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' "v\ p..- ' THE-$5 Mill"!!!WHIHHHU!IIUIHIIIHUII(IIUHWIIHHM 300794 9765 This is to certify that the thesis entitled An Evaluation of the Influence of Temperature on the Growth of Brook Trout in the Ford River, Dickinson County, Michigan from 1984 to 1991 presented by Melissa Kay Treml has been accepted towards fulfillment of the requirements for Master of Science degree in Fisheries & Wildlife Major professor / Date 451/92 //;L 0-7639 MS U is an Affirmative Action/Equal Opportunity Institution 1' l LIBRARY Hickman State University PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINES return on or before date due. — ————_——u—i—_——I DATE DUE DATE DUE DATE DUE Jl [__ MSU Is An Affirmative ActionlEqual Opportunity Institution cmmwn . ll AN EVALUATION OF THE INFLUENCE OF TEMPERATURE ON THE GROWTH OF BROOK TROUT IN THE FORD RIVER, DICKINSON COUNTY, MICHIGAN FROM 1984 TO 1991 BY MELISSA KAY TREML A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF.SCIENCE Department of Fisheries and Wildlife 1992 'ABSTRACT AN EVALUATION OF THE INFLUENCE OF TEMPERATURE ON THE GROWTH OF BROOK TROUT IN THE FORD RIVER, DICKINSON COUNTY, MICHIGAN FROM 1984 TO 1991 BY Melissa Kay Treml The influence of late spring and summer water temperatures on brook trout growth and age structure was evaluated from 1984 to 1991 in the Ford River, Dickinson County, Michigan. Brook trout were sampled from late May through September using fyke nets and weirs at four locations within a 25.8 river km section of stream. Scale analysis was used to determine age, to estimate past length at age, and to estimate relative annual growth rates. Late spring and summer temperature patterns varied between years. Most variability occurred in May and June. Age and size structure also varied between years and was related to yearly temperature differences. Years with temperatures near 16 C in mid June were dominated by older, larger brook trout, while years already above 16 C by mid June were dominated by younger, smaller brook trout. Temperature had a significant negative affect on brook trout growth from age 2 on. Growth rates were negatively related to the number of days which had temperatures greater than 20 C and to the rate at which the water warmed. Consequently, trout stream managers must consider the thermal regime of a stream when setting management goals. ACKNOWLEDGMENTS The ELF project provided the funding for this study. I would like to thank all of the people who worked on the ELF project during the past nine years, especially Bill Lavoie, Steve Mero, and Tim Nuttle. I would especially like to thank Steve Marod, who through his help and friendship made the many years spent collecting this data a truly memorable experience. I would also like to thank Dr. Taylor and my committee members, Dr. Kevern and Dr. Merritt, for all of their years of help and guidance. A special thanks to Dr. Taylor for his patience and understanding during my times of turmoil and doubt. Many thanks to all of graduate students in Dr. Taylor's lab, who have been more than just my colleagues. They have been my friends. I would especially like to thank Paola Ferreri for all of her help and support both in school and in life. My deepest appreciation goes to my parents (Bob and Joanne), and sister (Chris) who have always stood beside me and encouraged me to reach for my goals, regardless of what they might be. Last of all, I would like to thank my husband, Jim, for all of his love and encouragement. ii TABLE OF CONTENTS LIST OF TABLES . . . . . LIST OF FIGURES . . . . . LIST OF APPENDICES . . . INTRODUCTION . . . . . . DESCRIPTION OF STUDY SITE METHODS . . . . . . . . . Fish Collection. . . Temperature Monitoring . . . . . . . . . . . . Age Determination and Size Structure . . . . . Annual Growth Rates Temperature Patterns Length of Growing Season . . . . . . . . . . . Effect of Temperature on Growth . . . . . . . RESULTS 0 O O O O O O O 0 Fish Collection . . Temperature . . . Age Determination and Size Structure . . . . . Length at Age and Age Specific Growth Rates Relationship between Growth . . . . One Year Olds . Two Year Olds . Three Year Olds DISCUSSION . . . . . . . LITERATURE CITED . . . . Temperature, Length, and iv vi vii 10 10 11 13 16 16 17 17 20 25 27 27 27 29 33 37 60 Table 1. Table 