for the estimated values. Bond (1972) reported white sucker fecundity ranging from 15 983 to 60 242 with an average of 34 502 per female. Length-relative fecundity for white suckers ranged from 539.1 to 1222.4 ova per em of fork length while weight­ relative fecundity varied from 22.7 to 41.1 eggs per g of body weight. The right ovary contained more eggs than the left in nine out of 10 cases. Regression analysis indicated a significant (p< 0.01), positive correlation between fecundity and fork length (n = 10; r=0.887). The mathematical relationship between fecundity and fork length for Muskeg River white suckers is expressed by the equat ion: 10gloFecundity 2.9541 og loFo rk Length (mm) + 3.260;

sb 0.543 Fecundity also correlated positively with body weight (r=0.866, range 800 to 2680 g). The mathematical relationship between fecundity and body weight is described by the equation:

logloFecundity = 1.182log 10Body Weight (g) + 2.280; sb 0.241

5.4.1.13 Age and growth. Age and growth results from 1977 (Tables 14 and 15) were simi lar to those of 1976 (Bond and Machniak 1977)(Figure 15). Muskeg River suckers grew more slowly than those from George Lake, Ontario (Beamish 1970) but faster than those in the Bigoray River, Alberta (Bond 1972). Suckers from Lake, Wisconsin (Spoor 1938) grew more rapidly than Muskeg River suckers during their first few years but more s10wly after age four (Figure 15). 59

Table 13. Fecundity estimates for white suckers sampled during the 1977 Muskeg River spawning migration.

Number of Eggs Relative Fork Weight Fecundity Length (g) Left Right (mm) Ovary Ovary Total (cm) (g)

495 2000 19 268 26 200 45 468 918.6 22.7

497 1820 20 695 23 316 44 011 885.5 24.2

525 2680 31 609a 32 566 64 175 1222.4 23.9 (+3.2%)b a Actual egg counts. b Deviation of estimated counts from actual number. Table 14. Age-length relationship (derived from fin rays and oto1iths) for white suckers captured in the Muskeg River watershed, 1977, sexes separate and combined sample (includes unsexed fish).

Males Females All Fish Age t-test N Mean S.D. Range N Mean S.D. Range N Mean S.D. Range

1 17 49.8 10.63 38-82 15 50.5 9.36 35-75 43 48.7 9.30 35-82 0.196 2 0 1 107.0 1 107.0 3 5 179.4 13.74 157-193 3 187.7 11. 175-197 12 184.8 11 .88 157-197 0.875 4 19 232.4 28.96 177-293 21 234.1 34.61 182-264 48 230.4 30.23 177-293 o. 168 5 8 313.0 24.83 268-343 6 304.7 54.28 237-381 17 309.8 42. 14 237-381 0.386 6 3 346.0 67.67 296-423 10 386.2 47.10 315-447 13 376.9 52.32 296-447 1 . 187 7 8 411 .6 48.48 347-442 7 406.3 14.59 382-422 15 409.1 35.69 347-442 0.277 8 11 434.7 37.23 366-494 13 452.6 33.75 375-508 24 444.4 35.77 366-508 1 .236 0 9 15 457. 1 29.31 409-495 15 480.9 23.16 437-530 30 469.0 28.64 409-530 2.467a '" 10 9 463.8 27.73 408-498 16 478.4 35.46 415-527 25 473.2 33.07 408-527 1 .062 11 14 467.4 28.42 425-514 20 498.4 30.84 424-547 34 485.6 33.25 424-547 2.976a 12 11 468.4 24.70 423-505 6 524.7 11 .08 510-540 17 488.2 34.48 423-540 5.242a 13 1 509.0 5 510.6 38.71 443-539 6 510.3 34.63 443-539 14 2 465.0 12.73 456-474 7 530.9 18.78 504-562 9 516.2 33.59 456-562 15 0 1 525.0 1 525.0 16 0 2 526.5 2.12 525-528 2 526.5 2.12 525-528 Totals 123 148 297 a Indicates significant difference between means for males and females (Studentls t-test, P< 0.05). Table 15. Age-weight relationship for white suckers captured in the Muskeg River watershed, 1977, sexes separate and combined sample (includes unsexed fish).

Males Females All Fi sh Age t-test N Mean S.D. Range N Mean S.D. Range N Mean S.D. Range

1 17 1 .59 1 .48 0.5-6.8 15 1.47 0.98 0.3-4.5 43 1. 38 1. 13 0.3-6.8 0.136 2 0 1 13.3 1 13.3 3 5 68.0 17.89 60-100 3 80.0 20.0 60-100 12 74.6 17.25 60-100 0.882 4 19 147.9 75.76 60-300 21 165.7 103.36 75-220 48 150.8 84.47 60-300 0.616 5 8 385.0 124.10 240-520 6 393.3 231 . 14 150-780 17 396.5 173. 13 150-780 0.087 6 3 610.0 334.51 360-990 10 865.0 371 .99 420-1600 13 806.2 367.34 360-1600 1 .060 7 8 1018.8 390.33 520-1360 7 874.3 126.34 700-1060 15 951 .3 297.63 520-1360 0.934 8 1 1 1270.0 363.76 710-1660 13 1430.8 433.62 720-2280 24 1357. 1 402.90 710-2280 0.973 9 15 1525.3 410.12 1040-2400 15 1564.0 179.91 1280-1780 30 1544.7 311 . 79 1040-2400 0.335 -'" 10 9 1488.9 311 .27 940-1820 16 1603.8 385.88 890-2100 25 1562.4 358.51 890-2100 0.762 11 14 1587.9 477.71 1060-2870 20 1935.5 481 .54 790-2920 34 1792.4 503.56 790-2920 2.078a 12 11 1536.4 282.60 1150-1980 6 2055.0 268.46 1780-2540 17 1719.4 371.09 1150-2540 3.676a 13 1 1914.0 5 1794.0 347.10 1200-2100 6 1814.0 314.30 1200-2100 14 2 1460.0 113. 14 1 380-1 540 7 2211 .4 120.06 2020-2360 9 2044.4 349.58 1380-2360 15 0 1 1880.0 1 1880.0 16 0 2 2410.0 381 .84 2140-2680 2 2410.0 381 .84 2140-2680

Total 123 148 297 a Indicates significant differences between means for males and females (Student's t-test, P< 0.05). 600

E 500 ., E i ::c ///////lrt~tJ-1·:t-r t- 400 (!) Z W ,/ T l f .-J ),//_3 300 ~ c:::: i 0 LL i 200 ;" • Muskeg R 1976 '"N o Muskeg R 1977 100 i",,/ .,," ~,/ 7 8 9 10 II AGE (YEARS)

Figure 15. Growth in fork length for white suckers from the Muskeg River and from several other areas: 1 George Lake, Onto (Beamish 1970); 2. Bigoray River, Alta. (Bond 1972); and 3. Muskellunge Lake, Wis. (Spoor 1938). Circles represent means and vertical 1 ines the ranges in fork length within age groups for Muskeg River fish. 63 Muskeg River suckers added length at a relatively constant rate during their first eight to 10 years, after which age little length increase occurred. Females were generally longer than males of the same age. Bond and Machniak (1977) found this difference between the sexes to be significant only at age 14 in 1976, but in 1977, significant differences occurred in age groups 9, 11, and 12 (Table 14). Female suckers also tended to be heavier than males of equal age but significant differences occurred only in age groups 11 and 12 during 1977 (Table 15). Females outnumbered males in age groups 13 to 16 in our 1977 sample, suggesting that they tend to 1 ive longer than males. Other inves­ tigators have also reported that female white suckers grow larger and live longer than males (Spoor 1938; Raney and Webster 1942; Smith 1952; Hayes 1956; Lalancette 1973). The maximum age for Muskeg River suckers was 17 years in 1976 (Bond and Machniak 1977) and 16 years in 1977. Maximum fin ray ages reported by Verdon (1977) were 19 to 25 years for white suckers in the James Bay area of Quebec.

5.4.1.14 Sex and maturity. Age and sex were determined for 271 white suckers in 1977, of which 148 (55%) were females (Table 16). The sexes were equally represented in the younger age classes. However, females made up 60% of all fish age 10 and older. The youngest mature white sucker observed in the Muskeg River was a three year old male captured in 1976 (Bond and Machniak 1977). The earliest age of maturity observed in 1977 was four years in both sexes. Spoor (1938) reported that, in Wisconsin, males matured at age 5 or 6 and females at age 6 or 7. Geen (1958) stated that, in British Columbia, white suckers do not spawn before age 6. Bond (1972) captured no spent suckers less than six years old in the Bigoray River, Alberta. Muskeg River data, collected over two years, show that 36, 56, 67, and 86% of male white suckers were mature at ages 5, 6, 7, and 8 respectively. For females at the same age, the corresponding values were 37, 58, 61 , and 88%. A major discrepancy appears to exist, however, when the maturity data for four year old fish are examined. Table 16 shows that, in 1977, 47% of male and 5% of female white suckers were 64

Table 16. Age-specific sex ratios and maturity for white suckers from the Muskeg River drainage, 1977~ Sex ratios were based only on fish for which sex was setermined.

Females Males Age Unsexed Total % % Fish N % Mature N % Mature

15 47 a 17 53 a 11 43

2 100 a a 0 a a 3 3 38 a 5 62 a 4 12 4 21 53 5 19 47 47 8 48 5 6 43 50 8 57 38 3 17 6 10 77 90 3 23 67 a 13 7 7 47 86 8 53 100 a 15 8 13 54 100 11 46 100 a 24 9 15 50 100 15 50 100 0 30- 10 16 64 100 9 36 100 a 25 11 20 59 100 14 41 100 a 34 12 6 35 100 11 65 100 a 17 13 5 83 100 17 100 a 6 14 7 78 100 2 22 100 a 9 15 100 100 a a a a

16 2 100 100 a 0 a a 2

Totals 148 55% 123 45% 26 297 65 mature at four years of age whi Ie in 1976, the corresponding figures are 77 and 30%. Such values for four year old suckers seem too high. Although some males in the 200 to 300 mm size range appeared to be ripe at the time of their upstream migration (i .e. they were running mi It and had developed small tubercles), the virtual absence of fish this size from the downstream results (Figure 13) suggests that most of these small fish did not spawn. During 1976 (Bond and Machniak 1977), a few white suckers with undeveloped ova were noted among the older age classes. The presence of such fish suggests that some white suckers do not spawn every year.

5.4.1.15 Length-weight relationship. The following length- weight relationships were determined from white suckers captured during the 1977 counting fence operation on the Muskeg River. Both upstream and downstream fish were included in the calculations. The mathematical relationship between fork length and bod y we i g h t for ma I e s uc ke r s (n = 106 , r = o. 993, ran g e 157 to 5 14 mm) is described by the equation: loglOW = 3.338 (lo9IOL)- 5.832; sb = 0.040 The equivalent expression for female white suckers (n = 134, r = o. 985, ran gel 75 to 562 mm) is: lo9lOW 3.177 (lo9IOL) - 5.316; sb = 0.048 Analysis of covariance indicated a significant differ­ ence (p > 0.05) between the slopes (F = 9.653), but not the adjusted means (F = 0.084) of the length-we i ght re 1at i onsh ips of ma 1e and female white suckers.

5.4.1.16 Growth of young-of-the-year. Information on first year growth of white suckers in the Muskeg River is presented by Bond and Machniak (1977). At age 1, white suckers ranged in fork length from 35 to 82 mm and weighed from 0.3 to 7.5 g. White suckers in the Bigoray River had a mean fork length of 42.2 mm and a mean weight of 0.89 g at the end of their first year (Bond 1972). 66

5.4.1.17 Food habits. Sixty-three white sucker stomachs were examined in the field during 1977 and most (93%) contained no food. Similar observations were recorded in 1976 (Bond and Machniak 1977). The stomachs of six adult white suckers examined in the

laboratory contained insect remains, Gastropoda, Pelecypoda 1 digested material, and debris (Table 17). Debris (sand) made up

48 0 9% of the total volume of material found in sucker stomachs. Bond (1972) found that adult suckers fed almost exclusively on immature insects. The stomachs of 14 young-of-the-year and juvenile white suckers contained mostly digested material' (Table 18). Small suckers in the Bigoray River (Bond 1972) fed mainly on chironomid larvae, small Crustacea, Rotifera, diatoms, and desmids.

5.4.1.18 Rearing area. Young-of-the-year suckers (two species) were first captured in the Muskeg River on 30 May 1977 By 15 June they could be found in large numbers from the mouth of the tribu­ tary to Site 4 (Figure 2) approximately 35 km upstream. Throughout this section of river these small fish were concentrated in small back eddies near shore. Although most young-of-the-year drifted out of the Muskeg River during June, July, and August, the entire lower section must be considered important in terms of rearing of this species. Young-of-the-year were still common at Site 3 in lake August although they were obviously less abundant than they had been earlier in the year.

5.4.1.19 Overwintering. While most young-of-the-year suckers leave the Muskeg River during their first summer, a small percent­ age probably remains in the tributary over the winter. Yearl ing white suckers were captured at Sites 7, 8, and 9 (Figure 2) in May and June 1977, and at the mouth of Kearl Creek (Site 9) in June 1976 (Bond and Machniak 1977). Small numbers of two and three year old white suckers may also overwinter in the Muskeg River drainage. Tagging results suggest that the larger and older fish overwinter in Lake Athabasca. Table 17. Food habits of adult longnose suckers, white suckers and lake whitefish captured from the Muskeg River, 1977.

Longnose Suckers White Suckers Lake Wh i tefi sh Food Items % Freq. a % No. % Vo 1. % Freq.Cl % No. % Vol. % Freq.a % No. % Vo 1 .

Class Insecta

Diptera S i mu 1 i i dae 1a rvae 16.7 74.9 6.5 0.0 0.0 0.0 0.0 0.0 0.0 Trichoptera 33.3 2.7 3.6 ·0.0 0.0 0.0 0.0 0.0 0.0 Plecoptera 0.0 0.0 0.0 0.0 0.0 0.0 100.0 50.0 57. 1 Ephemeroptera 0.0 0.0 0.0 0.0 0.0 0.0 100.0 50.0 42.9 0' Insect Remains 16.7 0.0 2.0 16.7 0.0 3.8 0.0 0.0 0.0 ""-..I

Miscellaneous

Gastropoda 0.0 0.0 0.0 33.3 4.9 7.6 0.0 0.0 0.0 Pelecypoda 33.3 22.5 80.0 50.0 24.4 19.8 0.0 0.0 0.0 Vegetat i on 33.3 0.0 4.0 0.0 0.0 0.0 0.0 0.0 0.0 Digested Matter 16.7 0.0 1 .9 50.0 0.0 19.8 0.0 0.0 0.0 Debris (gravel, sand) 16.7 0.0 1.9 66.7 70.7 48.9 0.0 0.0 0.0

Total Stomachs 7 6 Empty (% of Total) 14.3 0.0 a Percentage frequency of occurrence, based on stomachs that contained food. Table 18. Food habits of young-of-the-year and juven les of the larger species captured in the Muskeg River, 1977.

Species

Food Items White Suckers longnose Suckers Northern Pike Lake Wh i tef i sh Yellow Perch Burbot

% Frequencya % Frequencya % Freq!3 % No. Freq.a % No. % Freq.a J~ No. % Freq.a % No.

Class Insecta

Diptera Chironomidae 0.0 33.3 0.0 0.0 0.0 0.0 33.3 40.0 0.0 0.0 Unidentified Dipterans 0.0 66.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Trichoptera 0.0 0.0 0.0 0.0 50.0 50.0 33.3 10.0 0.0 0.0 Plecoptera 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ephemeroptera 0.0 0.0 40.0 44.4 0.0 0.0 66.7 50.0 100.0 100.0 Hemiptera 0.0 33.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Q"\ Insect Remains 10.0 0.0 20.0 0.0 25.0 0.0 0.0 0.0 0.0 0.0 00

Mi sce 1 J aneous

Nematomorpha 0.0 0,0 0.0 0,0 25.0 50.0 0.0 0.0 0.0 0.0 Fish Longnose suckers 0,0 0.0 20.0 22.2 0.0 0.0 0.0 0.0 0.0 0.0 White suckers 0.0 0.0 20.0 22.2 0.0 0.0 0,0 0.0 0.0 0.0 Cyprinids 0.0 0.0 .0 11. I 0.0 0.0 0.0 0.0 0.0 0.0 Digested Matter 80.0 66.7 0.0 0.0 50.0 0.0 0.0 0.0 100.0 0.0 Debris (sand, gravel) 20.0 0,0 0.0 0.0 0.0 0.0 0,0 0.0 0,0 0.0

Total stomachs 14 3 6 3 2 Empty (% of Total) 28,5 0.0 16,7 20.0 0.0 50.0 a Based on stomachs that contained food. 69

5.4.2 Longnose Suckers

5.4.2.1 Seasonal timing of upstream migration. Longnose sucker spawning migrations appear to be initiated by increasing water temperatures following the spring break-up. Geen et al. (1966) observed that the spawning migration was associated with a water temperature of 5°C in British Columbia. Bailey (1969) reported that, in the Brule River, Wisconsin, spawning runs (over a seven year period) peaked at an average water temperature of 13°C (range 10.9 to 14.4°C). Longnose suckers were moving upstream in the Muskeg River in late April 1976 when the daily maximum water temperature was 9.5°C (Bond and Machniak 1977). However, the most intensive portion of that run occurred on 9 and 10 May when daily maximum water temperatures were 12 and 14°C respectively. The 1977 Muskeg River longnose sucker run began on 2 May at a water temper­ ature of 7°C (Table 8, Figure 16). Most upstream movement (63.9%) took place between 2 and 10 May with peak migrations occurring on 4 and 5 May when daily maximum water temperatures were 9 and 9.5°C respectively. The longnose run in the Steepbank River commenced 25 April but the largest portion of the migration took place on 2 to 4 May when stream temperatures were between 10 and 12°C (Machniak and Bond in prep.). The 1977 longnose sucker run into the Muskeg River apparently involved considerably fewer fish (n= 1641) than did the 1976 run (n 2837). Since the ice is known to have left the Muskeg River between 20 and 22 April 1977, it is possible that some upstream movement occurred prior to the installation of the counting fence. It seems more 1 ikely though, considering the small numbers of fish taken during th~ first few days of the study, that the lower numbers observed in 1977 were simply a reflection of natural year to year fluctuations that might be expected to occur.

5.4.2.2 Diel ti~ing of upstream migration. The majority of longnose suckers (83.1%) moved upstream between noon and 0300 h with maximum movement usually occurring between 2100 and 2400 h (Table 19). Similar results have been observed for other longnose LONGNOSE SUCKER 15

"- Q,) ....c 300 ~U ~o 10 ==c ~. o a. (f)J: 200 .E lJ.... o Males tj~ ""'-J ~ 0 lJ.... IZa Females r o 100 o All Fish a:: w 5

~ 0 Inn H II H 14 H II tn! Ii II I ! g IIII R II n Pi "- J 55 Z ~-C E 45 ~ 0 100 ~- 35 "0 Q) > Downstream o Q,) )(.....J n =1004 25 C ~ 28 15 31 I 15 APR. MAY

Figure 16. Seasonal timing of the longnose sucker migration in the Muskeg River, 1977. 71 Table 19. Summary of die1 timing of the upstream migration of longnose suckers in the Muskeg River, 1977. Fish that were counted at times other than those ind icated were included in the next time check.

Number of Fish Counted at Each Check Oate 0900 1200 1500 1800 2100 2400 Total (0300)a

28 Apri 1 NO trap opened 5 2 NO 7 29 5 NO 0 0 3 NO 8 30 NO 0 NO 0 0 NO 0 1 May NO 0 NO NO 3 10 13 2 0 0 1 8 49 34 92 3 5 NO 0 0 0 59 64 4a 36 4 4 91 35 71 241 5a 129 NO 9 44 NO 76 258 6a 32 7 1 30 24 22 116 7a 19 16 2 4 5 25 71 8a 29 36 10 11 7 9 102 9a 11 13 1 5 17 16 63 lOa 17 3 1 3 0 18 42 11 a 11 1 1 0 2 34 49 12a 10 4 NO 6 2 trap closed 22 13 NO 40 NO 71 NO 37 148 14 NO 25 NO 1 4 12 42 15 NO 19 NO 0 3 11 33 16 NO 0 NO 0 NO 9 9 17 NO 1 NO NO 2 0 3 18 NO 1 NO NO 0 4 5 19 NO 0 NO 1 13 3 17 20 NO 3 NO NO 16 9 28 21 NO 1 NO NO 0 15 16 22 NO 1 NO 2 2 12 17 23 NO 3 NO 7 NO 5 15 24 NO 3 NO 9 1 9 22 25 NO 2 NO 0 NO 11 13 26 NO 17 NO NO NO 3 20 27 NO 0 NO NO 6 2 8 28 NO 0 NO 0 NO 2 2 29 NO 0 NO NO NO NO 0 30 1 NO 12 NO NO 0 13 31 NO 1 NO NO 0 1 2 1 Junea 6 8 NO 9 NO 4 27 2 NO 10 NO NO NO 1 11 3 NO 0 NO 1 NO 4 5 4 NO 1 NO 1 NO 6 8 5 NO 8 NO 5 NO 0 13 6 NO 1 NO 0 NO 0 1 7 NO 1 NO 2 NO 2 5 8 NO 0 NO 1 NO 0 1 9 NO 2 NO 0 0 1 3 10 NO 0 0 0 0 1 1 11 NO 0 NO 0 NO 0 0 12 NO 1 0 NO 0 0 1 13 NO 0 0 0 0 1 1 14 NO 0 NO NO 1 1 2 15 NO 1 ope rat ions te rmi nated

Totals 311 234 42 317 197 540 1641 % Grand Total 18.9 14.3 2.6 19.3 12.0 32.9

a Checks were made at 0300 h rather than 0900 h during the peak of the runs. 72 sucker runs, both within the AOSERP study area (Machniak and Bond in prep.) and elsewhere (Geen et al. 1966).

5.4.2.3 Spawning period. The 1977 spawning period for longnose suckers in the Muskeg River probably Jasted from one to two weeks. Ripe males were first noted on 2 May and most females were ripe by 4 May. Ripe males and females were captured as late as 14 and 15 May respectively but virtually all fish observed were spent by 13 May. Most fish entering the upstream trap after 13 May were spawned out and had probably been recently passed through the downstream trap. Geen et al. (1966) reported a spawning period of short duration with some adults leaving the spawning stream as early as five days after the migration began. The 1977 spawning period occurred at about the same time as in 1976, but because the timing of this event is temperature dependent, it can be expected to vary considerably from year to year.

.....,,:....------'='---s areas, Spawning of longnose suckers was not observed in the Muskeg River desp te da i ly s ur- ve ill ance by field personne 1 . They apparently did not spawn down- stream of the fence site "".there white suckers spawned despite the fact that the two species have rather simi lar spawning requirements .. No attempts were made to locate fish on spawning grounds in the upstream areas. However, on 3 May 1976, a fish fitting the description of a male longnose sucker in spawning colouration was observed in Hartley Creek (Ore R. Hartland-Rowe, University of Calgary, verbal communication with W. A. Bond, 4 May 1976), From the distribution of young-of-the-year suckers (two species) we conclude that longnose suckers do not util ize areas upstream of Site 4 in the Muskeg River or upstream of Site 8 in Hartley Creek (Figure 2) for spawning purposes. Young-of-the-year suckers were abundant at Sites 3 and 4 by mid-June 1977 and small numbers were captured in the lower reaches of Hartley Creek in mid-June 1976. 73

5.4.2.5 Length of time spent in Muskeg River. By 15 June 1977,61.2% of the longnose suckers counted through the upstream trap had returned downstream. Bond and Machniak (1977) observed that downstream movements continued until at least 30 July, by which time 77.2% of the migrants had returned downstream. Their results showed that the length of time spent in the tributary can vary greatly (from two to 87 days) although the majority of fish that had moved downstream by 30 July (81.6%) had been in the Muskeg River less than 30 days. Machniak and Bond (in prep.) recaptured longnose suckers in a downstream trap in the Steepbank River and suggested that some individuals may stay in the tribu­ tary throughout the summer. They stated that immature longnose suckers tend to remain in the tributary longer than the spawners. A similar situation may occur in the Muskeg River as well.

5.4.2.6 Seasonal and diel timi of downstream mi ra ion. The first spent fish were observed upstream of the fence on 12 May 1977, on which date the downstream trap was opened. This was eight to 10 days after the beginning of the main upstream run. Longnose suckers continued to move downstream through 15 June (Table 8, Figure 16) when operations were terminated. However, the majority of fish taken at the downstream trap (72.7%) were captured prior to 1 June. As mentioned previously, longnose suckers continued their downstream movement through 30 July in 1976, although 66.9% of them had passed the fence by 31 May (Bond and Machniak 1977). The downstream migration of longnose suckers took place mainly at night as only 30% of the fish were captured between noon and 2100 h (Table 20). Geen et al. (1966) reported that down­ stream movement of spent longnose suckers ceased in the early morning when water temperatures reached their daily minimum.

5.4.2.7 Spawning mortal lty. Prior to the 15 June termination date in 1977 only a few longnose suckers were found dead in the Muskeg River. Bond and Machniak (1977) reported finding 63 dead longnose suckers between 18 June and 30 July. Geen et al. (1966) 74

Table 20. Summary of diel timing of the downstream migration of longnose suckers in the Muskeg River, 1977. Fish that were counted at times other than those indicated were included in the next time check.

Numbe r of Fish Counted at Each Check Oate 1200 1800 2100 2400 Total

12 May trap opened 1 a 2 3 13 2 4 NO 2 8 14 a 7 4 13 24 15 8 26 12 21 67 16 2 a NO 8 10 17 a NO 23 12 35 18 a NO 43 24 67 19 3 40 a 26 69 20 a NO 1 72 73 21 a NO a 6 6 22 20 50 10 34 114 23 8 a NO 21 29 24 a 2 2 37 41 25 32 13 NO trap closed 45 26 20 NO NO 4 24 27 26 NO 2 2 30 28 28 a NO a 28 29 20 NO NO NO 20 30 a 16 NO 10 26 31 11 NO a a 11 1 June 53 a NO 1 54 2 6 NO NO 1 7 3 8 a NO 5 13 4 21 a NO 1 22 5 8 2 NO 4 14 6 36 6 NO a 42 7 a a NO a a 8 19 1 NO 3 23 9 4 2 2 3 11 10 5 5 2 2 14 11 4 9 NO 7 20 12 4 1 1 3 9 13 7 2 3 2 14 14 4 NO 8 5 17 15 14 ope rat ions terminated 14 Total 373 187 113 331 1004 % Grand Total 37.2 18.7 11 03 33.0 75

produced mortality estimates of from 11 to 28%, and considered survival of spawning longnose suckers to be very high.

5.4.2.8 Size composition of migrant longnose suckers. Longnose suckers measured during the 1977 upstream migration ranged in fork length from 120 to 514 mm, although the majority (88.3%) were between 320 and 449 mm (Table 21 and Figure 17). Within this length range, females were clearly longer in fork length than males (Figure 17). This situation was practically identical to that observed in 1976 in the Muskeg River (Bond and Machniak 1977) and in 1977 in the Steepbank River (Machniak and Bond in prep.). Unl ike the situation observed for white suckers, where many juveni le fish took part in the run, the longnose migration was comprised mainly of adult fish (spawners).

