PB81-198954 Natural Variation in Abundance of Salmonid Populations in Streams and Its Implications for Design of Impact Studies. A Review Oregon State Univ. Corvallis Prepared for Corvallis Environmental Research Lab., OR Feb 81 U.S. DEPARTMENT OF COMMERCE National Technical Information Service ------- EPA 600/3-81-021 Febraury 1981 NATURAL VARIATION IN ABUNDANCE OF SALMONID POPULATIONS IN STREAMS AND ITS.IMPLICATIONS FOR DESIGN OF IMPACT STUDIES A Review by James D. Hall and Ned J. Knight Department of Fisheries and Wildlife Oregon State University Corvallis, OR 97331 Project Officer Jack H. Gakstatter U.S. Environmental Protection Agency 200 S.W. 35th Street Corvallis, OR 97330 ------- . NOTICE THIS DOCUMENT HAS BEEN REPRODUCED . FRO M THE BEST COP Y F URN ISH ED USB Y THE SPONSORING AGENCY. ALTHOUGH IT . . 1S RECOGNIZED THAt CE~TAIN PORTIONS ARE ILLEGIBLE, IT IS BEING RELEASED '. . IN THE INTEREST OF MAKING AVAILABLE AS MUCH INFORMATION AS POSSIBLE. ------- BIBLIOGRAPHIC INFORMATION PB81-198954 Natural Variation in Abundance of Salmonid Populations in ,Streams and Its Implication's for Design of Impact Studies. A Review, , Fe b 81 JamesD. Hall, and Ned J.Knight., PERFORMER: Oregon State Univ., Corvallis. Dept. of Fisheries and wildlife. EPA~600/3-81-021 . SPONSOR: Corvallis Environmental Research Lab., .OR. ," ------ .----. -- -- - - -'------.. --~ Literature on stock size and production of salmonid populations in streams has been reviewed. The objective is to bring together data on the magnitude of natural variation in population size and to relate this variability to environmental conditions where possible. Recommendations are presented for the use of this information in designing studies to ~stimate the impact of non-point source pollutiion. A partially annotated bibliography of 260 relevant reference is included. KE YWO RDS: *Ecology, *Salmon, *Water pollution. Available from the National Technical Information Service, Springfield, Va. 22161 PR ICE CODE: PC A05/MF AO 1 i ------- DISCLAIMER This report has been reviewed by the Corvallis Environmental Research Laboratory, U~S. Environmental Protection Agency, and approved for pub- lication~ Approval does not signify that the contents necessarily reflect the views and policies of the U.S. Environmental Protection Agency, nor does mention of trade names or commercial products constitute endorsement or recommendation for use. ii ------- ABSTRACT , ' Literature on stock size and, production of salmonid populations in streams has been reviewed. The objective is to bring together data on the magnitude of natural variation in population size and to relate this vari- ability to environmental conditions where possible. Recommendations are presented for the use of this information in,designing studies to estimate the impact of non-point source pollution. A partially annotated bibliography of 260 ,relevant references is included. ' A number of long-term studies, some up to 15-20 years, have provided useful data on temporal variation in population abundance. Other studies have examined spatial variation. Data from the best examples of both kinds of variation are presented in, Appendix Tables. Temporal and spatial varia- tion may be as high as several orders of magnitude in the extreme, and even at the least are sufficient to mask very significant perturbations caused by non-point source pollutants. Environmental variables most closely associated with spatial variation are those relating to the quality of salmonid habitat, particularly physical characteristics such as cover in its many forms. ' Streamflow and food abundance have been associated with both temporal and ~patial variation. In general, physical characteristics of, habitat seem most promising as descriptors of variability. Systems of rating habitat quality should receive considerable emphasis in attempts to minimize the effects of natural variation in the evaluation of impacts of non-point source pollutants. First priority should be placed on assessment of physical features. This approach has been used so far mainly to explain spatial variation,' but has promise of explaining temporal variation as well, particularly in reference to fluctuation in streamflow. . The other major emphasis should be in further development of systems of stream and watershed classification. The most useful of these devised to date take a perspective from geomorphology and focus on the potential of a stream system for oiological production. More emphasis on study of basic processes in stream ecosystems and more extensive use of paired comparisons in design of impact studies are also suggested as means of more clearly separating natural variation from damage caused by non-point source pollutants. . ., ~ III ------- CONTENTS Abstract. . . . . . . . . . . . . . . . . . . . . Tables. . . . -- . . . . . . . . . . . . . . . . . Appendix Tables. Acknowledgment. . . . . . . . . . . . . . . . . . . . . ; . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. 2. 3. 4. Introduction. . . . . . . . . '. ; . . '. . . . . . . . . . . . Conclusions and Recommendations. . . . . . . . . . . . . . . Studies of Variability. . . . . . . . . . . . . . Factors Affecting Natural Variability. . . . . . . . . . . . Physical factors. . . . . . . . . . . . . . Biological factors. . . . . . . . . . . . . . . . . . . Other factors~ . . . . . . . ," . . . . . . . . . Minimizing the Effects of Variability in Impact Studies Habitat quality rating systems. . . . . . . . . . Process studies. .'. .'. . . . . . . . . . . . . . . . . Stream class{fication. . . . . . . . . . . . . . . Improved study design. . . . . . . . . . . . 5. Bibliography. . . . . . Appendix. . . . . . . . . . .8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . --- - - - - - . - Preceding page blank v iv vi vii x 1 2 3 5 5 15 20 22 22 24 24 25 30 60 ------- Number TABLES 1 Long-term Studies of Stream Salmonid Populations. . . . . . . . 2 Correlations Between Mean Monthly Discharge and Annual Smolt Count and Mean June-April Biomass for Coho Salmon, Alsea Watershed Study, June 1960-May 1969 . . . . . . 3 Correlations Between Mean Monthly Discharge and Mean September Biomass in G/M2 for Cutthroat Trout, Alsea Watershed Study, October 1961-September 1972. . . . . . . . . 4 Correlations Between Mean Monthly Discharge and Mean Annual Biomass of Brown, Brook, and Rainbow Trout, Sagehen Creek, California, 1954-1961. . . . . . . . . . . . . 5 Summary of Advantages and Disadvantages of the Four Maj or Approaches to. Watershed Stream Analysis. . . . . . . . vi Page 4 6 7 9 27 ------- Number A-I A-2 A-3 A-4 A-5 A-6 A-7 A-8 A-9 A-10 A-ll A-12 APPENDIX TABLES , , Potential Egg Depos1tion and Freshwater Survival of Pink Saimon, Sash in Creek, A!aska, 1940-1959. . . . . . . . Weir Counts of Coho Salmon Fry and Smolts, Sashin Creek Alaska, 1956-1968. . . . . . . . . . . . . .' . . . . . Weir Counts of Downstream Migrating Pink and Chum Salmon Fry, Hooknose Creek, British COlumbia, 1947-1956 . . . Biomass of Coho Salmon and Rainbow and Cutthroat Trout in Streams in the Vicinity of the Carnation Creek Watershed, British Columbia, 1970-1977 . . . . . . . : . . . Biomass of Cutthroat Trout, Coho Salmon, and Sculpin in , Different Habitats of Six British,Columbia Streams, 1973-1976. . . . . . . . . . . . ',' . . . . . . . . . . . . Escapement, Potential Egg Deposition, and Freshwater Survival of Wild Coho Salmon, Minter Creek, Washington, 1938-1953. . . . . . . . . . . . . . . . . : . . . . . . . . Counts of Spawning Coho Salmon and Smolts at Downstream Weir on Gnat Creek, Oregon, 1954-1959. . . . . . . . . . . . Escapement, Potential Egg Deposition, and Freshwater Survi val of Coho Salmon, Deer Creek, Oregon, 1959-1971 Estimated Biomass of Juvenile Coho' Salmon, Deer Creek, Oregon, 1959-1968. . . . . . . . . . . . . . . . . . . Escapement, Potential Egg Deposition, and Freshwater Survival of Coho Salmon, Flynn Creek, Oregon, 1959-1971. . . Estimated Biomass of Juvenile Coho Salmon, Flynn Creek, Oregon, 1959-1968. . . . . -'-~ ,- -' . . . . . . . . . . . . . Escapement, Potential Egg Deposition, and Freshwater Survival of Coho Salmon, Needle Branch, Oregon, 1959-1971. . . . . . . . . . . . . . . . . . . . . . . . . . vii Page 61 62 62 63 64 65 65 66 66 67 67 68 ------- 'Number A-i3 A-14 A-15 A-16 A-17 . A-18 A-19 A-20 A-21 A-22 A-23 A-24 A-25 A-26 A-27 APPENDIX TABLES (continued) Estimated Biomass of Juvenile Coho Salmon, Needle Branch, Oregon, 1959-1968. . . . . . . . . . . . . Biomass of Cutthroat Trout, Alsea Watershed Study, 1962-1973. . . . . . . . . . . . . . Biomass of SalmonidSpecies in Three Northern California Streams, 1967-1969 . . : . . . . . . . . . . . . . . . .Biomass of Brook Trout in 10 Sections of Sagehen Creek, California, 1952-1961. . . . . . . . . . . . . . . . . . . . Biomass of Brown Trout in 10 Sections of Sagehen Creek, California, 1952-1961. . . . . . . . . . . . . . . . . . . . Biomass of Rainbow Trout in 10 Sections of Sagehen Creek, California, 1952-1961. . . . . . . . . . . . . . . . . Escapement, Potential Egg Deposition, and Freshwater . Survival of Coho Salmon, Waddell Creek, California, 1933-1940. . . . . . . . . . . . . . . . . . . . . . . Downstream Trap Counts of Steelhead Trout by Age Group, Waddell Creek, California, 1933-1942 .. . . . . . Biomass of Brook, Rainbow, and Brown.Trout, Trout Creek, Montana, 1950-1951 . ... . . . . . . . . . . . . . . . Biomass of Brown, Rainbow, and Brook Trout in 11 Sections of Little Prickly Pear Creek, Montana, Summer 1966 . . Number of Cutthroat Trout. Trapped, and Number Remaining in Stream After Trap was Removed, Arnica Creek, Yellowstone Park, Wyoming, 1950-1958 . . . . . . . . . . . " Biomass of Brook Trout, Lawrence Creek, Wisconsin, 1953-1957. . . . . . . . . . . . . . . . . . . . . . . . . . September Population Estimates of Ages 0 and I Brook Trout in Lawrence Creek and Big Roche-A-Cri Creek, Wisconsin, 1953-1964 . . . . . . . . . . . . . . . . . Biomass of Brook'Trout by Age Group in April and September, Lawrence Creek, Wisconsin, 1960-1970 . . . . . . . . . . . . Annual Production of Brook Trout by Section and Age Group, Lawrence Creek, Wisconsin, 1960-1970 . " . . . . . . . . . . viii Page 69 69 70 70 71. 71 72 72 73 73 74 75 75 76 77 ------- 'Number A-28 A-29 A-30 A-31 A-32 A-33 A-34 A-35 A-36 A-37 APPENDIX TABLES ( continued) Physical Characteristics and Biomass of Brook, Brown,'and Rainbow Trout in Sections of Three Michigan Streams, 1937 . . . . . . . . . . . . . . . . ." . . . . Number of Brook Trout Present in September in Hunt Creek, Michigan by Age-Group. ',' . . . . . . . . . . . . . . Mean Annual Biomass of Brook Trout in Streams in Matamek Watershed" Quebec, 1971-1973 . . . . . . . . . . . . . Numbers of Brook Trout in a 4l1~m Section of Hayes Brook, Prince Edward Island, 1947-l~60. . . . . . . . . . . . Counts of, Atlantic Salmon Smo1ts and Seaward Migrating Brook Trout, Little Codroy River, Newfoundland, 1954-1963. . . . . . . . . . . . .,. . . . . . . . . . . . . Biomass of Brown Trout in Tributaries and the Main Stem of the Upper River Tees System, England, ,1967-1970 . . . . . Mean Biomass of Brown Trout in Five Tributaries of the River Tees System, England, in May, August, and October. . . '. . . . . . . . . . . . . . . . . . . e. . Production of At1antic Salmon and Brown Trout in Three Sections of Shelligan Burn, Scotland, 1966-1968. . . . . . . Biomass of Atlantic Salmon and Brown Trout at the End of the Growing Season, She11igan Burn, Scotland, 1966-1975. Biomass of Brown Trout in Six Sections of Horokiwi Stream, New Zealand, 1940-1941 . . . .- . . . . . . . . . . . . . . . ix Page 78 79 80 81 81 82 83 84 85 - 85 ------- ACKNOWLEDGMENT We appreciate the advice of Mr. Michael Crouse, who provided the original impetus for the review, and that of Dr. Jack Gakstatter, Project Officer. Drs. Hiram Li and Terry Finger reviewed an early version of the manuscript. Dr. Scott Overton a4ded useful comments to the section on ' study design. Dr. Richard Gard graciously provided unpublished data from Sagehen Creek, California. The p~rmission of copyright holders and publish- ers to reprint data tables is gratefully acknowledged. One of us (J.D.H.) is especially appreciative of space and facilities made available by Dr. 'W. C. Clark, Zoology Department, University of Canterbury, and Dr. R. M. McDowall, Fisheries Research Laboratory, Ministry of Agriculture and Fisheries, Christchurch, New Zealand, where final work on the manuscript' was completed. We would also like to acknowledge the secretarial staff of the Department of Fisheries and Wildlife for exceptional service under difficult conditions, and Marthanne Norgren for skillfully coordinating final production of the manuscript. This is Technical Paper No. 5608, , Oregon Agricultural Experiment Station. ' ' x ------- SECTION I INTRODUCTION Assessment of impacts on streams caused by non-point source pollutants is now receiving increasing atte~tion. Salmonids are the principal fish species of economic importance affected in the western United States.Assess- ment of damage to these populations cannot be undertaken without some under- standing of natural variation in abundance within and between populations. Strategies of analysis' must be devised. that will separate natural variation from effects due to disturbance~ It is the purpose of this review to bring' together literature and unpublished data on the natural variation in abundance of salmonid populations in streams and to attempt to relate this variation to environmental variables--physical, chemical, and biological. There are two kinds of variability to be considered, spatial and temporal. Spatial variability can be studied at several levels of resolution, ranging from microhabitat preferences to variability within and between streams. . Tempor~l fluctuations in abundance can occur on a diel,. seasonal, or .annual scale. . .' . This paper will concentrate on studies ofsalmonid spec1es during that part of their lives spent in the stream environment. The species include the coho salmon (Oncorhynchus kisutch), chinook salmon (0. tshawytscha), pink salmon (0. gorbuscha), chum salmon (0. keta), brown trout (Salmo trutta), rainbow trout(S. gairdneri), steelhead trout (S. gairdneri-gaLrdneri), cut- throat trout (S~ clarki), Atlantic salmon (S. salar), brook.trout (Salvelinus fontinalis), and Dolly Varden (S~ malma). We began this review with the intention of emphasizing studies on the west coast of North America. However, we found that most of the quantitative data on variability in resident salmonid populations came from other areas, and much of that information has been included. . Much less information is available on population levels of the fish species associated with salmonids. Though not included here, the importance of this element of the aquatic system should be emphasized and steps taken to fill this gap in our knowledge of stream fish communities. "-- .