2. Table 3. Table 4. Table 5. Table 6. Table 7. Table 8. Table 9. Table 10. Table 11. Table 12. LIST OF TABLES The total number of net days, total catch, CPUE, and sampling time for the sampling periods from 1984 to 1991. . . . . . . . . . Total net days each year at sites 1, 2, 3, and 4 from 1984 to 1991. Total annual catch at sites 1, 2, 3, and 4 from 1984 to 1991. . . . . . . The mean daily temperature between May 1 and September 30 for each year, the relative rate at which temperatures warmed, and the number of days within the temperature range of optimal growth, poor growth, and no growth for each year. . . . . . . . . . . . . . . . The mean monthly temperatures for May through September from 1984 to 1991. . . . . . . . . The percent age composition of age classes 1, 2, and 3 in the total annual catch from 1984 to 1991. . . . . . . . . . . . . . . . . . . The regression equations describing the body- scale relationship for the 1983 through 1990 cohorts (L = length at age 1, S = scale radius at age i). . . . . . . . . . . . . . . Mean back-calculated length (mm) at age 1 for brook trout from the 1983-1990 cohorts. . . Mean annual growth rates of young of the year brook trout from the 1983-1990 cohorts. . . . Mean back-calculated length (mm) at age 2 for brook trout from the 1983-1989 cohorts. . . . . . . . . . . . . . . . . . . The mean annual growth rates of yearling brook trout from the 1983-1989 cohorts.. . . Mean back-calculated length (mm) at age 3 for brook trout from the 1983, 1984, 1986, 1987, and 1988 cohorts. . . . . . . . . . . iv 18 19 21 23 26 28 30 31 32 34 34 Table Table Table Table Table Table Table Table Table 13. 14. 15. 16. 17. 18. 19. 20. 21. The mean annual growth rates of 2 year old brook trout from the 1983, 1984, 1986, 1987, and 1988 cohorts. . . . . . . . . . . . . . Occurrence of Lee's Phenomenon and Reverse Lee's Phenomenon during the second and third year of life for the 1983-1989 cohorts. . . The average length (mm) at capture for age classes 1, 2, and 3 for each cohort. . . . . Results of the analysis of variance between mean annual growth rate of young of the year brook trout and temperature. . . . . . . . . Results of the analysis of variance between mean annual growth of 1 year old brook trout and temperature. . . . . . . . . . . . . . Results of the analysis of variance between mean annual growth rate of 2 year old brook trout and temperature. . . . . . . . . . . . Results of the analysis of variance between mean length of brook trout at age 1 and temperature. . . . . . . . . . . . . . . . Results of the analysis of variance between mean length of brook trout at age 2 and temperature. . . . . . . . . . . . . . . . Results of the analysis of variance between mean length of brook trout at age 3 and temperature. . . . . . . . . . . . . . . . 36 36 45 54 55 56 57 58 59 Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure 10. 11. 12. 13. 14. LIST OF FIGURES Location of fyke nets and weirs in the study section of the Ford River and Two Mile creek 0 O O O O O O O O O O O O O O O O O O 0 Mean daily discharge for late spring and summer at site 3 calculated on a weekly basis from 1984 to 1991. . . . . . . . . . . Mean daily temperature for late spring and summer at site 3 presented as a weekly average from 1984 to 1991. . . . . . . . . . Fyke nets arranged in tandem across a river. . . . . . . . . . . . . . . . . . . . Twenty-four hour daily temperature pattern for Two Mile Creek determined at 10 minute intervals with a Ryan Tempmentor. . . . . Cumulative mean daily temperature distribution from May 1 to September 30 from 1986 through 1991. O O O O O O O O O O C O 0 Length frequency distribution of the total annual catch in 1984 . . . . . . . . . . . Length frequency distribution of the total annual catch in 1985. . . . . . . . . . . . Length frequency distribution of the total annual catch in 1986. . . . . . . . . . . . Length frequency distribution of the total annual catch in 1987. . . . . . . . . . . . Length frequency distribution of the total annual catch in 1988. . . . . . . . . . . . Length frequency distribution of the total annual catch in 1989. . . . . . . . . . . . Length frequency distribution of the total annual catch in 1990. . . . . . . . . . . . Length frequency distribution of the total annual catch in 1991. . . . . . . . . . . . vi