5.4.2.9 Age composition of migrant longnose suckers. Age was determined for 132 longnose suckers captured during the 1977 migration, of which sex was determined in 108 cases (Figure 18). Migrant suckers ranged in age from four to 13 years with the major­ ity being seven to 11 years old inclusive. Similar results were obtained in 1976 (Bond and Machniak 1977). Although our age sample was not drawn at random and cannot, therefore, be sa i d to descr i be accurate 1y the age compos i­ tion of the population, an analysis of the length frequency distribution of the run (Table 21) in the light of our knowledge of the age and growth characteristics of this population supports the conclusion that the migration consisted largely of fish between seven and 11 years of age.

5.4.2.10 Sex ratio of migrant longnose suckers. Sex was deter- mined for 1,130 longnose suckers during the upstream migration, of which 599 (53%) were males (Table 22). This is a significant deviation from the expected 1:1 ratio (X 2 =4.10, P< 0.001). The main upstream movement of spawning longnose suckers occurred between 2 and 10 May. During this period the sex ratio was not constant as reported by Geen et al. (1966) for longnose Table 210 Length-frequency distribution of longnose suckers during the upstream migration in the Muskeg Rive r, 1977.

Fork Length Fork Length Male Female Unknown Total Male Female Unknown Total (10 mm intervals) (10 mm intervals)

120 - 129 0 1 0 1 360 - 369 69 14 32 115 160 - 169 0 0 1 1 370 - 379 95 20 40 155 170 - 179 0 0 0 0 380 - 389 79 33 33 145 180 - 189 0 0 0 0 390 - 399 63 55 38 156 190 - 199 0 0 6 6 400 - 409 45 53 25 123 200 - 209 1 2 11 14 410 - 419 19 57 21 97 210 - 219 2 1 6 9 420 - 429 7 59 7 73 220 - 229 1 0 6 7 430 - 439 5 53 0 58

230 - 239 0 1 2 3 440 - 449 0 21 0 21 "'-.J 240 - 249 2 0 7 9 450 - 459 1 11 0 12 '" 250 - 259 0 1 3 4 460 - 469 1 4 0 5 260 - 269 2 0 5 7 470 - 479 0 0 0 0 270 - 279 4 0 3 7 480 - 489 1 1 0 2 280 - 289 3 0 9 12 490 - 499 1 0 0 1 290 - 299 0 0 14 14 500 - 509 1 0 0 1 300 - 309 0 1 19 20 510 - 519 0 1 0 1 310 - 319 1 0 11 12 320 - 329 3 4 18 25 Totals 482 407 371 1260 330 - 339 7 2 19 28 340 - 349 21 5 16 42 "'7 350 359 48 J 19 74 LONGNOSE SUCKERS 300l A All Fish n = 1260 6 Males n=482 o Females n=407

::r:: en 200 G: LL. 0 0:: ""-J W I / \ ~ ""-J m ::E 100 z:::>

120 200 300 400 500 FORK LENGTH (mm)

Figure 17. Length-frequency distribution for longnose suckers during the upstream migration in the Muskeg River, 1977. 20 I n LONGNOS SUCKER Males n = 47 c· :c 15 CJ) 1 D Females n = 61 lL.. LL 10 cr W ""'-.l m I I I I Q..j ~ z 5

)

Figure 18. composition for longnose suckers sampled during the counting operation Muskeg River, 1977. 79

Table 22. Sex ratio for longnose suckers during upstream migration, Muskeg River, 1977.

Number of Fish Pe rcent Date a Males Females Unknown Total Males

28 Apri 1 0 0 7 7 0 29 1 0 7 8 100 30 0 0 0 0 0 1 May 5 1 7 13 83 2 49 35 8 92 58 3 17 17 30 64 50 4 107 114 20 241 48 5 93 124 41 258 43 6 47 53 16 116 47 7 22 28 21 71 44 8 32 49 21 102 40 9 17 32 14 63 35 10 24 15 3 42 62 11 17 5 27 49 78 12 7 5 10 22 58 13 49 12 87 148 80 14 14 6 22 42 78 15 12 2 19 33 86 16 0 0 9 9 0 17 3 0 0 3 100 18 3 0 2 5 100 19 8 2 7 17 80 20 19 2 7 28 90 21 6 1 9 16 86 22 9 0 8 17 100 23 3 0 12 15 100 24 6 1 15 22 86 25 6 2 5 13 75 26 4 0 16 20 100 27 3 0 5 8 100 28 0 1 1 2 0 29 0 0 0 0 0 30 3 0 10 13 100 31 1 0 1 2 100 1 June 4 12 11 27 25 2 1 2 8 11 33 3 1 1 3 5 50 4 1 3 4 8 25 5 2 5 6 13 29 6 0 0 1 1 0 7 0 3 2 5 0 8 0 0 1 1 0 9 1 0 2 3 100 10 0 0 1 1 0 11 0 0 0 0 0 12 1 0 0 1 100 13 0 1 0 1 0 14 0 0 2 2 0 15 1 0 0 1 100 Totals 599 531 511 1641 % 53 47

a Based on fish of known sex. 80

suckers in Frye Creek. Rather, males tended to outnumber females on the first few days, while females were more numerous in the later stages of the run ble 22). The sex ratio was also observed to vary with the time in the Steepbank River with males dominating the early stages of the upstream run (Machol nd Bond j prep.).

5.4.2.11 Homi suckers. Geen et a . (1966) ----~~----~~------indicated that 10ngnose suckers in Frye Creek tended to return each year to the same spawning stream. Bailey (1 demonstrated a similar tendency in Brule Creek, Wisconsin. During 977, evidence from tag returns clearly indicated t longnose suckers of the Hus River return to that tributary to spawn n subsequent years in preference to other tributaries. Floy tags were applied to 1267 10ngnose suckers during their 1976 m gration into the Muskeg River (Bond and Machniak 1977). Ten of these fish were known to have been dead prior to the inning of the 1977 run. Of the remainder, 270 (20.7%) were recaptured in the Muskeg River during the 1977 study. If, as we believe Mus River 10ngnose suckers are part of the Lake Athabasca popula these f sh had undertaken a round ip in excess of 500 km the over- wintering area and the spaltJn i ng grounds As was the ca th white suckers 10ngnose suckers demonstrated considerabl de 1 i ty the Mus River At the 1977 Steepbank fence ope ra t ion, for example, on y nine ongnose suckers, ta during the Muskeg River st were recovered out of 3811 ish counted One fish was recovered fence operation on the MacKay River in 978 (Appendi 8.2).

5.4.2.12 Fecundi Fecundity was estimated gray metrically for 13 female 10ngnose suckers from the Muskeg River The data in Table 23 represent addit ons to those given by Bond nd Machniak (1977). Considering the fecundity data r both years, the estima total number of eggs per female (fork length to 440 mm) ra from 6 068 to 33 060 with an average 23 639 per fema e Actual counts on eight ovaries led errors of from • 2% to 4% for the estimated values . 81

Table 23. Fecundity estimates for longnose suckers sampled during the 1977 Muskeg River spawning migration.

Number of Eggs Relative Fork Weight Fecundity Length (g) Left Right (mm) Total Ova ry Ovary (cm) (g)

370 700 11 218a 10 567a 21 785 588.8 31.1 {-1.9%)b {-4.2%)b

407 1120 15 310a 17 750 33 060 812.3 29.5 {+2.0%)b

411 820 1 1 125 12 500 23 625 574.8 28.8

419 950 15 143 12 260 27 403 654.0 28.9

420 930 9 680 12 785 22 465 534.9 24.2

423 111 0 14 303 16 250 30 553 722.3 27.5 a Actual egg count. b Deviation of estimated counts from actual number. 82

Length-relative fecundity for 10ngnose suckers ranged from 390.0 to 812.3 ova per cm of fork length, while weight­ relative fecundity varied from 17.9 to 33.2 eggs per g of body weight. These values are similar to those reported for this spec i es by McCart et a l. (1977), Machn i ak and Bond (i n prep.) and Bond and Berry (in prep.b) in other studies within the AOSERP area. Although a positive correlation was found between fecundity and fork length, this correlation was not statistically significant (n= 13, r= 0.233). The mathematical relationship between fecundity and fork length is described by the equation:

loglOFecundity = 1.408 10g 10 Fork Length (mm) + 0.677; sb 1.775 Whereas fecundity correlated poorly with fork length a better but still insignificant positive correlation was seen between fecundity and body weight (r= 0.633). The relationship between fecundity and body weight for the above 13 fish (range 700 to 1120 g) is described by the equation:

logloFecundity 1.195 10g 10 Weight (g) +0.830; sb == 0.441

5.4.2.13 rowth. Age and growth results from 1977 (Tables 24 and 25) were ilar to those of 1976 (Bond and Machniak 1977) (Figure 19). Most growth in length was achieved during the first eight years of 1 ife. After age 8 the rate of growth decreased considerably. Growth in length for Muskeg River fish is identical to that reported for 10ngnose suckers by other studies in the AOSERP area (McCart et a1. 1977; Machniak and Bond in prep; Jones et al. 1978; Bond and Berry in prep.a, in prep.b). Muskeg River suckers (Figure 19) grow faster than those from Pyramid Lake, Alberta (Rawson and Elsey 1950), but more slowly than suckers from Yellowstone Lake (Brown and Graham 1954), Great Slave Lake (Harris 1962), and (Bailey 1969). Female 10ngnose suckers from the Muskeg River were generally longer than males of equal age. This difference was significant in age groups 7 to 11 both in 1976 (Bond and Machniak 1977) and in 1977 (table 24). Also as reported in 1976, Table 24. Age-length relationship (derived from fin rays and otol iths) for longnose suckers captured in the Muskeg River watershed, 1977, sexes separate and combined sample (includes unsexed fish).

Males Females All Fi sh Age t-test N Mean S.D. Range N Mean S. D. Range N Mean S. D. Range

0 53.0 2 50.5 3.54 48-53 2 82.0 0 82.0 3 0 0 0 4 0 193.0 5 191 .8 15.51 165-204 5 6 228.7 25.18 203-270 5 221 .4 22.77 200-253 27 218.2 18.74 200-270 0.499 00 6 5 27302 29.76 240-321 0 10 277.8 23.25 240-300 w a 7 4 281.3 7.41 271-288 8 358.4 43.82 293-420 13 329.2 51 . 15 271-420 3.414 a 8 8 365.9 16.81 345-397 7 386.0 10.94 371-396 15 375.3 17.33 345-397 2.699 a 9 13 383.2 17.62 356-411 19 403.2 20.25 358-438 32 395. 1 21 .41 356-438 2.885 10 5 378.0 12.90 357-391 10 401 .3 25.84 369-450 15 393.5 24.62 357-450 1 .878a a 11 3 381.7 25.38 361-410 9 414.2 20.99 385-445 12 406.1 25.59 361-445 2.223 12 430.0 411 .0 2 420.5 13.44 411-430 13 381.0 0 381 .0

Totals 47 61 135

a Indicates significant difference between means for males and females (Student's t-test, P< 0.05). Table 25. Age-weight relationship for longnose suckers captured in the Muskeg River watershed, 1977, sexes separate and combined sample (includes unsexed fish).

Males Females All Fi sh Age t-test N Mean S.D. Range N Mean S.D. Range N Mean S.D. Range

0 1 .55 2 1 .33 0.32 1.10-1.55 2 6.55 0 6.55 3 0 0 0 4 0 110.0 5 100.0 15.81 80-120 5 6 131 . 7 56.01 80-220 5 124.0 43.36 80-180 27 115.0 36.82 80-220 0.250

00 6 5 256.0 129.73 150-480 0 10 257.0 92.26 150-480 ,J.::'- 7 4 365.0 210.16 250-680 8 592.5 210.76 290-930 13 498.5 229.70 250-930 1 .764 8 8 585.0 78.92 520-770 7 697.1 31 .99 640-740 15 637.3 83. 11 520-770 3.502a a 9 13 684.2 106.85 540-840 19 822.1 152.81 560-1120 32 766.1 150.70 540-1120 2.811 a 10 5 610.0 70.00 540-720 10 750.0 131 .66 590-1060 15 703.3 131.19 540-1060 2.199 a 11 3 716.7 193.48 600-940 9 966.7 .68 640-1140 12 904.2 211 .98 600-1140 1 .994 12 920.0 820.0 2 870.0 70.71 820-920 13 640.0 0 640.0

Totals 47 61 135 a Indicates significant difference between means for males and females (Student's t-test, p:< 0.05). 600 I .. _2 /""-- 500 E E 400 I I- (!) z 300 w " ...... J ,," I- Muskeg R 1976 ~ ". .... " ". o Muskeg R 1977 co 0:: 200 ". V1 0 LL

100

5 6 7 8 9 10 AGE (YEARS)

Figure 19. Growth in fork length for longnose suckers from the Muskeg River and from several other areas: 1. Yellowstone Lake, Wyo. (Brown and Graham 1954); 2. Great Slave Lake (south), N.W.T .. (Harris 1962); 3. Lake Superior (Bai ley 1969); and 4 .. Pyramid Lake, Alta. (Rawson and Elsey 1950). Circles represent means and vertical 1 ines the ranges in fork length within age groups for Muskeg River. 86

the 1977 data indi te females to be significantly heavier than males of the same age in age groups eight to eleven inclusive (Table 25). The divergence in growth rate between males and females apparently commences at about the age of first sexual maturity. Brown and Graham (1954) and Lalancette and Magnin (1970) also reported that female longnose suckers grew faster than the males. However, Harris (1962) found no difference in growth rate for longnose suckers in Great Slave Lake. On the basis of our Muskeg River data for both years of the study we can detect no tendency for one sex to live longer than the other as suggested by Scott and Crossman (1973)0 The maximum age of 13 years recorded in the present study was a1so reported for longnose suckers in the AOSERP area

by Machni ak and Bond (i n prep_) and by Jones et al. (1978) c Bond and Berry reported a 19 yea r old longnose sucker (aged from fi n rays) from the Athabasca de 1 ta _ Tri pp and McCart (1974) found that most spawn i ng run suckers in the Donnelly River, N.W.T. were 11 to 18 years old with a maximum age of 22 years.

5.4.2.14 Sex and maturity. Of 108 longnose suckers for which both age and sex were determined, 56% were females (Table 26). The 1976 sample of 182 fish contained 53% males (Bond and Machniak 1977) . The youngest mature longnose sucker observed during the two years of the study was a five year old male. However, most suckers probably do not spawn until seven or eight years of age (Table 26). Hayes (1956) stated that longnose suckers reach sexual maturity at the age of two years in Colorado, while in the Northwest Territories, suckers do not mature unti 1 age nine (Harris 1962; Tripp and McCart 1974).

5.4.2.15 Length-weight relationship. Analysis of covariance indicated no significant difference (P>0 .. 05) between adjusted means (F=0.914) or slopes (F=0.066) of the length-weight regressions for male and female longnose suckers sampled in 19770 The mathematical re1ationship for the combined sample (including 87

Table 26. Age-specific sex ratios and maturity for longnose suckers from the Muskeg River drainage, 1977. Sex ratios were based only on fish for which sex was determined.

Females Males Age Unsexed Total Fish % % N % Ma ture N % Mature

100 0 0 0 0 2

2 0 0 0 100 0 0

3 0 0 0 0 0 0 0 0 4 100 0 0 0 0 4 5

5 5 45 0 6 55 0 16 27

6 0 0 0 5 100 20 5 10

7 8 67 88 4 33 25 13

8 7 47 100 8 53 88 0 15

9 19 59 100 13 41 100 0 32 10 10 67 100 5 33 80 0 15

1 1 9 75 100 3 25 67 0 12 12 50 100 50 100 0 2

13 0 0 0 100 100 0

Tota 1s 61 56% 47 44% 27 135 88 unsexed fish) between fork length and body weight for longnose suckers from the Muskeg River as determined from 1977 data (n = 131, r = o. 984 , ran gel 20 to 450 mm) i s des c rib e d by the eq ua t ion: lo910W = 3.103 (lo910L) - 5.179; sb 0.048

For male longnose suckers (n=47, r=0.963, range 203 to 430 mm) the calculated values for the slope (b), intercept (a) and standard deviation of b (sb) were 3.119, -5.225 and 0.130 respect­ ively. The corresponding values for females (n = 63, r = 0.987,. range 120 to 450 mm) were 3.085, -5.126 and 0.064.

5.4.2.16 Growth of young-of-the-year. Although young suckers were abundant throughout the lower 35 km of the Muskeg River by mid-June in both years of the study, the two species were indistinguishable at that time of the year. By the time young suckers were large enough to be identified to species, most of the fish present appeared to be white suckers. Few verified young-of­ the-year longnose suckers were captured in the Muskeg River either in 1976 or 1977. Bailey (1969) also reported difficulty in locating longnose sucker fry in spawning streams in western Wi scons i n and suggested that they dri ft to the lake soon after hatching. Perhaps young-of-the-year longnose suckers in the Muskeg River behave sim larly. On the other hand, Machniak and Bond (in prep.) reported no difficulty in locating young longnose suckers in the Steepbank River. The available information suggests that longnose suckers that remain in the Muskeg River attain a fork length of approximately 50 mm by the end of their first year (Bond and Machniak 1977). However, the growth rate of such "residentll young suckers may differ considerably from that of those that drift back to Lake Athabasca.

5.4.2.17 Food habits. Field analysis of longnose sucker stomachs during the spring spawning migration indicated that suckers fed little at that time (Bond and Machniak 1977). Suckers whose stomach contents were examined in the laboratory (n = 7) had fed primari lyon Simul i idae larvae~ Trichoptera larvae, Pelecypoda, and vegetable matter (Table 17). Young-of-the-year had consumed mainly small aquatic insects (Table 18). The diet of longnose suckers is known to be highly variable consisting largely of benthic invertebrates (Scott and Crossman 1973).

5.4.2.18 Reari areas. The lower 35 km of the Muskeg River ---~--- appear to be important as a rearing area for white suckers during June, July, and August. However, as few young-of-the-year suckers verified to be longnose were captured in the Muskeg River either in 1976 or 1977, no definite statement can be made concerning this species. Longnose sucker fry may leave the tributary more rapidly after hatching than white sucker fry. Alternatively, 1976 and 1977 may have been sub-normal years for longnose sucker spawning, with poor reproductive success.

5.4.2.19 Overwinteri No winter sampl ing was done in the Muskeg River during this study. However, small numbers of year­ lings were captured at Site 7 Hartley Creek and Site 9 the mouth of Kearl Creek in mid-June 1976, suggesting that some young-of­ the-year spend at least one winter in the Muskeg River water­ shed. Tagging results suggest that larger and older longnose suckers overwinter in Lake Athabasca.

5.4.3 Arctic Grayling

5.4.3.1 Spring movements. Arctic grayl ing spawning migrations appear to be initiated by increasing water temperatures and often begin with ice break-up (Brown 1938; Rawson 1950; Reed 1964; Schallock 1966; Bishop 1971). Tack (1972) reported that, in Alaska, the first grayl ing arrived on the spawning grounds when the water temperature was OoC. Ice left the Muskeg River between 20 and 22 April 1977, and the daily maximum water temperature exceeded 8°c on 24 April. An upstream migration of Arctic grayl ing was in progress in the Muskeg River on 28 April when the counting fence operation began. By 6 June, 161 grayling had passed through the upstream trap with 73.3% of them moving up prior to 15 May. 90

This represented approximately half the number of grayling counted in 1976 (Bond and Machniak 1977). Only 11 grayl ing were captured in the downstream trap (Table 8). As in 1976, most grayl ing (86.1%) moved upstream between noon and midnight with the heaviest movements occurring between 1500 and 2100 h (56.7%). The 1977 Steepbank River migra­ tion also occurred mainly during the daytime as 70% of the grayl ing moved upstream between 0900 and 2100 h (Machniak and Bond in prep.). Fai lure to catch the entire upstream grayling migration in 1976 was the major reason for repeating the fence study in 1977. Angl ing results during the summer of 1976 suggested that the Muskeg River supported considerably more grayling than our fence results had indicated, and it was hoped that more accurate counts could be achieved in 1977. Unfortunately we were unable to do this. An attempt to fulfill this objective by operating a downstream trap in the fall of 1978 was also thwarted, this time by extremely high water (Figure 8)0

5.4.3.2 Size of migrant grayl ing. Fork lengths were taken from 149 grayling captured during the 1977 upstream migration. These fish ranged in length from 175 to 389 mm (Table 27) with the length-frequency distribution exhibiting three modes (Figure 20). These three modes represent fish of age groups 2, 3, and 4 as indicated by age and growth information given by Bond and Machniak (1977) and in a later section of the present report. The length-frequency distribution did not remain constant during the period of fence operation in 1977. The early stages of the migration (25 April to 5 May) were dominated by grayl ing smaller than 260 mm fork length while most grayl ing passing upstream after this date exceeded 260 mm (Figure 20). The initial phase of the upstream grayl ing migration in the Steepbank River consisted of large, mature fish which were followed by smaller, immature fish in the later stages of the run (Machniak and Bond in prep.). Craig and Poul in (1975) demonstrated a simi lar pattern of upstream movement for grayl ing in northern streams. The small grayl ing captured in the Muskeg River between 91

Table 27. Length-frequency distribution of Arctic grayling during the upstream migration in the Muskeg River, 1977.

Fork Length Male Female Unknown Total (10 mm intervals)

170-179 0 2 180-189 0 0 1 190-199 2 5 8 200-209 2 3 5 10 210-219 3 8 12 220-229 0 6 7 230-239 0 6 7 240-249 2 0 10 12 250-259 1 3 4 8 260-269 1 2 11 14 270-279 1 6 8 280-289 0 8 9 290-299 2 0 3 5 300-309 3 6 2 11 310-319 3 4 3 10 320-329 5 4 10 . 330-339 6 0 7 340-349 3 0 4 350-359 2 0 0 2 360-369 0 0 0 0 370-379 0 0 0 0 380-389 2 0 0 2

Totals 38 28 83 149 92

20- 28Apr - 5 May n=73

10

::c 20 6May-6June en n= 76 LL LL 0 10 0:: W CD :E z:::> Total n= 149 20

10

200 300 400 FORK LENGTH (mm)

Figure 20. Length-frequency distribution for Arctic grayl ing during the upstream migration in the Muskeg River, 1977. 93

28 April and 5 May are thought to represent the late stages of the main upstream run. Most spawners are believed to have passed the counting fence prior to 28 April. The larger fish taken after 5 May may have spawned in other tributaries before entering the Muskeg or they may have spawned in the Muskeg River below the fence site and then moved upstream to summering areas.

5.4.3.3 Spawning. Grayling usually spawn over gravel or rocky bottom with water depth appearing not to be an important factor (Fabricius and Gustafson 1955; Kruse 1959; Bishop 1971). Grayling in tributaries of the southern Athabasca River drainage spawn in May at stream temperatures of 4.5 to 11°C (Ward 1951). Tack (1972) noted that, in Alaska, spawning was first observed when the stream temperature was 4°c and that by 10°C spawning was completed. Records over a 10 year period at Black Lake, a shield lake in northern Saskatchewan, indicate that, although spawning occurs over a three week period, the peak spawning period lasts only three days to a week (Johnston 1971; Kratt and Smith 1977). Spawning of Arctic grayling was not observed in the Muskeg River either in 1976 or 1977. However, the lower 35 km of the main river and the lower reaches of Hartley Creek provide many areas that appear suitable for this purpose. Spawning probably occurred during the last week of April and first week of May in both years. Young-of-the-year were taken on 15 June 1976 (range 32 to 42 mm) and on 3 June 1977 (range 18 to 24 mm) at Site 3 (Figure 2). As well, fry were captured between 16 and 21 June 1976 (range 27 to 38 mm) and on 19 June 1977 (38 mm) at Site 7 in Hartley Creek. While most grayling spawning is believed to occur upstream of our fence site, there is a possibility that some fish spawn below this site. Grayling fry were captured between the fence site and the tributary mouth on 15 June 1976 (range 32 to 42 mm) and on 7 June 1977 (range 26 to 27 mm). As mentioned previously, the capture of spawning size grayling in the 1977 upstream trap after 6 May (Figure 20) suggests either that these fish had spawned in other tributaries and were now moving into the Muskeg River or that they had spawned in the Muskeg River below the fence and were now moving into upstream areas. 94

5.4.3.4 Summer residence of Arctic grayl ing. Arctic grayling did not leave the Muskeg River after spawning in 1976 and 1977 but remained in the tributary throughout the summer. This is unlike the situation in most northern streams (Craig and Poulin 1975) but similar to that reported by Ward (1951) and Machniak and Bond (in prep.) for other streams in the Athabasca River drainage. During summer 1976, angl ing produced considerable numbers of grayl ing in the lower 10 km of the Muskeg River. Ten angler hours, applied to this area on 8 to 10 August, produced 28 Arctic grayling, aged one to four years (Bond and Machniak 1977). Dr. D. Barton (University of Waterloo, verbal communication with W. A. Bond, April, 1978) reported angl ing grayl ing from the area just downstream of the mouth of Hartley Creek (Site 4) throughout the summer of 1977 as well as in Hartley Creek itself ite 7) and at Site 3 (Figure 2) on the Muskeg River. Dr. Barton tated that most grayl ing occurred in areas where water up to 1 m deep flowed with a moderate current over beds of macrophytes and sand Within the canyon portion of the Muskeg River, most grayling were found near the upstream ends of pools, just below riffles Grayl ing were never observed in the Muskeg River upstream of Ha tley Creek. Grayling were still abundant in the Muskeg River on October 1977, when anglers captured 28 fish at Site 3 (Figure 2) These fish had a mean fork length of 308.9 mm, ranging from to 355 mm. The situation in the Muskeg River is probably similar to that described by Machniak and Bond (in prep.) in the adjacent Steepbank River. In that tributary an upstream run of Arctic grayl ing took place in April and May. The grayl ing remained in the tributary through the summer, returning to the Athabasca River between 6 and 15 October, just prior to freeze-up. AOSERP fishery crews working on the Athabasca River captured few grayl ing during the summer, but reported fish showing up in their catches between 6 and 20 October 1977 (Bond and Berry in prep.b). Jones et al. (1978) took 25 gray1 ing during their study on the Athabasca River upstream of Fort McMurray, but none was captured prior to mid-October. 95

Although a fence operation was not possible during autumn 1978, a 1 imited amount of angl ing was conducted in the lower 5 km of the Muskeg River between 25 September and 13 October in an attempt to verify the presence of large grayling. During this period the overnight low water temperature decreased from 6.7 to 3.9°C, but these temperatures were warmer than those recorded at the time of the downstream grayl ing run in the Steepbank River (Machniak and Bond in prep.). Two grayling, a male and female measuring 385 and 360 mm respectively in fork length, were captured on 30 September. Another male (342 mm) and female (328 mm) were captured on 13 October. The 1978 grayl ing migration out of the Muskeg River probably took place between 15 October and freeze-up, which occurred on 5 November (Figure 8).