~- I ------- SECTION 2 . . CONCLUSIONS AND RECOMMENDATIONS The standing stock biomass of salmonid fishes in streams shows great natural variation, both in time ~d space. 2Reported levels of biomass vary from zero or just above to just over 60 g/m. This variation is sufficient to mask large-scale perturbations caused by non-point source pollutants, such as result from logging and agricultural practices. Among the most important ,causes of variation are differences in physical characteristics of streams, inCluding streamflow and habitat quality, particularly cover. Biological factors, such as food abUndance and predation, may sometimes influence abund- ance, but their mode of action is less clear and the case for their involve- , ment more equivocal than that of the physical elements of the habitat. We recommend several courses of action that will help to minimize the effects of this natural variation when attempts are made to evaluate impacts of a particular non-point source pollutant. Habitat quality rating systems are being developed that show promise of explaining much of the spatial varia- tion in salmonid populations in streams. These rating systems are based pri- marily on assessment of physical features. They may also help to explain temporal variation caused by changes in streamflow, but other influences on temporal variation need further study. The other major approach that may aid impact assessment is development of schemes of , stream and watershed classifi- cation, such as those of Platts (1974) and Warren (1979). The latter is particularly promising in that it focuses' on the potential of a system for biological production, rather than a particular value of the moment, and takes a biogeoclimatic perspective. Continuing emphasis on study of the basic physical and biological processes that lead to growth, mortality, and produc- tion of. stream salmonids is another promising approach to understanding natural variation in abundance. Finally, new approaches to the design of impact studies are suggestd that may aid in more clearly separating natural variation from that c~used by non-point source pollutants and in monitoring the time required for biological systems to recover from perturbation. 2 ------- SECTION 3 STUDIES OF VARIABiLITY Natural variability of salmonid populations in streams has been measured by two principal methods. In some streams, weirs or traps have been con- structed to get reliable counts of migrating fish. Other studies have examined standing crops in the stream by electro shocking, netting, or angling. There have beert a number of important long~term studies on natural variation in abundance ofanadromous and resident species, which are briefly described in Table 1. As an aid to. further analysis, data from these studies and others of shorter duration that deal with spatial variation have been compiled from original sources and are included in tables in the Appendix. Further description of many studies is included in the annotated bibliography. We performed some preliminary analyses on the data in the Appendix Tables and in other publications, in search of general patterns in variation ..- over the species and geographical areas included. We used the range in abundance as a fraction of mean abundance for a measure of relative vari- ability, rather than the coefficient of variation, owing to small sample sizes. Not surprisingly, the extremes of temporal variation occur in pink and chum salmon fry; their numbers may vary over several orders of magnitude. The most stable populations are those of brook trout in Wisconsin and . Michigan, where the range is in the order of only one-half the mean abundance. . Notably, two of the most useful analyses of variation and its causes were from these two populations (McFadden et al. 1967; Hunt 1974). Where good compari- sons of both temporal and spatial variation could be made in the same stream system (Sagehen Creek, Calfiornia and Lawrence Creek, Wisconsin), spatial variation was the greater, by a significant margin. This, again, may not be a surprising result, but is one with important implications for impact studies. . It appears that inferences about natural fluctuation in abundance and its causes may best be found in detailed analyses of individual research studies, including information on as many relevant environmental variables as possible. Thus the bulk of this review is concerned with attempts to relate variation in abundance to the environmental factors with which it may be associated. 3 ------- 'TABLE 1. /. i. LONG-TERM STUDIES 9F STREAM SALMONID POPULATIONS. J Location Sashin Creek, Alaska Hooknose Creek, British Columbia Carnation Creek, British Columbia Minter Creek, Washington A1se.a River, Oregon Waddell Creek, California Sagehen Creek, California Lawrence Creek, Wisconsin '. Hunt Creek, Michigan Au Sable River, Michigan Hayes Brook, Prince Edward Island Little Codroy Ri ver , Newfoundland Shelligan Burn, Scotland Species pink salmon coho salmon pink salmon chum salmon coho salmon cutthroat trout stee1head trout coho salmon coho. salmon cutthroat trout coho salmon stee1head trout rainbow trout brown trout brook trout brook trout brook trout . brook trout brown trout brook trout Atlantic salmon brook trout Atlantic salmon brown trout I.i" InclUSive Dates II. for Data Presented . 1940-1959 1956-1968 1947-1956 1970-1977 (con tinuing) 1938-1953 1959-1973 1933-1944 1952-1961 1953-1970 1949-1962 1957-1967 1947-1960 1954-1963 1966-1975 Principal Reference Merrell (1962) Crone and Bond (1976) Hunter (1959) "Narv-er.and':Arraersen (1974) Sa10 and Bayliff (1958) Moring and Lantz (1975) Knight (1980) Shapova1ov and Taft (1954) Gard and F1ittner (1974) Hunt (1974) McFadden et al. (1967) Alexander (1979) Saunders and Smith (1962) Murray (1968) Egg1ishaw and Shackley (1977) 4 ------- SECTION 4 FACTORS AFFECTING NATURAL VARIABILITY One approach to listing the important factors or variables in the stream environment that can affect abundance of sa1monid populations is the following: A. Physical factors 1. 2. Streamflow Habitat quality 'B. Biological factors 1. 2. 3. Food abundance Predation Movement and migration In most instances these variables may interact to influence a popula- tion, and the classification is inevitably artificial. For example, habitat preferences are often related to food availability. Under natural conditions, it is often difficult to measure the effect of one factor independently. However, the variables will be considered separately in this discussion, with an attempt to show how interactions may be involved. PHYSICAL FACTORS Streamflow , ' One of the earliest studies that attempted to relate streamflow to sa1monid abundance was conducted by McKernan et al. (1950). They found that low summer flows correlated with subsequent low returns of adult coho salmon in the Si1etz River, Oregon from 1924 to 1945. No relation was apparent in the Coquille River from 1923 to 1948. Scarnecchia (1978) found a significant correlation (r = 0.68) between total streamflow in the 17-month period of stream residence of juvenile coho and the commercial troll catch of adult salmon 2 years later. These data came from five Oregon rivers from 1942 to 1962. In addition, there was a significant correlation (r = 0.56) between total annual flow and catch 2 years later. Smoker (1955) obtained an even higher correlation (r = 0.91) in the same analysis (total annual flow vs. catch of adult coho 2 years later) for Puget Sound streams from 1935 to 1954. In Cowichan Bay, B. C., a lower availability of coho to the sport fishery was noted for year classes that experienced low summer streamf10ws in their juvenile stages (Neave 1949). In Nile Creek, B. ~., from 1946 ,through 1949 5 ------- the output of coho smolts varied directly with the minimum mon/thly rai.nfall during the previous summer (Wickett 1951). These studies sho~that stream- flow ~uring some part of the. freshwater phase of coho life hi~tory can influ- . ence J.ts level of abundance J.n the catch. ! '..;" , . \~ ~ :t..\ . We carried' out a s~milar analysis for juvenile coho salmon in two of the streams that were part of the Alsea Watershed Study in Oregon.. Mean monthly and seasonal discharge were correlated with mean June-April biomass and also with the smolt count in the same period, from June 1960 through May 1968. In . both streams the few significant correlations were mostly in the spring (Table 2) . TABLE 2. CORRELATIONS BETWEEN MEAN MONTHLY DISCHARGE AND ANNUAL SMOLT COUNT AND MEAN. JUNE-APRIL BIOMASS FOR COHO SALMON, ALSEA WATERSHED STUDY, JUNE 1960-MAY 1969. . Deer Cr. . Flynn Cr. Period Smo1t Biomass Smo1t Biomass June -0.'080 -0.080 -0.307 -0.321 July 0.095 -0.442 -0.004 -0.126 August 0.050 -0.199 -0.155. -0.267 September 0.098 -0.045 -0.206 -0.221 October 0.431 0.661 -0.122 0.145 November -0.076 -0.461 0.132 0.032 December 0.153 0.003 0.291 -0.373 January -0.398 -0.218 0.531 -0.654 February -0.099 . -0.006 0.042 0.088 March -0.687* -0.426 0.099 -0.055 April . 0.630 0.076 0.931** 0.936** May 0.569 -0 . 162 0.714 * 0.694.* June-May -0 . 350 -0 . 507 0.150 -0.364 Nov-Apr -0.482 -0.544 -0.209 -0 . 446 Jan-Apr -0.691* -0.448 -0.097 -0.279 Mar-Apr -0.344 -0.352 0.518 0.385 June-Sept -0.021 -0.125 -0.262 -0.230 * P <0.05 ** P <0.01 6 ------- Knight (1980) used a longer series of data on smo1t abundance alone' and found significant negative corre~ations between. mean January discharge and total November-May smo1t count for Deer Creek and Flynn Creek (r = -0.64 and -0.65 . respectively) for the 1959-1960 through 1972-1973 seas~:ms" We performed a similar analysis for cutthroat trout from September biomass data and mean monthly discharge data (October 1961-September 1972) for all three streams in the A1sea Watershed Study. 'Generally, 'corre1ations were' negative, but nonsignificant, in the winter months 'in all three streams (Table 3). TABLE 3. CORRELATIONS ~1WEEN MEAN MONTIfLY DISCHARGE AND MEAl'l SEPTEMBER BIOMASS IN G/ FOR CUTTHROAT TROUT, ALSEA WATERSHED STUDY, OCTOBER 1961-SEPTEMBER 1972. . . Deer Cr. Flynn Cr. Needle Br. Month(s) All ye ars All years All years Pre-logging a Post-logging b October -0.119 -0.014 -0.221 0.829 0.586 November -0.133 0.085 -0.060 -0.393 0.150 December 0.222 -0.174 -0.384 -0.382 0.292 January -0.061 -0.139 -0.538 -0 . 804 -0.309 February -0.136 -0.311 -0.269 0.645 0.686 March 0.209 0.304 -0.302 0.646 -0.144 April .0.571 0.399 -0.071 0.335 -0.256 May 0.415 0.423 0.589 0.558 0.277 . June -0.212 -0.210 -0.095 0.993** 0.720 July -0.092 0.094 -0.158' 0.335 0.465 August -0.367 0.013 0.063 0.002 0 . 650 September O. 097 0.117> -0.262 0.264 0.718 Oct-Sept 0.214 0.006 -0.565 -0 . 906 0.336 Nov-Apr 0.195 -0.027 -0.574 -0.867 0.109 Jan-Apr 0.186 0.058' -0.711* -0.654 -0.085 Mar-Apr 0.419 0.417 -0.249 0.669 -0 . 202 June-Sept -00173 -0.100 -0.135 0.551 0.755 a 1962-1965 b 1967-1972 . * P <0.05 ** P <0.01 7 ------- In all three of 'these analyses the lack of consistency in the correla- tions was notable. Although one can attach plausible explanations to the statistically significant correlations, there were hardly more of them than . might be expected due to chance in a series of that many analyses. Our conclusion is that there is no solid basis for a relationship between stream- flow and abundance of coho salmon and cutthroat trout in these streams, a surprising result in the face of so much other evidence for such a relation. The small size of the streams involved and the resultant low numbers of juvenile fish may have reduced the power of the analysis, however. In several western Oregon streams, Pearson et ale (1970) did a prelim- inary short-term study of the effects of streamflow on juvenile coho salmon during the summer low-flow season: From 1962 to 1965, thZy found a signifi- cant positive relationship between coho density (number/m) and minimum streamflow in McKay Creek, a tributary to the Tualatin River. They also found a significant positive relationship between mean water velocity in pools and coho density for 50 pools in fi ve ,streams of the Nehalem River system. Preliminary data also indicated that streams with higher flows sup- ported coho of larger sizes. In streams in Maine, Havey and Davis (1970) found through multiple regression analysis of several environmental variables that rainfall in July and August, presumed to be an index of streamflow during the dry season, was the single most important factor influencing survival of Atlantic salmon from age 0+ to age 1+. Their multiple regression analysis was weakened, however, by a small sample size. Wickett (1958) reviewed the effects of low water levels on adult migra- tion and egg deposition by pink and chum salmon in British Columbia streams. Low flows result in excessive spawning density, leading to. superimposition of redds and crowding of eggs. Adult migration is inhibited by low stream- flow; other consequences include failure of egg deposition and increased predation on spawning fish crowded in shallow water. The effects of streamflow on survival of pink and chum salmon in spawning beds were studied by McNeil (1966;1968) in streams in southeastern Alaska. Below normal streamflow, both in summer and winter, caused significant mortality of eggs and alevins in the gravel. In summer,. low streamflow acted by causing low levels of dissolved oxygen inintragravel water. In winter, low streamflow led to freezing of eggs and alevins, especially in streams subject to greatly fluctuating flows. High streamflow during winter caused mortality by displacement of eggs and alevins from spawning gravel. Studies have. also been undertaken on the influence of streamflow on resident populations of salmonids. In Big Roche-a-Cri Creek, Wisconsin, brook trout biomass fluctuated greatly with streamflow. White (1975) found that from 1958 through 1966, biomass was significantly correlated with mean January-February discharge (r = 0.867). - Using data from Sagehen Creek, California, kindly provided by Dr. Richard Gard, we correlated mean monthly and seasonal discharge with mean annual bio- mass of brook, brown, and rainbow trout (Table 4).: Brown trout biomass was 8 ------- best correlated with December flows ~dbrook trout biomass showed the best correlation with February discharge. Neither of these was statistically significant, however. Rainbow trout biomass showed significant negative . correlations with discharge in January, 'April,. and June. Again, the total number of significant correlations among the 51 comparisons is very close to the number that would be expected by chance. However, the predominance of negative correlations for brook and rainbow trout and positive correlations for brown trout is in itself a significant result that deserves further analysis.' . TABLE 4. CORRELATIONS BETWEEN MEAN M)NTHLY DISCHARGE AND MEAN ANNUAL BIOMASS OF BROWN, BROOK-, AND RAINBOW TROUT. SAGE HEN CREEK, CALIFORNIA, 1954-1961. DATA FROM DR. RICHARD GARD (PERSONAL COMMUNICATION). . Month(s) Brown Brook Rainbow January 0.049 -0.142 -0.806* February -0.174 -0.632 -0.495 March -0.168 0.072 -0.449 April 0.067 0.023 -0.746* May 0.183 -0.488 -0.595 June 0.123 -0.547 -0. 