Page 15 22 46 47 48 49 50 51 52 53 Appendix A Appendix B LIST OF APPENDICES vii INTRODUCTION Brook trout are highly regarded game fish and are a favorite among anglers in the eastern United States and Canada (Power 1980). Their range extends from Northern Quebec to Georgia with the northern portion of the range stretching from the Atlantic Ocean to Manitoba and the southern portion of the range confined to the Appalachian ridge (Scott and Crossman 1985, Meisner 1990). They are endemic to North America and are found under conditions which are generally described as clean, pure, and aesthetically desirable (Scott and Crossman 1985, Power 1980). Typical brook trout habitat conditions are those associated with a cold temperate climate, cool spring-fed ground waters, and moderate precipitation (Raleigh 1982). The rate of growth and maximum size of brook trout varies significantly throughout the native range, depending on local habitat conditions (Scott and Crossman 1985). Brook trout have a greater cold water tolerance than other trout with positive growth occurring at temperatures between 5 C and 20 C (Powers 1980) and with the upper lethal temperature being 25.3 C (Fry et. al. 1946). Brook trout growth tends to be optimal between 11 C to 16 C with 2 temperatures warmer or colder reducing growth (Raleigh 1982). Consequently, marked seasonal changes in brook trout growth rates coincide with seasonal changes in water temperature (McFadden et. a1. 1967) with growth rates increasing with temperature and reaching a maximum at 16 C and then progressively decreasing at higher temperatures (Hokanson et. a1. 1973). In Michigan most brook trout growth occurs from March to June with little growth occurring from July through September with the exception of age 0 brook trout who have been noted to grow throughout their first winter (Cooper 1953). In northern Michigan, brook trout are generally slow growing (average 3 year old is approximately 201 mm long) when compared to other stream populations reported in Carlander (1969) and relatively short lived with few fish surviving past their third year (McFadden 1961, Wydoski and Cooper 1966, Cooper 1967). The short life span is most likely a function of high natural mortality and/or exploitation rates from age 2 (150 mm) on (McFadden 1961, Wydoski and Cooper 1966, Flick and Webster 1975, Cooper 1967). Annual growth rates and the length of the growing season have been found to be positively related (Gerking 1966). The growing season of a fish is defined as the period of time where the water temperature remains within the range where positive growth can occur, for brook trout 3 this range is between 5 C and 20 C (Powers 1980). Gerking (1966) reports that populations with rapid growth rates had longer growing seasons than those with slower growth rates. Consequently, it is important to take into account the length of the growing season when comparing annual growth rates of different fish populations (Conover 1990). In addition, since growing season is a function of temperature, temperature variations must also be considered when examining annual growth rates of a single population. The purpose of this study was to determine the growth patterns of brook trout in the upper Ford River from 1984- 1991 and to determine its relationship with growing season and summer water temperatures. This was accomplished by: (1) determining the age and size structure of brook trout in the upper Ford River, (2) determining the annual growth rate for each age class from each cohort of brook trout in the upper Ford River, (3) examining the temperature patterns of the upper Ford River during late spring and summer from 1984-1991, and (4) evaluating the relationship between late spring and summer water temperatures and the annual growth rates of brook trout in the Upper Ford River. DESCRIPTION OF STUDY SITES The Ford River is a fourth order stream in northern Dickinson County, Michigan. Its source is near Sagola in 4 the northwestern corner of Dickinson County. Two Mile Creek is a tributary flowing from southern Marquette County into the Ford River from the north. The Ford River flows into northern Green Bay south of Escanaba, Michigan. The Ford River is classified as a blue ribbon trout stream because of its domination by wild brook trout, stream size and depth, diverse insect