5.4.3.5 Age and growth. Growth of Arctic grayl ing in the Muskeg River is described in detail by Bond and Machniak (1977). Addition­ al growth information gathered in 1977 is presented in Tables 28 and 29. Muskeg River grayling have almost identical growth patterns to those reported for populations in southern tributaries of the Athabasca River (Ward 1951) and in other tributaries of the AOSERP study area (Griffiths 1973; Machniak and Bond in prep.). Muskeg River grayl ing grew at a rate similar to that reported for grayl ing from Great Bear Lake (Falk and Dahlke 1974), Great Slave Lake (Bishop 1967), and the Mackenzie River (Hatfield et al. 1972) for their first year or two, but thereafter, they grew more slowly than the lake populations but faster than Mackenzie River fish (Figure 21). Grayl ing from the Muskeg River grew considerably faster than those from the Kavik River (Craig and Poulin 1975). The maximum scale age recorded for Arctic grayl ing in the Muskeg River is seven years (Bond and Machniak 1977). This was also the maximum age observed in the Steepbank River (Machniak and Bond in prep.). The oldest grayl ing reported to date from the AOSERP study area is a 12 year old male, aged from otoliths (Jones et al. 1978). Grayling appear to live longer in the northern Tab 1e 28. Age-length relationship (derived from scales) and age-weight relationship for mountain whitefish and Arctic grayling captured in the Muskeg River, 1977, sexes combined (includes unsexed fis

Fa rk Length (mm) We i gh t (g) Age Male Female Total Mean s. D. Range Mean s. D. Range

Arcti c gray1 ing

2 204.8 14.19 175-231 91.2 18.67 60-140 7 9 17

3 251. 0 4.85 247-258 182.0 44.94 140-240 2 2 5

4 312.5 13.44 303- 322 330.0 42.43 300-360 2

Totals 10 12 24 \..0 0"

Mountain whitefish

3 245.3 16.86 226-257 173.3 46.19 120-200 2 3

5 342.4 11.04 327- 358 551.4 43.66 475-600 2 5 7

6 368.5 7.05 364- 379 580.0 59.44 510-650 2 4

7 392.0 1000.0 0

Totals 5 9 15 Tab 1e 29. Size and weight relationships for young-of-the-year and juveniles of larger fish species coll- ected from the Muskeg River, 1977.

Fork Length (mm) Species/Age Weight (g) Total Mean S.D. Range Mean S.D. Range

Yellow perch 0+ (July-Aug.) 39.2 6.62 23-49 0.7 0.30 0.2-1.2 33 Arctic grayl ing 0+ (June) 23.4 4.50 18-38 o. 1 0.10 0.1-0.5 17 (July) 79.0 3.6 1 (Aug. ) 89.0 8.0 1 Lake wh i tef ish

0+ (June-July) 24-38 16 \..0 29.5 4.99 0.3 O. 15 0.1-0.5 ...... Northern pike 0+ (June) 26.5 4.95 23-30 0.2 0.00 2 ~Ju1Y) 107.0 15.56 96-118 6.9 1 .20 6.0-7.7 2 Aug. ) 126.0 4.24 123-129 14.0 2.12 12.5-15.5 2 1+ (May) 167.0 34.0 1 Burbot a 0+ (July) 54.0 0.8 1+ (May) 107.5 3.54 105-110 7.0 0.42 6.7-7.3

a Tota 1 1ength. 98

400 _0."",4 3 ...... "".. .. _0' ,,' ."" .'

~ E E '-' 300 I I-- (!) I Z / W ...J .I .I ~ I c:: 200 : ~ I I • Muskeg R 1976 o Muskeg R 1977

100

0+ I 2 3 4 5 6 7 8 9 10 II 12 AGE (YEARS)

Figure 21. Growth in fork length for Arctic grayling from the Muskeg River and from several other areas. 1. Great Bear Lake (Falk and Dahlke 1974); 2. Great Slave Lake (Bishop 1967); 3. Mackenzie River (Hatfield

eta 1. 1972 ) ; 4 0 Ka v i k Ri ve r ( Cr a i g and Po u 1 i n 19 75) . Circles represent means and vertical 1 ines the ranges in fork length within age groups for Muskeg River fish. 99

part of their range than in the south. A maximum scale age of 12 years is reported from Great Slave Lake (Bishop 1967) and Great Bear Lake (Falk and Dahlke 1974). Craig and Poulin (1975) recorded an otolith age of 22 years in the Firth River, Yukon Territory.

5.3.4.6 Length-weight relationship. Comparison of length- weight relationships in 1976 indicated no significant difference (p > 0.05) between the regressions for male and female grayl ing from the Muskeg River and the data for the two sexes were combined (Bond and Machniak 1977). Length and weight data are available for only 24 g ray lin g f ro m the 1977 stu d y (r = o. 971, ran ge 175 to 322 mm). The mathematical relationship between fork length and body weight for this sample is expressed by the equation: 0.162

5.4.3.7 Sex and maturity. Sex and age were determined for only 22 Arctic grayl ing in 1977, of which 10 were males (Table 28). Males accounted for 62% of the sample in 1976 (Bond and Machniak 1977) . Bond and Machniak (1977) reported that, for Muskeg River grayling, the earl iest age of sexual maturity was two years for males, three years for females, and that 50% of both sexes were mature at age three. By age four virtually all grayl ing were sexually mature. Similar findings were reported by Ward (1951) and Machniak and Bond (in prep.)o Craig and Poulin (1975) reported that grayl ing in Alaska reached sexual maturity between age five and age eight, the oldest age of maturity for grayling in North Amer i ca.

5.4.3.8 Fecundity. Total egg counts were performed on two Muskeg River grayling in 1976 (Bond and Machniak 1977). One fish

( fork 1eng t h 225 mm) co n t a i ned 27 19 ova, wh j 1e the 0 the r (fo r k length 308 mm) contained 6971 eggs. Fecundity in Steepbank River grayl ing varied from 2206 to 8546 (mean 4.689) for seven fish 100

ranging in fork length from 275 to 365 mm (Machniak and Bond in prep.). The average fecundity for this species is probably between 4000 and 7000 (Scott and Crossman 1973), although counts on individual fish have ranged from 574 (Ward 1951) to 15 907 (Bishop 1971) .

5.4.3.9 Food habits. Studies of the food habits of Arctic grayling indicate that this species is extremely opportunistic, feeding on a great variety of food items. Many authors have stressed the importance of aquatic insects in the diet (Kruse 1959; Bishop 1967; Reed 1964), while others (Miller 1946; Rawson 1950; Wojcik 1955; Schallock 1966) have found terrestrial insects to make up a large proportion of the food. Fish, fish eggs, lemmings, and amphipods have also been found in grayling stomachs (Miller 1946; Reed 1964; McPhail and Lindsey 1970). Sixty grayl ing stomachs were examined in the field during 1976 of which only 10 "Jere empty (Bond and Machniak 1977). Most were one-quarter to one-half full, the contents consisting mainly of aquatic insects. During 1977, 21 additional stomachs were examined in the field, of which only one, a ripe female, contained no food. The remaining stomachs were one-half full to full, the principle food being aquatic insects of the orders Trichoptera, P1ecoptera, Odonata and Hemiptera. The stomach contents of 20 adult gray1 jng were examined in more detail in the laboratory. The results of this analysis (Table 30) showed that aquatic insects occurred in all stomachs examined, with immature stages of the orders Diptera, P1ecoptera, Trichoptera, Hymenoptera, and Odonata accounting for most of the food. The stomachs of four adult grayling captured from the Muskeg River in September and October 1978 were gorged with Plecoptera nymphs and Corixidae adults, and also contained small numbers of Trichoptera and Diptera larvae, Odonata nymphs, and Coleoptera adults. The stomachs of young-of-the-year grayl ing also con­ tained mostly immature insects (Table 30) (Bond and Machniak Table 30. Food habits of Arctic grayling collected from the Muskeg River during 1976 and 1977.

May (1976/77) July (1976) August (1976) Y-O-Y (1977) Food Items % F req •a % No. % Vol. % Freq.a % No. % Vo 1. % F req .a % No. % Vo 1 . % F req . a % No.

Class Insecta Diptera Ch i ronomi dae 1a rvae 0.0 0.0 0.0 0.0 0.0 0.0 26.7 5.6 0.8 33.3 40.9 pupae 0.0 0.0 0.0 0.0 0.0 0.0 13.3 9.9 0.7 33.3 11.4 Unidentified Dipterans 1a rvae 66.7 25.4 4.8 0.0 0.0 0.0 86.7 36.0 11 .0 33.3 4.5 adults 0.0 0.0 0.0 0.0 0.0 0.0 6.7 1.2 O. 1 0.0 0.0 Trichoptera 33.3 31. 7 4.3 0.0 0.0 0.0 40.0 13.7 4.6 16.7 2.3 P1ecoptera 41 .8 100.0 19. 1 0.0 0.0 0.0 60.0 14.3 14.5 33.3 22.7 0 Coleoptera 0.0 0.0 0.0 100.0 66.7 20.2 6.7 0.6 0.6 0.0 0.0 Hemiptera 0.0 0.0 0.0 0.0 0.0 0.0 13.3 2.5 1 .5 16.7 2.3 Corixidae 33.3 3.2 2. 1 0.0 0.0 0.0 6.7 0.6 0.3 0.0 0.0 Notonectidae 33.3 3.2 1.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Hymenoptera Formicidae 33.3 1 .6 1 . 1 50.0 16.7 4.2 40.0 6.2 2.6 0.0 0.0 Odonata 66.7 9.5 37.9 0.0 0.0 0.0 26.7 7.5 3.7 16.7 203 Lepidoptera 33.3 1.6 4.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Ephemeroptera 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 16.7 13.6 Insect Remains 0.0 0.0 0.0 0.0 0.0 0.0 73.3 0.0 35.9 16.7 0.0 Mi sce 11 aneous Arachnida 100.0 6.3 2.6 50.0 16.7 8.4 13.3 1.2 2.2 0.0 0.0 Vegeta t i on (a 1gae, seeds) 0.0 ND ND 0.0 ND ND 6.7 ND 3.7 16.7 ND Fish 0.0 0.0 ND 0.0 0.0 ND 6.7 0.6 9.3 0.0 ND Di gested Matter 0.0 ND ND 0.0 ND ND 13.3 ND 7.4 16.7 ND Debris (sticks, stones) 0.0 ND ND 50.0 ND 67.2 6.7 ND 0.6 16.7 ND Number of stomachs 3 2 15 6 Empty (% of Total) 0 0 0 0

a Percentage frequency of occurrence, based on stomachs that contained food. 102

1977). Scott and Crossman (1973) state that young grayling initially feed on zooplankton but undergo a shift to immature insects as they increase in size.

5.4.3.10 Rearin areas. The entire lower 35 km of the Muskeg River, and the lower 10 km of Hartley Creek are utilized as rearing areas by Arctic grayling. Many young-of-the-year were observed throughout the summer of 1977 in the area just down­ stream from Hartley Creek as well as near Site 3 (Figure 2). As mentioned previously, fry were captured at Site 7 as well as at Sites 2 and 3 during both summers. In the canyon section of the Muskeg River, approximately 6 to 8 km upstream from the mouth of the tributary, many young-of-the-year were observed on 10 August 1976. These sma1 1 fish occupied long shallow pools where a moder­ ate current flowed over a very uniform gravel bottom.

5.4.3.11 Overwinteri We believe, for several reasons, based on our observations on the Muskeg and Steepbank rivers, that young­ of-the-year grayling in both streams remain in the tributaries over their first winter, and do not join the migrant population until the autumn of their second year. Firstly, despite intensive sampling with small mesh beach seines, few young-of-the-year gray- 1 ing have been captured in the Athabasca River within the AOSERP study area (Bond and Berry in prep.a, in prep.b). Secondly, upstream grayl ing runs in the Muskeg River in 1976 and 1977 and in the Steepbank River in 1977 (Machniak and Bond in prep.) included no one year old fish. Although such fish could pass through the 2.54 x 2.54 cm mesh used in the fence, some, had they been present, would have been captured, if not in the fence at least by seines or minnow traps. Thirdly, the downstream migration in the Steep­ bank River in October 1977 (Machniak and Bond in prep.) included fish in the 130 to 230 mm size range that are thought to have been age 1+ fish. These small fish were the last to leave the Steep­ bank River. Lastly, D. Barton (University of Waterloo, verbal communication with W. A. Bond, December 1976) reported sighting six to 10 juvenile gray1 ing through the ice at Site 7 Hartley 103

Creek, on 30 October 1976. Overwintering within the tributaries would appear to enhance the survival chances of young fish which would not be exposed to the rigors of migration or to predation by piscivorous fish. Migrant grayl ing (age 1+ and older) probably overwinter in the Athabasca or Clearwater rivers upstream of Fort McMurray.

5.4.4 Northern Pike

5.4.4.1 Movements and distribution. A total of 433 northern pike were passed through the upstream trap of the 1977 Muskeg River counting fence (Table 8, Figure 22). The majority of pike (76.4%) passed the fence between 30 April and 12 May, during which period the daily maximum water temperature increased from 5 to 14°c. Pike demonstrated a pronounced diel periodicity during their upstream run as 62% of those captured through 12 May were taken between 1200 and 2100 h, the largest movements occurring after 1500 h. Frankl in and Smith (1963) reported that pike began moving into spawning streams or flooded marshes at water temperatures of 1 to 4.5°C, and that most movement takes place at night. This represents a sharp contrast to the situation observed in the Muskeg River. By 15 June, 59 pike had moved through the downstream trap, leaving 374 still upstream of the counting fence. Pike continued passing through the downstream trap through 30 July 1976 (Bond and Machniak 1977). During the summer months, northern pike seem to be confined largely to the lower reaches and mouth area of the Muskeg River, although individual fish may ascend the tributary for considerable distances. Angl ing results in 1976 suggested that most pike do not ascend more than 6 or 7 km upstream (Bond and Machniak 1977). However, in 1977, Ilquite a few" pike were angled in the vicinity of small fish collection Site 3 (13 km upstream) (Dr. D. Barton, University of Waterloo, verbal communi­ cation with W. A. Bond, April 1978). One young-of-the-year 96 mm in fork length, was captured in a seine at Site 4 (Figure 2) on 18 July 1977.

105

Although there is no direct evidence, most pike probably left the Muskeg River before freeze-up to overwinter in the Athabasca River. Most pike in the Steepbank River had appar­ ently left that stream during the summer, and 42 fish were were captured moving out of the watershed during September and October (Machniak and Bond in prep.). Within the AOSERP study area, northern pike appear to move around very 1 itt1e. Tagging results from the present study (Bond and Machniak 1977) (Appendix 8.2) show that most pike recap­ tured outside the Muskeg River watershed had travelled less than 20 km from the point of tagging. Similar results were obtained by Machniak and Bond (in prep.) and Bond and Berry (in prep.a, in prep.b).

5.4.4.2 Spawning. Northern pike usually spawn in April and early May immediately after ice breakup at water temperatures of 4.4 to 11. 1°C (Scott and Crossman 1973). While pike may spawn in a variety of habitats, a requirement of the spawning site appears to be the presence of vegetation (Machniak 1975). Marshes or marsh-like conditions along small streams seem to be preferred areas. Such areas are uncommon in the Muskeg River and it is felt that any spawning that does occur in this tributary is minor. This belief is supported by the fact that most of the pike passed through the upstream trap are thought to have been immature or maturing fish that would not have spawned in 1977. Of 40 fish whose gonads were inspected, only four were ripe. Machniak and Bond (in prep.) also reported many immature and spent fish in the Steepbank River migration. Despite its probable minor nature, some northern pike spawning apparently does occur in the Muskeg River. Two small young-of-the-year pike (23 and 30 mm) were captured on 12 June 1977 approximately 0.5 km downstream of the fence site. Four others (96 to 129 mm) were captured in the Muskeg River between 18 July and 13 August, one of which was taken at Site 4 (Figure 2). Four pike fry were captured on 22 June 1978 near the upper part of the canyon, approximately 9 km upstream from the mouth of the tributary. 106

Bond and Berry (in prep.b) captured ripe pike in the Athabasca River from 27 April to 9 May 1977, and spent fish from 7 May on. Young-of-the-year measuring 19 to 36 mm were captured between 16 and 18 June.

5.4.4.3 Length-frequency distribution. Pike captured during the 1977 upstream run in the Muskeg River ranged in fork length from 244 to 788 mm, with the majority (86.3%) being in the 300 to 524 mm size range (Figure 23). Male pike varied from 244 to 565 mm while females had fork lengths between 330 and 788 mm. Fish less than 400 mm in fork length made up 59.2% of the total sample, whereas in the Steepbank River, 66.7% of all pike measured exceeded 400 mm fork length (Machniak and Bond in prep.).

5.4.4.4 Age and growth. Northern pike examined during 1977 ranged in age from 0+ to seven years, with most fish (75.0%) being three to five inclusive. Seven was also the maximum age reported during the 1976 study (Bond and Machniak 1977). The maximum age reported for pike in the AOSERP study area is 13 years (McCart et al. 1977). Pike increased in fork length at a constant rate throughout life. Females tended to be longer than males of the same age with significant differences (p< 0.05) occurring in age groups 3, 4, and 5 lTable 31). The growth rate of northern pike compares favourably with that reported in previous studies on pike in the AOSERP study area (Griffiths 1973; McCart et ala 1977;

Machniak and Bond in prep.; Bond and Berry in prep.a~ in prep.b; Jones et al. 1978). Muskeg River pike grew more rapidly than those from Lake Athabasca and Great Bear Lake (Miller and Kennedy 1948) and the Kakisa River (Falk and Dahlke 1975), but slower than was reported by Pinsent (1967) for pike from Beaver Lake, Alberta (Fi gure 24). Northern pike gained weight slowly up to age 3, but more rapidly thereafter. Female pike were significantly heavier (p > 0.05) than males in age groups 4 and 5 (Table 32). 250 300 400 500 600 800 FORK LENGTH (mm)

Figure 23. Length-frequency distribution for northern pike during the upstream migration in the Muskeg River, 1977. Table 31. Age-length relationship (derived from scales) for northern pike captured in the Muskeg River, 1977, sexes separate and combined sample (includes unsexed fish).

Males Females All Fi sh Age t-test N Mean S.D. Range N Mean 5 0 Range N Mean S.D. Range

0+ 129.0 123.0 6 86.5 47.84 23-129

0 167.0 167.0

2 0 2 275.0 26 256-294 2 275.0 26.87 256-294

3 13 337.4 14.78 315-364 9 35201 19.76 320-378 342.7 18.08 315-378 2.000a

0 4 6 384.2 28.74 356-434 5 8.8 26. 469 11 404.5 35. 14 356-469 2.657a co a 5 8 469.9 16.92 442-497 6 524.7 45 3 448-560 14 493.4 41.58 442-560 3.184

6 522.0 599.0 2 560.5 54.45 522-599

7 0 648.0 648.0

Totals 29 26 60 a Indicates significant difference between means r males and females (Student1s t-test, P< 0.05). 109

NORTHERN PIKE

700

600

E E 500 I r ~ 400 w ..-I

~ 0::: 300 ~

200

100 T I 0+1 2 3 4 5 6 7 8 9 10 II 12 AGE (years)

Figure 24. Growth in fork length for northern pike from the Muskeg River and from several other areas. 1. Beaver Lake, Alta. (Pinsent 1967); 2. Kakisa River, N.W.T. (Falk and Dahlke 1975); 3. Lake Athabasca, Alta. (Miller and Kennedy 1948); and 4. Great Bear Lake, N.W.T. (Miller and Kennedy 1948). Circles represent means and vertical 1 ines the ranges in fork length within age groups for Muskeg River fish. Table 32. Age-weight relationship for northern pike captured in the Muskeg River, 1977, sexes separate and combined sample (includes unsexed fish).

Ma les Females All Fi sh Age t test N Mean S.D Range N Mean S.D Range N Mean S.D. Range

0+ 15.5 2,5 6 7.0 6. 0.2-15.5

0 .0 34.0

2 0 2 .35 2 205.0 134.35 110-300 3 13 .8 • 18 180- 9 50 220-370 23 267.8 48.52 180-370 o.

4 6 39607 79 41 340-540 5 .0 09.91 420-720 11 463.6 117.92 340-720 2. 0

5 8 713.8 111.73 600-860 6 916.7 . 6 700-1160 14 800.7 172.20 600-11 2.633a

6 1320.0 1220.0 2 1270.0 .71 1220-1320 7 0 2330.0 2330.0 Totals 29 26 60 a Indicates significant difference between means for males and females (Student's t-test, P< 0.05). 111

Growth data for young-of-the-year pike are presented in Table 29. Two fish, captured 12 June, measured 23 and 30 mm in fork length, two taken in July had lengths of 96 and 118 mm, while two pike caught in August measured 123 and 129 mm. A one year old pike captured 3 May 1977 had a fork length of 167 mm. Bond and Berry (in prep.b) reported young-of-the-year pike to range in fork length from 19 to 36 mm in mid-June, from 39 to 44 mm at. the end of June, and to reach a max i mum size of 185 mm and 41.6 g by mid-October.

5.4.4.5 Length-weight relationship. For male northern pike taken from the Muskeg River in 1977 (n = 30, r = 0.973, range 129 to 522 mm), the length-weight relationship is described by the equation: lo910W = 3.084 (lo910L) - 5.372; sb = 0.444 The corresponding equation for female pike (n = 26, r = 0.986, range 123 to 648 mm) is: lo910W = 2.899 (lo910L) - 4.913; sb = 0.099 No significant difference was found between the slopes of the regression lines (F = 0.905) for male and female pike.

5.4.4.6 Sex and maturity. Age and sex were determined by by gonadal inspection for 55 pike of which 29 (53%) were males (Table 33). Bond and Berry (in prep.a) found that male and female pike in the Athabasca River occurred in equal numbers, while Jones et a1. (in prep.) reported female pike (58%) outnumbered males in late fall. Both sexes appear to achieve sexual maturity for the first time at age 3 (Table 33). Over two years, 22% of females and 33% of males were mature at this age. The earl iest age of sexual maturity for pike in the Steepbank River was three years for males and four years for females while Bond and Berry (in prep.a, in prep.b) reported that some pike in the Athabasca River may spawn at age 2.

5.4.4.7 Fecundity. Total egg counts were performed for two northern pike captured at the Muskeg River counting fence in 1977. 112

Table 33. Age-specific sex ratios and maturity for northern pike from the Muskeg River, 1977. Sex ratios were based only on fish for which sex was determined.

Fema 1es Males Age Unsexed Total % % Fish N % Mature N % Mature

0+ 50 0 50 0 4 6

100 0 0 0 0 0

2 2 100 0 0 0 0 0 2

3 9 41 22 13 59 46 23

4 5 45 60 6 55 83 0 11

5 6 43 83 8 57 100 0 14

6 50 100 50 100 0 2

7 100 100 0 0 0 0

Totals 26 47% 29 53% 5 60 113

One fish (fork length 599 mm) contained 26 155 eggs while the other (648 mm) had 36 763 (Table 34). Bond and Berry (in prep.b) reported pike from the Athabasca River C5,44 to 656rnmfork length) to have an average fecundity of 28 896 eggs per female (range 17 764 to 42 962). Fecundit!es reported for Athabasca River pike by McCa rt et a 1. (1977) ranged from 20 267 to 53 295 wi th a mean of 32 452 eggs per female (fork length 528 to 710 mm).

5.4.4.8 Food habits. During 1977, the stomachs of 42 northern pike were examined in the field, of which 25 (59.5%) contained no food. The remaining stomachs all contained fish, the food species including white sucker, burbot, brook stickleback, Arctic gray­ ling, and northern pike. Bond and Berry (in prep.b) also report AOSERP area pike to be mainly piscivorous with the major food species being flathead chub, suckers, and trout-perch. These authors determined that insects, Plecoptera, Odonata and Lepidop­ tera accounted for 1.4% of the food volume, while frogs and mice made up 4.1%. Jones et al. (1978) found remains of rodents in 5% of the stomachs they examined. Young-of-the-year pike from the Muskeg River (Table 18) had fed mainly on Ephemeroptera nymphs and fish, including white suckers, longnose suckers, and cyprinids.

5.4.4.9 Rearing and overwintering. Some pike do uti 1 ize the Muskeg River for rearing purposes as small numbers of young-of­ the-year were captured in 1977 and 1978. These small fish were taken in quiet, weedy areas along the stream margin, out of the main current. One such area was found to be near the top of the canyon, 9 km upstream of the river mouth, while another was a side slough situated approximately 0.5 km downstream of the fence site. Whether or not northern pike overwinter in the Muskeg River is unknown. It is bel ieved, however, that the larger pike that participated in the spring upstream migration left the tributary prior to freeze-up to overwinter in the Athabasca River. 114

Tab1e 34. Actual egg coun ts 0 f two northern pike females sampled during the 1977 spawning period, Muskeg River.

Number of Eggs Fork Relative Weight Fecundity Length (g) Left Right (mm) Total Ovary Ovary (cm) (g)

599 1220 11 483 14 672 26 155 436.6 21.4 648 2330 18 613 18 150 36 763 567.3 15.8 115

5.4.5 Mountain Whltefish

5.4.5.1 Spring movements and distribution. Approximately 50 mountain whitefish were counted through the upstream trap in 1977, with 68.4% of them passing the fence prior to 14 May (Table 8). Seventeen mountain whitefish had returned downstream by 15 June. During the 1976 study, only 33 mountain whitefish were captured moving upstream, but 101 were taken in the downstream trap (Bond and Machniak 1977). The suggestion was that mountain whitefish did not spend the summer in the Muskeg River but returned to the Athabasca River. The 1977 fence was not operated long enough to detect such a downstream run in the second year. In the Steepbank River, a spring run of mountain whitefish had not returned down­ stream by 29 May (Machniak and Bond in prep.). The counting fence in that study was not operated during the summer but a fall oper­ ation failed to detect any downstream movement of whitefish between 12 September and 15 October. Whether mountain whitefish left the Steepbank River during the summer or remained in the tributary beyond 15 October was unknown. Davies and Thompson (1976) observed a complex movement pattern for mountain whitefish in the Sheep River, Alberta, involv­ ing spring feeding, summer feeding, prespawning, spawning, and post-spawning-overwintering movements. They found that white- fish enter tributaries during the spring to feed and leave in June, returning to the larger rivers when water levels decl ine and water temperatures rise in the smaller tributaries. A similar pattern of movement may also occur in the AOSERP study area, although it should be noted that some stream-dwell ing populations remain in tributaries all summer. In Idaho, mountain whitefish are reported to move into tributaries during late spring and early summer, remain in the upper reaches until spawning in November, and then return to the large rivers to overwinter (Pettit and Wallace 1975). The locations occupied by mountain whitefish in the Muskeg River are unknown as no specimens were captured in the watershed during the present study apart from those taken at the 116

counting fence. Shell (1975) reported capturing no mature mountain whitefish in the Muskeg River during their study, although eight juvenile fish were taken in Hartley Creek during August 1974.

5.4.5.2 Spawning. Mountain whitefish usually spawn in October and early November, the young hatching about March (Paetz and Nelson 1970). No young-of-the-year mountain whitefish \lIJere taken in the Muskeg River in 1976 or 1977, and spawning is not believed to occur in the tributary. Machniak and Bond (in prep.) found no evidence that this species spawns in the Steepbank River. Griffiths (1973), however, reported finding large numbers of young-of-the-year mountain whitefish in the High Hill River as well as the Clearwater River in late August and September. Tripp and McCa t (in prep.) located young-of-the-year mountain whitefish at the mouths of tributary streams in the Athabasca River upstream of the Cascade Rapids"

5.4.5.3 Length-frequency distribution. Mountain whitefish captured in the upstream trap in 1977 varied in fork ength from 198 to 395 mm vvith the majority (57%) being in the 320 to 379 mm size range (Figure ).