727* July 0.109 -0.494 -0. 649 Augus t 0.259 -0.408 -0. 592 September 0.354 -0.275 -0.468 October -0.416 -0.199 0.373 November -0.050 -0.209 -0.127 , , December 0.514 0.146 0.147 Jan-Dec 0.208 -0.461 -0.294 Jan-June 0.137 -0.448 -0.305 March-June 0.151 -0.449 -0.339 March-April 0.164 -0.510 -0.454 July-Dee 0.426 -0.184 -0.080 * P < 0.05 Floods can have a very severe impact on salmonid and other fish popula- tions. Wickett (1958) reported that floods are a ',major cause of mortality in 9 ------- pink and chum salmon streams in British Columbia and have often reduced the size of succeeding runs. The principal cause of mortality is scouring of eggs and alevins from the gravel. In Nile Creek, B.C., chum salmon survival , was considerably reduced in years of severe floods. In 1945-46, there were no floods and the fry had a 3% survival rate. There were several severe floods in 1946-47 and 1947-48, with survival rates dropping to 0.44% and 0.38%, respectively. ' There was high water but no severe flooding in 1948-49 and fry survival increased to 6.0% (Neave and Wickett 1953)~ In the Horokiwi stream, New Zealand, severe flooding occurred between May and October 1941. Based on studies over the previous year, Allen (1951) estimated the effects of these floods on the streambed, the benthic fauna, and the brown trout population. The bottom fauna was reduced to 40-50% of ' levels of the previous year. The estimated number of most age classes present in October 1941 was only 25-50% the number present of the same age in October 1940. Destruction of eggs by flooding represented 80-90%, compared to a negligible loss the year before. The reduction in bottom fauna resulted 'in a higher percentage of this food resource being required by the remaining reduced trout population just for maintenance. This reduction, left a lower proportion of the food for growth. Thus the floods caused a reduction in the bottom fauna that limited the trout stock to a lower biomass and production. This effect occurred independently of the direct reduction in numbers of trout caused by the floods. Although the study terminated at that time, the limitation was presumed to be only temporary, with both benthos and trout populations returning to original levels in periods of normal rainfall.' In Valley Creek, Minnesota, four severe floods were recorded in 1965 and 1966. Two year classes of brook trout were nearly eliminated from the population. The older age groups were reduced as a result of changes in habitat caused by flooding (Elwood and Waters 1969). A later study showed that the brook trout population made a substantial recovery in 4-5 years. S~anding.crop increased from 418.fish/ha in 1966 t02l?,882 fish/ha in 1969. B~omass ~ncreasedfrom 2.5 g/m ~n 1966 to 14.8 g/m ~n 1970 (Hans02 and Waters 1974),.still somewhat lower than the average of about 25 g/m from 1961 to 1965. In Sagehen Creek, California, survival of spring-spawned rainbow trout fry increased in years following winter floods (Seegrist and Gard 1972). This increased survival of age-O rainbow trout was presumed to be caused by reduced competition from young brook trout, a consequence of brook trout eggs being destroyed by flooding. When floods occurred in May, rainbow trout eggs were destroyed and survival of young brook trout was improved. Adult trout were less affected by flooding than were the young. These studies illustrate the impacts that floods can have on sa1monid populations. Generally, they affect the eggs and young, older fish being somewhat more resistant. The magnitude of the impact, however, can vary according to the severity of the storm, the particular species, the time of year, and the physical characteristics of the stream. ' 10. ------- Habitat Quality Salmonids are not uniformly distributed within a stream reach. If , habitat preference or use can be defined for a species, the potential exists for prediction of spatial variation in abundance based on measurement o'f habitat qualityo It may also be possibie to relate temporal variability in . abundance to seasonal changes in habitat caused by changing streamflow or other variables. A number of studies have attempted qualitative or quantita- tive des~ription of habitat use by stream salmonids. Juvenile coho salmon in their first summer prefer a pool environment. Emerging fry in Waddell Creek,California, initially utilized shallow gravel areas, particularly those near the stream margin (Shapovalov and Taft 1954). 'The youngest fry tended to school, but as the fish grew larger, these schools broke up and individuals took up territories, which they defended. The larger fry moved into deeper water and by July and August were mainly found in the ,deeper pools. Chapman (1962) further defined this territorial and aggressive behavior and related it to habitat utilization. Ruggles (1966) found that over twice as many fry remained in a pool-like environment than a riffle-like condition in stream channels in British Columbia. In Oregon streams, Nickelson and ~eisenbichler (1977) described characteristics of prime habitat for juve- nile. coho salmon as having water depth of at least 30 cm, velocity of less than 30 cm/sec, a cobble substrate, and cover consisting of undercut banks and submerged roots. In the South Fork system of the Salmon River, Idaho, juvenile summer chinook salmon rear primarily in the main stem. Platts and Partridge (1978) recently reported significant use of many tributaries as well. In these tributaries the juvenile salmon preferred high quality pools iri the larger streams that had lower channel gradients and grassy streambanks. Yet 59% of all the salmon were found in stream reaches where less than 20% of the channel consisted of pools. This distribution was presumably the result of . the fact that most of the juvenile chinook in the tributaries occupied stream reaches within 400 m of the main river, where there was naturally a low pool/ riffle ratio. . , ' Ina north central Colorado stream, Stewart (1970) sampled 41 sections four times from June through September. He found mean depth and underwater, overhanging rock cover to be the most important variables determining the density of brook and rainbow trout larger than 18 em. Undercut banks and areas of deep turbulent water seemed to be related to brook trout density, but not that of rainbow trout. He also presents,useful data on spatial and temporal variation in biomass of the 41 sections, along with the physical data. Biomass of brook trout >18 em varied from 0 to 63.9 g/m, rainbow trout o to 8103 g/m, and combined trout 0 to 117.5 g/meter of stream (data on area not presented). In Little PrickJy Pear Creek, Montana, Lewis (1969) conducted a similar study involving 19 sections. He found that cover was the most important factor determining the density of brown trout. Increased stream velocity was associated with increases in density of both brown and rainbow trout per unit area of pool surface and per unit area of cover. :The most stable trout 11 ------- populations occurred in deep. slow pools with extensive cover; brown trout showed greater stability than rainbow trout. Current velocity was the most important factor determining density of rainbow trout. Useful data on spatial , variation in density are presented. but there is no information on biomass. Use of habitat by steelhead trout was studied by Shapovalov and Taft (1954) in Waddell Creek. California. Young fry showed similar tendencies to coho fry. initially congregating in schools and later setting up territories. However. unlike coho. steelhead fry inhabited riffles in late summer rather than deep pools. Dolly Varden fry in Hood Bay Creek. Alaska. were found in quiet water near stream banks and in small pools. The fry were usually inactive and found in or on the substrate. in contrast to the more aggressive coho fry. often found in the same habitat. The coho were actively swimming and feeding from the water surface (Blackett 1968).' Species interaction can have a strong influence on habitat utilization. Apparent preferences shown in the presence of a competing species may change if that species is absent. or if another is present. so care must be used in interpreting results from field studies of species interaction. Careful field observation coupled with experimental analysis is needed to define these interrelations. . . Seasonal habitat preferences and behavior of juvenile coho salmon and steelhead trout were studied by Hartman (1965) in British Columbia. In spring and summer coho occupied pools and steelhead occupied riffles. Both were aggressive in defending their respective habitats. This behavior is similar to that observed in Waddell Creek, California. discussed earlier. In winter, however, both species inhabited pools. Low population numbers, low aggres~ . siveness. and different microhabitat preferences were thought to be responsi- ble for this coenstence. Glova (1978) examined sympatric and allopatric populations of coho salmon and cutthroat trout in six British Columbia streams. In each. three habitats were defined in terms of stream veloci ty--pools « 8 cm/ sec), glides (8-20 cm/sec), and riffles (> 20 cm/sec). . In sampling during 1973 in Bush and Holland creeks, where both species occurred, coho salmon dominated the salmonid biomass in pools. composing 53-91% of the combined biomass. compared to 9-47%' made up by trout. In riffles trout were dominant, making up 63-88% of the combined total biomass. Glides were areas of intermediate biomass for both species. although coho also tended to dominate here, with 52-81% of total biomass. Above barrier falls. where they were found alone. cutthroat trout utilized pools more so than riffles, possibly due to the absence of coho. His analysis of diets suggested that coho were more specialized feed~rs. relying mainly on drifting foods. whereas cutthroat were more generalized. utilizing both drift and benthos. Glova (1978) noted that cutthroat emerged much later in the ye~r than did coho salmon. into an environment that may already be saturated by coho fry.: As a result of aggressive interaction with young coho. the trout would be largely restricted. to riffle areas during summer and early fall. and this habitat type is usually less abundant than pools at this time. He concluded that production:of sympatric trout may be 12 ------- limited by interspecific interaction, although total fish production may be greater..in multi-species streams. Glova (1978) also found that pools were more extensively utilzed by the, total fish species complex than were riffles. There was a strong negative correlation (r = -0.92) between the biomass of all fish species combined (coho salmon, cutthroat trout, and Coastrange sculpin, Cottus aleuticus) and mean stream velocity in Holland Creek during September. Based on behavioral studies he postulated that large pools would be less densely populated by , salmonids than small ones, owing to competition near the heads of pools for incoming food and resultant low densities of fish in the downstream ends of the larger pools. In support of this hypothesis, he found a significant negative correlation (r = -0.40) between logarithms of salmonid biomass and pool surface area, based on,data from a total of 37 pools in three streams. In British Columbia, Bustard and Narver (1975b) found in experiments that overwintering coho s.almon and cutthroat trout strongly preferred side- pools with overhanging bank cover to those without such cover. Given a choice between clean rubble substrate and silted rubble, they preferred the side- pools with clean rubble. In a natural stream studied'during winter, age 1+ coho and steelhead were found mainly at greater depths and in deeper water' than age 0 fish of either spec~es. As stream temperature dropped below 90 C, coho and age 1+ steelhead occupied progressively de'eper water 'and both' species moved closer to cover (gustard and Narver 1975a). Logs and upturned roots were the most commonly used cover. Steelhead were more closely asso- ciated with the substrate than were coho. Habitat utilization by sympatric populations of coho and chinook salmon fry was studied by Lister and Genoe (1970) in the Big Qualicum River, British Columbia~ At emergence, fry of both species were found along stream margins in association with streambank cover. As the young fish grew they moved into' areas of, faster velocity. Spatial segregation soon occurred between the two species because chinook fry emerged about one month earlier than coho fry and grew at a faster rate. As a result chinook prefe.rred higher current veloc- ities than did coho fry at a given date. .' Somewhat different results, invol v- ing more overlap of distribution and more interspecific interaction, were noted in an Oregon river where the two species emerged more nearly at the same time (Stein et al. 1972). . Diel variability in habitat preferences of juvenile chinook salmon and steelhead trout in Idaho streams was shown by Edmundson et al. (1968). Both species tended to move inshore at night to areas of quieter and shallower water than those occupied during the day. Steelhead used areas of faster velocity during the day than did chinook. Everest and Chapman (1972) found that most age 0 steelhead trout and chinook salmon in two Idaho streams lived in water velocities of less than 0.15 m/sec during summer. However, chinook occupied areas of finer substrate and deeper water than did steelhead. There is little interaction for living space between the two species because they spawn and emerge at different times; steelhead spawn in spring and chinook spawn in early fall. The larger juveniles tend to occupy deeper water, and the size differences resulting from these different spawning periods thus reduce compe- tition for food and space between the two species (Chapman and Bjornn 1969). 