life and fly hatches, pure water conditions, and reputation for quality trout fishing (Fisheries Division, Michigan Department of Natural Resources). Four study sites on the upper Ford River were used to collect information on brook trout age and growth from 1984- 1991 (Figure 1). The first three sites were located on the mainstream of the Ford River and the fourth site was located on Two Mile Creek, a tributary. Site 3, the downstream site, was approximately 1.62 river km upstream of Ralph, Michigan. Site 2 was approximately 14.7 river km upstream of site 3. Site 1 was 11.1 km upstream of site 2. Site 4 was located on Two Mile Creek, approximately 11.5 km upstream of site 2. The Ford River typically has high spring discharge and low summer discharge (Figure 2). Temperatures rise during the spring and reach a high anywhere from late June to late July and remain high through August (Figure 3). The downstream region of the study section is characterized by a sandy bottom. The upstream region of the study section of river has areas with substrate ranging from pebbles to large rocks, intermitted .xoouo mafia 039 can um>flm puom may no :o«uomm >psum on» Ca muwo3 can mum: oxau no :ofiuoooa .H ouzmfim mtm em; I mEm .52 5;”. 4 mm>_m GEO”— $i N v xmmmo Guzman... ¥mm¢0 mumm; xwwmo m.=s_ 03... 12 I _ g; --1984 ------ 1985 ------------ 1986--- 1987 10 " l '00-. O -- 11111.1111141 >r >— >_ 3.. g- g.— g- g— L.— L— JJJJJAAAAASSSSOOO 12 _ ———-1988 ------ 1989 ------------ 1990--- 1991 I 10 MEAN DISCHARGE (cu. m/sec) l l l l l l l l l l l l..l. l l l l J l l 1 l AAAMMMMJJJJMJMJAAAAASSSSOOO WEEK Figure 2. Mean daily discharge for late spring and summer at site 3 calculated on a weekly basis from 1984 to 1991. .Hmma ou emma souu ommuu>m >qum3 m we cousomoum m mufim um Mozasm can msfiumm mama you musumuomsou hafimp cam: .m ousmfim 2mm; OOmmmm<<<<<=.=._—.=._a—.fi_.EEEE<< u d u u q d Ommmm<<<<<__..=a=.=._._._.—.2225<< - q q q d ( o) aunivuadwal with regions of sand. § METHODS Fish Collection Brook trout were generally collected with passive gear at the four study sites from at least mid-May to mid- September from 1984 to 1991. Passive gear was used to take advantage of the movement patterns of Ford River brook trout noted by other researchers (Marod and Taylor 1991). Sites 2 and 3 were fished with 1/2 inch bar mesh fyke nets arranged in tandem with one net facing upstream and one net facing downstream (Figure 4). Sites 1 and 4 were fished with 1/2 inch bar mesh hardware cloth weirs arranged in tandem. All gear was fished 7 days/week until the mean daily catch of brook trout fell below 1 fish/day, after which all gear was fished continuously from Monday morning through Friday evening. Nets were checked once daily. All wild brook trout captured were anesthetized with MS-222 at a 500 mg/l of water dosage in order to reduce handling stress (Meister and Ritizi 1958 and Schoettger and Julin 1967). Fish were then measured for total length (nearest 1 mm), weighed on a calibrated Ohaus Port-o-Gram scale (nearest 0.1 gram), and given a site specific fin clip. Additionally, in May and June a scale sample was taken above the lateral line and anterior to the dorsal fin for age and growth determination. After recovery in fresh water, all fish were released in .uo>flu m mmouom Eoccmu ca pomcwuum mum: mxxm .v ousvfih \ 30...... so: 10 their original direction of travel. Recaptured fish were again measured, weighed, checked for a fin clip, and released. Temperature Monitoring Late spring and summer water temperatures were monitored (half hour intervals) with Omnidata data pods using thermistors at sites 2 and 3 from mid-April to October (Burton 1991). Temperature was monitored at site 4 using Ryan Tempmentors in 1988 (10 minute intervals), 1990 (10 minute intervals), and 1991 (30 minute intervals). Temperature was not monitored at site 4 in 1989 because of equipment failure. The Ryan tempmentors were installed from late June to mid—September in 1988 and 1991 and from early May to mid-August in 1990. In addition, Wecksler max-min thermometers calibrated daily with a laboratory thermometer were used to monitor maximum and minimum temperature at sites 2, 3, and 4 for all net days in all years. MW A stratified random subsample of brook trout were aged from each year's total catch by counting scale annuli as described