5.4.5.4 --==:---.:=----rowth. During 1977, only 15 mountain white- fish were sacrificed for biological analysis (Table 28). These fish ranged in age up to seven years, which appears to be the maximum age for mountain whitefish in the Muskeg River. Machniak and Bond (in prep.) recorded a maximum age of eight years in the Steepbank River. Eight was also the maximum age for stream popu­

lations in Montana (Brown 1971) ~nd in the Sheep River, Alberta (Thompson and Davies 1976). Maximum ages reported for lake populations of mountain whitefish are 17 to 18 in Bow Lake, Alberta (McHugh 1942) and 16 in Rock Lake, Alberta (Lane 1969). The growth rate observed for mountain whitefish from the Muskeg River (Figure 26) was similar to that reported for the Steepbank River (Machniak and Bond in prep.) and ranks among the fastest reported for this species. Stream dwel ling mountain 10

Mountain Whitefish I n = 53 CJ) lL. lL. o 5 0::: W a:l '-l ~ :::) Z

200 220 240 260 280 300 320 340 360 380 400 FORK LENGTH (mm)

Figure 25. Length-frequency distribution for mountain whitefish during the upstream migration in the Muskeg River, 1977. 118

400

E 300 E

I t­ <.9 Z W ~ 200 ~ 0::: MUSKEG RIVER E2 l'l • 1976 // o 1977

1// I 100 I I I

0+ I 2 3 4 5 6 7 8 9 10 II 12 AGE (years)

Figure 26. Growth in fork length for mountain whitefish from the Muskeg River and from several other areas. 1. Montana streams (Brown 1971); 2. Sheep River, Alta. (Thompson and Davies 1976);3. RockLake,Alta. (Lane 1969);and 4 . PY ram i d La ke , Alta. ( Raw son and E1 s e y 1950). Ci r c 1e s represent means and vertical lines the ranges in fork length within age groups for Muskeg River fish. 119 whitefish (this study; Brown 1971; Thompson and Davies 1976) appear to grow faster than most reported lake populations in Alberta (Lane 1969; Rawson and Elsey 1950) although the latter achieve greater maximum ages.

5.4.5.5 Sex and maturity. Sex was determined for only 14 mountain whitefish in 1977, of which five were males (Table 28). Fish of both sexes a ppea r to reach sexua 1 ma tu r i ty a t age 3 in the Muskeg River (Bond and Machniak 1977). Machniak and Bond (in prep.) reported that, in the Steepbank River, the earl iest age of maturity was two years for males and three years for females.

5.4.5.6 Length-weight relationship. Based on 1976 data (Bond and Machniak 1977), the length-weight relationship for Muskeg Rive r mo un t a i n whit e f ish (n = 23, r = O. 977, ran ge 159 to 353 mm) is described by the equation: 10910W = 2.751 (10910L) - 4.301; sb O. 131

5.4.5.7 Food habits. Twenty-nine mountain whitefish stomachs were examined in the field during the two years of the study, of which 21 contained no food 0 The remainder contained traces of insects but only Hemiptera (Corixidae) could be identified. Mountain whitefish are usually reported to be bottom feeders consuming a variety of organisms but mainly immature aquatic insects (Scott and Crossman 1973).

5.4.6 Other Large Fish Species

5.4.6.1 Lake whitefish. Small numbers of lake whitefish were taken at the counting fence in 1976 (Bond and Machniak 1977) and again in 1977 (Table 8). Machniak and Bond (in prep.) also report the movement of small numbers of lake whitefish into the lower Steepbank River during the spring migrations of other species. No young-of-the-year lake whitefish were reported from the Muskeg River in 1976. During 1977, however, 16 whitefish fry 120 were collected in the lower reaches (Table 6). Fourteen of these fish were collected at Site 2 on 12 and 13 June at a time when the Athabasca River was backed up into the Muskeg River. Whitefish fry ranged in fork length from 24 to 38 mm with a mean of 29.5 mm (Table 19) and had fed mainly on aquatic insects (Table 18). Lake whitefish are known to migrate through the AOSERP study area in large numbers during late summer and fall (Bond and Berry in prep.a, in prep. b). Spawning occurs below Mountain and Cascade rapids, upstream of Fort McMurray in the Athabasca River (Jones et al. 1978). The mouth of the Muskeg River is known to be used as a resting area by lake whitefish in September during the spawning migration as AOSERP fishery crews, working on the Athabasca River, reported capturing large numbers in the river mouth at that time. There is no evidence, however, that lake whitefish util ize the Muskeg River for spawning purposes. Machniak and Bond (in prep.) reported no fall migration of lake whitefish into the adjacent Steepbank River.

5.4.6.2 Burbot. Only one burbot was captured at the counting fence in 1977 (Table 8) while three were taken during the 1976 operation (Bond and Machniak 1977). One young-of-the-year burbot (total length 54 mm) was seined from Site 2 (Figure 2) on 10 July 1977, while two yearl ings (105 and 110 mm) were collected from the same area on 25 and 29 May. During May 1976, three immature burbot were captured in minnow traps at Site 2 (Bond and Machniak 1977). Bond and Berry (in prep.b) found large burbot to be common in the Athabasca River during the early spring and reported fry appearing in June. They speculated that burbot utilize the Mildred Lake area of the Athabasca River or areas upstream of it for spawning purposes. Recent evidence suggests that burbot may spawn in the Clearwater River (Figure 1) upstream of its junction with the Christina River (Tripp and McCart in prep.). 121

5.4.6.3 Walleye. Although large numbers of walleye pass through the AOSERP study area in April on their way to spawning grounds (Bond and Berry in prep.a, in prep.b), they do not appear to util ize the Muskeg River for this purpose. Eight walleye were captured in the Muskeg River upstream trap between 10 May and 10 June 1977 while five were taken moving downstream between 12 May and 4 June (Table 8). Small numbers of walleye were also taken during the 1976 fence operation (Bond and Machniak 1977). Post-spawning movements of immature and spent male walleye have been observed in the Steepbank River (Machniak and Bond in prep.) and the MacKay River (Machniak et al. in prep.).

5.4.6.4 Dolly Varden. Three Dolly Varden were recorded at the upstream trap of the Muskeg River counting fence in 1977 (Table 8). The one fish sampled was an immature, three year old male, with a fork length of 202 mm and a weight of 77.0 g. Dolly Varden are common in the headwaters of the Peace, Athabasca, Red Deer, Bow, and Oldman drainages (Paetz and Nelson 1970). Their occurrence in the AOSERP area is rare although several were taken in the Steepbank River (Machniak and Bond in prep.) and Athabasca River (Bond and Berry in prep.b) during 1977.

5.4.6.5 Lake cisco. One lake cisco was captured at the upstream trap on 2 May 1977. Cisco are common in lakes of the Birch Mountains (Turner 1968) and lakes in the southern Athabasca River drainage such as Lesser Slave Lake and Lac la Biche (Paetz and Nelson 1970).

5.4.6.6 Yellow perch: A total of 33 young-of-the-year yellow perch were captured at Sites 1 and 2 of the Muskeg River between 18 July and 13 August 1977. These fish ranged in size from 23 to 44 mm and had a mean fork length of 39.2 mm (Table 29). Of 18 fish for which sex was determined, 10 were males. The stomachs of three young-of-the-year perch contained chironomid and Trichoptera larvae and Ephemeroptera nymphs (Table 18). 122

Perch are thought to have originated from headwater lakes of the Athabasca River drainage and drifted down to the study area. They are commonly found around tributary mouths in the AOSERP study area during July and August (McCart et ale 1977; Bond and Berry in prep.a, in prep.b).

5.4.7 Brook Stickleback

5.4.7.1 Distribution and relative abundance. A total of 308 brook stickleback were collected from the Muskeg River in 1977. Excluding suckers, this species accounted for 48% of all small fish taken (Table 6). Although captured at nine of the small fish collection sites, brook stickleback were particularly abun­ dant in the upper reaches of the watershed. Over two years this spec i es accounted for 84% of a 11 sma 11 fish captu red ups t ream of Site 3 (excluding suckers).

5.4.7.2 rowth. Brook stickleback, captured from the Muskeg River watershed in 1977, ranged from 15 to 71 mm in total length. However, the length-frequency distribution (Figure 27) varied throughout the summer. Stickleback captured in June were predominantly one year old fish, most of which ranged from 36 to 49 mm in total length. Young-of-the-year first appeared in late June and fish of this age class (15 to 35 mm) made up 83.7% of the total catch during July and August. Age-length and age-weight relationships (Table 35) are similar to those reported in the 1976 results (Bond and Machniak 1977) and to those presented by Machniak and Bond (in prep.) for brook stickleback from the Steepbank River. The maximum age recorded for brook stickleback in the Muskeg River is three years. Eight age 3 stickleback were captured during 1976 and 1977 of which only one was a female.

5.4.7.3 Sex and maturity. Female brook stickleback comprised 54% of all fish for which sex was determined (n= 160). However, the sex ratio was not significantly different from unity (X 2 =0.90, P > 0.05) • 123

45 -

- ~ 40 Brook Stickleback n = 262 35 - l::: :- :: 30 - :c (J) LL 25 - I LL 0 ~ :::: 0:: W 20 - :: tIl ~ :::: I:: :::> ::::: z ::::: ::::::::: 15 - :::: :::::::: .:::::: ::::::: 1° < .? 1':::::::,:: 10!- ::. ,:::>;: i':ii }::::: 7: ::: -:.:::. '7:' 5 - ~ r:::::

(;>\\\1[\,:: ::::::~ ~ :;:::: :\\:.\\:::::::}:>:::. ::::::: rm I I , I 10 20 30 40 50 60 70 TOTAL LENGTH (mm)

Figure 27. Length-frequency distribution for brook stickleback from the Muskeg River, 1977. Table 35. Age-length (derived from otol iths) and age-weight relationships, age-specific sex ratios and maturity of small fishes captured in Muskeg River, 1977.

Fork or Total a Length (mm) Weight (g) Females Males Unsexed Samp 1e Species/Age Fish Size % % N % Mature N % Mature Mean S.D. Range Mean S.D. Range

Brook sticklebacka 0+ 6 50 0 6 50 0 10 22 25.3 5.80 15-36 0.2 0.13 0.1-0.5 1 14 48 79 15 52 80 2 31 39.3 5.49 29-49 0.6 0.27 0.2-1.3 2 5 33 100 10 67 100 0 15 51.1 4.18 47-58 1 .4 0.32 1.2-2.0 N 3 1 25 100 3 75 100 0 4 60.0 7.62 54-71 2.3 1.22 1.5-4.1 ~ Totals 26 34 12 72

Lake chub 0+ 2 100 0 0 16 18 28.3 5.39 19-36 0.3 0.16 0.1-0.6 1 13 45 0 16 55 0 3 32 41 . 1 5.60 33-58 0.7 0.42 0.3-2.4 2 1 100 0 0 1 2 60.0 0.00 70-81 2.2 o. 14 2.1-2.3 3 1 50 100 1 50 100 0 2 75.5 7.78 4.3 1 .48 3.2-5.3 4 1 100 100 0 0 1 93.0 10.2 Totals 18 17 20 55

Slimy sculpina 0+ 0 1 100 0 8 9 28.2 6.32 17-36 0.3 0.16 0.1-0.5 1 7 54 0 6 46 0 1 14 43. 1 3.99 37-50 0.9 0.35 0.4-1.7 continued Table 35. Continued.

Fork or Total a Females Males Length (mm) Weight (g) Species/Age Unsexed Sample Fish Size % N % Mature N % Mature Mean S.D. Range Mean S.D. Range

Slimy sculpina 2 2 67 0 33 0 0 3 55.0 1 .00 54-56 2.2 o. 15 2.1-2.4 3 0 100 100 0 1 76.0 4.4 Totals 9 9 9 27 Longnose dace N 0+ 1 100 0 0 3 4 22.3 5.38 16-29 0.2 0.06 0.1-0.2 U"1 1 5 56 0 4 44 0 0 9 43. 1 5.60 34-49 0.9 0.44 0.3-1.5 2 2 100 50 0 0 2 59.0 2.83 57-61 2.3 0.42 2.0-2.5 Totals 8 4 3 15 Pea r 1 dace 1 3 50 0 3 50 0 1 7 38.0 7.28 27-47 0.6 0.27 0.2-1.0 4 0 1 100 100 0 1 103.0 10.9 Totals 3 4 8 Trout-perch 0+ 0 4 100 0 1 5 38.0 2.55 35-41 0.6 0.12 0.4-0.7 1 0 100 0 0 1 43.0 0.9 continued Table 35. Concluded.

Fork or Total a Females Males Length (mm) Weight (g) Unsexed Sample Species/Age Fish Size % % n % Mature N % Mature Mean S.D. Range Mean S.D. Range

Trout-perch 2 0 100 100 0 54.0 1.0 Totals 0 6 7 Ninespine a stickleback N 0" 0+ 100 0 0 0 35.0 0.3 Total 0 0

Fathead minnow 2 0 100 0 56.0 1 .8 Total 0 0 127

The smallest mature fish were observed in the 25 to 29 mm length range for males and in the 30 to 34 mm range for females (Table 36). Results from 1976 showed the smallest mature stickleback to be males in the 20 to 24 mm range while all fish were mature in the 40 to 44 mm group (Bond and Machniak 1977). Sexual maturity is first achieved at age 1 in both sexes (Table 35).

5.4.7.4 Length-weight relationship. The common length-weight relationship for brook stickleback captured from the Muskeg River in 1977 (n::::: 262, r::::: 0.962, range 15 to 71 mm) is descri bed by the equation: lo910W::::: 2.764 (lo910L) - 4.628; sb = 0.049 The value of the exponent (2.764) is intermediate between that calculated for Muskeg River stickleback in 1976 (3.0435) and that reported by Machniak and Bond (in prep.) for Steepbank River fish (2.4260).

5.4.7.5 Spawning. Most brook stickleback in Alberta spawn in late spring and early summer (Paetz and Nelson 1970). Mature and ripe females were captured between 3 and 19 June 1977 at Site 3 (Muskeg River) and Site 7 (Hartley Creek). The first young-of-the­ year (19 mm total length) were collected at Site 10 (Kearl Creek) on 19 June. Many young-of-the-year stickleback were observed in Kearl Creek on 22 June 1978. A sample of 39 fish captured on this date ranged in total length from 11 to 18 mm. By 18 July 1977, young-of-the-yearstickleback, captured at Site 10, ranged in total length from 19 to 27 mm while fish taken 30 July at Site 6 had total lengths of 15 to 24 mm. Stickle­ back fry captured 13 October at Site 6 ranged from 23 to 36 mm in tota 1 length.

5.4.7.6 Food habits. Brook stickleback from the Muskeg River watershed had fed primarily on immature aquatic insects of the orders Diptera, Trichoptera, Plecoptera, Ephemeroptera, and Hemiptera. Other food items included Ostracoda, Nematoda, and Nematomorpha (Table 37). Scott and Crossman (1973) report brook Table 36. Sex and maturity ratios, by size class, for brook stickleback captured from the Muskeg River watershed, 1977. Sex ratios were based only on fish for which sex was determined.

Maturity Total Sex Rat i 0 Length Sample % Unsexed Males Females (mm) Size % Female % Male % Immature % Mature % Immature % Mature

0-14 0 15-19 17 100 0 100 0 76 50 50 20-24 75 100 0 100 0 72 86 14 25-29 47 83 17 100 0 60 63 37

30-34 26 92 8 90 10 15 45 55 N 00 35-39 16 56 44 20 80 13 36 64 40-44 38 0 100 33 67 3 63 34 45-49 25 0 100 0 100 48 52 50-54 8 0 100 0 100 55-59 9 0 100 0 100 22 8 60-64 0 65-69 0 70-74 0 100 100 0

Totals 262 39% 54% 46% Table 37. Food hab its of sma 11 fishes collected from the Muskeg River, 1977.

Species

Food Items Brook stickleback Lake Chub Sl imy Sculpin Longnose Dace Pearl Dace

% Freq.a % No. % Freq.a % No. % Freq.a % No. % Freq.a % No. % Freq.a % No.

Class Insecta

Diptera Chironomidae larvae 40.0 66. I 0.0 0.0 33.0 72.7 28.6 89.3 0.0 0.0 pupae 20.0 16.1 5.6 5.9 0.0 0.0 0.0 0.0 0.0 0.0 Unidentified Dipterans 5.0 1.7 11. I 23.5 13.3 9.1 0.0 0.0 16.7 50.0 Tri choptera 10.0 2.5 0.0 0.0 13.3 3.9 0.0 0.0 0.0 0.0 Plecoptera 5.0 0.8 0.0 0.0 13.3 9.1 14.3 10.7 0.0 0.0 N Ephemeroptera 10.0 3.4 0.0 0.0 6.7 1.3 0.0 0.0 0.0 0.0 \.0 Hemiptera 10.0 1.7 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Insect Remains 30.0 64.7 40.0 42.9 16.7 Hi sce II aneous

Nematoda 5.0 2.5 16.7 41.2 13.3 2.6 0.0 0.0 0.0 0.0 Nematomorpha 5.0 1.7 0.0 0.0 6.7 1.3 0.0 0.0 16.7 50.0 Ostracoda 10.0 3.4 5.6 11.8 0.0 0.0 0.0 0.0 0.0 0.0 Vegetation (algae, seeds) 0.0 5.6 17.6 0.0 0.0 0.0 0.0 Digested Hatter 20.0 27.8 20.0 14.3 0.0 33.3 Debris (sand, gravel) 5.0 5.6 6.7 0.0 0.0 16.7

Total stomachs 24 24 15 8 8 Empty (% of Total) 16.7 25.0 0.0 12.5 25.0 a Percentage frequency of occurrence, based on stomachs that contained some food. 130 stickleback to consume aquatic insects and crustacea as well as Gastropoda, Oligochaetes, Arachnida, and fish eggs. Some individ­ uals are also reported to feed on algae (Winn 1960).

5.4.8 lake Chub

5.4.8.1 Distribution and relative abundance. Excluding suckers, lake chub made up 26% of all small fish captured in the Muskeg River watershed during 1977 (Table 6). This species was present throughout the watershed and was taken at eight of the 10 collec­ tion sites. Chub were taken at Sites 2, 3, 4, 6, 7, 8, and 9 during both years of the study, but were never captured at Sites 5 and 10 (Figure 2). The largest catches of this species were made at Site 7 of Hartley Creek during 1976 (Bond and Machniak 1977) and at Site 3 in 1977. Lake chub occurred in association with brook stickleback at Site f.

5.4.8.2 Age and growth. lake chub, captured from the Muskeg River watershed in 1977, ranged in fork length from 19 to 93 mm (Figure 28), with those in the 25 to 44 mm size range accounting for 82% of the total sample. The length-frequency distribution was not constant throughout 1977, but varied, largely as a result of the disappearance of one year old fish (mostly 34 to 45 mm) in late June and the appearance and subsequent growth of young-of­ the-year in July and August. Young-of-the-year captured in July had a mean fork length of approximately 27 mm (range 21 to 31 mm) while those taken in August averaged about 30 mm in fork length (range 19 to 41 mm). One year old chub, captured in June, ranged from 31 to 49 mm in length. Otolith ages were determined for 55 lake chub, the oldest being a four year old female, 93 mm in fork length (Table 35). Five years appears to be the maximum age for lake chub in the Muskeg River (Bond and Machniak 1977) and in the adjacent Steepbank River (Machniak and Bond in prep.). lake chub live to age 4 in western Labrador (Bruce and Parsons 1976) and age 5 in British Columbia (Geen 1955). 20

Lake Chub I 15 n = 160 en LL LL o 10 a::: w en w ~ ::>z 5

20 30 40 50 60 70 80 90 100 FORK LENGTH (mm) Figure 28. Length-frequency distribution for lake chub from the Muskeg River, 1977. 132

Growth rates, determined from 1977 data (Table 35), are similar to those produced in 1976 (Bond and Machniak 1977) for the Muskeg River as well as to those for lake chub in the Steep­ bank River (Machniak and Bond in prep.). Geen (1955) reported that females grow faster and live longer than males.

5.4.8.3 Sex and maturity. Sex was determined for 85 lake chub, of which 51% were females. Most chub examined were immature. Only five fish, two males and three females, were judged to be sexually mature. Sexual maturity in Muskeg River lake chub appears to occur at age 3 for both sexes (Table 35) (Bond and Machniak 1977). The smallest size at sexual maturity was 62 mm for males and 70 mm for females. Bond and Machniak (1977) found the smallest mature males and females to be in the 55 to 59 and 70 to 74 mm size classes respectively.

5.4.8.4 Length-weight relationship. The length-weight rela- tionship for lake chub (sexes combined) captured from the Muskeg Ri ve r d r a ina g e d uri n g 1977 (n = 146, r = O. 96 1, ran gel 9 to 33 mm) is described by the equation: l0910W = 2.950 (l0910L) -4.892; sb = 0.071

5.4.8.5 Spawning. Lake chub probably spawn in late Mayor early June in the Muskeg River watershed. A mature male (62 mm) was captured on 3 June 1977 at Site 3 (Figure 2) and a spent male (81 mm) and female (70 mm) were taken 19 June at Site 7 (Hartley Creek). The first young-of-the-year were captured on 18 July at Sites 2 (21 to 29 mm) and 3 (27 to 31 mm). Young-of-the-year lake chub were also captured later in the year at Sites 1, 6, and 8 (Figure 2). Machniak and Bond (in prep.) report capturing a ripe lake chub on 7 May in the Steepbank River. Lake chub in the Montreal River, Saskatchewan, spawn in shallow water (about 5 cm) amongst and underneath large rocks at water temperatures of 10°C (Brown 1969). 133

5.4.8.6 Food habits. Muskeg River lake chub fed primarily on aquatic insects, Nematoda, Ostracoda, and plant materials (Table 37). A similar diet was reported for lake chub in the Steepbank River (Machniak and Bond in prep.). Stomachs of lake chub from the Athabasca River contained immature insects of seven orders as well as some fish remains (Bond and Berry in prep.b).

5.4.9 Sl imy Sculpin

5.4.9.1 Distribution and relative abundance. The sl imy sculpin is a common forage fish in gravelly areas of the lower Muskeg River and Hartley Creek. It was taken at Sites 1,2, 3, and 7 during both years of the study, but was never captured at sites upstream of Hartley Creek (Sites 4, 5, 6, 9, and 10). Excluding suckers, this species accounted for 6% of all small fish captured in the Muskeg River in 1977. However, in 1976, sculpins made up 26% of all small fish captured (Bond and Machniak 1977) and are probably more abundant than our 1977 data indicate.

5.4.9.2 Age and growth. Sl imy scu1pins captured in the Muskeg River watershed during 1977 ranged from 7 to 76 mm in total length (Figure 29). Otolith ages were determined for only 27 sculpins, the oldest of which was a three year old male, 76 mm in total length. The maximum age reported for sl imy scu1pins in the AOSERP area is four years (Bond and Machniak 1977; Machniak and ond in prep.). Growth patterns for sl imy sculpins in the Muskeg River (Table 35) (Bond and Machniak 1977) are similar to those reported for the Steepbank River (Machniak and Bond in prep.), the Chandalar River, Alaska (Craig and Wells 1976), and the Mackenzie Delta (de Graaf and Machniak 1977).

5.4.9.3 Sex and maturity. Sex was determined for 18 sl imy sculpins, of which 50% were males. Only one mature sculpin was captured during 1977, that being a three year old male, 76 mm in total length (Table 35). Most sl imy sculpins captured in 1976 5. Slimy Sculpin n = 38 I 4 J: (J)- LL3 LL 0 0::

--> 2l \.AJ ::g r»t~W? rtl .t:- ::E ::>z

10 20 30 40 50 60 70 80 TOTAL LENGTH (mm) Figure 29. Length-frequency distribution for sl imy sculpin from the Muskeg River, 1977. 135 were also immature fish. The sma1 lest size at sexual maturity was 60 to 64 mm for males and 75 to 79 mm for females (Bond and Machniak 1977). In the Steepbank River, the smallest mature scu1pins were males in the 45 to 49 mm size class and both sexes matured at age 2 (Machniak and Bond in prep.).

5.4.9.4 Length-weight relationship. The mathematical relation- ship between total length and body weight for sl imy sculpins (sexes combined) captured in the Muskeg River during 1977 (n = 34, r=0.941, range 17 to 76 mm) is described by the equation: 10g 10W = 3.059 (10910L) - 5.059; sb = 0.194

5.4.9.5 ~pawning. Sl imy sculpins spawn in the early spring over rocky bottoms. Spawning occurred in late April in Valley Creek, Minnesota fry were first observed in June (Petrosky and Waters 1975). Craig and Wells (1976) estimated that sl imy sculpins spawned a week after spring breakup in the Chandalar River drainage, Alaska. Scott and Crossman (1973) state that scu1pins spawn at water temperatures of SoC in northern Saskat­ chewan with the eggs hatching in about four weeks. Slimy scu1pins spawned between late April and mid-May in the Muskeg River, both in 1976 and 1977. In the first year, ripe sl imy sculpins were captured on Sand 9 May at Site 2 and young-of-the-year, 11 mm in total length, were taken on 9 June in Hartley Creek (Bond and Machniak 1977). Young-of-the-year (n=4), 7 to 9 mm in total length, were captured in a drift net near the fence site on 6 and 7 June 1977.

5.4.9.6 Food habits. Slimy scu1pins had fed primarily on Chironomidae and other Diptera larvae, Trichoptera, P1ecoptera, and Ephemeroptera (Table 37). Petrosky and Waters (1975) indicated that the most important foods of Minnesota sculpins we re GammaPU3 (Amph i poda), 0 i pte ra and Tr i chopte ra 1arvae, and Gastropodn. Other food items included Ephemeroptera, I sopoda , Coleoptera adults and larvae, Annel ida, Ostracoda, Nematoda,and sculpin eggs. 136

5.4.10 Longnose Dace

5.4.10.1 Distribution and relative abundance. Longnose dace are characteristic of gravel or bouldery areas of swift-flowing streams (Scott and Crossman 1973). Because adults live in crannies between stones, they are difficult to capture (McPhail and Lindsey 1970) and, therefore, are probably under-represented in seine catches. Only 28 longnose dace were captured from the Muskeg River in 1977. They were most commonly found in the lower reaches of the Muskeg River (Site 2), where 19 specimens were collected (Table 6). Of 73 longnose dace captured in 1976, 72 were found at Site 2 (Bond and Machniak 1977). Over two years, six longnose dace were taken at Site 1, while single specimens were captured at Sites 3, 7, and 8 (Figure 2).

5.4.10.2 Age and growth. Longnose dace captured in the Muskeg River in 1977 ranged from 15 to 62 mm in fork length (Figure 30) and from 0+ to two years in age (Tab 1e 35) . The largest dace taken in 1976 was 89 mm long and three years old (Bond and Machniak 1977).