13 ------- Platts (1974) conducted an extensive study of fish habitats in 291 , sites in 38 streams within the upper South Fork of the Salmon River system, Idaho. Geomorphic characteristics were an important determinant of popula- , tion abundance. He found the highest fish population densities in channels having 30-50% pools. Total density of the fish populations was positively correlated with width and depth of the sampled streams. Rainbow trout and chinook salmon dominated the populations. Rainbow trout were predominantly found in riffles that were combined with shallow pools. Juvenile chinook were found most abundant in high quality pools. ' In the Miramichi River, New Brunswick" Keenleyside (1962) studied habitats and feeding behavior of Atlantic salmon and brook trout. Salmon , fry were most abtmdant in the upper reaches, where rapids and riffles were conunon. The fry were most abundant in fast water over substrate composed of small gravel and stones. Salmon parr (1-4 years old) were also more abundant in the upper sections of the river than the lower, but were found in de~per water and over larger substrate. Brook trout were found only in the upstream areas. Fry were most common in shallow slow-moving water , along the margin. Older fish were found in deeper water that was often swift or turbulent. Keenleyside (1962) noticed feeding segregation between the species. Salmon fry and parr fed on benthic fauna and surface'organ- isms, whereas trout fed almost exclusively on surface foods, possibly because they held positions further above the substrate than the salmon. In the Indalsalven River, Sweden, brown trout and Atlantic salmon were found together (Lindroth 1955). The young trout (age 0+) occupied shallow water near the stream margin. The trout were territorial and aggressive, actively chasing salmon fry away from these areas. In Scottish streams, Mills (1969) fotmd varying degrees of dominance between Atlantic salmon and brown trout. In some streams he found salmon fry and parr and juvenile trout living together in the same pools and riffles. All possible combinations were noted, from predominance of trout in some streams through to predominance of salmon in others. ' , Additional evidence that habitat quality is an important determinant of salmonid biomass comes from efforts to improve the quality of existing stream habitat. Although much of this work has gone unevaluated, a number of careful studies have shown population response to habitat development. Among the best documented is the work of Hunt (1971) at Lawrence Creek, Wisconsin. Habitat development in one 0.7 km section of the stream in 1964 increased permanent bank cover by 416% and pool area by 289%. As a result, total brook trout biomass increased from a mean of 59 kg in 1961-63 to 110 kg in 1965-67. In a follow-up study, Hunt (1976) fotmd the mean tota~ biomass in 1968-70 to have increased even further, to 165 kg (21.9 glm ). One of the earliest studies on habitat development in the West was conducted by Tarzwell. (1938) in two Arizona streams. In Horton Creek, small log dams, deflectors, and artificial bank cover were added to one section. A section of nearby Upper Tonto Creek was left unimproved as a control. From 1932 to 1937, 25,150 brook, brown, and rainbow trout were stocked in Horton Creek and 46,190 trout were stoc~ed in Upper Tonto Creek. 14 ------- A complete creel 'census was conducted in both streams. In 1936 and 1937, following improvement in Horton Creek, that stream yielded more trout to the angler ,and a greater weight of trout per hectare than did Upper Tonto Creek" ,in spite of the much heavier stocking of the latter stream. The effects of cover manipulation on trout abundance were studied by Boussu (1954) in Trout Creek, Montana. Four inventories were carried out before alteration of habitat (June, December, March, and June); three inven- tories were made after the alterations (September, December, and March). Rainbow and brook trout comprised about 98% of the sa2monids, the remainder being a few brown trout. 'Brush cover totaling 14.42m was added to four sections of the stream having a total area of 263 m. Following the cover addition, t2tal trout bi2mass in those sections increased from 1.13 kg to 4.04 kg (4.3 g/m to 15.4 g/m). Trout biomas2 in three un2ltered control 2 ' sections increased only 22% (from 8.5 g/m to 10.4g/m). When 11.9 m 2 of natu:al brush cover were removed from two sectionswit2 an area of 208 m , trout bl.Omass decreased from 3.83 kg to 2.28 kg (35.5 g/m to 21.1 g/m). 2 At the same2time trout biomass in a controL section increased26% (38.1 g/m to 40.5 g/m ). The third treatment~nvo1ved removal of 1.4 m ' of undercut. bank from two sectionstota2ing 80 m . 2In this case biomass decreased from' 0.68 kg to 0.45 kg (8.5 g/m 2to 5.6 g/m )2 while biomass in a control area increased 20%, from 14.4 g/m to 17.3 g/m. In each of'the three treatments the response by legal-sized fish (>18 cm) to change in cover was greater than that of smaller fish. Another result of his work not explicitly presented was the finding of a very significant spatial variation in trout biomass. In t2e 13 sections used for the study t2e pre-alteration biomass averaged 16.4 g/m , but ranged from 0.11 to 46.7 g/m. Because the data are reported as averages for four sampling dates, actual variability was undoubtedly greater. It should also be noted that these data resulted from a single pass with an electroshocker through each section blocked with stop nets, rather than from a formal population estimate. Thirteen dams , 12 deflectors, and several covers were constructed in a 4ll-meter section of Hayes Brook, Prince Edward Island, in 1959. In the, . following year, the number of age ° brook trout increased to 526 ,compared, . with a 13-year pre-treatment mean of 482 (Saunders and Smith 1962). Numbers of older trout increased from a mean of 348 (1947-1959) to 611 in 1960. . Many of these studies have shown great variability in habitat preferences between species, at different times of the year, for different ages of fish, and in association with other species present. Knowledge of these preferences is an important concern in the design of a sampling program. ' BIOLOGICAL FACTORS Food Abundance There has been an enormous amount of work done on food habits and feeding' behavior of stream salmonids. However, very few of these studies bear directly on the matter at issue here: can differences in abundance or availability of food account for spatial or temporal variation in salmonid biomass in streams? 15 ------- The question is complicated by difficulty in defining an appropriate measure of food availability--benthos, drift, or some combination. Very few studies have focused on these important issues. A reorientation of feeding studies . is required before a definitive answer to the question of food limitation is possible. Our review will concentrat~ on the few studies relating food abun- dance to variation in salmonid abundance. ' A starting point is to examine the significance of invertebrate drift. Drifting invertebrates represent a potential food source of considerable magni- tude, but of variable availability. Of particular importance is a strong diel ' periodicity, most drift occurring during darkness. Other factors that may affect the rate of drift include water temperature, current velocity, stage of the life cycle, and population density (Waters 1969). Some studies (Mason and Chapman 1965; Elliot 1973; Gibson and Galbraith 1975) have shoWn greater fish standing crops in stream sections with greater incoming drift. Yet other studies have shown a significant part of the diet to be made up of non-drift benthic forms. For example, Warren et ale (1964) reported the greatest food consumption in stream sections with the least drift, possibly because of a much greater abundance of benthic fauna in these sections. Other work has shown little correlation between drift and diet. One such study was conducted by Mundie (1969) on coho salmon fry in ,British Columbia. In seeking an explan- ation for the lack of correlation he postulated diel and spatial variation in drift composition, and variation in fry behavior. It is clear that there is considerable variation in the degree to which drift is utilized as food by , stream dwelling salmonids. " There is evidence that food can be a limiting factor for some popula- tions of stream salmonids. One of the strongest cases was brought forward by Mason (1976). He found that food limited the stream production of juvenile coho salmon during the summer in Sandy Creek, B.C. Through supplemental feeding, the summer biomass was increased 6-7 fold compared with previous, levels. However, there was no significant increase in the number of smolts the following spring. The estimate of smolt yield under natural conditions, was 212 fish, and the February population estimate was 257+71 fish surviving from supplemental feeding the previous summer. Thus in thIs stream the winter carrying capacity appeared to be the ultimate limit to smolt production. , In the Horokiwi stream, New Zealand, Allen (1951) found evidence suggest- ing that the food supply of brown trout, primarily the benthic fauna, could play an important role in regulating the trout population. He found that an increase in trout abundance increased pressure on the food supply, decreasing the density of that supply. This resulted in a reduction in surplus food (the amount that could be used for growth and production). Consequently, there was a decrease ,in mean individual growth rate. This resulted in a feedback system that would tend to keep the population biomass relatively constant by changing growth rate in response to changes in population size. ' In a later review, Allen (1969) discussed the role of the benthic fauna in regulating production of stream salmonids as a group. He suggests that fish production can be limited by the density of the bottom fauna, which in turn may be controlled by consumption by fish. This interaction provides a mechanism for stabilizing the salmonid production rate. 16 ------- Ellis and Gowing (1957) examined bottom fauna and brown trout popula- tions above and below a domestic sewage outfall into Houghton Creek, Michigan. Although the biomasses of trout were similar above and below, there were' . significant increases in the benthic fauna and condition of trout below the point of sewage input., They also noted that trout below the outfall relied less heavily on terrestrial foods, and concluded that trout growth was strongly influenced by the quantity and kinds of food consumed. Symons (1971) experimented with effects of fluctuating food quantities on behavior and abundance of Atlantic salmon parr in a stream tank. He found that such fluctuations had little effect on'the abundance of socially dominant parr. Socially subdominant fish, however, seemed more abundant where food was plentiful than where it was scarce. Thus total fish' abundance was 'higher in channels where food was more abundant. Mason and Chapman (1965) studied behavior and abundance of juvenile coho salmon in two experimental stream channels. They found that one channel received about a third more. volume in potential food organisms, 'and'this was associated with about a two- thirds increase in total fish weight in that channel. However, there was no replication, and other causes may also have been involved. . Variation in food abundance was associated with spatial variation in abundance of cutthroat trout populations in the Oregon Cascades. One pair of open and shaded ,stream reaches was studied intensively for 4 years. Primary production and insect emergence were significantly 'greater in the open area compared to the forested section (Triska et ale 1980). Production, growth rate, and biomass of cutthroat trout were about twice as great in the' open area (Hall et al. 1978). Murphy (1979) expanded the study to include nine pairs of open and forested sites, the openings being the results of earlier clearcuts. He found the same general relations to hold, including increased abundance of primary producers, predatory insects, and cutthroat trout in the open areas. . Predation Although predation has been shown to cause some significant mortality in stream salmonids (Hunter 1959; Mills 1964; Tagmaz'yan 1971), there have been very few studies to support the position that variation in level of predation leads to ultimate variation in size of the salmonid population. One of the few studies to combine stream population studies with preda- tor manipulation was carried out over a number of years in New Brunswick. Elson (1962) reports investigations of predation on juvenile Atlantic salmon by mergansers and kingfishers from 1942 to 1953. , In a sample of 117 mergan- ser stomachs analyzed, an average of 42.1% of the number of items were salmon. These salmon comprised an average of 10.3% of the total fish numbers in the river, yielding a forage ratio of 4.1. Kingfishers also selectively fed on salmon and had a forage ratio of 3.1. Predator control was practiced from 1947 through 1950 and. the abundanc~ of mergansers and kingfishers was signifi- cantly reduced. Consumption of salmon by these two species of predators was estimated to have been reduced to about 10% of pre~control levels. Before control, smalt output ranged from approximately 1,000 to 5,000 each year. During predator control, output ranged from 14,000 :to 24,000 smolts. Elson 17 ------- (1962) concluded that predation by mergansers waS a limiting factor on Atlantic salmon smo1t production. Unforttmate1y, the study design was some- what flawed by differing levels of stocking in the pre-control and control 'years, and by lack of data on adult returns. One of the most detailed long-ferm studies on trout populations and predation has-been carried out in Michigan on the North Branch of ' the Au Sable River (Alexander 1979). Estimates of population size of brook and brown trout were made in spring and fall each year from 1957 to 1967. Catch by anglers was determined from a statistically designed creel census in two sections of the river, one in which normal angling regulations prevailed and another in w~ich angling was significantly restricted. Predators were collected for stomach analysis from 1960 to 197~. From these analyses Alexander concluded that the annual rate of mortality of both brook and brown trout was very high (average rates calculated by cnapman-Robson method for age groups O-IV from his data in Tables 2-5: brook - 0.84 and 0.82; bro\~ - 0.64 and 0.74, normal and special regulations respectively). Consumption by known predators (prin- cipally the American merganser, great blue heron, be 1 ted kingfisher, mink, otter, and large brown trout) accotmted for a large fraction of this mortality; their consumption was estimated between 43 and 46% of annual production. Anglers'took another 37 and 8% in the normal and restricted water respectively. Notwithstanding the sizeable mortality caused by predators, Alexander is of the opinion that reduction of their abundance, short of complete removal of all predators, would not, have a significant impact on salmonid abundance, owing to , a compensatory kill rate that would be demonstrated by the remaining predators. The fact that total annual mortality rates are similar for each age group in the two sections, in the face' of much less angling "predation" in the special water, supports this view. More effort must be put into