by Cooper (1951), McFadden (1959), and Van Oosten (1929). The criteria for an annuli were those given by Cooper (1951); the crowding of adjacent circuli, irregularity or incompleteness in circuli form, the cutting over of circuli in the posterio-lateral areas and the sudden change in the growth pattern of the circuli. Only scale 11 samples taken in May and June were used for age determination and annual growth determination because samples taken as close to annulus formation as possible have the least amount of variation in the body-scale relation (Carlander 1982, Weatherly and Gill 1987). The mean length at capture for age classes 1, 2, and 3 was then determined. The determined ages were used to construct an age-length key to estimate the age structure of the total brook trout catch each year. Due to small sample sizes (N < 6 for all years combined) of fish greater that 3+ years olds, only fish through age 3+ were included in my study. The size structure of each year's brook trout population was described by a length frequency distribution of the total yearly catch. For the 1988 total catch no age 3 (1985 cohort) fish were aged because no scales were collected from age 3 fish; consequently, the age-length key for the 1988 total catch only contains age 1 and 2 fish. WM Once aged, the fish were separated into cohorts to minimize error in the body-scale relation (Carlander 1981). For example, age 1 fish caught in 1984 belonged to the 1983 cohort as did age 2 fish caught in 1985 and age 3 fish caught in 1986. This was necessary because a sample taken at one time really represents a series of year classes. As a result, a single years catch cannot be used to determine the body-scale relationship and the back-calculation 12 equations for brook trout from different cohorts. The annual growth rates of the brook trout in the upper Ford River watershed were estimated by first back—calculating the previous lengths at age from scale analysis (Bagenal and Tesch 1978). The Fraser-Lee method of back-calculation was used to estimate past length at age. This method assumes that body growth of the fish is related to the proportional growth of its scale (Carlander 1981). The Fraser-Lee back-calculation formula is (Carlander 1981): s. Li-a+[Lc-a]*——l SC Where, 1% = length at capture a = Y-intercept of the body-scale regression 1% = length at age i Sc== scale radius at capture Si== scale radius at age i The back—calculated lengths were then compared to observed lengths at capture for each age class of the same cohort to determine if the back-calculated lengths were realistic. Relative annual growth rates were determined using the equation given by Ricker (1975): 13 where, G = annual growth rate 134 = length at age i-1 I8 = length at age i A relative growth rate was used because the effects of temperature on growth are dependent on the size of the fish (Baldwin 1956). In addition, annual growth rates were only calculated from the last complete year of growth for each age class, i.e. age 1 growth rate was determined only from age 2 fish and age 0 growth rates were determined from only age 1 fish. This method minimizes uncertainty due to Lee's Phenomenon and reverse Lee's Phenomenon (Gutreuter 1987), which was found to occur in several cohorts. For the annual growth rate of young of the year (YOY) fish, length at time i-l was assumed to be 22.86 mm which is the average size of brook trout in northern Wisconsin streams at swim-up and the onset of feeding (Avery 1983). The average size at swim-up for brook trout in northern Wisconsin streams was used rather than that of brook trout from streams in the Lower Peninsula of Michigan. This was because the Ford River is thermally and geologically more closely related to northern Wisconsin streams. W The mean daily temperature was calculated for each day at sites 2, 3, and 4. At site 4, the daily maximum, minimum, and current temperature when the weirs were checked were averaged and presented as the mean value. Due to the 14 cyclic nature of the daily temperature patterns, these three points were found to adequately estimate the mean daily temperature when compared to mean daily temperatures obtained from the tempmentors (Figure 5). Similarity in water temperature between sites was tested with Pearson's Correlation. Similarity between sites allowed the use of