5.4.10.3 Sex and maturi Only two mature longnose dace were captured in the Muskeg River during the two years of the study. Both were females, one two years old and the other age 3.

5.4.10.4 Length-weight relationship. The mathematical relation­ ship between fork length and body weight for longnose dace (sexes combined) captured in the Muskeg River during 1977 (n=28, r= 0.968, range= 15 to 62 mm) IS described by the equation: lo91oW = 2.827 (lo910L)- 4.722; sb = 0.143

5.4.10.5 Spawning. Bartnik (1970) reported that, in streams of southern Manitoba, longnose dace spawned in late May when daily maximum water temperatures exceeded 15°C, and that spawning occurred over a gravel substrate in water velocities greater, than 45 cm/s. However, in Alberta, dace are reported to spawn from 5. 1:-:-:.:-. Longnose Dace n = 28 I r']~~~~~:~ 4 I CJ) LL LL 3 0 0::w CD 21 \.,.to.)- F:q F",] r:\] -.....J ~ =>z

20 30 40 50 60 70 FORK LENGTH (mm) Figure 30. Length-frequency distribution for longnose dace from the Muskeg River, 1977. 138 early June to mid-August (Paetz and Nelson 1970). Spawning in the Muskeg River probably occurs between late May and early July. The only ripe longnose dace (not yet free running) was taken near the fence site (Site 2) in early May 1976. During 1977, young-of-the­ year (n = 4) were fi rst collected from the mouth of the Muskeg River on 16 July (range 15 to 23 mm). Dace fry were also captured at Site 2 during early August 1976. At that time fork lengths ranged from 18 to 37 mm (Bond and Machniak 1977). Young-of-the­ year dace captured in the lower Steepbank River between 18 July and 2 August 1977 had a mean fork length of 21.5 mm and varied from 17 to 25 mm (Machniak and Bond in prep.).

5.4.10.6 Food habi Stomach analysis of 8 longnose dace revealed the main food in the Muskeg River to be Diptera (Chironomidae) larvae which comprised 89.3% of the identifiable food items (Table 37).

5.4.11 Other Small Fish S ies

5.4.11.1 Pearl dace. Pearl dace are a common forage fish in the AOSERP study area. This species was found throughout the Steep­ bank River watershed and made up 31% of all small fish taken in that stream (Machniak and Bond in prep.). However, they do not appear to be abundant in the Muskeg River as sampling over a two year period produced only 15 specimens. Fourteen of these were captured at Site 2 while one specimen was taken at Site 10, the outlet of Kearl Lake (Table 6) (Bond and Machni~k 1977). The largest pearl dace taken was a four year old male, 103 mm in fork length (Table 35). Peail dace had fed primarily on aquatic insects.

5.4.11.2 Trout- rch Trout-perch are widely distributed through- out the AOSERP study area and are extremely abundant in the Athabasca River (Bond and Berry in prep a, in prep.b). Ripe males and females can be found in the Athabasca River in late April and early May. These fish move into tributaries during May to spawn 139 in late Mayor early June. Trout-perch are known to spawn in the lower reaches of the Ells River as W~ A. Bond collected ripe males and a 10 mm long fry there on 8 June 1977. Spawning also occurs in the lower Steepbank River (Machniak and Bond in prep.) and in the MacKay River (Machniak et al. in prep.). From our observa­ tions in 1976 and 1977, it would appear that the lower Muskeg River (downstream of the fence site) also serves as a spawning area for this species. During 1976, a ripe three year old female was captured near the fence site on 14 May and young-of-the-year (10 to 17 mm) were caught near the mouth of the tributary (Site 1) on 15 June (Bond and Machniak 1977). Five young-of-the-year (35 to 41 mm) were captured at Site 1 on 13 August 1977 (Table 35). The food habits of trout-perch were not examined during the present study. However, in the Athabasca River, trout-perch were observed to have fed primarily on immature aquatic insects, Copepoda and Ostracoda (Bond and Berry in prep.b).

5.4.11.3 Ninespine sticklebac~. One young-of-the-year ninespine st i ck 1eback (35 mm tota 1 1ength) was captured at Site 2 of the Muskeg River on 6 August 1977 (Table 35). This species is rarely found in the AOSERP study area, either in tributaries or in the Athabasca River. Intensive sampling of the Athabasca River with small mesh seines produced only two ninespine stickleback during 1977 (Bond and Berry in prep.b).

5.4.11.4 Fathead minnow. Fathead minnows are common in the upper Athabasca watershed and Wood Buffalo National Park (Paetz and Nelson 1970). They appear to be rare in the Athabasca River down­ stream of Fort McMurray (Bond and Berry in prep.a, in prep. b), although Tripp and McCart (in prep.) captured them in considerable numbers in the Athabasca River upstream of Fort McMurray. Only one fathead minnow was captured in the Muskeg River during the present study. That specimen was a two year old male, 56 mm in fork length (Table 35). 140

5.4.11.5 Spottail shiner. No spottail shiners were taken in the Muskeg River in 1977 and only one was captured during 1976 (Bond and Machniak 1977). They are reported to be common in the Athabasca River, and Bond and Berry (in prep.a, in prep.b) provide information relative to the 1 ife history of this species in the AOSERP study area.

5.5 HABITAT ANALYSIS

5.5. 1 River Mainstem

The mainstem of the ~uskeg River was divided into five reaches on the basis of gradient differences, flow characteristics, substrate, channel form, and other physical features (Table 38, Figure 31). Point samples taken at nine locations during June 1978 provided site-specific information with respect to certain physical and chemical parameters (Table 39) as well as additional information on fish (Table 40) and benthic macroinvertebrates (Table 41). This information collectively defines the aquatic habitat of the Muskeg River mainstem and permits an assessment of fish util ization in each reach.

5.5.1.1 Reach 1M. This reach extends from the confluence of the Muskeg and Athabasca rivers to approximately 0.5 km upstream. Because it 1 ies within the flood plain of the Athabasca River it is subject to periodic flooding by Athabasca River water which can greatly affect its width and depth. This mouth region has 1 ittle gradient (0.3 m/km) , low velocity, and pool-l ike conditions often prevail. The substrate is very homogeneous, comprised mainly of fines (90%) and small gravels (10%). Because Reach 1M is frequently inundated by Athabasca River water it is to be expected that, at some time or other, virtually all fish species occurring in the Muskeg or Athabasca rivers will be found in this area. That this is so is demonstra­ ted by Table 42 which shows that documented fish presence in this reach includes the adults of eight species and the fry and/or juveniles of 20 species. FIGURE 31. MUSKEG RIVER BIO-PHYSICAL MAP LEGEND

KILOMETRES I 0 I 2 3 l:a:a:::a:a:L I :l

REACH SYHBOLS SutsUatf' Hater! ills

GENERAL: fiSH SPECIES

CHANN~l SUBS TRATE I, fines naterials In 0-2 O\'TI siz!? class os SpCl.ic;, Abbreviations gravels - mtltcrldis jr, 2-64 Il\ITI s;ze class larges - l1'..aterials greater than 6lj mm In sl ze c I F ~s I./hi te longf1ose sucker Arctic grayl inc;; NP Northern pike Mountain whi lef!:.h

l~ who tefish

Yellow walleye Example OS Other speCies

Channel

1. long j tudinal P~ofi le::

2, Stope 'Site Spec i ric Stream Symbols

3. CrO~S-5c:ct ion: A point sample si tc WI th biophysical data available Beach seine sampl irtg 51 tc Gl t 1 net sampl ing Angl ing sampling si te Kick sa"'pl log site Beaver dall' ;('ot all shown)

R~ach OOundi) r'l wi th reach SJrvey boun1ary

WS LNS NP AG os BS,AS r 0.1 ul F WS LNS NP AG as K BS,DN r 0.1 u I F WS LNS AG os r 0.3 c I 63 BS,KS,AS

WS LNS AG os WS LNS AG as c I 73 r 0.5 c I 12 LONGITUDINAL PROFILE OF THe: LONGITUDINAL PROFILE OF MUSKEG RIVER HARTLEY CREEK

350 350

WS LNS AG os r <0.1 c I 81

300 300

>­ E Ct: BS.KS WS LNS NP AG as ;3 E Z :::l r 0.2 c I 45 z o o rn ti > W ...J I.IJ

>­ REACH NO. ill ti: 3H....j.. :::l (/) 4H 2H IH REACH NUMBER 3M ..... 5HI"-- ..... 5M 4M ...... WS LNS LW YW NP AG MW os 200 s 0.4 c I 15R I

r <0.1 100 75 50 25 0 ~ 0 DISTANCE FROM ATHABASCA RIVER ( Km) DISTANCE FROM MUSKEG RIVER ( Km ) Table 38. Physical characteristics of the Muskeg River mainstem, 22 to 23 June 1978.

Reach 1M Reach 2M Reach 3M Reach 4M Reach 5M

Distance Upstream of Confluence (km) o - 0.5 0.5 - 8 8 - 13 13 - 75 75 - 88a

Width (m) 16 14 15 13 9 Gradient (m/km) 0.3 4.1 1.5 0.4 2·9 Velocity (m/s)b NO 1.0 0.9 0.4 0.6 Mean Depth (ml b NO 0.6 0.5 0.9 0.2

90 10 40 90 90 10 50 50 5 5 0 30 10 5 5 0 10 0 0 0 .s:- w

) 5 0 10 -0 40 10 20 -5 50 80 65 j 5 10 5

sand clay, sand, gravel clay clay

River Channel Characteristics Thread single single single single single Form straight meander i ng straight, irregular irregular meander straight to irregular Flow Character placid sw i rl i ng to placid placid placid broken a Survey ended at km 88 - not necessarily end of reach. b Based on data from point samples. Table 39. Summary of physical and chemical informati collected at each sampl ing point in the Muskeg River watershed, 22 to 23 June 1

Muskeg River Mainstem Ha rt I ey Creek Kearl Creek Pa rameter Ml M2 M3 M4 M5 M6 M7 M8 M9 Hl H2 H3 Kl K2

Stream width (m) 15 13 14 15 44 13 16 11 9 8 10 5 6 16 Mean depth (m) 0.6 0.6 0.7 0.5 0.3 1.0 0.7 1.5 0.2 0.7 0.4 0.7 0.5 0.4 Velocity (m/s) 1.0 1.0 1.0 0.8 0.9 0.5 0.5 0.2 0.5 0.3 1.1 0.4 0.3 Ni I Substrate (%) Fines «2 I'IITI) 20 o 5 20 30 80 90 100 o 100 5 10 100 100 Grave I s (2 to 64 mm)m 80 50 75 80 70 o o o 10 o 20 15 o o Larges (>64 I'IITI) 0 50 20 o o 20 10 o 90 o 75 75 o o Bedrock o o o o o o o o o o o o o o L-Stable L-Unstable . Bank Stabi Ii ty Stable Stable Stable R-Unstable R-Stable Stable Stable Stable Stable Unstable Stab I e Stab Ie Uns tab J e Q.uaki ng Clay Sand Gravel Gravel L-gravel Sand Clay Clay Clay Sand,cJay Clay,larges Clay Clay Ground Bank Materials Larges Gravel Large~Larges R-sand,clay Gravel .J:'" Riparian vegetation ..s:- Grasses + + + + + + + + + + + + + + Dogwood + + + + + Typhaceae Wi I lows + + + + + + + + + + + + + Alder + + + + + + + Aspen + + Aquatic vegetation Algae Cladophora Cladophora Cladophora Cladophora + + Vascular + + + + + + + + + Time of day (hr) 0900 1145 1215 1245 1315 1445 1600 1130 1030 1430 1600 1630 1300 0930 Water temperature (OC) 14.0 NO 14.5 NO 14.0 NO 14.0 13.0 13.0 14.0 14.0 13.0 14.0 16.0 pH 8.5 8.5 NO 8.5 NO 8.0 8.0 8.5 8.5 8.0 8.0 8.0 8.0 Conductivity (~mhos/cm@25°C) 305 NO 305 NO 310 NO 300 300 260 200 190 195 220 160 oi sso I ved oxygen (mg/ I) 9 NO 9 NO 9 NO 8 7 8 9 10 9 8 8 Table 40. Number of fish captured in small mesh seines at each sampl ing point in the Muskeg River watershed, 22 to 23 June 1978.

Muskeg River Mainstem Hartley Creek Kearl Creek Total Species M1 M2 M3 M4 M5 M6 M7 M8 M9 Hl H2 H3 K1 K2 Numbe r %

Arc tic gray 1 in g 0 21 4 22 7 NO 0 NO 0 0 0 0 0 0 54 23.8

Northern pike 0 0 4 0 0 NO 0 NO 0 0 0 0 0 0 4 1 .8

Trout-perch 0 0 0 0 NO 0 NO 0 0 0 0 0 0 0.4

Longnose dace 2 0 0 0 NO 0 NO 0 0 0 0 0 0 3 1.3 .:::- White sucker 0 2 NO NO 0 0 0 0 0 0 6 2.6 V1

Longnose sucker 0 0 0 0 0 NO 0 NO 0 0 0 0 2 0.9

Brook stickleback 0 0 0 0 NO NO 23 30 0 21 39 116 51 . 1

Lake chub 0 0 0 0 20 NO NO 18 0 0 0 0 40 17.6

Pea rl dace 0 0 0 0 NO 0 NO 0 0 0 0 0 0 0.9

Total 5 21 11 23 29 NO 3 NO 24 49 0 22 39 227 99.9 Table 41. Percentage composition by numbers for the major benthic macro-invertebrate groups taken at each sampl ing point in the Muskeg River watershed, 22 to 23 June 1978

Muskeg River Mainstem Ha rt 1ey Creek Kea rl Creek

M1 M2 M3 M4 M5 M6 M7 M8 M9 H1 H2 H3 K1 K2

Chironomidae 25 6 10 ND 10 ND ND 40 92 18 47 46 29 44

Epheme ropte ra 11 20 47 ND 66 ND ND 0 4 19 7 8 < 1 2

Pl ecoptera 0 0 0 ND 0 ND ND 0 0 0 0 0 0 0

Tri choptera 9 21 14 ND 3 ND ND 0 2 13 21 7 2 < 1

..t:- Simuli idae 0 0 O ND 0 ND ND 0 0 0 0 0 47 0 (j'\

Oligochaeta < 1 < 1 ND 1 ND ND 0 < 1 2 7 12 5 7

Other taxa 55 53 27 ND 20 ND ND 60 48 18 27 16 48

Number Counted 453 683 985 ND 897 ND ND 10 1393 738 431 303 154 657

Percentage of Sample Counted 50 25 13 ND 25 ND ND 6 13 25 13 13 25 25 Table 42. Documented distribution of adult and young fish in the Muskeg River mainstem based on catch data obtained in 1976-1978, and on reports by other individua1so

Reach 1M Reach 2M Reach 3M Reach 4M Reach 5M Species Fry/ Fry/ Fry/ Fry/ Fry/ Adults J 01 J J J Adults J 01 uven I es Adults uvenl°1 es Adul ts uvenl°1 es Adults uvenl°1 es uvenl es

Lake whitefish + + + + + Mounta in wh i tefi sh + +a + + Lake cisco + Arct i c grayl i ng + + + + + + + + Dolly Varden + + .,J:- Northe rn pike + + + + + + +c + ~ Pearl dace + Redbe 11 y dace +b Lake chub + + + + + + + + Longnose dace + + + + Emerald shiner +a Spottai 1 sh i ne r + Fathead minnow +b + White sucker + + + + + +c + + Longnose sucker + + + + + +c + con t i nued •. 0 Table 42. Concluded.

Reach 1M Reach 2M Reach 3M Reach 4M Reach 5M Species F-ry/ Fry/ Fryl Fry/ Fry/ Adults Juveniles Adults Juveniles Adults Juveniles Adults Juveniles Adults Juveniles

Trout-perch +c + + Bu rbot + + Broo k s tic k 1e ba c k + + + + + + + + + Ninespine stickleback + Slimy sculpin + + + + Walleye + + + + J:- Ye llow pe rch + + ex> a Reported by Bond and Berry (inprep.a). b Reported by Bond and Berry (in prep.b). c Reported by Shell Canada Ltd. (1975) in mouth of Hartley Creek. 149

Reach 1M has little potential for spawning by any species encountered within the Muskeg River. Although white suckers, longnose suckers, northern pike, Arctic gray1 ing, and (probably) trout-perch are known to migrate through this reach in April and May in order to reach spawning areas upstream in the Muskeg River, none of these species, with the possible exception of northern pike, is bel ieved to spawn in Reach 1M. The placid water conditions of this reach appear to provide excellent rearing habitat for the young of most species. Young-of-the-year suckers, brook stickleback, sl imy sculpin, yellow perch, mountain whitefish, Arctic grayl ing, and ninespine stickleback have all been captured from this reach. These young-of-the-year probably came from upstream areas of the Muskeg River as well as from the Athabasca River. Although no benthos was collected from Reach 1M during the study, the small particle size of the substrate and the lack of substrate diversity would indicate a sparse bottom fauna. Thus, poor feeding conditions obtain for such fish as lake whitefish, mountain whitefish, white ~nd longnose suckers, that feed predomi­ nantly on benthic invertebrates. On the other hand, the large numbers of young-of-the-year fish (especially suckers) found in the mouth area of the Muskeg River, would provide excel lent forage for piscivorous species such as northern pike and walleye. Reach 1M appears to be of importance to lake whitefish, and probably also walleye, in terms of providing resting areas during upstream spawning migrations in the Athabasca River (Bond and Berry in prep.a, in prep.b). The potential of Reach 1M as an overwintering area is unknown.

5.5.1.2 Reach 2M. Reach 2M, the canyon of the Muskeg River, extends from km 0.5 to approximately km 8 (Figure 31). Through this region the stream has cut a deep, narrow, tortuously meandering channel into Waterways 1 imestone, by which it is largely confined. The gradient in this reach is generally steep (4.1 m/km) and the water velocity is rapid (1.0 m/s). However, the river is stepped in such a way that riffles and pools alter­ nate providing a wide range of habitat types. In the upper 150 canyon (Figure 6), short rapid riffles alternate with long shallow pools while in the lower end of Reach 2M (Figure 5) the pools are generally smaller and separated by long gravel riffles. The substrate in Reach 2M is mostly gravels (50%) and 1arges (30%) but some silt occurs with gravel in pool areas and the stream passes occasionally over 1 imestone ledges. Substrate particle size tends to decrease from the upper to the lower end of this reach. Riparian vegetation was estimated as 65% deciduous trees (aspen, alder), 20% spruce, and 10% deciduous shrubs (willows). In many areas, however, where the river contacts the limestone cl iffs 1 ittle riparian vegetation is found. At low water levels, much gravel and bounder is exposed and the riparian vegetation seldom overhangs the stream. Benthic samples were taken at two sites (M1 and M2) within Reach 2M. At the lower site, where the substrate consisted primarily of small gravels (Table 39), the most abundant benthic forms were Chironomidae, Ephemeroptera, and Trichoptera which made up 25, 11, and 9% of the sample respectively (Table 41). Trichop­ tera (21%) and Ephemeroptera (20%) were the most common inverte­ brates taken at point M2 where the substrate consisted of gravels and 1arges in approximately equal amounts. The diversity of substrates found in Reach 2M and the combination of pools and riffles lead one to expect a great diversity of benthic forms in this region. This is confirmed by the work of Barton and Wallace (in prep.) who identified 166 invertebrate taxa within Reaches 2M and 3M. The documented fish fauna of Reach 2M includes adult fish of 11 species and fry and/or juveniles of 17 species (Table 42). Spawning potential in Reach 2M is excellent for white suckers which have been seen spawning over gravel riffles at the lower end of the reach. The riffle areas of this reach also provide excellent spawning areas for longnose suckers, Arctic grayl ing, longnose dace, sl imy scu1pins, lake chub, and trout­ perch. Areas suitable for northern pike and brook stickleback spawning are few and limited to side sloughs out of the main current. Although the Muskeg River in Reach 2M contains areas 151 suitable for spawning of mountain whitefish, lake whitefish, and walleye, these species are not believed to spawn within this watershed. Rearing for Arctic grayling is excellent in the shallow gravel riffles and shallow pools found in Reach 2M. Although young-of-the-year grayling have been taken all along this reach, they are apparently most abundant in the middle and upper portion. At low water levels, small back eddies develop along the shoreline of 2M in which large numbers of sucker fry are found during June, July, and August. Rearing in this reach is also good for longnose dace and sl imy sculpins. Within the canyon, large grayl ing were found immediately below riffle areas at the extreme upstream ends of pools. Northern pike are extremely limited by the current in this reach and usually occur in quiet pools and back eddies. Most pike that enter the Muskeg River in the spring probably remain in the downstream areas of Reach 2M, although some are known to traverse the canyon. The extremely abundant bottom fauna in Reach 2M provides an excellent food source for Arctic grayl ing, longnose and white suckers, and virtually all fish species found in this section of the river. Forage for northern pike and walleye is also excellent in the form of sucker fry. Of the fish species captured at the Muskeg River upstream trap, only Arctic grayl ing, northern pike, white suckers, and longnose suckers have been recorded upstream of Reach 2M (Table 42). This suggests that such species as lake whitefish, walleye, lake cisco, Dolly Varden, and burbot, although entering the Muskeg River in small numbers, probably do not ascend the tributary for any great distance. The extent of overwintering in Reach 2M is unknown. Measurements taken near the downstream end of this reach in February 1975 (NHCL 1975) indicate a water depth of 0.1 m under 0.4 m of ice and a discharge of 0.3 m3/s suggesting that over­ wintering may be possible for some fish in this reach.

5.5.1.3 Reach 3M. Reach 3M comprises approximately 5 km of stream between km 8 and km 13 (Figure 31), and is a transitional zone between the low gradient Reach 4M and the steep gradient Reach 2M. The top of this reach marks the approximate 152

point at which the Muskeg River leaves the flat central portion of its watershed and begins to cut through the McMurray Oil Sands formation that overlies the Waterways 1 imestone referred to in Reach 2M. Although the gradient is moderate (1.5 m/km) and the

stream velocity rapid (0.9 m/s) 9 the river flows fairly smoothly over a uniform substrate of fines (40%, mostly sand) and small gravels (50%). The current increases and the substrate material gets coarser in the vicinity of point sample M3 (Figure 31) as the stream begins to enter the canyon. Riparian vegetation in Reach 3M was mostly deciduous trees (40%) and shrubs (50%). The stream banks in the upper parts of the reach were largely sand and clay and were easi ly eroded. However, at the downstream end of the reach, stable banks of gravel and larges occurred. Benthic samples taken at sites M3 and M5 revealed a diverse invertebrate fauna consisting largely of Ephemeroptera, Trichoptera, and Chironomidae (Table 41), As mentioned previously, Barton and Wallace (in prep.) identified 166 invertebrate taxa between point M5 and point Ml. Because the substrate particle size is smaller in most of the region between sites M5 and M3 than it is at those sites, the invertebrate fauna is probably less diverse through much of this reach than is indicated by Barton and Wa 11 ace. The fish fauna of Reach 3M appears to be limited to eightor nine species (Table 42). Spawning potential for Arctic grayling is excellent in the coarse gravel areas found at the upstream and downstream ends of this reach, but is low in areas of sandy substrate. Gravel areas also provide good spawning sites for longnose and white suckers, longnose dace, lake chub and slimy sculpins. The occasional area of quiet water with emergent vegetation provides good spawning habitat for pike and brook stickleback, but such areas are few in number. Rearing for Arctic grayling, sl imy sculpin, and longnose dace is good in shallow. gravelly areas of Reach 3M. Back-eddies, out of the main channel, which become choked with weeds afford protection for young-of-the-year suckers, pike, and brook stickleback. The abundant benthic fauna, especially at the upper and lower ends 153 of the reach, provides an excellent food source for all fish species. The only piscivore taken in Reach 3M was northern pike for which abundant forage occurs during the summer, especially in the form of young suckers. No winter flow data are available for Reach 3M and it is not known if overwintering conditions exist in this area.

5.5.1.4 Reach 4M. Reach 4M extends from km 13 to km 75, running through the large central area of the watershed. This area is flat, poorly drained, and covered with marshland and treed muskeg. This low gradient (0.4 m/km) reach is typically deep, slow moving (0.4 m/s) and cluttered with large numbers of beaver dams. The occurrence of such dams increases greatly upstream of Hartley Creek (Figure 31). Through most of the reach the substrate is composed of sand, silt, mud, and organic debris with occasional boulderyareas. The low clay banks are vegetated with willows and grasses. During periods of flood, water overflows these banks through much of the reach and the limits of the stream become almost impossible to define. Benthos was sampled only at point M8 (Figure 31) in which area the maximum depth exceeds 1.5 m. Chi ron­ omidae comprised 40% of this sample (Table 41). Barton and Wa1 lace (in prep.) identified 81 invertebrate taxa from the same area and found that Chironomidae (56%) and 01igochaeta (11%) accounted for the majority of animals taken. They captured virtually no Ephemer­ optera, Plecoptera, or Trichoptera. Thus, the very uniform physical conditions to be found in Reach 4M are reflected in the benthic community by a greatly reduced species diversity. The fish community in Reach 4M is also severely restrict­ ed. There appears to be little spawning potential in this area for any species other than brook stickleback. Although large numbers of sucker fry have been captured in Reach 4M downstream of, and for approximately 300 m upstream of the confluence of Hartley Creek, it seems more likely that these fish were spawned in Hartley Creek than in the Muskeg River itself. Ripe pike and longnose and white suckers have been reported from the mouth of Hartley Creek (Shell Canada Ltd. 1975). Reach 4M of the Muskeg 154

River, with its reduced current and abundance of small chironomid larvae provides very good rearing habitat for young-of-the-year fish. However, the only fry captured in large numbers in this area were suckers and brook stickleback. Small numbers of young­ of-the-year grayling, pike, and lake chub have been taken from Reach 4M downstream of Hartley Creek. The lower regions of Reach 4M, between Hartley Creek and the lower boundary of the reach, provide summer feeding grounds for adult Arctic grayl ing. Grayling in this region are said to be found where water up to 1 m deep flows with moderate current over beds of macrophytes and sand (Dr. D. Barton, University of Waterloo, verbal communication with W. A. Bond, April 1978). Winter measurements taken just upstream of the mouth of Hartley Creek and near point 8M (Figure 31) by NHCL (1975) indicate a water depth of from 0.5 to 0.7 m in mid­ February. This would suggest the potential exists for overwinter­ ing in Reach 4M, depending on dissolved oxygen levels. Although no winter fish sampl ing was conducted in this study, yearl ing white and longnose sucker~ have been captured in early spring at the mouth of Kearl Creek which suggests overwintering by the young of these species. Brook stickleback, the dominant fish species in Reach 4M, is undoubtedly a year round resident of this reach.