well-designed stream studies such as this one before a definitive conclusion on the significance of predation to population abundance of stream dwelling salmonids can be provided. Movement and Migration Nearly all salmonid species undergo varying degrees of movement in their lifetime. Some non-anadromous species undergo annual migrations within the same stream system for the purpose of spawning. Others remain in the same general area, undergoing local movements motivated by food, temperature, , streamflow, or other factors. Movement and migration can be considered a form of temporal variability. The timing and magnitude of these movements need to be understood in order to know what age and size range of fish to expect from sampling ata particular time of the year. A comprehensive review of migra- tory strategies of freshwater fishes and their significance to fish production is provided by Northcote (1978). Migrations of anadromous species are so conspicuous and generally well known that it seems unnecessary to include them in this review. One caution- ary example is perhaps in order, however. Conventional wisdom for many years held that juvenile faU chinook salmon migrated to saltwater shortly after emergence from the gravel, whereas juvenile spring chinook resided in fresh water for a full year before migrating to the ocean. More recent studies have indicated considerable variation from this pattern, both within and between stocks of fall and spring chinook (Reimers:and Loeffel 1967; Reimers 18 ------- 1973; Schlueter and Lichatowich 1977). These results indicate the impor- tance of careful studies of the migratory pattern in each stock of fish. Most studies of resident salmonids have found their movement to be quite restricted, with the exception of some activity associated with. spawning. In Kettle Creek, Pennsylvania, Watts et al. (1942) observed an upstream migration of brook trout into. colder tributaries in late May and early June. Spawning .took place in the fall, after which the trout moved downstream once again. Resident trout in this watershed, however, moved little between tributaries. There have been several studies of the movements of resident brown trout. Solomon and Templeton (1976) studied a population in a 7.5 km section of a chalk stream in. England, from which they recognized five life history stages with respect to movements and migration. The first was a downstream movement from hatching to nursery areas. Fish stayed in these areas for about 6.months. Then came a second movement further downstream to areas of adult growth, where the trout remained until they were about 15 months old. Following this was .a period of very limited adult movement . until maturation..Then came an upstream spawning migration followed by downstream movements after spawning. . In the Pine River, Michigan, Mense (1975} studied effects of varying brown trout densities on movement. Among fish >15 em, he found no change in movement patterns in a comparison of densities of 209 and 87 trout/hao He does not przsent data on biomass, but we have made a rough estimate of 3.8 and 3.3 gfm , based on his data for the two respective years. Both values for biomass are rather low, and the fact that the average size of fish was much larger in the year of lower density reduced the power of his test of the hypothesis. In Convict Creek, California, Needham and Cramer (1943) found extensive downstream movement of brown trout during spring. The peak coincided with rising, but not maximum, streamflows, although flow was not felt to be a causative factor. Most migrants were sexually immature. The downstream migration may have been initiated by lack of adequate food and shelter in the upper reaches of the stream. Little migration of rainbow or cutthroat trout was noted. Movement into and out of an intermittent tributary was shown to be "an important feature in the life history of rainbow trout in Sagehen Creek, California. From 39 to 47% of the spawning adults used this tributary from 1972 to 1975. Two possible reasons were given for this high use of an inter- mittent stream while permanently flowing tributaries were. used by only a small percentage of the spawning fish. Peak runoff from snow melt is much greater and occurs earlier in the year in the intermittent tributary. In addition, there is no competition from brook trout.because they cannot spawn there in the fallowing to insufficient flow (Erman and Hawthorne 1976). The rainbow trout fry from this tributary showed a .diel periodicity in down- stream movement that differed between a dry and wet year. In 1973, the dry year, fry moved downstream mainly during the day. In 1974, when the tribu- tary retained permanent flow, fry migrated d~wnstream mainly at night. In 19 ------- that year many fry remained in the tributary throughout the summer (Erman and Leidy,1975). In another population of rainbow trout, movement was not extensive. In Elder Creek, Oregon, Osborn (1967) made 755 observations of rainbow trout movement, based on recaptures of marked fish larger than 75 mID. Less than 4% of the fish had moved more than 91 m. ' Several studies have indicated that resident cutthroat trout undergo relatively limited movements. In Gorge Creek, Alberta, Miller (1957) found that of 58 tagged fish recaptured, 32 (55%) were recovered in the same pool in which they were tagged. He concluded that most cutthroat in this stream had a home territory less than 18 m long. In Lookout Creek, Oregon, restricted'home ranges were also found for cutthroat trout. Wyatt (1959) noticed no general downstream movement, but he did obs~rve two periods of limited ,upstream movement. From October through January some trout made $cattered vislts to tributaries. Then from the end of March to early June there was a spawning migration, with a peak in April. . OTHER FACTORS There are a number of other factors that may affect natural variation in abundance of salmonids. This section includes consideration of those factors that are worthy of mention but have not been studied in enough detail to warrant discussion in separate sections. In the Pigeon River, Michigan, Benson (1953b) studied the effects of ground water on brook and brown trout populations. Spawning of brook trout occurred only in sections, with considerable ground water seepage. Brown trout spawned in more widely scattered areas, but the greatest concentration of redds was located where ground. water was abundant. In turn, these areas of greater spawning produced higher population estimates. In a later study in'the same river system, Latta (1969) found that numbers of brook trout fry were directly correlated with ground water levels. He suggested that the relation would be' stronger in lower reaches of streams than in headwaters. Ice formation can have substantial effects on overwintering salmonid populations in high mountain streams or high latitudes. In Sagehen Creek, California, Needham and Jones (1959) noted that anchor ice, which forms under- water in riffle areas, is an important ecological factor in that it can raise the water level in pools and reduce streamflow over riffles. The breakup or melting of anchor ice can dislodge the benthic fauna, making more food avail- able to trout. In British Columbia, Bustard (1974) found collapsing snow and subsurface ice to be two major causes of winter mortality in salmonids. Beaver dams can significantly alter physical characteristics and carry- ing capacity of salmonid streams. .In Sagehen Creek, California, the balance of abundance of brook, brown, and rainbow trout was shifted by the presence or absence of dams (Garq 1961). 20 ------- Chemical properties of stream water may influence salmonid abundance'and growth rate. In New South Wales, Lake (1957) examined brown and rainbow trout populations in 1~0 streams. He found a strong correlation between water , chemistry and growth rate. Streams with the hardest water and highest pH had the most abundant bottom fauna and produced trout with the greatest length at a given age. Kennedy and Fitzmaurice (1971) examined over 40 streams arid rivers in Ireland. They found the largest and fastest growing ,brown trout in streams having a high calcium content. The smallest and slowest growing ones were in lime deficient waters draining acid rocks. These results are supported by Thomas (1964) from rivers in west Wales. He found that the growth rate of brown trout in waters having a pH of 7 or more with a high ion and calcium content was greater than that in more acid waters. Brown trout populations in six streams of varying hardness- in Pennsylvania were sampled by McFadden . (196la). There was no consistent difference in trout density between hard and soft water streams, yet brown trout growth rate was consistently greater in hard water streams. Fish of similar size had greater fecundity in hard water. . . , Stream gradient usually operates to limit distribution of salmonids, rather than abundance. However, in transitional areas" where two species are involved, consideration of gradient may help to explain variation in abundance. In the ClearWater Rive.r system" Idaho, Griffith (1972) f<:>und evidence suggesting that stream gradient may influence the relative abundance of brook and cutthroat trout in streams inhabited, by both species. In some parts of Idaho cutthroat live in slow water. « 6 em/see) when not associated with brook trout~ but they did not occupy this habitat in association with brook trout in Crystal Creek. Brook trout were found in the low gradient sections of Crystal Creek, whereas cutthroat were more abundant upstream iri areas of higher gradient. The same distribution of the two species was also found in a tributary of the St. Joe River. . 21 ------- SECTION 5 MINIMIZING THE EFFECTS OF VARIABILITY IN IMPACT STUDIES The temporal and spatial variability in populations of stream salmonids are clearly sufficient to mask very significant man-caused changes in these populations. This is especially true for damage done by non-point source pollutants. If we are to effectively monitor impacts of such perturbations, means must be found to minimize the effects of natural variability in detect- ing these effects.. It now seems. clear that the traditional watershed study design, with its long-term pre-treatment calibration and post-treatment evalu- ation, is not adequate for such analysis (Hall et al. 1978). After reviewing. existing approaches to the problem, we present several interrelated ideas that may improve sensitivity of future studies. HABITAT QUALITY RATING SYSTEMS . Models that quantitatively describe the qu~lity of salmonid habitat promise to significantly reduce the amouht of unexplained va,riabili ty in population abundance. The principal stimulus for the development of many of these models has been concern about loss of water from streams caused by irrigation or other appropriation. Hence the focus has been on determination of minimum streamflow requirements and on ch~ges in habitat quality and . quantity with changing streamflow. A good review of the historical basis for this work is provided in proceedings of the Symposium on Instream Flow Needs (Orsborn and Allman 1976).'. . One of the early attempts to develop such a model was made by Wesche. (1973), who combined hydrologic parameters, surface area, and available trout cover to define available habitat for brown trout. Continuation of this work extended the analysis to a cover rating system that provided a signifi- cant linear predictor of brown trout biomass in several stream systems (Wesche, 1976) . Another early study was initiated by the Oregon Department of Fish and Wildlife, mentioned earlier. They began with an attempt to relate habitat quantity and quality to streamflow by manipulating flow in a natural stream channel through a diversion (Keeley and Nickelson 1974; Nickelson 1975). Though initial work was marred by technical difficulties in establishing the diversion, recent results have been quite promising. Two types of models are presently being developed. One describes the relation between stream habitat and rearing potential of salmonids during the low flow period. Another is designed to predict the amount of habitat for an/ value of streamflow (Nickel- son and Reisenbichler 1977; Nickelson and Hafele 1~78). Pool volume alone 22 ------- explained 93.5% of the variation in summer standing crop of juvenile coho salmon in 12 sections of three coastal streams. For cutthroat trout a habitat' quality rating (HQR) is computed as 'a product of a cover value, velocity .preference factor, and wetted area. The cover value is a combination of depth, escape cover, overhanging cover, turbulence, and velocity shelter. Two alternative formulations of the HQR explained 91 and 87% of the variation in cutthroat trout standing crop in 31 sections. of six streams' (Nickelson and Hafele 1978). A related HQR for steelhead trout, involving cover, depth and velocity, . and wetted area, explained 79% of the variation in standing. crop of juveniles in 23 sections of four streams. Further work is underway to validate these models. . A related approach has been taken in a follow-up of work done in Wyoming streams by Wesche (1976). Binns and Eiserman (1979) developed a habitat. quality index for trout from analysis of 22 physical, chemical, and biological at.tributes in a sample of 36 streams. Using a multiple regression approach for'selection2of model attributes, they constructed an index (Model I) that produced an R value of 0.955 for the .initial 20 streams sampled. When 2 this model was used to predict trout standing crop at 16 new stations, R dropped to 0.594. A new model was developed for 'all 36 sites, based. on only. nine habitat attributes, all physical and chemical (late summer flow, annual flow variation, maximum stream temperature, and a food index and, a cover index that are combinations of nitrate nitrogen, cover, eroding stream banks, sub- state, water velocity, and stream width). This new model (Mo~l II) explained 97% of the variation in trout standing crop at the 36 sites (R = 0.