data from only one site when evaluating the relationship between growth patterns and late spring and summer temperatures. Only temperatures between May 1 and September 30 of each year were used because temperatures were below the range for optimal growth prior to May 1 and after September 30 in all years of my study. Cumulative mean daily temperature distributions from May 1 to September 30 were used to describe the temperature patterns of each year. In addition, I measured the relative rate at which the water warmed each year by counting the number of days from May 1 that it took to reach a mean weekly temperature of 11 C (lower end of range for optimal growth), 16 C (optimal growth), and 20 C (upper bound on positive growth). I also measured the number of days the water was within the range of optimal growth, poor growth (greater than 16 C but less than 20 C), and no positive growth (greater than 20 C). 15 .uoucoamsoe cm>m c nuw3 mam>uousfl ouasfia ca um DUCMEumumc xomuu mafia 038 now sumuumm musuouomemu mawmc Mao: u50u1>ucmsa .m ousmflh 855E o : ._<>mm:.z_ moszmsmzm: 00 w O¢_. ON P 00F 00 00 0? ON C _ _ _ _ _ _ _ N. F ( o) aanlvuadwal VN l6 Len o row'n Season The growing season is the time when water temperatures support positive growth (5 C to 20 C). Since the water temperatures reached 5 C prior to the onset of temperature monitoring in my study, the length of the growing seasons had to be estimated indirectly. One indirect way of comparing the effects of different growing seasons on brook trout growth was to compare the number of days during the summer that had temperatures too high for positive growth (number of days greater than 20 C) to the mean annual growth rates and lengths at age of brook trout. An alternative method was to compare the relative rate at which water temperatures reach a mean temperature that is best suited for the overall welfare of brook trout (11 C) to the lengths at age and age specific growth rates for each cohort. Since 11 C is at the lower end of the temperature range best for the overall welfare of brook trout (Raleigh 1982), the number of days from May 1 that it took to reach a weekly mean temperature of 11 C were used to evaluate the effects of the relative rate at which temperatures rise in the spring on annual growth rates and length at age. Eii2QE_2£_IQEEQEQEBES_QD_QLQEED The effect of temperature on growth was evaluated by comparing the following temperature conditions with the mean length at age and the mean age specific growth rate for each age class for each cohort: 1) the mean daily temperature 17 between May 1 and September 30, (2) the cumulative temperature distribution (3) the rate at which temperatures rise (4) the number of days with temperatures within the range of optimal growth (5) the number of days with temperatures greater than those for optimal growth but still with the positive growth range, and (6) the number of days with temperatures higher than the upper bound on positive growth. RESULTS Fish Collection During the sampling periods from 1984 to 1991, the total number of net days varied from 197 in 1986 to 335 in 1984 (Table 1). The number of net days at each sampling site varied between sites and between years (Table 2). The mean annual catch was 590.6 fish and ranged from 317 in 1986 to 1186 in 1984 (Table 1). In addition, the total number of fish captured at each site varied among sites and years (Table 3). Site 4 had the highest annual catch every year except 1987 where site 2 had the highest annual catch. In 1989 site 1 and 4 had equal annual catches. Mean catch per unit effort (CPUE - mean daily catch per net per day) was 2.40 and ranged between 1.28 in 1989 and 3.54 in 1984 (Table 1). Sampling had began by the third week of May in all years except 1987 in which sampling did not begin 18 Table 1. The total number of net days, total catch, CPUE, and sampling time for the sampling periods from 1984 to 1991. Year Number of Total CPUE Sampling Time net days catch 1984 335 1186 3.54 5/14 to 11/10 1985 214 616 2.88 5/22 to 9/18 1986 197 317 1.61 5/21 to 9/19 1987 201 673 3.35 6/16 to 10/10 1988 218 333 1.53 5/19 to 10/6 1989 253 324 1.28 5/23 to 10/13 1990 265 400 1.51 5/30 to 9/17 1991 253 876 3.46 5/17 to 9/11 19 Table 2. Total net days each year at sites 1, 2, 3, and from 1984 to 1991. 