5.5.1.5 Reach 5M. Upstream of Reach 4M the gradient of the Muskeg River increases (2.9 m/km) , but the flow is reduced in most areas by beaver dams. The substrate in Reach 5M consists mainly of sands and silts (90%) and the clay banks are well vegetated with willows (80%) and grasses. The only point sample in this reach was taken at the upper 1 imit of our survey (Figure 3, Table 39) at a site that was atypical of the reach as a whole. At this site (Figure 3) the stream bed was strewn with moss­ covered boulders and thick mats of CZadophora were found. The benthic fauna was dominated numerically by Chironomidae which made up 92% of our sample (Table 41). Barton and Wallace (in prep.) identified only 78 invertebrate taxa from the vicinity of point M9. The samples showed Chironomidae (31%) and Oligochaeta (27%) 155

to be the most abundant benthic invertebrates in this area. The fish fauna at point M9 is restricted to brook stickle­

back and lake chub which are~ undoubtedly, year round residents of this reach. Movement of fish between Reach 5M and downstream areas is restricted by beaver dams, although one young-of-the-year sucker fry did manage to ascend as far as site M9.

5.5.2 Hartley Creek and Kear1 Creek Hartley Creek was divided into five reaches on the basis of gradient differences, flow characteristics, substrate, and channel form (Table 43, Figure 31). Point samples were taken at only three sites, one each in Reaches 1H, 3H, Gnd 5H (Figure 31). At these locations, site-specific information was collected with respect to certain physical and chemical parameters (Table 39), fish (Table 40), and -thic macro-invertebrates (Table 41). This information, in combination with fish data gathered in 1976 and 1977, permits an assessment of fish utilization in each reach and defines the aquatic habitat of this tributary. Point samples were also taken at two locations on Kear1 Creek (Figure 31). One of these was situated just upstream from the Muskeg River and the other at the outlet from Kearl Lake.

5.5.2.1 Reach 1H. The lower 3.0 km of Hartley Creek are charac- terized by relatively low gradient (1.0 m/km) , slow current (0.3 m/sl~ and sandy substrate (Table 43). The riparian vegeta­ tion is mostly willows and grasses, and beaver activity is evidenced by the presence of several beaver dams (Figure 31). The most abundant invertebrate groups in samples taken at point H1 (Figure 31, Table 41) were Chironomidae (18%), Ephemeroptera (19%), and Trichoptera (13%). Six species of fish have been documented from Reach 1H (Table 44). Ripe Arctic grayl ing, white suckers, and longnose suckers have been reported from this reach by Shell Canada Ltd. (1975) but were not captured during the present study. The spawning potential of Reach lH is considered to be poor for these species. However, they probably migrate through it to reach Table 43. Physical characteristics of Hartley Creek, 22 to 23 June 1978.

Reach 1H Reach 2H Reach 3H Reach 4H Reach 5H

Distance Upstream of Confluence (km) 0-3.0 3.0-6.7 6.7-8.2 8.2-14.7 14.7- 17. Ob Width (m)a 8 NO 10 NO 5 Grad i ent (m/km) 1.0 2.9 5.1 2.3 NO Ve10ci ty (m/s)a 0.3 ND 1 • 1 NO 0.4 Mean Depth (m)a 0.7 NO 0.4 NO 0.7 Substrate Composition (%) Fines « 2 mm) 100 60 10 70 80 Gravels (2 to 64 mm) o 30 20 30 10 La rges (> 64 mm) o 10 70 o 10 Bedrock o o o o o Riparian Vegetation (%) 10 10 \n Coniferous Trees o 10 o (T'\ Dec i d uous Trees 10 60 70 60 10 Deciduous Shrubs 70 20 20 30 80 Grasses 20 10 o o 10

Bank Materials clay, sand sand, clay clay, gravel sand, clay clay I arges River Channel Characteristics Thread single single single single single

Form irregular t irregular, straight irregular, i rregu 1ar, meandering meandering meander i ng meander i ng Flow Character placid moderate swift moderate placid a Based on data from single point samples. b End of survey, but not necessari Iy end· of reach. Tab1e 44. Documented distributjon of adult and young fish in Hartley Creek and Kear1 Creek based on catch data obtained in 1976-1978, and on reports by other individuals.

Hartley Creek Kearl Creek Reach 1H Reach 2H Reach 3H Reach 4H Reach 5H Point K1 Point K2 Fry/ Fry/ Fry/ Fry/ Fry/ Fry/ Fry/ Adults Juveniles Adults Juveniles Adults Juveniles Adults Juveniles Adults Juveniles Adults Juveniles Adults Juveniles

Mountain whltefisha Arctic grayl ing +b + + + + + + Northern pike +b

Pearl dace + \on -...... J Lake chub + + + + + + + Longnose dace + + + White sucker +b + + + + + Longnose sucker +b + + +c + + Brook stickleback + + + + + + + + + + + + Slimy sculpin + + + + + a Reported by Shell Canada Ltd. (1975) but location not given. b Ripe specimens trapped near mouth of Hartley Creek in May 1973 (Shell Canada Ltd. 1975). c Reported by Dr. R. Hartland-Rowe (University of Calgary, verbal communication with W. A. Bond, May 1976). 158 suitable spawning areas further upstream. Shell also reported ripe northern pike from the mouth of Hartley Creek. Some pike spawning may occur near the mouth of the tributary but beaver dams probably 1 imit the upstream movement of this species. Reach lH has good spawning potential for brook stickleback and lake chub. The beaver ponds and placid water conditions of Reach lH provide favourable rearing conditions for young fish but the fry of only four species were captured in this region (white and longnose suckers, lake chub, and brook stickleback). Winter measurements, taken in Hartley Creek just upstream of its con­ fluence with the Muskeg River in February 1975, showed a water depth of 0.5 m (NHCL 1975). This would suggest some overwintering potential in this area, depending on oxygen levels.

5.5.2.2 Reaches 2H, 3H and 4H. Upstream of Reach lH the gradient of Hartley Creek increases for approximately the next 12 km. This region has been divided into three reaches because of the extremely steep nature of the stretch between km 6.7 and km 8.2 (Table 43). The stream gradient within Reach 3H is 5.1 mlkm and the water velocity is more than 1 m/s. The substrate consists mainly of larges with smaller areas of gravel. Because of the very rapid current and large substrate size the spawning potential of much of Reach 3H may be limited, although some Arctic grayl ing may spawn there. No fish were collected from most of the reach because the nature of the substrate and the water velocity made seining very difficult. The larger boulders in this reach, however, do provide holding places for Arctic grayl ing which can be captured from this area throughout the summer by angling (Mr. R. Crowther, International Environmental Consultants, Calgary, Alberta, telephone communication with W. A. Bond, January 1979). Upstream and down­ stream of Reach 3M, the stream gradient is 2.3 to 2.9 m/km and the velocity is somewhat less than 1 m/s. Conditions in Reaches 2H and 4H are very similar. The substrate is largely sand (60 to 70%) with gravel riffles accounting for about 30%. The gravel in Reaches 2H and 4H provide excellent spawning potential for Arctic grayl ing, suckers, sl imy sculpins, and longnose dace. Although no 159 fish sampling was conducted in Reach 4H, fry of seven species were taken in Reaches 2H and 3H (Table 44). Sucker, lake chub~ and brook stickleback fry were captured in quiet back eddies and in beaver ponds, while young longnose dace, slimy sculpin, and Arctic grayling were taken in gravel riffles. Overwintering potential apparently occurs within Reaches 2H, 3H, and 4H, Not only have yearling and two year old white and longnose suckers been captured in this area, but young-of-the-year grayl ing were observed through the ice at point H2 on 31 October 1976 (Dr. D. Barton, University of Waterloo, verbal communication with W. A. Bond, December, 1976).

5.5.2.3 Reach 5H. Upstream of Reach 4H, Hartley Creek flows through flat, poorly drained terrain in which there is little gradient and pool conditions prevail. Beaver activity is extensive through this reach. Reach 5H was sampled only at its extreme downstream end (Figure 31) and the fish fauna at this point (Table 44) is probably more similar to that in Reach 4H than to that in most of Reach 5H.

5.5.2.4 Kearl Creek. Point samples were taken at two locations on this tributary (Figure 31). Point Kl was situated near the mouth of the stream, a short distance upstream of the confluence of the tributary with the Muskeg River (Figure 4). This part of Kearl Creek 1 ies within the flat, poorly drained area typical of most of Reach 4M of the Muskeg River. There is little gradient at this point. The stream is fairly deep (0.5 m), slow flowing (0.3 m/s), and typical pool conditions prevail. The clay banks are vegetated with willows and grasses and show some signs of erosion. The substrate consists of clay and silts and is 1 ittered with organic debris. Only four species of fish were taken at point K1 (Table 44). The capture of yearl ing white and longnose suckers in June suggests that such fish can and do overwinter in this region. However, the area appears to have little potential for sucker spawning. Fish movement into the area from downstream is probably restricted by the increased incidence of beaver dams 160 upstream of Hartley Creek (Figure 31). Brook stickleback and lake chub, the only resident fish in the upper t1uskeg River watershed, were also captured at point Kl. Kearl Creek was also sampled at its upstream end where it exits from Kearl Lake (Figure 31). Marsh-l ike conditions occurred at this site as the stream passed through an area of reeds and sedges. There was no perceptible current. and the substrate consisted of silts covered with organic debris. The benthos collected was largely Chironomidae and 01 igochaeta (Table 41). Batton and Wallace (in prep.) identified 87 inverte­ brate taxa from this location with 01 igochaeta accounting for 48% of their sample. The fish fauna at this site consists almost exclusively of brook stickleback which inhabit Kearl Lake in large numbers. The reedy area of point K2 is bel ieved to function as a spawning and nursery area for this species as large numbers of newly-hatched fry were observed here on 23 June 1978. The only other fish captured at this location was a 103 mm, four year old pearl dace taken in 1977 (Tab le 35). 161

6. CONCLUSION The Muskeg River provides spawning habitat for white suckers, 10ngnose suckers, and Arctic grayling that migrate into the tributary from the Athabasca River in late April and early May. White suckers were observed spawning over gravel riffles in the lower 1 km of the Muskeg River. No other precise spawning areas were located although potential spawning sites for these species occur and young-of-the-year of these species were captured throughout the lower 35 km of the Muskeg River and in the lower 15 km of Hartley Creek. Northern pike and trout-perch from the Athabasca River also appear to spawn within the Muskeg River watershed to a limited extent. White and longnose suckers began to leave the Muskeg River by mid-May, approximately two to three weeks after the commencement of the upstream runs. This exodus continues at least through July and probably throughout the summer. Sucker fry began to emerge by the end of May and drifted out of the watershed during the summer. Small numbers of young-of-the-year suckers apparently overwintered within the Muskeg River watershed. However, most fry, as well as the adults, probably overwinter in Lake Athabasca. Suckers are seldom highly ranked when considered in terms of their direct importance to man. However, they occur in large numbers in the lower Athabasca drainage and are known to spawn in several other tributaries in addition to the Muskeg River. Because of their high fecundity, an enormous amount of sucker biomass is contributed annually to the system. Although the significance of this contribution has not been quantified, it is i likely that such piscivorous fishes as pike, walleye, burbot, and goldeye depend on young suckers for a large part of their annual food intake. Arctic grayling, unl ike suckers, did not leave the Muskeg River following the spawning period but remained in the tributary throughout the summer to feed. Grayling were never observed in the Muskeg River upstream of the confluence of Hartley Creek. However, they are known to occur as far upstream 162 as Slte 8 in Hartley Creek. Between Sites 3 and 4 of the Muskeg River, grayling occurred in areas where water up to 1 m deep flowed with a moderate current over beds of macrophytes and sand. Within the canyon, most grayl ing occupied the upstream end of the pools. Although it was not possible to monitor the fall down­ stream migration this event probably occurs just prior to freeze­ up as was reported in the Steepbank River (Machniak and Bond in prep.). Large grayl ing still occurred in the Muskeg River on 13 October 1978. Young-of-the-year grayl ing are bel ieved to over­ winter within the Muskeg River watershed and join the migrant population at the end of their second summer. Thus, the Muskeg River provides not only spawning habitat for Arctic grayl ing, but also summer feeding areas for adults and juveniles, rearing and overwintering sites for young-of-the-year. It is also possible that, in the tributary, grayl ing (especially young-of-the-year) are less susceptible to predation and severe environmental fluctuations than would be the case in the Athabasca River, thereby increasing their sU,rvival rate. The grayl Ing population in the Muskeg River does not appear to be as large as that in the Steepbank River. Its proximity to the proposed Alsands project places this population in considerable jeopardy, This species is highly susceptible to habitat disturbances and is easily over-exploited by angl ing. Without adequate protection of its habitat and appl ication of a sound fisheries management program, this population will be quickly lost. The Muskeg River provides some summer feeding for northern pike and small numbers of mountain whitefish, walleye, and lake whitefish. Pike have been observed as far upstream as Hartley Creek, but most are bel ieved to remain in the lower reaches of the tributary throughout the summer. We found no evidence to suggest that mountain whitefish, lake whitefish, or walleye uti 1 ize the Muskeg River for spawning purposes and most of these fish are thought to leave the Muskeg River before freeze-upo The mouth region of the Muskeg River may be important as a resting area for walleye and lake whitefish during spawning migrations 163

in the Athabasca River, and may provide nursery areas for young­ of-the-year fish of several species. The resident fish fauna of the Muskeg River watershed consists largely of four species of forage fish. The fauna of the upper watershed is dominated by brook stickleback, a large population of which occupies Kear1 Lake Lake chub are most abundant in the mid-reaches of the watershed (Sites 3 and 7) and were found in association with brook stickleback at Site 6. 51 imy sculpin and longnose dace were most common in gravelly areas of the lower Muskeg River and Hartley Creek (Sites 1, 2, 3, and 7). Pearl dace, which are abundant in the adjacent Steepbank River, apparently occur only in small numbers in the Muskeg River watershed. Several species of fish considered to be more typical of the Athabasca River than of the Muskeg are sometimes taken in the extreme lower reaches of the tributary. Their presence in the Muskeg River is probably incidental and it is felt that they seldom proceed more than 1 km upstream in the tributary. 164

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McCart, P.J., D. Tripp, P.T.P. Tsui, W. rant and R. Green, in prep. Basel ine study of the water qual ity and aquatic resources of the MacKay River Albe tao Prep. for Syncrude Canada Ltd. by Aquat c Environments Ltd. 200 pp.

McHugh, J.L. 1942. Growth of Rocky Mountain whitefish. J. Fish. Res. Board Can. 5: 7-343.

McPhail, J.D., and C C Lindsey. 970. reshwater fi hes of northwestern Canada and Alaska. Bull. ish Res. Board Can. No, 173. 381 pp.

Machniak, K. The effects of hydroe ectric deve opment on the bio ogy of northern fishes (reproduction and population dynamics). I. Northe pike Esox (Linnaeus). A 1 terature rev ew and bibl iography. Fish. Mar Servo Res. Dev. Tech, rt. 528. 82 pp.

Machniak, K., and W.A. Bond. in prep. An nten ive tudy of the fish fauna of the Stee k River wa shed of no th- eastern Alberta. rep. fo the Albe Oil Sands Environmental rch Program by Isner sand Environment F sheries and rine Service, AOSERP P 5.2. 191 pp.

Machniak, K., W~ M R Or D. , D. M in prep. Fisher es and aquatic habitat in the MacKay Rive watershed of northweste Prep. for Alberta Oil Sands Environmenta Research Program by Fisheries and Environment Ca da Fisheries and Marin~ Service AOSERP Project WS .3. 1

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Northwest Hydraul ic Consu tants Ltd. 1974. Northeast Alberta regional plan, water resources study, sector 1. Prep. for Ekistic Des Consultants Ltd. 28 pp. and appendices. 169

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Olsen, D.E., and W.J. Scidmore. 1963. Homing tendency of spawning white suckers in Many Point Lake, Minnesota. Trans. Amer. Fish. Soc. 92:13-16.

Paetz, M.J., and J.S. Nelson. 1970. The fishes of Alberta. Queen1s Printer, Edmonton. 282 pp.

Pennak, R.W. 1953, Freshwater invertebrates of the . The Ronald Press Co., New York. 769 pp.

Petrosky, C.E., and T.F. Waters. 1975. Annual production by the slimy sculpin in a small Minnesota trout stream. Trans. Amer. Fish. Soc. 104:237-244.

Pettit, S.W., and R.L. Wallace. 1975. Age, growth, and movement of mountain whitefish (Prosopium ~iZZiamsoni Girard) in the North Fork Clearwater River, Idaho. Trans. Amer. Fish. Soc. 104:68-76.

Pinsent, M.E. 1967. A comparative limnological study of Lac la Biche and Beaver Lake, Alberta. M.Sc. Thesis. Univ. Alberta. 147 pp.

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8 APPENDICES

8.1 MAXIMUM AND MINIMUM DAILY WATER TEMPERATURES RECORDED AT THE MUSKEG RIVER COUNTING FENCE, 1977. 173

Table 45. Maximum and minimum daily water temperatures recorded at the t4uskeg River counting fence, 1977.

Daily Water Temperatures (OC) Date Maximum Minimum Mean

Apri I 28 9.0 29 8.0 30 5.0 4.0 4.50 May 1 6.5 4.0 5.25 2 7.5 4.0 5.75 3 7.0 5.5 6.25 4 9.0 6.0 7.50 5 9.5 6.5 8.00 6 10.0 6.0 8.00 7 11.5 5.0 8.25 8 11.5 8.0 9.75 9 11.5 7.0 9.25 10 14.0 9.0 11.50 11 14.0 10.0 12.00 12 12.0 10.0 11.00 13 12.0 8.0 10.00 14 12.0 7.5 9.75 15 11.0 7.0 9.00 16 10.5 9.0 9.75 17 8.5 5.5 7.00 18 7.0 5.0 6.00 19 7.0 5.5 6.25 20 8.5 6.0 7.25 21 9.5 5.0 7.25 22 10.0 6.0 8.00 23 12.0 7.0 9.50 24 12.0 8.5 10.25 25 11.5 9.0 10.25 26 13.0 10.0 11.50 27 13.0 10.0 11.50 28 13.5 8.5 11.00 29 13.5 10.0 11.75 30 13.0 9.0 11.00 31 13.0 9.5 11.25 June 1 13.0 10.0 11.50 2 12.0 9.0 10.50 3 11.5 9.0 10.25 4 12.0 9.0 10.50 5 14.0 10.0 12.00 6 15.0 10.5 12.75 7 15.5 11.5 13.50 8 15.5 12.5 14.00 9 14.0 11.0 12·50 10 12.5 10.5 11.50 11 13.0 9.5 11.75 12 14.0 10.0 12.00 13 13.0 11.0 12.00 14 13.0 12.0 12.50 15 15.0 12.5 13.75 174

8.2 DATES OF TAGGING AND RECAPTURE, LOCATION OF RECAPTURE, DISTANCES TRAVELLED, AND ELAPSED TIME BETWEEN RELEASE AND RECAPTURE FOR FISH TAGGED AT THE MUSKEG RIVER COUNTING FENCE IN 1976 AND 1977, AND SUBSEQUENTLY RECAPTURED OUTSIDE THE MUSKEG WATERSHED IN 1977 AND 1978. . Table 46. Dates of tagging and recapture, location of recapture, distances travelled, and elapsed time between release and recapture for fish tagged at the Muskeg River counting fence in 1976 and 1977, and subsequently recaptured outside the Muskeg watershed in 1977 and 1978.

Distance Date Location Date a Elapsed Species Travelled T- (D ) Tagged Recaptured Recaptured (km) I me ays

White sucker 20 May!76 Muskeg River Upstream Trap 8 May!77 0 353 Second Recap. Athabasca De 1 ta 21 June!77 -224 399 19 May!76 Mouth MacKay River 24 Apri 1!77 -5 340 10 May!76 Mouth MacKay River 27 April!77 -5 352 19 May!76 Mouth MacKay River 2 May!77 -5 348 24 May!76 Mou th MacKay Rive r 14 May!77 -1 355 20 May/76, MacKay River Upstream Trap 14 May!78 -13 724 -...... J 16 July!76 Steepbank River Upstream Trap 4 May!77 +18 282 V1 6 May!76 Muskeg River Downstream Trap 20 May!76 0 14 Second Recap. Muskeg River Upstream Trap 8 May!77 0 367 Third Recap_ Near GCOS Intake (Athabasca River) 28 June!77 +16 418 9 May!77 Mouth Muskeg River 19 June!77 -1 41 14 May!77 Mouth Athabasca Riverb 19 June!77 -250 36 18 May!77 Old Fort Bayb 19 June!77 -272 32 24 May!77 Mouth Fletcher Channe1 b 16 June!77 -250 23 11 May!77 Muskeg River Downstream Trap 27 May!77 0 16 Second Recap_ MacKay River Upstream Trap 15 May!78 -13 369 Third Recap_ Mouth Athabasca Riverb June!78 -250 385-415 25 May!77 MacKay River Upstream Trap 15 May!78 -13 355 4 May!77 MacKay River Upstream Trap 3 May!78 -13 364 17 May!77 Mouth Athabasca Riverb June!78 -250 380-410 6 May!77 Muskeg River Downstream Trap 26 May!77 0 20 Second Recap. Mouth Athabasca Riverb June!78 -250 369-399 continued ... Table 46. Cont i nued.

Distance Date Location Date Elapsed Species Trave 11 T~~~~,.I Recaotured Recaptured Ti me (Days ) (km)

White sucker 6 May/77 Mu River Downstream Trap 22 May/77 0 16 Second Recap. Mouth Athabasca Riverb June/78 -250 369-399 10 May/77 Muskeg River Downstream Trap 25 May/77 0 15 Second Recap. Mouth Athabasca Riverb June/78 -250 386-416 13 May/77 Mouth Athabasca Riverb June/78 -250 389-419 13 May/77 Mouth A habasca Riverb June/78 -250 389-419 3 May/77 MacKay Rver Upstream Trap 30 Apr i 1/78 -13 362 11 May/77 Muskeg River Downstream Trap May/77 0 14 Second Recap. MacKay River Upstream Trap 15 May/78 -13 369 ""'-J 0' Longnose sucke rs May/76 Mouth Ri ver 23 Apri 1/77 -1 331 4 May/76 Steepbank River Upstream Trap 4 May/77 +18 365 22 May/76 Steepbank River Downstream Trap 14 May/77 +18 357 22 May/76 Steepbank River Upstream Trap 1 May /77 +18 344 29 May/76 Steepbank Rver Upstream Trap 8 May/77 +18 344 Second Steepbank Rver Downstream Trap 24 May/77 +18 360 29 May/ Steepbank River ream Trap 2 May/77 +18 338 5 June/ Steepbank Rive Upstream Trap 2 May/77 +18 331 Second Reca p. Steepbank River Downstream Trap 24 May/77 +18 341 10 June/76 Steepbank River Downstream Trap 15 +18 339 28 June/76 Steepbank River Upstream Trap 7 +18 313 16 July/76 Steepbank River Upstream Trap 4 +18 292 13 July/76 Muskeg River Upstream Trap 6 May.77 0 358 Second Muskeg River Downstream Trap 9 June/77 0 361 Third Mouth Clark Creek 23 June/77 +46 375 continued Table 46. Concluded.

Distance Date Locat ion Date a Elapsed Species Travelled T' (D ) Tagged Recaptured Recaptured (km) I me ays

Longnose sucker 29 May/76 Lower Beaver River 14 May/77 +2 350 15 May/77 Goose Islandb 22 June/77 -264 38 28 May/77 Old Fort Bayb 19 June/77 -280 22 25 May/77 Old Fort Bayb 31 May/77 -296 6 18 May/77 MacKay River Upstream Trap 1 May/78 -13 348

Northern pike 21 July/76 Mouth Muskeg River 15 Sept/76 -1 56 Second Recap. Mouth Muskeg River 24 Ju.y/77 -1 369 24 JLlly/76 Mouth Muskeg River 27 Sept/77 -1 430 ...... 14 May/76 Mouth Poplar River 8 May/77 +29 360 ...... 2 May/77 Mouth Muskeg River 24 JUly/77 -1 83 2 May/77 Mile 27 (km 43) Athabasca River 28 Sept/77 +13 149 3 May/77 Mile 60 (km 96) Athabasca River 5 Oct/77 -41 155 3 May/77 Muskeg River Downstream Trap 13 June/77 0 41 Second Recap, Mouth MacKay River 21 Aug/77 -5 110 3 May/77 Mouth MacKay River 22 Sept/77 -5 142 4 May/77 Mouth MacKay River 16 June/77 -5 43

Arct i c grayl i ng 30 Apri 1/76 Steepbank River 10 Oct/77 +18 528 a Distance shown is approximate distance from counting fence to recapture point and + or - designates upstream or downstream from Muskeg River in the Athabasca River. On occasion movement was upstream or downstream in the Athabasca and then upstream in a tributary. bLake Athabasca. 178

9. AOSERP RESEARCH REPORTS

1- AOSERP First Annual Report, 1975 2. AF 4. 1 . 1 Walleye and Goldeye Fisheries Investigations in the Peace-Athabasca Delta--l 3. HE 1. 1 • 1 Structure of a Traditional Baseline Data System 4. VE 2.2 A Preliminary Vegetation of the Alberta Oil Sands Environmental Research Program Study Area 5. HY 3.1 The Evaluation of Wastewaters from an Oil Sand Extraction Plant 6. Housing for the North--The Stackwal1 System 7. AF 3.1.1 A Synopsis of the Physical and Biological Limnology and Fisheries Programs whithin the Alberta Oil Sands Area 8. AF 1 .2. 1 The Impact of Saline Waters upon Freshwa er Biota (A Literature Review and Bibliography) 9. ME 3.3 Prel iminary Investigations into the Magnitude of Fog Occurrence and Associated Problems in he Oil Sands Area 10. HE 2.1 Development of a Research Design Related to Archaeological Studies in the Athabasca Oil Sands Area 11- AF 2.2.1 Life eyc 1es of Some Common Aquat i c I nsects of the Athabasca River, Alberta 12. ME 1.7 Very High Resolution Meteoro ogical Satell ite Study of 0 i 1 Sands \>Jeathe r: IIA Feas i b 1 ty t udy" 13. ME 2.3.1 Plume Dispersion Measurements f an Oil Sands Extraction Plant, March 1976 14. 15. ME 3.4 A C1 ima of Low Level Air T ector es in the Alberta Oi Sands Area 16. ME 1 .6 The Feasi i ity of a Weather Radar near Fort McMurray, Alberta 17. AF 2. 1 . 1 A Survey of Baseline Levels of Contaminants in Aquatic Biota of the AOSERP Study Area 18. HY 1.1 Interim i1ation of Stream Gauging Data to December 1976 for the Alberta Oil Sands Environmental Research Program 19. ME 4.1 Calculations of Annual Averaged 5 lphur Dioxide Concentrations at Ground Level in the AOSERP Study Area 20. HY 3. 1 . 1 Characterization of Organic Constituents in Waters and Wastewaters of the Athabasca Oil Sands Mining Area 21. AOSERP Second Annual Report, 1976- 22. HE 2.3 Maximization of Technical Training and Involvement of Area Manpower 23. AF 1. 1 .2 Acute Lethality of Mine Depressurization Water on Trout Pe and Rainbow Trout 24. ME 4.2. 1 Air System Winter Field Study in the AOSERP Study Area, February 1977. 25. ME 3.5. 1 Review of Pollutant Transformation Processes Relevant to the Alberta Oil Sands Area 179