~66). However,. this analysis highlights a frequent misinterpretation of R as a measure of reliability of the model (W.S. Overton, Department of Statistics, Oregon State Univ., personal communication). One vrlue of standi~g crop is . more than twice as large as the next largzst (63.4'V vs 28.4 g/m ). This one point tends to inflate the value of R by its large contribution to the sum of squares for standing crop. A more valid measure of the goodness of fit is the relative predict~on error. The authors noted ~hat no prediction was in error more than 5.5 g/m and that an error of 5.4 g/m at Sand Creek (the highest trout population) was within 9% of the measured value. However, the percent error at many stations with lower biomass was substantially higher than that, and averaged 32.4% for the 36 stations with Model I (range 0-179%) and 26.2% (range 0-157%) for Model II: Nonetheless, this approach is a very useful one that promises to increase the precision of impact evaluation. The most extensive development of indices to habitat quality has been undertaken by the Cooperative Instream Flow Service Group of the U.S. Fish and Wildlife Service (Bovee and Cochnauer 1977, Bovee 1978). Their general approach has been to couple information on the state of several hydraulic parameters of the stream environment with a "probability of use" for a com- bination of these parameters. A weighted usable area is then calculated for each level of discharge for the various life history stages of each species 1 It is noteworthy that this estimate appears to be one of the ever reported for sa1monid biomass in streams, especially in resulted from only a single pass through the study section. largest that it 23 ------- of interest. This effort, focused on the effects of,incremental losses of streamflow on reduction in quality and quantity of. fish habitat, has been substantially influenced by the thinking of physical scient:l.sts, primarily . hydrologists. The input from physical scientists has been a significant feature of the program and one that should be encouraged. Addition of the perspective of geomorphology (Platts 1974; Swanson and Lienkaemper1978) could significantly improve the generality of the approach. PROCESS STUDIES Unde~standing of the basic physical and biological processes that lead to biological production and eventually to fish production will provide a much sounder basis for assessment than has been available through the case history approach. One particularly relevant example is found in the analysis of temperature changes following logging in the Alsea Watershed Study (Brown 1967; Brown and Krygier 1970). By developing a model of the heating and cool- ing process in an undisturbed stream and quantifying each element in the energy budget, Brown was able to identify direct solar radiation as the pri- mary source of warming in streams. This procedure allowed a prediction to be made of the potential impact before timber was cut, and thereby provided a basis for planning necessary buffer strips to minimize adverse effects caused by warming of stream water. The process study provided an energy budget approach that is general enough to be applied in most watersheds. It is probably more feasible to carry out such studies of the physical processes in streams than those of the biological components. Additional work on physical process is now underway, for example, in suspended sediment and bedload transport (Beschta 1978; Beschta and Jackson 1979). Nonetheless, studies of biological process are essential to an understanding of variability in stream salmonid populations, and further emphasis must be placed there. Though far from complete, the work in Mack Creek carried out under the Coniferous .Forest Biome Study and mentioned earlier (Triska et al. 1980) provided some evidence of the validity of this approach. Knowledge of pri- mary production, insect abundance, and trout production provided evidence that the higher trout biomass in streams flowing through clearcut areas was a real phenomenon rather than simply the result of movement of trout in response to preference for open areas. It also provided some evidence of at least one pathway through which the increase in trout production might have been achieved. Much more work will be necessary in many more systems, however, before models of biological processes will achieve the same level of under- standing and predictability now enjoyed by models of physical processes in streams. '. . _._~- STREAM CLASSIFICATION It seems clear that some sort of classification of streams and their watersheds will be an essential element of future impact assessment (W.S. Overton, pers. corom.). Classification has had a long history, especially'in Europe, where it has been incorporated in management schemes (cf. Huet 1959). 24 ------- However, the perspective of those involved in classification has often seemed to focus on differences rather than similarities of stream ecosystems, thus leading to unmanageable complexity in the system of classification (cf. Pennak . 1971) . . One of the approaches most adaptable to the present problem is that of Platts (1974). His classification is based on stream order and a ,small 'number of geomorphic characteristics and provides a manageable and quantifiable system. Application to a stream ecosystem encompassing 220 km of the South Fork of the Salmon River in Idaho provided significant explanation of vari- ability in distribution and abundance of nine fish species. A recent synthesis by Warren -(1979) forwards a more inclusive classifi- cation scheme, based on a biogeoclimatic perspective. It takes the promising approach of classification based.on capacity or potential of a system rather than its present state. This potential would be indexed solely by geomorphic characteristics of the stream habitat and the watershed system within which it is imbedded. The scheme thus avoids much of the complexity inherent in meas- uring both taxonomic and quantitative variability in biological components within and between stream ecosystems. Further development of this concept should provide a much more solid basis for impact assessment in the future. IMPROVED STUDY DESIGN Another source of improvement in efficiency of detection of impacts appears available through modifications in the way in which observational data are gathered. Field observations will probably always be the major basis for impact assessment. As a consequence, 'much of the body of experi- ence and theory in the field of experimental design will not be directly applicable to such analysis. A sampling perspective is more appropriate, and Overton (1978) provides a useful discussion of three levels at which sampling questions can be addressed, along with general guidelines on study design. Eberhardt (1978) provides a valuable review of the problems of appraising variability in population studies, one that should be required reading for anyone beginning a study to assess impacts of non-point source pollution. A related article (Eberhardt 1976) provides further detail, particularly on his suggestions for handling the "single-site problem" that is often a character- istic of impact assessment. He proposes substitution of repeated observa- tions in time or space for true replication. The ratio of population density in the affected area to that in the "control" site(s) would be the measure of impact. He is cautious, indicating potential problems and suggesting the whole approach as a "pseudodesign." Nonetheless, these two papers are a very significant contribution to the topic under review here. A number of different approaches to field observation are possible, and appropriate combinations may lead to more fruitful results than will a single approach. These possible approaches have been cl~ssified in two ways by Hall et al. (1978). In a review of effects of watershed perturbations on streams, they grouped studies according to whether they bracketed (before-after) or 2S ------- followed (post-) treatment. The other level of classification was based on whether detailed studies were made on one or very few streams (intensive) compared to less detailed work on many streams, including a wide range of . habitat types (extensive}. This two-level classification results in four categories, which are evaluated for efficiency and sensitivity of impact detection. An expanded listing.of advantages and disadvantages of each type (Table 5) reveals that no one design is optimum. The extensive post- , treatment approach does have a number of advantages over the cl~ssical water- shed study (intensive before~after). The best approach appears to be a . combination of ,extensive post-treatment analysis with carefully designed process studies carried out at one or more locations. ' Pairing of treatment and control is proposed to improve sensit~vity of detection (Hall et al. 1978). This procedure places an upstream control very close to a treatment area on each stream. It proved to be a sensitive design to investigate changes in both predator populations and their habitat in small clearcutsin the western Cascades in Oregon (Murphy 1979; Murphy and Hall MS.)~ By inclusion of watersheds that had been harvested up to 35 years' earlier, it also provided some insight into the rate of change of physical and biological characteristics following treatment. This approach does have the limitation that it can detect only those effects that occur in the immedi- ate stream reach affected by the treatment. It is relatively insensitive to downstream effects or those that accumulate over the larger watershed. ' A modification that would provide some in~ight into effects on that scale would pair watersheds, treated and untreated. However, it would often be difficult to find untreated watersheds adjacent to treated areas, and such pairs would undoubtedly be more unlike than adjacent reaches of the same stream. Nonetheless Welch et al. (1977) used a variation on this approach to document effects of forestry and agriculture on streams in New Brunswick, examining a total of 34 watersheds, all smaller than about 1000 ha. ~ Erman et al. (1977) used an innovative' form of this approach in a study of effects of clearcutting on invertebrate populations in Northern California streams. They sampled a total of 62 streams, all in small watersheds «800 ha). There were two objecives: to test effects of various widths of buffer strips in preventing changes in invertebrate populations, and to examine localized effects of point disturbances such as road-related landslides. For the latter purpose their design was to sample upstream of the landslide as a control, at the disturbance point, and downstream where no visual evidence of the disturbance remained. To evaluate the role of bufferstrips, they used a design that employed two controls for each logged section, one upstream from the treatment and another in an adjacent untreated watershed. The hypothesis tested was that if effects occurred, the two control streams should be more similar than either control and the treated section. Various measures of similarity were compared and nonparam~tric ranking tests were used in the statistical anal- ysis. They found significant effects on community composition in unbuffered streams and found no significant differences between controls and streams with wide bufferstrips (Newbold et al. in press). 26 ------- TABLE 5. SUMMARY OF ADVANTAGES AND DISADVANTAGES OF THE FOUR MAJOR APPROACHES TO WATERSHED STREAM ANALYSIS. A. Advantages Intensive Before-After (10-15 years; 5-7 years before ,and after treatment). Disadvantages 1) Possible to assess year-to-year variation and place size of impact in context of that variation. 2) Can assess short-term rate of - recovery (ca. 5 years). 3) No assumptions required about initial conditions. 4) Possible to monitor whole water- shed impacts (provided substan- tial investment in facilities such, as flow and sediment sampling wiers, fish traps). , ' , 5) Long time frame provides format for extensive process s4udies. 1) No replication; results must be viewed as a case study. '2) Results not necessarily applicable' elsewhere (areas of different soils, geology, fish species, etc.) 3) Results vulnerable, to unusual . climatic events (e.g. high or low rainfall season(s) immediately following treatment). ' 4) Final results and management recom- mendations require exceptionally long time to formulate - up to 15 yrs after initial planning stage. 5) Difficult to maintain intensity of investigation and continuity of investigators over such a long period.. 6) Must rely on outside agencies or firms to complete treatments as scheduled - considerable coordina- tion required. . B. 'Advantages Extensive Before-After (2-4 years; 1 year before treatment, 1 year after). Disadvantages 1) Provides broader perspective across geographical area than (A). 2) Larger number of streams examined lessens danger of extreme case. 3) Increased generality of results allows some extrapolation to other areas.' 4) Relatively short time to achieve results (3-4 years from planning stage) . 1) Lack of long-term perspective-- little opportunity to observe year-to-year variation. 2) Able to assess only immediate results, which may not be repre- sentative of longer time sequence. 3) Treatment vulnerable to unusual weather (if all treatments in same year) . . 4) Must rely on outside agency (see (A) above). 27 ------- TABLE 5. (Continued. ) C. Intensive Post-Treatment following treatment). Advantages (One Watershed--Paired Sites) (4~5 years, Disadvantages 1) Shorter tlme for results than (A) . 2) Moderate ability to assess year-to~year variation. 3) Provides opportunity for moderate level of effort on process studies. 1) Provides no strict control-- requires assumption that upstream control was identical to treated area prior to. treatment. 2) "Control" most logically must be. located upstream of treatment. Strong downstream trend in any feature would confound analysis. 3) Provides no spatial perspective-- results of limited' application elsewhere. (or more); all observations in D. Extensive Post-Treatment, 10-30 Watersheds 1~2 years (vaFiable time' after treatment). Advantages Disadvantages 1) Wide spatial perspective allows extrapolation to, other areas. 2) Long temporal perspective is possible--can assess recovery for as many years as past treat- ments have occurred. 3) Provides ability to assess interaction of physical setting and treatment effects (e.g. effects of sediment input at different stream gradients). 4) Requires least time of all four designs to get results-- as little as 2 years. 5) Probably most economical of all four approaches per unit of information. 1} No data available on pre-treatment conditions--forces assumption that control and treatment were identi~ cal (on average). 2) Control predominately upstream. 3) Total cost concentrated in very short period--requires extensive planning. 4) Not as effective as (A) in assessing whole watershed effects. 5) Methods used in early treatments may. not be comparable to later ones. ---~~._- ~.