4 Year Site 1984 1985 1986 1987 1988 1989 1990 1991 1 44 53 52 48 56 52 69 69 2 78 42 42 57 55 69 69 59 3 92 58 52 59 53 61 58 52 4 121 61 51 37 54 71 69 73 Table 3. Total annual catch at sites 1, 2, 3, and 4 from 1984 to 1991. Year Site 1984 1985 1986 1987 1988 1989 1990 1991 1 180 62 34 16 28 O 139 2 313 147 84 148 72 94 33 3 170 97 77 357 47 51 89 4 523 310 122 152 186 179 139 90 123 109 554 20 until mid June due to the late arrival of necessary equipment (Table 1). Temperature The temperature at sites 2, 3, and 4 were all highly correlated during the late spring and summer each year (minimum p = .758, maximum p = 0.999). Consequently, site 3 was chosen for all remaining temperature calculations. The mean daily temperatures for each year between May 1 and September 30 are given in Table 4, along with temperature variables used to describe the relative rate at which water temperatures rise, the number of days within the range of optimal growth, poor growth, and no positive growth. The average for the late spring and summer of all years was 16.26 C and ranged from 15.35 C in 1985 to 17.67 C in 1988. One way analysis variance detected significant differences between the means ( F = 7.14, df = 7, P < 0.05). Fisher's Least Significant Difference (LSD) multiple comparison test (P < 0.05) revealed that the mean temperatures during the study period in 1985 (15.4 C) and 1990 (15.4 C) were significantly lower than the mean temperatures in 1988 (17.7 C) and 1991 (16.8 C), and the means of the remaining years fell in between. The temperature patterns for the periods from May 1 to September 30 for each year are depicted by the cumulative mean daily temperature distributions in Figure 6. The Kolmogorov Smirnov test (P < 0.05) detected two distinct cumulative temperature distributions. The distributions for 21 Table 4. The mean daily temperature between May 1 and September 30 for each year, the relative rate at which temperatures warmed, and the number of days within the temperature range of optimal growth, poor growth, and no growth for each year. Mean Daily Temperature Days to 11C1 Days to 16C1 Days to 20c1 Days > 16C Days > 20C Days Between 11C & 16C Days Between 16C & 20C 1984 1985 1986 Year 1987 1988 1989 1990 1991 15.9 15.4 16.4 14 32 97 8O 13 18 65 7 53 153 55 46 100 0 27 78 69 20 27 51 16.7 17.7 15.8 15.4 16.8 5 2 13 O 10 39 27 50 42 23 45 29 63 58 75 67 81 66 67 90 28 48 20 11 21 34 25 37 42 13 TMean weekly temperature on umnfimummm on H has Eouu :oflusnfluumfic muaucummamu aafimc Emma m>fluoazano CON 2 .62 65:99 > 16C 0.0782 0.5435 Days > 20c 0.0212 0.7553 Days Between 110 & 16C 0.0350 0.6878 Days Between 16c & 20C 0.0727 0.5586 TvaIean weekly temperature 55 Appendix B Table 17. Results of the analysis of variance between mean annual growth of 1 year old brook trout and temperature. 33 Probability Mean Daily Temperature 0.1268 0.5564 Days to 11c1 0.0007 0.9547 Days to 1601 0.1137 0.4595 Days to 20c1 0.0144 0.7976 Days > 16C 0.2593 0.4431 Days > 200 0.0658 0.5786 Days Between 11c & 16c 0.1386 0.4108 Days Between 16C & 200 0.0526 0.6208 1 Mean weekly temperature 56 Appendix B Table 18. Results of the analysis of variance between mean annual growth rate of 2 year old brook trout and temperature. RE Erobability Mean Daily Temperature 0.1268 0.5564 Days to 1101 0.8847 0.0172 Days to 1601 0.5547 0.1487 Days to 20c1 0.0285 0.7861 Days > 16C 0.1604 0.5041 Days > 20c 0.0533 0.7086 Days Between 110 & 16c 0.1175 0.5723 Days Between 16C & 20c 0.0001 0.9872 1 Mean weekly temperature 57 Appendix B Table 19. Results of the analysis of variance between mean length of brook trout at age 1 and temperature. 33 a Mean Daily Temperature 0.0278 0.7209 Days to 11c1 0.0452 0.6473 Days to 1601 0.0907 0.5117 Days to 20c1 0.0284 0.7181 Days > 16C 0.0812 0.5355 Days > 20C 0.0219 0.7514 Days Between 11C & 16C 0.0376 0.6768 Days Between 16C & 20C 0.0744 0.5541 1 Mean weekly temperature Table 20. Results of the analysis of variance between 58 Appendix B mean length of brook trout at age 2 and temperature. 83 22223211152 Mean Daily Temperature 0.6004 0.0408 Days to 1101 0.1536 0.3845 Days to 16c1 0.2245 0.2823 Days to 20c1 0.4284 0.1107 Days > 16C 0.0441 0.6512 Days > 20C 0.5051 0.0734 Days Between 110 & 16C 0.0738 0.5557 Days Between 16C & 20C 0.3493 0.1623 T*Mean weekly temperature Table 21. 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