26. AF 4.5.1 Interim Report on an Intensive Study of the Fish Fauna of the Muskeg River Watershed of Northeastern Alberta 27. ME 1. 5. 1 Meteorology and Air Quality Winter Field Study in the AOSERP Study Area, March 1976 28. VE 2. 1 Interim Report on a Soils Inventory in the Athabasca Oil Sands Area 29. ME 2.2 An Inventory System for Atmospheric Emissions in the AOSERP Study Area 30. ME 2.1 Ambient Air Quality in the AOSERP Study Area, 1977 31. VE 2.3 Ecological Habitat Mapping of the AOSERP Study Area: Phase I 32. AOSERP Third Annual Report, 1977-78 33. TF 1.2 Relationships Between Habitats, Forages, and Carrying Capacity of Moose Range in northern Alberta. Part I: Moose Preferences for Habitat Strata and Forages. 34. HY 2.4 Heavy Metals in Bottom Sediments of the Mainstem Athabasca River System in the AOSERP Study Area 35. AF 4.9.1 The Effects of Sedimentation on the Aquatic Biota 36. AF 4.8.1 Fall Fisheries Investigations in the Athabasca and Clearwater Rivers Upstream of Fort McMurray: Volume 37. HE 2.2.2 Community Studies: Fort McMurray, Anzac, Fort MacKay 38. VE 7.1.1 Techniques for the Control of Small Mammals: A Review 39. ME 1.0 The Climatology of the Alberta Oil Sands Environmental Research Program Study Area 40. WS 3.3 Mixing Characteristics of the Athabasca River below Fort McMurray - Winter Conditions 41. AF 3.5. 1 Acute and Chronic Toxicity of Vanadium to Fish 42. TF 1 .1.4 Analysis of Fur Production Records for Registered Traplines in the AOSERP Study Area, 1970-75 43. TF 6.1 A Socioeconomic Evaluation of the Recreational Fish and Wildlife Resources in Alberta, with Particular Reference to the AOSERP Study Area. Volume I: Summary and Conclusions 44. VE 3. 1 Interim Report on Symptomo10gy and Threshold Levels of Air Pollutant Injury to Vegetation, 1975 to 1978 45. VE 3.3 Interim Report on Physiology and Mechanisms of Air-Borne Pollutant Injury to Vegetation, 1975 to 1978 46. VE 3.4 Interim Report on Ecological Benchmarking and Biomonitoring for Detection of Air-Borne Pollutant Effects on Vegetation and Soils, 1975 to 1978. 47. TF 1. 1. 1 A Visibil ity Bias Model for Aerial Surveys for Moose on the AOSERP Study Area 48. HG 1.1 Interim Report on a Hydrogeological Investigation of the Muskeg River Basin, Alberta 49. WS 1 .3.3 The Ecology of Macrobenthic Invertebrate Communities in Hartley Creek, Northeastern Alberta 50. ME 3.6 Literature Review on Pollution Deposition Processes 51. HY 1.3 Interim Compilation of 1976 Suspended Sediment Date in the AOSERP Study Area 52. ME 2.3.2 Plume Dispersion Measurements from an Oil Sands Extraction Plan, June 1977 53. HY 3. 1 .2 Baseline States of Organic Constituents in the Athabasca River System Upstream of Fort McMurray 54. ws 2.3 A Preliminary Study of Chemical and Microbial Characteristics of the Athabasca River in the Athabasca Oil Sands Area of Northeastern Alberta 55. HY 2.6 Microbial Populations in the Athabasca River 56. AF 3.2.1 The Acute Toxicity of Saline Groundwater and of Vanadium to Fish and Aquatic Invertebrates 57. LS2.3.1 Ecological Habitat Mapping of the AOSERP Study Area (Supplement): Phase I 58. AF 2.0.2 Interim Report on Ecological Studies on the Lower Trophic Levels of Muskeg Rivers Within the Alberta 0,. 1 Sands Envi ronmental Research Program Study Area 59. TF 3. 1 S~mi-Aquatic Mammals: Annotated Bibliography 60. WS 1. 1 . 1 Synthesis of Surface Water Hydrology 61 . AF 4.5.2 An Intensive Study of the Fish Fauna of the Steepbank River Watershed of Northeastern Alberta 62. TF 5.1 Amphibians and Reptiles in the AOSERP Study Area 63. An Overview Assessment of In Situ Development in the Athabasca Deposit ------64. LS 21.6. 1 A Review of the Baseline Data Relevant to the Impacts of Oil Sands Development on Large Mammals in the AOSERP Study Area 65. LS 21.6.2 A Review of the Baseline Data Relevant to the Impacts of Oil Sands Development on Black Bears in the AOSERP Study Area 66. AS 4.3.2 An Assessment of the Models LIRAQ and ADPIC for Application to the Athabasca Oil Sands Area 67. WS 1. 3.2 Aquatic Biological Investigations of the Muskeg River Watershed 68. AS 1.5.3 Air System Summer Field Study in the AOSERP Study Area, AS 3.5.2 June 1977 69. HS 40. 1 Native Employment Patterns in Alberta's Athabasca Oil Sands Reg ion 70. LS 28. 1 .2 An Interim Report on the Insectivorous Animals in the AOSERP Study Area 71. HY 2.2 Lake Acidification Potential in the Alberta Oil Sands Environmental Research Program Study Area 72. LS 7.1.2 The Ecology of Five Major Species of Small Mammals in the AOSERP Study Area: A Review 73. LS 23.2 Distribution, Abundance and Habitat Associations of Beavers, Muskrats, Mink and River Otters in the AOSERP Study Area, Northeastern Alberta Interim Report to 1978 74. AS 4.5 Air Qual ity Modelling and User Needs

These reports are not available upon request. For further information about availabil ity and location of depositories, please contact:

Alberta Oil Sands Environmental Research Program 15th Floor, Oxbridge Place 9820 - 106 Street EDMONTON~ Alberta T5K 2J6 This material is provided under educational reproduction permissions included in Alberta Environment and Sustainable Resource Development's Copyright and Disclosure Statement, see terms at http://www.environment.alberta.ca/copyright.html. This Statement requires the following identification:

"The source of the materials is Alberta Environment and Sustainable Resource Development http://www.environment.gov.ab.ca/. The use of these materials by the end user is done without any affiliation with or endorsement by the Government of Alberta. Reliance upon the end user's use of these materials is at the risk of the end user. AR 139/2007 SPECIFIED GAS EMITTERS REGULATION http://www.qp.alberta.ca/574.cfm?page=2007_139.cfm&leg_ty...

(Consolidated up to 127/2011)

ALBERTA REGULATION 139/2007

Climate Change and Emissions Management Act

SPECIFIED GAS EMITTERS REGULATION

Table of Contents Part 1 Interpretation and Application 1 Definitions 2 Application

Part 2 Emissions Intensity Limits, True‑up, Emission Offsets, Fund Credits, Emission Performance Credits 3 2007 emissions intensity limits 4 Emissions intensity limits for 2008 and subsequent years 5 True‑up 6 Duty to comply 7 Emission offsets 8 Fund credits 9 Emission performance credits 10 Nature of emission offsets, fund credits and emission performance credits

Part 3 Reporting, Records, Confidentiality and Third Party Auditors 11 Compliance report 12 Further information, verification, resubmission 13 Access to application for baseline or compliance report 14 Publishing application for baseline or compliance report 15 Retention of records 16 Request for confidentiality 17 Annual report to Information and Privacy Commissioner 18 Qualifications of third party auditors 19 Prescribing forms

Part 4 Emissions Intensity Baselines 20 Application for establishment of baseline emissions intensity

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21 Determination of baseline emissions intensity 22 Establishment of baseline emissions intensity 23 Establishment of new baseline emissions intensity

Part 5 Exemptions 24 Application for exemption

Part 6 Enforcement 25 Inspections, investigations, audits 26 Order where net emissions intensity limit exceeded 27 Offences 28 Penalties 29 Due diligence

Part 7 Expiry 30 Expiry

Schedule

Part 1 Interpretation and Application

Definitions 1(1) In this Regulation,

(a) “Act” means the Climate Change and Emissions Management Act;

(b) “actual emissions intensity” means total direct emissions, not including industrial process emissions, per unit of production from a facility;

(c) “baseline emissions intensity” means the baseline emissions intensity for a facility established in accordance with Part 4;

(d) “CO2e” means the 100 year time horizon global warming potential of a specified gas expressed in terms of equivalency to CO2 set out in column 3 of the Schedule;

(e) “direct emissions” means the release of specified gases from sources actually located at a facility, expressed in tonnes on a CO2e basis;

(f) “emission offset” means

(i) a reduction in the release of specified gases, expressed in tonnes on a CO2e basis, that meets the requirements of section 7(1), but does not include an emission performance credit,

(ii) a geological sequestration of specified gases, expressed in tonnes on a CO2e basis, that meets the requirements of section 7(1.1), and

(iii) a capture of specified gases that are geologically sequestered that meets the requirements of section 7(1.2);

(g) “emission performance credit” means a reduction in the release of specified gases, expressed in

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tonnes on a CO2e basis, that meets the requirements of section 9(1);

(h) “emissions intensity” means the quantity of specified gases released by a facility per unit of production from that facility;

(i) “established facility” means, subject to subsection (2), a facility that

(i) completed its first year of commercial operation before January 1, 2000, or

(ii) has completed 8 years of commercial operation;

(j) “facility” means

(i) a plant, structure or thing where an activity listed in section 2 of the Schedule of Activities to the Environmental Protection and Enhancement Act occurs, and

(ii) a site or 2 or more contiguous or adjacent sites that are operated and function in an integrated fashion where an activity listed in any of sections 3 to 11 of the Schedule of Activities to the Environmental Protection and Enhancement Act occurs,

including all the buildings, equipment, structures, machinery and vehicles that are an integral part of the activity;

(k) “Fund” means the Climate Change and Emissions Management Fund established by the Act;

(l) “fund credit” means a fund credit described in section 8;

(m) “industrial process emissions” means direct emissions from an industrial process involving chemical or physical reactions other than combustion, and where the primary purpose of the industrial process is not energy production;

(n) “net emissions intensity” means the net emissions intensity for a facility determined in accordance with section 3(4) or 4(3), as the case may be;

(o) “net emissions intensity limit” means the applicable maximum net emissions intensity permitted under section 3 or 4;

(p) “new facility” means

(i) a facility that

(A) completed its first year of commercial operation on December 31 of 2000 or a subsequent year, and

(B) has completed less than 8 years of commercial operation, or

(ii) a facility designated as a new facility under subsection (2);

(q) “person responsible” means, where the release of the specified gas occurs

(i) at a facility that is the subject of an approval or registration under the Environmental Protection and Enhancement Act, the holder of the approval or registration,

(ii) at a facility that is not the subject of an approval or registration referred to in subclause (i) but is the subject of an approval or other authorization issued by the Energy Resources Conservation Board or the Alberta Utilities Commission, the holder of that approval or authorization, or

(iii) at any other facility, the owner of the facility;

(r) “production” means the quantity, expressed in the applicable unit of production, of

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(i) end product produced by a facility, or

(ii) any input, output or other thing specified under subsection (4);

(s) “specified gas” means a gas listed in column 1 of the Schedule;

(t) “third party auditor” means a person who meets the requirements set out in section 18;

(u) “total annual emissions” means total direct emissions in a year, not including industrial process emissions;

(v) “total half year emissions” means total direct emissions in the period July 1, 2007 to December 31, 2007, not including industrial process emissions;

(w) “unit of production” means the unit of measure of production of a facility, unique to the facility, as approved or determined by the director in establishing a baseline emissions intensity for the facility under Part 4;

(x) “year” means a calendar year unless otherwise specified.

(2) The director may designate an established facility as a new facility if the director considers it appropriate to do so.

(3) In determining whether it is appropriate to make a designation under subsection (2), the director may consider

(a) the nature and extent of any expansion or significant change undergone by the facility and the technologies employed in the expansion or significant change that affect specified gas emissions, and

(b) any other matter that in the director’s opinion is relevant.

(4) If a facility does not produce an end product, the director may specify an input, output or other thing as the standard of measurement of production of the facility for the purposes of this Regulation. AR 139/2007 s1;254/2007;127/2011

Application 2 Subject to section 3(1), this Regulation applies to a facility that has direct emissions totalling 100 000 tonnes or more in 2003 or any subsequent year.

Part 2 Emissions Intensity Limits, True-up, Emission Offsets, Fund Credits, Emission Performance Credits

2007 emissions intensity limits 3(1) This section applies to a facility that had direct emissions totalling 100 000 tonnes or more in a year of commercial operation in any of the years 2003, 2004, 2005 or 2006.

(2) The net emissions intensity for a facility that is an established facility on January 1, 2007 for the period commencing on July 1, 2007 and ending on December 31, 2007 shall not exceed 88% of the baseline emissions intensity for the facility.

(3) If the period commencing on July 1, 2007 and ending on December 31, 2007 is the last 6 months of

(a) the 4th year of commercial operation of a new facility, the net emissions intensity for the facility for that period shall not exceed 98% of the baseline emissions intensity for the facility,

(b) the 5th year of commercial operation of a new facility, the net emissions intensity for the facility

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for that period shall not exceed 96% of the baseline emissions intensity for the facility,

(c) the 6th year of commercial operation of a new facility, the net emissions intensity for the facility for that period shall not exceed 94% of the baseline emissions intensity for the facility,

(d) the 7th year of commercial operation of a new facility, the net emissions intensity for the facility for that period shall not exceed 92% of the baseline emissions intensity for the facility, or

(e) the 8th year of commercial operation of a new facility, the net emissions intensity for the facility for that period shall not exceed 90% of the baseline emissions intensity for the facility.

(4) For the purposes of subsections (2) and (3), the net emissions intensity for a facility must be determined by the following formula:

NEI = (THYE ‑ (EO+FC+EPC)) P

where

NEI is net emissions intensity for the facility;

THYE is total half year emissions from the facility;

EO is allowable emission offsets applied by the person responsible;

FC is allowable fund credits applied by the person responsible;

EPC is allowable emission performance credits applied by the person responsible;

P is production for the period July 1, 2007 to December 31, 2007.

Emissions intensity limits for 2008 and subsequent years 4(1) Commencing with the year 2008, the net emissions intensity for a year

(a) for an established facility shall not exceed 88% of the baseline emissions intensity for the facility, and

(b) for a new facility shall not exceed

(i) 98% of the baseline emissions intensity for the facility, in the case of the 4th year of commercial operation of the facility,

(ii) 96% of the baseline emissions intensity for the facility, in the case of the 5th year of commercial operation of the facility,

(iii) 94% of the baseline emissions intensity for the facility, in the case of the 6th year of commercial operation of the facility,

(iv) 92% of the baseline emissions intensity for the facility, in the case of the 7th year of commercial operation of the facility, and

(v) 90% of the baseline emissions intensity for the facility, in the case of the 8th year of commercial operation of the facility.

(2) The Minister may, by order, establish net emissions intensity limits in addition to or in substitution for those set out in subsection (1).

(3) For the purposes of this section, the net emissions intensity for a facility must be determined by the following formula:

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NEI = (TAE ‑ (EO+FC+EPC)) P

where

NEI is net emissions intensity for the facility;

TAE is total annual emissions from the facility;

EO is allowable emission offsets applied by the person responsible;

FC is allowable fund credits applied by the person responsible;

EPC is allowable emission performance credits applied by the person responsible;

P is production for the year.

True-up 5(1) The emission offsets, fund credits and emission performance credits that may be subtracted from total half year emissions under section 3(4) or from total annual emissions under section 4(3) are those emission offsets, fund credits and emission performance credits that are described in sections 7, 8 and 9 respectively and that are available to the person responsible on the earlier of the date the compliance report required by section 11 is submitted and the deadline for submitting the compliance report.

(2) The determination required by section 3(4) or 4(3) must be made on or before the deadline for submitting the compliance report required by section 11.

Duty to comply 6(1) The person responsible shall comply with the net emissions intensity limits established by sections 3(2) and (3) and 4(1) and, if applicable, under section 4(2).

(2) If there is more than one person responsible for a facility for the period referred to in section 3(2) or (3) or in the year 2008 or a subsequent year, subsection (1) applies only to the person who is the person responsible on

(a) December 31, 2007, in the case of the period referred to in section 3(2) or (3), or

(b) December 31 of the relevant year, in any other case.

Emission offsets 7(1) The following requirements must be met in order for a reduction in specified gas emissions to constitute one or more emission offsets:

(a) the specified gas emissions reduction must occur in Alberta;

(b) the specified gas emissions reduction must be from an action taken that is not otherwise required by law at the time the action is initiated;

(c) the specified gas emissions reduction must

(i) result from actions taken on or after January 1, 2002, and

(ii) occur on or after January 1, 2002;

(d) the specified gas emissions reduction must be real and demonstrable;

(e) the specified gas emissions reduction must be quantifiable and measurable, directly or by accurate estimation using replicable techniques.

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(1.1) The following requirements must be met in order for a geological sequestration of specified gas to constitute one or more emission offsets:

(a) the specified gas that is geologically sequestered must be captured through a dedicated process from sources located at a facility in Alberta;

(b) the specified gas must be stored in a geological formation that is located wholly or partly in Alberta;

(c) the geological sequestration of the specified gas must not be required by law at the time geological sequestration of specified gas is initiated;

(d) the construction of the infrastructure used to geologically sequester the specified gas must have been initiated on or after January 1, 2002;

(e) the geological sequestration of the specified gas must occur after January 1, 2002;

(f) the quantity of specified gas that is geologically sequestered must be quantifiable and measurable, directly or by accurate estimation using replicable techniques.

(1.2) The following requirements must be met in order for a capture of specified gas to constitute one or more emission offsets:

(a) the specified gas must be captured through a dedicated process from sources located at a facility upgrading or refining bitumen in Alberta;

(b) the capture of the specified gas must not be required by law at the time capture of specified gas is initiated;

(c) the construction of the infrastructure used to capture the specified gas must have been initiated on or after January 1, 2012 and use of the infrastructure to capture specified gas must have been initiated before December 31, 2017;

(d) the specified gas must be

(i) captured by infrastructure capable of capturing, and

(ii) stored in geological formations capable of storing

1 000 000 tonnes of specified gas per year, expressed on a CO2e basis;

(e) at least 51% of the volume of specified gas captured through the dedicated process from sources located at a facility upgrading or refining bitumen in Alberta in a year must be sequestered in a geological formation in respect of which a pore space tenure agreement has been entered into with the Government of Alberta on or after January 1, 2011;

(f) the quantity of specified gas that is captured must be quantifiable and measurable, directly or by accurate estimation using replicable techniques;

(g) the captured specified gas must be geologically sequestered in accordance with subsection (1.1);

(h) the amount established under section 8(2) must be $80 or less at the time the captured specified gas is geologically sequestered.

(1.3) The calculation of the amount of

(a) a reduction in specified gas emissions, or

(b) specified gas that is geologically sequestered

must accord with any Ministerial guidelines issued under section 62 of the Act.

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(1.4) A one tonne reduction in specified gas emissions, expressed on a CO2e basis, that meets the requirements of subsection (1) constitutes one emission offset.

(1.5) A geological sequestration of one tonne of specified gas, expressed on a CO2e basis, that meets the requirements of subsection (1.1) constitutes one emission offset.

(1.6) The quantity of emission offsets constituted by a capture of specified gas that meets the requirements of subsection (1.2) is determined in accordance with the following:

(a) if the amount established under section 8(2) is equal to or less than $40 at the time that the captured specified gas is geologically sequestered, the quantity of emission offsets constituted is calculated in accordance with the following formula:

A x 1

where

A equals the emission offsets constituted by the geological sequestration of the captured specified gas in accordance with subsection (1.1);

(b) if the amount established under section 8(2) is more than $40 but not more than $80 at the time that the captured specified gas is geologically sequestered, the quantity of emission offsets constituted is calculated in accordance with the following formula:

A x (80-B)/40

where

A equals the emission offsets constituted by the geological sequestration of the captured specified gas in accordance with subsection (1.1);

B equals the amount established under section 8(2).

(2) An emission offset may be used in meeting net emissions intensity limits under section 3 or 4 subject to the following rules:

(a) an emission offset must be held by the person responsible using it;

(b) an emission offset may only be used once;

(c) if an emission offset is jointly held, each holder may only use a portion of the offset on a pro rata basis;

(d) the use of an emission offset must accord with any Ministerial guidelines issued under section 62 of the Act. AR 139/2007 s7;127/2011

Fund credits 8(1) A person responsible may obtain fund credits by contributing money to the Fund.

(2) The Minister may, by order, establish the amount of money that a person responsible must contribute to the Fund to obtain one fund credit equal to a one tonne reduction in emissions, expressed on a CO2e basis.

(3) A fund credit may be used in meeting net emissions intensity limits under sections 3 and 4 subject to the following rules:

(a) a fund credit obtained on or before March 31, 2008 may only be used in meeting net emissions intensity limits applicable to the period commencing on July 1, 2007 and iending on December 31, 2007;

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(b) except as provided in clause (a), a fund credit obtained on or before March 31 in a year may only be used in meeting net annual emissions intensity limits for the previous year;

(c) a fund credit obtained after March 31 in a year may only be used in meeting net annual emissions intensity limits for that year;

(d) a fund credit may not be used by more than one party;

(e) the use of a fund credit must accord with any Ministerial guidelines issued under section 62 of the Act. AR 139/2007 s8;127/2011

Emission performance credits 9(1) When a facility to which this Regulation applies achieves actual emissions intensity for a period that is less than the applicable net emissions intensity limit for that period, the reduction in specified gas emissions that is not used in meeting the net emissions intensity limit constitutes an emission performance credit or credits.

(2) An emission performance credit may be used in meeting net emissions intensity limits under sections 3 and 4 subject to the following rules:

(a) an emission performance credit created at a facility in a year may be used in meeting net emissions intensity limits

(i) for another facility for that year, or

(ii) for the facility at which it was created or for another facility, for a subsequent year;

(b) an emission performance credit must be held by the person responsible using it;

(c) an emission performance credit may only be used once;

(d) if an emission performance credit is jointly held, each holder may only use a portion of the credit on a pro rata basis;

(e) the use of an emission performance credit must accord with any Ministerial guidelines issued under section 62 of the Act.

Nature of emission offsets, fund credits and emission performance credits 10(1) For greater certainty, emission offsets, fund credits and emission performance credits are revocable licences authorizing persons responsible, subject to this Part, to use the quantity of specified gas emission reductions from or represented by the emission offsets, fund credits and emission performance credits in meeting net emissions intensity limits under sections 3 and 4.

(2) Nothing in this Regulation ensures or guarantees the availability of emission offsets or emission performance credits.

Part 3 Reporting, Records, Confidentiality and Third Party Auditors

Compliance report 11(1) The person responsible for a facility on December 31 of a year shall submit to the director a compliance report with respect to that facility for that year by March 31 of the following year.

(2) The report must contain the information and data required in a form prescribed by the director.

(3) The person responsible shall report by electronic means as prescribed by the director.

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(4) The report must

(a) either confirm that the net emissions intensity limit for the facility has been met or provide an acknowledgement that the net emissions intensity limit for the facility has not been met, with an explanation and proposal to address and remedy the non‑compliance,

(b) be certified by a person and in a manner required by the form, and

(c) be verified by a third party auditor.

(5) The person responsible may submit one report in a form prescribed by the director for all of the facilities in respect of which the person is the person responsible if, in the report, the information required by subsection (4) is provided in respect of each facility.

Further information, verification, resubmission 12 The director may do one or more of the following regarding a report or information submitted to the director:

(a) require that additional information or data be provided;

(b) require verification or further verification by a third party auditor of any information or data;

(c) collect any additional information or conduct any review that the director considers necessary;

(d) direct the person responsible to resubmit information in accordance with any directions that the director considers necessary.

Access to application for baseline or compliance report 13(1) Within a reasonable time after receiving a request in writing to review an application for the establishment of a baseline emissions intensity or a compliance report, the director shall, except with respect to prescribed information within the meaning of section 16(6) and (7) and information that is the subject of enforcement proceedings under the Act or this Regulation,

(a) make the application or report available for review by the person requesting it during normal business hours at the location where the application or report is kept, and

(b) provide a copy of the application or report free of charge to the person requesting it.

(2) The director may refuse to comply with subsection (1) unless the director is satisfied that the person making the request to inspect has first made a request to obtain a copy of the application or report from the appropriate person responsible and that the request

(a) was refused, or

(b) was not satisfied within 30 days after the request was made.

Publishing application for baseline or compliance report 14 Subject to section 59 of the Act and any order made under section 16(4)(a) of this Regulation, the director may publish an application for the establishment of a baseline emissions intensity or a compliance report or information in an application or a report in any form and manner the director considers appropriate.

Retention of records 15(1) A person responsible who submits an application for the establishment of a baseline emissions intensity or compliance report shall, for at least 7 years following the submission of the application or report, retain

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(a) a copy of the application or report, and

(b) the records, information and data on which the application or report was based.

(2) A person responsible shall retain all records, information and data respecting emissions intensity for at least 7 years after the date of their creation.

(3) The material retained under subsections (1) and (2) must be located

(a) at the head or principal office, in Alberta, of the person responsible, or

(b) at the facility to which the application, report, records, information or data relate.

Request for confidentiality 16(1) A person responsible who submits an application for the establishment of a baseline emissions intensity or a compliance report may include a written request that certain information in the application or report be kept confidential for a period of up to 5 years after the date of submission on the basis that the information is commercial, financial, scientific or technical information that would reveal proprietary business, competitive or trade secret information about a specific facility, technology or corporate initiative.

(2) The director shall have regard to the following when making a decision on a request for confidentiality made under subsection (1):

(a) whether disclosure of the information could reasonably be expected to harm significantly the competitive position of the person responsible;

(b) whether disclosure of the information could reasonably be expected to interfere significantly with the negotiating position of the person responsible;

(c) whether disclosure of the information could reasonably be expected to result in undue financial loss or gain to any person or organization;

(d) the availability of the information or the means to obtain the information from other public sources;

(e) whether there are any other competing interests that would suggest that disclosure of the information is warranted.

(3) The director may require a person responsible to provide additional reasons, in writing, in support of a request for confidentiality made under subsection (1).

(4) The director shall

(a) if the director considers that the request is well founded, approve the request and order that the information to which the request relates be kept confidential and not be disclosed for the period prescribed by the director, or

(b) refuse the request if the director considers that the request is not well founded.

(5) The director shall, in writing, notify the person responsible of the director’s decision under subsection (4) within 150 days after receiving the request.

(6) If the director is considering a request for confidentiality under this section, the information to which the request relates is prescribed information for the purposes of section 59 of the Act until a decision is made.

(7) If the director makes an order under subsection (4)(a), the information that is the subject of the order is prescribed information for the purposes of section 59 of the Act for the period prescribed in the order.

Annual report to Information and Privacy Commissioner

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17 The director shall provide annually to the Information and Privacy Commissioner, in the form and manner the director considers appropriate, a report setting out the following:

(a) the number of requests received by the director under section 16(1) in the year;

(b) the number of requests approved by the director under section 16(4)(a) in the year;

(c) the period prescribed by the director under section 16(4)(a) for each approved request.