- - 28 ------- Although much variability will undoubtedly remain in any study of natural populations in field situations, the ideas discussed above should help to resolve some of the uncertainty that has been present in past . analyses. A good deal of ingenuity and insight will be needed in making the right choices of habitat parameters and in devising methods of quanti- fying them. Choosing the appropriate variables for watershed classification will likewise be a formidable task. Hopefully, however, some judicious' com9ination of these approaches should make the task of assessing and con- trolling ,non-point source pollution a more effective and rational one. 29 ------- BIBLIOGRAPHY As part of a development of strategy for analysis of impacts of non-point source pollutants on streams, we have searched the literature on' population dynamics of stream salmonids. The following list is not exhaustive, but we have put emphasis on studies that ~eport quantitative estimates of population size, and those that deal with spatial and temporal variability in such data. In addition we have included studies that may be useful in interpreting this variability. Some emphasis is placed on work on the west coast of North America, but many valuable contributions from other areas have also been included. The more important contributions in the following list are annotated; other reports of interest are listed only by title. Aho, R.S. 1976. A population study of the cutthroat trout in and shaded section of stream. M.S. thesis. Oregon State Corvallis. 87 pp. an unshaded Univ. , Alexander, G.R. 1979. Predators of fish in coldwater streams. Pp. 153-170 in Henry Clepper Ced.), Predator-prey systems in fisheries management. Int. Symp. on Predator-Prey Systems in Fish Communities and their Role in Fisheries Management. Sport Fishing Institute, Washington, D.C. ' Allen, K.R. 1940. Studies on the biology of the early stages of the salmon, (Salmo salar). 1. Growth in the River Eden. J. Anim. Ecol. 9:1-23. ' An investigation of the juvenile Atlantic Salmon population in the River' Eden, England. Growth is considered in terms of length and weight. In addition, data are presented on seasonal variability of the major' benthic organisms, and a study is also made of those consumed by the salmon. Allen, K.R. 1951. The Horokiwi stream: a study of a trout population. New Zealand Mar. Dept. Fish. Bull. No. 10. 231 pp. A classic study of the brown trout and benthic fauna in a New Zealand stream. Spatial variability in mean estimated trout biomass in six sections of the stream is presented for 1940 and 1941~ Movements, mortality, production, growth, length-weight relationships, and angling effects are also analyzed. The bottom fauna is sampled in each section and total numbers and weights are estimated. The availability of these organisms in the-stream is compared to the consumption by the trout and is expressed as forage ratios. The'role of the food supply as a pos- sible limiting factor in trout production is discussed. The effects of floods on the invertebrate fauna arid trout population are also examined. 30 ------- Allen, K.R. 1969. Limitations on production.m salmonid populations in streams. pp. 3-18 in T.G. Northcote (ed.), Symposium on salmon and trout in st~eams'; H:"'R. MacMillan Lectures in Fisheries.' Uni v. British Columbia, Vancouver. . A review of factors that can limit salmonid production in streams. In. the egg stage in gravel, stream discharge and gravel permeability are critical. Predation, territoriality, and food availability are other stream factors of importance. The extent to which these factors can be limiting depends on the species, the amount of.time spent in fresh water, the stream's physical characteristics, and climatic condi tions. Andersen, B.C. 1978. Fish Barkley So~d streams, 118 pp. . populations of Carnation Creek and other 1975-1977. Fish~ Mar. Servo Data Rep. 89. Andersen, B.C., and D.W. Narver. 1975. Fish populations of Carnation Creek and other Barkley Sound streams 1974: data record and progress report. Fish. Res. Bd. Can. MS Rep. 1351. 73 pp. . ~trong, R.H. 1970. Age, food, and migration of Dolly Varden smolts in southeastern Alaska. . J. Fish. Res. Bd. Can. 27:991-1004. Armstrong, R.H. 1974. Migration .of anadromous Dolly Varden (Salvelinus malma) in southeastern Alaska.. J. Fish. Res. Bd. Can. 31:435-444. Au, D.W.K. 1972. Population dynamics of the coho salmon and. its response to logging in three coastal streams. Ph.D. thesis. Oregon State Univ., Corvallis. 245 pp. An encompassing study of coho salmon in the Alsea Watershed Study~ in which orie watershed was clearcut down to the streambank in 1966, another watershed was clearcut but buffer strips were left, .and a third was left unlogged as a control. Fish traps enabled total counts to be made of upstream migrating adults and downstream migrating juveniles. Data on mean monthly biomass of juveniles are presented for the 1963-1968 year classes in each stream. Spawning migrations are analyzed in relation to streamflow. Sex ratios, redd location, fecundity, redd survival, and fry emergence are also discussed. Fry dispersal, colonization, behavior, and migration are also investigated. Population estimates, survival, growth, net production, and smolt yields are determined. No significant long-term effects of logging were noted in these populations. See also Chapman (1961; 1965) and Moring and Lantz (1975). Becker, C.D. 1973. Food and growth parameters of juvenile chinook salmon, Oncorhynchus tshawytscha, in central Columbia River. Fish. .Bull. n : 387 -400. Benson, N.G. 1953a. Seasonal fluctuations in the feeding of brook trout in the Pigeon River, ~lichigan. Trans. Amer. Fish. Soc. 83:76-83. 31 ------- Benson, N.G. 1953b. The importance of ground water to, trout populations in the Pigeon River, Michigan. Trans. N. Amer. Wildl. Conf. l8:26~-28l. . Benson, N.G. 1960. Factors influencing production of immature cutthroat trout in Arnica Creek, Yellowstone Park. Trans. Amer. Fish. Soc. 89:168-175. Presents data on the number of cutthroat trout in age groups 0, I, and II from 1950 through 1958. Production of these year classes is also investigated. Factors that may be related to production, include migration down into Yellowstone Lake, streamflow, photoperiod, predation, number of spawners, and timing of the spawning run. ' Beschta, R.L. 1978. Long-term patterns of sediment production following road construction and logging in the Oregon Coast Range. Water Resource Res. 14:1011-1016. Beschta, R.L., and W.L; Jackson.. 1979. The intrusion of fine sediments into a stable gravel bed. J. Fish. Res. Bd. Can; 36:204-210. Binns, N.A., and F.M. Eiserman. 1979. Quantification of fluvial trout , habitat in Wyoming. Trans. Am. Fish. Soc. 108:215-228. Bjornn, T.C. ,1965. The production of juvenile stee1head trout in an Idaho stream. ' Proc. 45th Ann. Conf. West. Assoc. State Game Fish Commrs. 45:210-216. Bjornn, T.C. 1968. Survival and emergence of trout and salmon frY in various gravel-sand mixtures. pp. 80-88 in Proc. Forum on the relation between logging and salmon. Feb. 8-9, 1968. Alaska Distr. Amer. Inst. Fish. Res. BioI., Juneau. Bjornn, T.C. 1971. Trout and salmon movements in two Idaho streams as related to temperature, food, streamflow, cover and population density. Trans. Amer. Fish. Soc. 100:423-438. Bjornn, T.C. 1978. Survival, production, and yield of trout and chinook salmon in the Lemhi River, Idaho. Univ. Idaho ColI. For., Wild1., Range Sci. Bull. No. 27. 57 pp. A long-term study of chinook salmon, stee1head trout, and resident trout from 1962 through 1975. Stocking the river with different densities of stee1head fry and subsequent yields of subyear1ings are discussed. Survival, growth, mortality, and predation are examined. The effects of introducing anadromous fish on resident trout and interactions between stee1head trout and chinook salmon are also investigated. Blackett, R.F. 1968. Spawning behavior, fecundity, and early life history of anadrornous Dolly Varden Sa1velinus'ma1ma (Walbaum) in southeastern Alaska. Alaska Dept. Fish Game Res. Rep. No.6. 85 pp. 32 ------- Bohlin, T. 1977. Habitat selection and intercohort competition of juvenile sea-trout Salmo trutta. Oikos 29:112-117. A study of juvenile brown trout in a small Swedish stream from 1971 through 1975. Seven study sections were established to determine densities of ages 0+ and I+ trout in relation to water depth and substrate type. Differences in trout lengths in the various habitats are also investigated. Competition between year classes is shown in stream tank experiments. Additional.field studies show biennial fluctuations in densities of the two groups. The role of intercohort competition as a possible cause of these variations is discussed. Boussu, M.F. 1954. small stream. - . Relationship between trout populations and cover on a J. Wildl. Mgmt. 18:229-239. Bovee, K.D. 1978. Probability-of-use criteria for the family Salmonidae. Instream Flow Info. Pap. 4. FWSjOBS-78j07. 80 pp. . . Bovee, K.D., and T. Cochnauer. ~977. Development and evaluation of weight~d criteria; probabi1ity-of-use curves for instream flow assessments: fisheries. Instream Flow Info. Pap. 3. FWSjOBS- 77j63. 39 pp. Brown, G.W. 1967. Temperature prediction using energy budget techniques on small mountain streams. Ph.D. thesis. Oregon State Univ., Corvallis. 120 pp. Brown, G.W., and J.T. Krygier. 1970. temperature. Water Resource Res. Effect of clearcutting on stream 6: 1133-1139. Burnet, A.M.R. 1959. Some observations on natural fluctuations of trout populations numbers. New Zealand J. Sci. 2 :410-42l. Burns, J.W. 1970. Spawning bed sedimentation studies in northern California streams. Calif. Fish Game 56:253-270. 1 ) Burns, J.W. 1971. The carrying capacity for juvenile salmonids in some northern California streams. Calif. Fish Game 57:44-57. Presents 3 years of data on salmonid biomass (1967-196Q) from seven northern California streams. An attempt is made to define the natural carrying capacity of these streams. Biomass per unit of surface area seems to be the best method of expressing this capacity. The use of streambed sediments, total dissolved solids, alkalinity, and total phosphate in predicting carrying capacity is also discussed. Changes in annual carrying capacity up to 50% are attributed to natural variation. Bustard, D. 1974. Some possible effects of logging on wintering juvenile salmonids. In Stream ecology: a symposium, Parksvi11e, B.C. Assoc. British Columbia Prof. Foresters. 19 pp. . 33 ------- Bustard, D.R., and DoW. Narver. 1975a.. Aspects of the winter ecology of juvenile. coho salmon (Oncorhynchus kisutch) and steelhead trout (Salmo gairdneri). J. Fish. Res. Bd. Can. 32:667-680. Bustard, D.Ro, and D.W. Narver. 1975bo Preferences of juvenile coho . salmon (Oncorhynchus kisutch) and cutthroat trout (Salmo clarki) relative to simulated alteration of winter habitat. J. Fish. Res. Bd. Can. 32:681-687. Chapman, DoW. 1957. Studies on age, growth, and migration of steelhead trout, Salmo gairdneri gairdneri, in the Alsea River, Oregon. M.S. thesis. Oregon State Univ., Corvallis. 96 pp. Chapman, D.W. 1961. Factors determining production of coho salmon, Oncorhynchus kisutch, in three Oregon streams. Ph.D. thesis. Oregon State Univ., Corvallis. 214 pp. A comprehensive study of coho salmon production and ecology in the three streams of the AlseaWatershed Study from 1958 to 1961. Physical factors investigated include streamflow, rainfall, water temperature, suspended sediment loads) light incidence, water chemistry, and stream area measurements. Biomass, yield, and growth rates of juvenile coho salmon are examined in determining production estimates. Behavior patterns of fry are also studied, including dominance, hierarchy formation, hiding, territorial defense, and aggression. Food habits of the fry are also analyzed. In addition, the aquatic insect food habits are examined to determine th~ proportion of coho food that comes either directly or indirectly from allochthonous sources. Chapman, D.W,. 1962. Aggressive behavior in juvenile coho salmon as a cause of emigration. J. Fish. Res. Bd. Can. 19:1047-1080. Chapman, D. IV. 1965. Oregon streams. 0 Net production of juvenile coho salmon in three Trans. Amer. Fish. Soc. 94:40-52. ,i Presents data on mean monthly biomass, production, and instantaneous growth rate of the 1959-1962 year classes in all three streams of the Alsea Watershed Study. The smolt yield for these years is also presented. Relatively large freshets seem to increase downstream movements of juveniles. Downstream drift of fry and smoltmigrations are discussed as possible sources of bias in estimates of growth and production in the residual populations. Chapman, D.W. 1966. Food and space as regulators of salmonid populations . in streams. Amer. Nat. 100:345-357. An important synthesis emphasizing the importance of social behavior in population regulation of stream salmonids. No new data are included, but many earlier studies are reviewed. The author makes the case for a food-space interaction mediated through social behavior as the important mechanism controlling population size in streams. 34 ------- Chapman, D.W., and T.C. Bjornn. 1969. Distribution of salmonids in streams, with special reference to food and feeding. pp. 153-176 in T.G. Northcote (ed.), Symposium on salmon and trout in streams; H.R. MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. Chapman, WoM., and E. Quistorff. 1938. The food of certain fishes of north central Columbia River drainage, in particular, young chinook salmon and steelhead trout. Wash. Dept. Fish. BioI. Rep. No.. 37A. 14 pp. Clary, JoR. 1970. Predation on the brown trout (Salmo trutta) by the slimy sculpin (Cottus cognatus). Ph.D. thesis. Pennsylvania State Univ., University Park. 50 pp. Cordone, A.J., and. D.W. Kelley. 1961. .The influences of i~organic sediment on the aquatic life of streams. Calif. Fish Game 47:189-228. Creaser, C.W. 1930. Relative importance of hydrogen-ion concentration, . temperature~ dissolved oxygen, and carbon-dioxide tension, on habitat selection by brook-trOut~ Ecology' 11:246-262. Crisp, D.T., R.HoK~ Mann, and J.C. McCormack; 1974. The populations of fish at Cow Green, Upper Teesdale, before impoundment. J. Appl. Ecol. 11:969-996. . . . A study of fish populations in several tributaries and the main stem of the upper River Tees, England from August 1967 through May 1970. Species studied are brown trout, bullhead (Cottus), and minnow. Data are presented on physical and chemical characteristics of each stream study site. Biological parameters examined for each species include distribution, length, weight, density, biomass, growth, and production. These parameters are also considered by age group for trout and bullheads. Initial effects of dam construction on these parameters are also investigated. Crisp, D~T., R~H.K. Mann, and J. C. McCormack. 1975. The populations of fish in the River Tees system on. the Moor House National Nature Reserve, Westmorland. J. Fish BioI. 7:573-593. A study of brown trout and bullhead (Cottus) populations in tributaries and the main stem of the River Tees system, England. Mean density and biomass from 1968 through 1972 are presented for May, August, and October for five tributary sites. Growth rates, survival, production, and recruitment are also investigated. Crone, R.A. 1968. Behavior and survival of coho salmon, Oncorhynchus kisutch (Walba~), in Sashin Creek, southeastern Alaska. M.S. thesis. Oregon State Univ., Corvallis. 