Qualifications of third party auditors 18(1) A person is eligible to be a third party auditor under this Regulation if the person

(a) is

(i) registered as

(A) a professional engineer under the Engineering, Geological and Geophysical Professions Act, or

(B) a chartered accountant under the Regulated Accounting Profession Act,

(ii) a member of a profession that has substantially similar competence and practice requirements as a profession referred to in subclause (i)

(A) in a province or territory of Canada, or

(B) approved by the director, in a jurisdiction outside of Canada,

(b) has technical knowledge of

(i) specified gas emission quantification methodologies,

(ii) audit practices, and

(iii) any other matters considered relevant by the director,

and

(c) has any other qualifications that the director considers necessary.

(2) A person is not eligible to be a third party auditor for a facility if

(a) that person is the person responsible for the facility or is a director, officer or employee of the person responsible for the facility or of an affiliate, within the meaning of section 2 of the Business Corporations Act, of the person responsible, or

(b) the person is an employee or agent of the Government.

(3) The director may request evidence of a person’s qualifications and eligibility as a third party auditor and may determine that the person is not eligible to perform the functions of a third party auditor if the director is not satisfied that the person possesses the necessary qualifications or that the person is eligible.

Prescribing forms 19 The director may prescribe forms for the purposes of this Regulation.

Part 4 Emissions Intensity Baselines

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Application for establishment of baseline emissions intensity 20(1) The person responsible for a facility shall apply for the establishment of a baseline emissions intensity by

(a) December 31, 2007, in the case of a facility that is subject to an emissions intensity limit under section 3,

(b) the later of June 1, 2008 and June 1 of the 4th year of commercial operation of the facility, in the case of a new facility that

(i) has direct emissions of 100 000 tonnes or more in any of its first 3 years of commercial operation, and

(ii) is not subject to an emissions intensity limit under section 3,

or

(c) June 1 of the year following the year of commercial operation of a facility in which the facility first has direct emissions totalling 100 000 tonnes or more, in any other case.

(2) An application for the establishment of a baseline emissions intensity for a facility must

(a) be submitted by the person responsible to the director on a form prescribed by the director,

(b) include the information and supporting data required by the form, and

(c) include the verification by a third party auditor of the information and data provided with the application form as required by the form. AR 139/2007 s20;156/2007;210/2007

Determination of baseline emissions intensity 21(1) The baseline emissions intensity for a facility that is an established facility on January 1, 2007 must be determined by one of the following methods:

(a) by calculating the average of the ratio of total annual emissions to production for the years 2003, 2004 and 2005, as expressed in the following formula:

where

BEI is baseline emissions intensity;

TAE is total annual emissions for the year indicated;

P is production for the year indicated;

(b) by an alternative method specified in writing by the director where the director determines that the method in clause (a) is not appropriate.

(2) The baseline emissions intensity for a new facility must be determined by one of the following methods:

(a) by calculating the ratio of total annual emissions to production for the 3rd year of commercial

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operation of the facility as expressed in the following formula:

where

BEI is baseline emissions intensity;

TAE3 is total annual emissions for the 3rd year of commercial operation;

P3 is production for the 3rd year of commercial operation;

(b) by an alternative method specified in writing by the director where the director determines that the method in clause (a) is not appropriate.

Establishment of baseline emissions intensity 22(1) On considering an application for the establishment of a baseline emissions intensity, the director may do one or more of the following:

(a) request additional information or data;

(b) require verification or further verification by a third party auditor of any information or data;

(c) collect any additional information or conduct any review that the director considers necessary in order to determine the baseline emissions intensity for the facility;

(d) direct the applicant to resubmit the application and give any directions about the resubmission that the director considers necessary.

(2) The director may establish a baseline emissions intensity for a facility

(a) as requested in an application, or

(b) that is different from the baseline emissions intensity requested in an application and may, for that purpose, determine the unit of production for the facility.

(3) The director shall give written notice of a decision under subsection (2) to the person responsible.

(4) In making a decision under subsection (2), the director may consider factors the director considers relevant, including, but not limited to,

(a) technologies that affect specified gas emissions that are in use at comparable facilities, and

(b) the best available technology economically achievable for the facility, integrating sustainability and economics, accounting for project technology and design characteristics.

Establishment of new baseline emissions intensity 23 The director may at any time review the baseline emissions intensity for a facility and establish a new baseline emissions intensity or direct the person responsible to apply for a new baseline emissions intensity if the director is of the opinion that

(a) the baseline emissions intensity is inaccurate,

(b) the facility has undergone an expansion or significantly changed, or

(c) for any other reason, a revised baseline emissions intensity is appropriate.

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Part 5 Exemptions

Application for exemption 24 The director may, on application, exempt the person responsible for a facility from the duties imposed by Parts 2 and 3 subject to any terms or conditions the director considers appropriate for a period not exceeding one year if the director is of the opinion that

(a) for a prolonged period the facility was operated under unusual conditions or was shut down, and

(b) the conditions or shutdown caused a material reduction in the specified gas emissions for the applicable period.

Part 6 Enforcement

Inspections, investigations, audits 25 An inspector or investigator may, in accordance with the Act, undertake an inspection, investigation or audit of a person responsible or a facility, or both, in respect of obligations under this Regulation.

Order where net emissions intensity limit exceeded 26(1) The director may issue an order to the person responsible for a facility requiring the person responsible to take the measures specified in the order to minimize or remedy the effects of the facility releasing specified gases into the environment in amounts in excess of those within the net emissions intensity limit for the facility where

(a) a compliance report indicates that the net emissions intensity limit for the facility has not been met,

(b) the director determines that the calculation of the net emissions intensity of the facility was incorrect or was based on inaccurate, incorrect or false information and that the net emissions intensity limit for the facility was exceeded, or

(c) the value for the emission offsets that was used to calculate the net emissions intensity of the facility for a year is no longer valid because some or all of the tonnes of specified gases which the emissions offsets represented as not being released into the environment have subsequently been released.

(2) An order under subsection (1) may require the person responsible to take the following measures:

(a) obtain emission offsets or emission performance credits;

(b) make contributions to the Fund;

(c) any other measures that the director considers advisable.

(3) An emission offset or emission performance credit obtained to comply with the terms of an order under this section may not be used under section 3 or 4.

(4) This section applies whether or not a person has been charged with or convicted of an offence or required to pay an administrative penalty in relation to the matter with respect to which the order is made.

Offences 27 A person who

(a) contravenes section 6,

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(b) contravenes section 11, 15 or 20,

(c) performs the functions of a third party auditor and does not meet the requirements set out in section 18,

(d) retains a person as a third party auditor who does not meet the requirements set out in section 18, or

(e) does not comply with an order issued under section 26

is guilty of an offence.

Penalties 28(1) A person who is guilty of an offence under section 27(a) is liable to a fine of not more than $200 for every tonne of CO2e by which the total release of specified gases exceeds the net emissions intensity limit for the facility established by section 3(2) or (3) or 4(1) or under section 4(2), as the case may be.

(2) A person who is guilty of an offence under section 27(b), (c), (d) or (e) is liable

(a) to a fine of not more than $50 000, in the case of an individual, or

(b) to a fine of not more than $500 000, in the case of a corporation.

Due diligence 29 No person shall be convicted of an offence under this Regulation if that person establishes on a balance of probabilities that the person took all reasonable steps to prevent its commission.

Part 7 Expiry

Expiry 30 For the purpose of ensuring that this Regulation is reviewed for ongoing relevancy and necessity, with the option that it may be repassed in its present or an amended form following a review, this Regulation expires on September 1, 2014.

Schedule

Specified Gases and Their Global Warming Potentials

Specified Gas Chemical Formula Global Warming Potential (100 year time horizon) Carbon dioxide CO2 1 Methane CH4 21 Nitrous oxide N2O 310 HFC-23 CHF3 11700 HFC-32 CH2F2 650 HFC-41 CH3F 150 HFC-43-10mee C5H2F10 1300 HFC-125 C2HF5 2800 HFC-134 C2H2F4 1000 HFC-134a CH2FCF3 1300 HFC-152a C2H4F2 140

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Specified Gas Chemical Formula Global Warming Potential (100 year time horizon) HFC-143 C2H3F3 300 HFC-143a C2H3F3 3800 HFC-227ea C3HF7 2900 HFC-236fa C3H2F6 6300 HFC-245ca C3H3F5 560 Sulphur hexafluoride SF6 23900 Perfluoromethane CF4 6500 Perfluoroethane C2F6 9200 Perfluoroproprane C3F8 7000 Perfluorobutane C4F10 7000 Perfluorocyclobutane c-C4F8 8700 Perfluoropentane C5F12 7500 Perfluorohexane C6F14 7400

17 of 17 9/26/12 10:42 PM Oil sands mining and reclamation cause massive loss of peatland and stored carbon

Rebecca C. Rooney, Suzanne E. Bayley, and David W. Schindler1

Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada T6G 2E9

Contributed by David W. Schindler, November 3, 2011 (sent for review August 30, 2011)

We quantified the wholesale transformation of the boreal land- ref. 6). They found that 64% of the land supported wetland vege- scape by open-pit oil sands mining in Alberta, Canada to evaluate tation, whereas only 23% of the land supported upland vegetation its effect on carbon storage and sequestration. Contrary to claims (Table 1). The most common land cover type was fen vegetation, made in the media, peatland destroyed by open-pit mining will whereas deep water, shallow open water, and marsh habitat were not be restored. Current plans dictate its replacement with upland scarce (Table 1). Due to the heterogeneous distribution of wetlands forest and tailings storage lakes, amounting to the destruction of in the region, the exact proportion of wetland habitat differs among over 29,500 ha of peatland habitat. Landscape changes caused by the 10 approved mining projects (Table 2). Generally, the east bank currently approved mines will release between 11.4 and 47.3 of the Athabasca River supports more wetland habitat than the drier million metric tons of stored carbon and will reduce carbon se- west bank (Fig. 1 and Table 2). questration potential by 5,734–7,241 metric tons C/y. These losses Despite efforts to standardize industry reporting (SI Text), have not previously been quantified, and should be included with many inconsistencies remain that impede assessment of cumula- the already high estimates of carbon emissions from oil sands tive effects and direct comparison between pre- and postmining mining and bitumen upgrading. A fair evaluation of the costs landscapes. For example, the taxonomic and spatial resolutions and benefits of oil sands mining requires a rigorous assessment used to make predictions about the postmining landscape are of impacts on natural capital and ecosystem services. coarser than those detailing baseline conditions, mainly due to difficulty in predicting drainage and nutrient conditions. wetland reclamation | tar sands A direct comparison of pre- and postmining landscapes is only SCIENCES possible for 4 of the 10 approved mines: The Horizon, Jackpine– ENVIRONMENTAL n area larger than the state of Rhode Island will eventually be Phase 1, Muskeg, and Kearl mine closure plans provided the Amined by oil sands companies in northern Alberta. These relative abundance of pre- and postmining vegetation cover (7– boreal lands must be reclaimed, but despite claims to the contrary 10) (Table 3 and Table S1). These four mines represent only 42% (1), operators are not required to return the land to its original of the area approved for mining, although they are representative state (2). This study was precipitated by the disparity between in their distribution: 59% on the wetter east bank and 41% on the statements made by the oil sands industry regarding the extent and drier west bank, compared with 61% and 39% of the total leased anticipated success of mine reclamation and their official closure area approved for mining on the east and west banks, respectively. plans, which serve as agreements between mine operators and the Suncor Energy Inc. and Syncrude Canada Inc. did not provide Alberta government regarding actual reclamation expectations. data on the relative abundance of vegetation covers for the six Oil sands deposits accessible by open-pit surface mining cover mines they operate, but did provide figures contrasting the pre- about 475,000 ha of boreal Alberta, 99% of which is already and postmining landscapes (e.g., Fig. 1 and Fig. S1). Thus, al- leased (3). Currently, 10 mines have government approval to though we cannot quantify changes to land cover across the entire operate, covering about 167,044 ha (Fig. 1). This is a conserva- region, we can make generalizations about vegetation changes tive estimate that excludes the pipelines, roads, seismic lines, and with confidence that they apply to all mines. other infrastructure that support the mines. It also excludes The most striking change to result from reclamation will be impacts from aerial deposition (4) and aquifer dewatering (5) the conversion of wetland habitat to upland forest. According to that extend off-site and the area of land associated with the three company closure plans, uplands will increase by 15,030 ha on the additional mines currently undergoing environmental review. leaseholds of the four mines, mainly at the expense of peatlands, Constraints imposed by the postmining landscape and the which will decrease by 12,414 ha (67% of their premining cov- sensitivity of peatland vegetation prevent the restoration of erage). Wetlands in general will decrease by 11,761 ha, with the peatlands that dominated the premining landscape. Mine pro- loss of peatlands slightly offset by the creation of marsh and ri- ponents are required to describe the premining landscape and parian shrublands (Table 3). Operators will create end-pit lakes produce closure plans that detail the postmining landscape. by capping tailings ponds with freshwater (SI Text), boosting the Current reclamation regulations do not require the restoration amount of deep water and littoral habitat (Table 3). End-pit of previous land covers or the restitution of lost carbon formerly lakes will be fed by extensive drainage networks (e.g., Fig. 1) that stored in soils and vegetation. In place of destroyed peatlands, may support riparian habitat (Table 3). Scaling up, assuming operators plan to construct upland forest with well-defined similar land conversion ratios for the additional six mines, about drainage channels and subsaline shallow open water wetlands 29,555 ha of peatlands will be lost as a result of currently ap- draining into large tailings ponds capped with freshwater. The proved mining (net wetland loss = 28,002 ha). net effect of this landscape transformation on biodiversity and ecosystem functions has not been assessed. Here we quantify the land cover changes that will result from approved oil sands mine Author contributions: R.C.R. designed research; R.C.R. performed research; R.C.R. contrib- projects and their impact on carbon storage. uted new reagents/analytic tools; R.C.R. analyzed data; and R.C.R., S.E.B., and D.W.S. wrote the paper. Pre- Versus Postmining Landscapes The authors declare no conflict of interest. The oil sands mining area was originally wetland-rich, covered in Freely available online through the PNAS open access option. forested and shrubby fens. In 2002, Canadian Natural Resources 1To whom correspondence should be addressed. E-mail: [email protected]. Ltd. mapped vegetation cover types within a 2,277,376-ha area This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. that encompasses the surface-mineable area (Fig. 1) (details in 1073/pnas.1117693108/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1117693108 PNAS Early Edition | 1of5 Fig. 1. Map of the surface-mineable area and the footprints of oil sands mining projects with approval to operate as of March 2011. Data are adapted from the Energy Resources Conservation Board’s online Scheme Approval Map Viewer. Gray lease areas are included in our detailed comparison of pre- and postmining land cover. Short arrows connect labels to smaller lease areas. (Insets titled Suncor 2009) Pre- and postmining land cover for the Millenium and North Steepbank mines, adapted from Suncor (11). An expanded version of the Insets is available as Fig. S1.

2of5 | www.pnas.org/cgi/doi/10.1073/pnas.1117693108 Rooney et al. Table 1. Vegetation cover within and surrounding the surface mineable oil sands area. Land cover class Total area (ha) Regional study area (%)

Terrestrial vegetation Coniferous 25,309 1 Deciduous 273,050 12 Mixedwood 217,990 10 Terrestrial vegetation subtotal 516,349 23 Water Deep water (>2 m) 13,352 1 Shallow open water (<2 m) 27,728 1 Water subtotal 41,080 2 Wetlands Graminoid fen 61,395 3 Marsh 41,320 2 Poor wooded fen/wooded bog 187,349 8 Shrubby fen 231,109 10 Wooded fen 923,895 41 Wetlands subtotal 1,445,068 64 Other Burn (within 20 y) 144,227 6 Cloud 25 <1 Cutblocks 57,648 3 Disturbances 63,492 3 Shrubland 8,619 <1 Urban/industrial 868 <1

Other subtotal 274,879 12 SCIENCES Total 2,277,376 100 ENVIRONMENTAL

Data are adapted from table B3-2 in Raine et al. (6).

In terms of the vegetation, reclamation will mean the re- No closure plan calls for the restoration of lost peatlands (7–9, placement of low-productivity tamarack (Larix larcina) and black 11, 14). Cattails and other marsh plants may tolerate the salt, spruce (Picea mariana) fens and bogs with higher-productivity metals, and naphthenic acids present in groundwater and surface forests of white spruce (Picea glauca), jack pine (Pinus banksiana), runoff in reclaimed areas (15), but peatland vegetation is very and trembling aspen (Populus tremuloides). Understory vegetation sensitive to high conductivity and ion concentrations (16). Two will change from sedges, ericaceous plants such as labrador tea pilot fen construction projects are under way to study survival of (Ledum groenlandicum), and mosses such as Sphagnum spp. and fen species in a tailings-contaminated environment and the ca- Drepanocladus spp. (which can deposit up to several meters of peat) pacity of reclamation materials to support fen-type hydrology. to blueberry (Vaccinium myrtilloides), dogwood (Cornus spp.), and Recreating fen-type hydrology in the postmining landscape is low-bush cranberry (Viburnum edule) (which accumulate much less possible, but requires a minimum 2:1 upland to peatland ratio for carbon in the soil). Reclamation will also mean a shift in age uplands to supply adequate seepage to maintain peat wetness structure, as reclaimed forests will begin as seedlings and will take (17). Thus, even if the entire closure landscape were designed to 50–70 y to reach harvestable age (11). The shift to a drier forest will maximize fen habitat, it could not recreate the area of fens that also mean a change in fire regime, as drier forest types are more was lost. Other considerations, such as the need for end-pit lakes susceptible to fire (12) and thus support younger stands than wetter and the limited availability of suitable substrate and vegetation forests on average. (e.g., pilot fens were constructed by transfer of live peat from natural fens), ensure that constructed fens will only constitute Impediments to Wetland Restoration a small fraction of the postmining landscape. There are several reasons closure plans favor the creation of well- drained habitat over wetlands (e.g., 8, 11). First, Alberta has no Implications wetland policy requiring compensation for wetland loss in No large-scale oil sands reclamation project has undergone in- the boreal region. Second, because the volume of tailings and dependent evaluation, and thus the ultimate success of closure upgrading by-products exceeds the size of mine pits, the closure plans remains uncertain (18). Upland habitat has been created landscapes will consist of hills instead of the level topography that (e.g., the 104 ha of Syncrude’s Gateway Hill certified as re- dominated the region before mining. Thus, wetlands will be re- claimed in 2008, representing 0.15% of land reported as dis- stricted to the depressions between hills and surrounding end-pit turbed by industry), but efforts to create marsh and shallow open lakes (e.g., Fig. 1, Inset). Third, to foster geotechnical stability, the water wetlands are less successful at restoring biological integrity closure landscapes are channelized to drain quickly (e.g., Fig. 1, (19, 20). Even if the goals outlined in closure plans are achieved, Inset). Creating wetland habitat that slows the flow of water can peatland loss will occur with substantial impacts to ecosystem result in soil saturation, gully formation, and landform collapse services, including carbon storage. (13). Fourth, end-pit lakes are designed to remediate tailings Oil sands mining is frequently criticized as a carbon-intensive water (SI Text), and extensive wetlands would increase the evap- means of acquiring oil. Its contribution to the global carbon im- orative surface area of the closure landscape, reducing end-pit balance has provoked numerous calls to slow oil sands de- lake function. Given that precipitation is less than potential velopment, including, most recently, a letter to Canada’sprime evapotranspiration in the oil sands-mineable area, water avail- minister signed by eight Nobel Peace Laureates. Greenhouse gas ability will limit wetland area in the reclaimed landscape. emissions from mining and upgrading oil sands bitumen are

Rooney et al. PNAS Early Edition | 3of5 Table 2. Summary of baseline vegetation cover within the development (DA) or local study areas (LSA) of mines with approval to operate granted by March, 2011 Mildred Jackpine Suncor Steepbank Horizon Lake and Suncor Muskeg mine– and Millenium Aurora mine expansion Basemine and expansion phase 1 Kearl mine mines Fort Hills mine North mine

Bank West West West West East East East East East Units ha in LSA ha in LSA ha in LSA ha in LSA ha in DA ha in LSA ha in DA ha in DA ha in LSA Terrestrial vegetation 17,040 14,662 16,745 2,775 4,408 15,416 2,806 3,350 17,733 Peatlands 5,355 1,870 16,813 3,075 1 9,986 6,422 751 19,714 Riparian communities 2,600 708 0 1,216 1,434 7,804 100 1,012 199 Graminoid marsh 318 0 0 36 523 1 19 6 435 Shallow open water 332 175 61 61 21 42 8 0 249 Wetlands subtotal 8,605 2,753 16,874 4,388 1,979 17,833 6,549 1,769 20,597 Lakes and rivers 267 175 61 43 1,359 561 0 0 580 Disturbed land 1,874 909 1,300 5,270 38 206 0 419 1,197 Total 27,786 18,499 34,980 12,476 7,784 34,016 9,355 5,538 40,107 % wetland 31 15 48 35 25 52 70 32 51 % terrestrial 61 79 48 22 57 45 30 60 44

The west bank is typically drier and supports more upland habitat relative to the east bank, which supports more wetland habitat. As a part of their environmental impact assessments (EIAs), mine operators designate DAs, which represent the footprint of all facilities directly associated with mining, i.e., mine pits, tailings storage, bitumen recovery plants, etc., and LSAs, which include both the DA and a buffer around the DA that is intended to accommodate any potential indirect effects of the proposed development. Baseline conditions are typically presented for either the DA or LSA, but not for both. The vegetation cover values were obtained through a review of baseline studies in EIAs and the most recently updated reclamation, conservation, and closure plans (see SI Text for references).

estimated at between 62 and 164 kg CO2 equivalents per barrel of The boreal forest is the world’s largest and most important oil produced, two to three times more than emissions from con- forest carbon storehouse (24), but its continued storage depends ventional oil production (21). With daily production of mined bi- on future land management practices (SI Text). Based on extensive tumen exceeding 1,142,000 barrels in 2010 (22), emissions add up work in the Mackenzie River Basin, the range in peatland carbon – quickly (>70,000 t CO /d) and hundreds of millions of dollars are storage is estimated at 530 1,650 metric tons (t) C/ha (25), 2 equivalent to 1,943–6,050 t CO /ha. The breadth of this range being invested in reducing and capturing CO (23). These tallies, 2 2 reflects uncertainties associated with variability in peat depth, however, completely neglect the carbon emissions resulting from composition, and bulk density. Unfortunately, this information is peatland loss, yet our analysis suggests that carbon storage loss not available from baseline studies, and we therefore chose to be caused by peatland conversion could be equivalent to 7-y worth of conservative and represent the effects of this uncertainty on the carbon emissions by mining and upgrading (at 2010 levels). range of C values. Reclamation prescriptions for postmining soils

Table 3. Net change in land cover types to result from oil sands mining reclamation based on baseline reports and closure plans for the Horizon, Jackpine–Phase 1, Kearl, and Muskeg mines Net change

Description Total pre (ha) Total post (ha) (ha) (%)

Upland forest 39,114 54,587 15,473 40 Meadow 1 0 −1 −100 Shrubland 524 82 −442 −84 Bog 5,179 1,320 −3,859 −75 Fen 13,238 4,683 −8,555 −65 Graminoid marsh 878 2,595 1,717 196 Swamp 13,054 9,795 −3,259 −25 Shallow open water 456 94 −362 −79 Lake 2,059 5,702 3,643 177 River 171 152 −19 −11 Riparian shrubland 1 2,327 2,326 232,600 Littoral zone 0 230 230 Infinite Clearcut 730 98 −632 −87 Disturbance 6,658 395 −6,263 −94 Peatland subtotal (bog and fen) 18,417 6,003 −12,414 −67 Wetland subtotal 32,806 21,045 −11,761 −36 (peatland, graminoid marsh, swamp, shallow open water, riparian shrubland, and littoral zone) Total 82,060 82,060 0 0

This constitutes 42% of the total area approved for mining as of March 2011, but is a representative sample of the region in terms of east and west bank distribution.

4of5 | www.pnas.org/cgi/doi/10.1073/pnas.1117693108 Rooney et al. contain much less carbon: between 50 and 146 t C/ha (26). Thus, peatlands, not newly sequestered carbon. Additionally, Turcotte’s the replacement of 12,414 ha of peatlands with reclaimed soils will study of soil organic matter in reclaimed land on oil sands mine result in the loss of 4.8–19.9 million t of stored carbon. Based on leases has demonstrated unexpectedly rapid decomposition of the the carbon value estimated by the Intergovernmental Panel on peat in soil amendments, even the relatively recalcitrant lignin Climate Change at $52/t of carbon sequestered (27), this equates phenols (32). This suggests that conversion of peatlands to uplands to a $248 million to $1 billion loss of natural capital, yet we have with peat soil amendments transforms a relatively permanent car- only considered 42% of the area currently approved for mining. bon storage pool (historical peatlands) to a temporary one that Scaling up, as we did with land cover, a loss of between 11.4 and leaks carbon rather than sequesters it. This is supported by Wel- 47.3 million t of stored carbon (between $590 million and $2.5 ham’s model, which predicts that reclaimed forests will require 15 y billion of carbon storage capital) will occur. Converting from units of growth before carbon sequestration by vegetation begins to ex- of carbon to CO2 equivalents, this is between 41.8 and 173.4 t of ceed the carbon emissions from decomposing peat amendments, CO2 lost, as much as 7-y worth of mining and upgrading emissions suggesting that for years following mining and reclamation, at 2010 production levels. reclaimed land will be a net carbon source (31). Peatland loss will also influence the region’s potential to se- quester carbon in the future. Vitt et al. (28) estimated that Conclusion western continental peatlands sequester 19.4 g C/m2 of peatland/y. Claims by industry that they will “return the land we use - in- Accounting for forest fires, Turetsky et al. (29) suggest that the 2 cluding reclaiming tailings ponds - to a sustainable landscape true rate of carbon sequestration is 24.5 g C/m of peatland/y. ” Thus, the loss of 12,414 ha of peatland translates into 2,408– that is equal to or better than how we found it (33) and that it “will be replanted with the same trees and plants and formed 3,041 t of annual carbon sequestration potential. Scaling up, as ” with carbon storage, this equates to 5,734–7,241 t C/y (21,025– into habitat for the same species (34) are clearly greenwashing. The postmining landscape will support >65% less peatland. One 26,550 t CO2/y) lost due to approved mines. The reclaimed landscape will sequester carbon at a much lower rate (28), de- consequence of this transformation is a dramatic loss of carbon termined by complex interactions between plant species (and the storage and sequestration potential, the cost of which has not chemical composition of their litter), climate, soils, management, been factored into land-use decisions. To fairly evaluate the costs and the fire regime (30). Looking at Imperial Oil’s Kearl Lake and benefits of oil sands mining in Alberta, impacts on natural

mine, Welham found that the vast majority of carbon seques- capital and ecosystem services must be rigorously assessed. SCIENCES

tered in the reclaimed landscape was derived from peat ENVIRONMENTAL amendments made to the soil during the first stages of reclama- ACKNOWLEDGMENTS. We thank Tanya Richens and John Keeler for assistance collecting data; Maria Strack for recommending carbon storage tion (31). Given that the peat used in these amendments is obtained studies; and Suncor Energy Inc. for permission to reprint the Inset of Fig. 1. by stripping and stockpiling peat from adjacent land in preparation Funding was provided by Killam Trusts and Alberta Innovates Technology for mining, this fraction is actually residual storage from historical Futures in the form of scholarships to R.C.R.

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