79 pp. 35 ------- A study of 'coho salmon from 1963 to 1966. Spawning adult investigations include number, timing, and age of returning migrants, distribution on the spawning grounds, effects on pink salmon~ redd life, fecundities, and egg retention. Juvenile investigations include population size, distribution in the stream, age composition, food habits, mortality, and timing and size of smolt migrations. Crone, R.A., and C.E. Bond. 1976. Life history of coho salmon, Oncorhynchus , kisutch, in Sashin Creek, southeastern Alaska. Fish. Bull. 74:897-923. Presents data on coho fry and smolt weir counts from 1956 through 1968. Number of adults, ti~ng of migration, age composition, fecundity, and spawner distribution and density are examined. Interspecific competition with pink salmon is also investigated. Freshwater survival, growth, mortality, and size and age of smolts are analyzed. In addition, the significance of fry migration and early salinity tolerance is discussed. ' Cummins, K.W. 1974. Structure and function of stream ecosystems. Bioscience 24:631-641. , , Davidson, F.A., and S.J. Hutchinson. 1942. Natural reproduction of pink salmon studied at Little Port Walter, Alaska. Ecology 23:234-235. Dodge, D.P., and H.R. MacCrimmon. 1971. Environmental influences on extended spawning of rainbow trout (Salmo gairdneri). Trans. ' Amer.Fish. Soc. 100:312-318. Drucker, B. 1972., Some life history characteristics of coho salmon of the Karluk River System, Kodiak Island, Alaska. Fish. Bull. 70: 79-94. Eberhardt, L.L. 1976. Quantitative ecology and impact assessment. Environ. Mgmt. 4:27-70. J. Eberhardt, L.L. 1978. Appraising variability in population studies. Wildl. Mgmt. 42:207-238. J. Edmundson, E., F.E. Everest, and D.W. Chapman. 1968. Permanence of station in juvenile chinook salmon and steelhead trout. J. Fish. Res. Bd. Can. 25:1453-1464. Egglishaw, H.J. Scotland. 1970. Production of salmon and trout in a stream in J. Fish BioI. 2:117-136. A study of Atlantic salmon and brown trout populations in Shelligan Burn, Scotland from 1966 through 1968. Population size, growth rates, biomass, production, and yield data are presented. Variability in production is ~hown in three stream sections for the 1963-1968 year classes. Contributions of each year class to the total production for both species are discussed. . 36 ------- Egglishaw, H.J.,' and P.E. Shackley. 1977.. Growth, ~urvival, and production of juvenile salmon and trout in a Scottish stream, 1966-1975. J. Fish BioI. 11:647~672. A long-term study (1966-1975) of Atlantic salmon and brown trout. populations in Shelligan Burn, ~cotland. Mean biomass of each age group and total biomass of each species every year for the 10-year period are presented. Seasonal variability in growth rates, mean size and weight, and population density are investigated. Variability within each cohort through its lifetime is also discussed. Elliott, J.M. 1973. The food of brown and rainbow trout (Salmo trutta and S. gairdneri) in relation to the abundance of drifting invertebrates in a-mountain stream. Oecologia 12:329-347. Elliott, J.M. 1976. Salmo trutta L. The downstream drifti~g of eggs of brown trout, J. Fish BioI. 9:45~50. Ellis, R.J., andH. Gowing. 1957. Relationship betweert food supply and condition of wild brown trout, Salmo trutta Linnaeus, in a Michigan stream. Lirnnol. Oceanogr.---Z:Z99-308. . Elser, A.A. 1968. Fish populations of a trout stream in relation to. o major habitat zones and channel alterations. Trans. Amer. Fish. Soc. 97:389-397. A study of brown, rainbow, and brook trout populations in Little Prickly Pear Creek, Montana, during the summers of 1965 and 1966. Spatial variability is shown through" data on biomass in 11 sections of the stream. Four of these sections have been altered due to adjacent railroad and highway construction. Major changes include straightening 'of the channel, loss of streambank cover, and changes in pool-riffle ratios. Comparisons are made between altered and unaltered sections in the same habitat zones in terms of physical stream characteristics and trout abundance. Elson, P.F. 1957. The importance of size in the change from parr to smolt in Atlantic salmon. Can. Fish Cult. 21:1-6. Elson, P.F. 1962. Predator-prey relationships between fish-eating birds and Atlantic salmon. Fish. Res. Bd. Can. Bull. 133. 87 pp. A study of relationships between mergansers, kingfishers, and juvenile Atlantic salmon in the Pollett River, New Brunswick, from 1942 to 1953. Control of the predators was applied .from 1947 to 1950, reducing pressure on the salmon by approximately 90%. Survival of planted underyearlings in years before and during control is examined. Data are presented on number of fish planted, number of parr one year later, and smolt output for each year of the ,study. The role of these predators as limiting factors for smolt production is discussed. Methods of predator control are also investigated. The optimum levels of each predator species for this river. section are determined. 37 ------- Elson, P.F. 1975. Atlantic salmon rivers, smolt production and optimal spawning: an overview of natural production. pp. 96-119 in J.R. Bohne ,and L. Sochasky (eds.), New England Atlantic salmon-restoration conference. Int. Atlantic Salmon Spec. Publ. Sere No.6. Elwood, J.W., and T.F. Waters. 1969. Effects of floods on food consUmption and production rates of a stream brook trout population. Trans. Amer. Fish. Soc. 98:253-262. Erman, D.C., and V.M. Hawthorne. 1976. The quantitative importance of an intermittent stream in the spawning of rainbow trout. Trans. Amer. Fish. Soc. 105:675-681. Erman, D.C., and G.R. Leidy. 1975. Downstream movement of rainbow trout fry in a tributary of Sagehen Creek, under permanent and intermittent flow. Trans. Amer. Fish. Soc. 104:467-473. ' Erman, D.C., J.D. Newbold, and K.B. Roby. 1977.' Evaluation of streamside buffers trips for protecting aquatic organisms. 165. Calif. Water'Resource Center, Davis. 48 pp. Contr. No. , , Everest, F.H. 1973. Ecology and management of summer steelhead in the Rogue River. Ore. State Game Comm. Fish. Res. Rep. No.7. 48 pp. Everest, F.H., and D.W. Chapman. 1972. Habitat selection and spatial interaction by juvenile chinook salmon and steelhead trout in two Idaho streams. J. Fish. Res. Bd. Can. 29:91-100.' , Foerster, R.E., and W.E. Ricker. 1941. The effect of reduction of , predaceous fish on survival of young sockeye salmon at Cultus Lake. J. Fish. Res. Bd. Can. 5:315-336. Foerster, R.E., and W.E. Ricker. 1953. The coho salmon of Cultus Lake and SweItzer Creek.J. Fish. Res. Bd. Can. 10:293-319. Gard, R. 1961. Effects of beaver on trout in Sagehen Creek, California. J. Wildl. Mgmt. 25:221-242. Gard, R.,and G.A. Flittner. 1974. in Sagehen Creek, California. Distribution and abundance of fishes J. Wildl. Mgmt. 38:347-358. A 10-year study of salmonid and other fish populations, from 1952 through 1961. Data on average biomass over the entire period in 11 sections of the stream for brook, brown, and rainbow trout show spatial variability and relative distributions of the species. Temporal variability in abundance is also shown. The effects of stream gradient, water temperature, elevation, floods, and beavers on these two kinds of variability are examined. 38 ------- Gard, R., and D.W. Seegrist. 1972. Abtn1dance' and harvest of trout in Sagehen Creek, California. 'Trans. Amer. Fish Soc. 101:463-477., 'Gibbons, D.R., and E.O. Salo. 1973. An annotated bibliography of the effects of logging on fish of the western United States and Canada. U.S.D.A. For. Servo Gen. Tech. Rep. PNW-10. 145 pp. Gibson, R.J. 1973. The interrelationships of brook trout, Salvelinus fontinalis (Mi tchill), and juvenile Atlantic salmon, Salmo salar L. Ph.D. thesis. Univ. Waterloo, Ontario. Gibson, R.J., and D. Galbraith. 1975. The relationships between invertebrate drift and salmonid populations in the Matamek River, Quebec, below a lake. Trans. Amer. Fish. Soc. 104:529-535. Gibson, R.J., and G. Power.' 1975. Selection by brook trout (Salvelinus fontinalis) and juvenile Atlantic salmon (Salmo salar) of shade' related to water depth. J. Fish. Res ~ Bd. Can. 32:1652-1656. Giger, R.D. Oregon. 1972. Ecology and management of coastal cutthroat trout in Ore. State Game Comm. Fish. Res., Rep. No.6. 61 pp. Glova, G.J. 1978~ Pattern and mechanism of resource p~rtitioning between stream populations of juvenile coho salmon (Oncorhynchus 'kisutch) and coastal cutthroat trout (Salmo clarki clarki). Ph.D. thesis. Univ. British Columbia; Vancouver. 185 pp. An investigation of sympatric and allopatric populations 'of coho salmon and cutthroat trout in six British Columbia streams from 1973 to 1976. Data are presented on biomass and fish length in pool, glide, and riffle habitats in each stream; Relationships between biomass and streamflow,pool area, and volume are examined. Spatial variability in salmonid density and biomass in three study sections in each of two streams is shown. Interactions between the two species are discussed in terms of abundances in different habitats. The importance of drift organisms in the diet of each species is investigated. Density and biomass of the sculpin (Cottus aleuticus) are estimated, and. its' relationships to the salmonids are studied. Griffith, J.S., Jr. 1972. Comparative behavior and habitat utilization of brook trout (Salvelinus fontinalis) and cutthroat trout (Salmo clarki) in small streams in northern Idaho. J. Fish. Res. Bd. Can. 29: 265-273. Griffith, J.S., Jr. 1974. Utilization of invertebrate drift by brook trout (Salvelinus fontinalis) and cutthroat trout (Salmo clarki) in small streams in Idaho. Trans. Amer. Fish. Soc. 103:440-447. Hall, J.D., M.L. Murphy, and R.S. Aho. 1978. An improved design for assessing impacts of watershed practices on small streams. Verh. Int. Ver. Limnol. 20:1359-1365. 39 . I ------- Four approaches to studying impacts of watershed management practices on streams are discussed. The long-term study is an intensive approach involving pre-treatment and post-treatment work on sites in the same stream. The extensive before-after approach is a short- term study involving sites in several streams. The intensive post- treatment analysis compares two adjacent sections of the same stream, one in a previously treated area and the other just upstream, over several years. The last technique is extensive post-treatment analysis, a short-term study involving comparisons between adjacent logged and forested stream sections in many watersheds. Advantages and disadvantages of each approach are discussed, as well as recommendations from statisti~al, practical, and economic viewpoints. Hallam, J.C. 1959. Habitat and associated fauna of four species of fish in ontario streams. J. Fish. Res. Bd. Can. 16:147-173. Hanson, D.L., and T.F. Waters. 1974. Recovery of standing crop and . production rate of a brook trout population in a flood-damaged stream. Trans. Amer. Fish. Soc. 103:431-439. Hanzel, D.A. 1960. The distribution of the cutthroat trout (Salmo . clarki) in Montana. Proc.Montana Acad. Sci. 19:32-71.----- Hartman, G.F. 1965. The role of behavior in the ecology and interaction of underyearling coho salmon (Oncorhynchus kisutch) and steelhead trout (Salmo gairdneri). J. Fish. Res. Bd. Can. 22: 1035-1081. Hartman, G.F., and C.A. Gill. 1968. Distributions of juvenile steelhead and cutthroat trout (Salmo gairdneri and~. clarki clarki) within streams in southwestern British Columbia. J. Fish. Res. Bd. Can. 25:33-48.' . Havey, K.A., and R.M. Davis. 1970. Factors influencing standing crops and survival of juvenile salmon at Barrows Stream, Maine. Trans. Amer. Fish. Soc. 99:297-311. Presents data on biomass of Atlantic salmon in this stream from 1960 through 1965. Biological and physical variables that could influence biomass and survival are studied. These factors include rainfall, streamflow, air temperature, water temperature, numerical standing crop, and biomass of other fish species in the stream. Regression analyses between these parameters and survival and biomass of age 0+ and 1+ salmon are performed to determine those of greatest significance. Hazzard, A.S., and M.J. Madsen. 1933. Studies of the food of the cutthroat trout. Trans. Amer. Fish. Soc. 63:198-207. Heiser, D.W. 1966. Age and growth of anadromous Dolly Varden char Salvelinus malrna (Walbaurn) in Eva Creek, Baranof Island, southeastern Alaska. Alaska Dept. Fish Game. Res. Rep. No.5. 29 pp. 40 ------- Hildebrand, S.G. 1971. The effect of coho spawning on the benthic invertebrates of the Platte River, Benzie County, Michigan. Trans. Amer. Fish. Soc. 100:61-68. Hoar, W.S. 1951. The behavior of chum, pink and coho salmon in relation to ~heir seaward migration~ J. Fish. Res. Bd. Can. 8:24l-263~ Hobbs, D.P. 1937. Natural reproduction of quinnat salmon, brown and rainbow trout in certain New Zealand waters. New Zealand Mar. Dept. Fish. Bull. No.6. 104 pp. Holton, G.D. 1953. A trout population study on a small creek in Gallatin County, Montana. J. Wildl. Mgmt. 17:62-82. Spatial variability in biomass of brook, brown, and rainbow trout is shown for four sections of Trout Creek, Montana in 1950 and 1951. This variability may be related tothe relative distribution of the species, stream section width and depth, and pool-riffle ratiQs. Survival, growth, reproduction, arid movements of these species are also examined. In addition, survival of planted brook trout is investigated. Hoopes, R.L. 1975. Flooding as the result of Hurricane Agnes, and its effect on a native brook trout population in an infertile headwater stream in central Pennsylvania. Trans. Amer. Fish. Soc. 104:96-99. Hopkins, C.L. 1970. Some aspects of the bionomics of fish in a brown. trOut nursery stream. New Zealand Mar. Dept. Fish. Res. Bull. No.4. . 38 pp. A study of brown trout and benthic invertebrate populations in two New Zealand streams. Seasonal trout population estimates from October 1964 through June 1966 are presented.. Population density, growth, and food of the trout are analyzed. The availability of food and living space are discussed as potential limiting factors for the trout. The extent to ~hich these factors are important with respect to size and age of the fish is also examined. In addition, data on biomass of the dominant benthic organisms in three study sections from 1964 through 1966 are presented. Horton, P.A. 1961. The bionomics of brown trout in a Dartmoor stream. Anim. Ecol. 30:311-338. J. Horton, P.A., R.G. Baily, and S.I. Wilsdon. 1968. A comparative study of the bionomics of the salmonids of three Devon streams. Arch. Hydrobiol. 65:187-204. Huet, M. 1959. Profiles and biology of Western European streams as related to fish management. Trans. Amer. Fish. Soc. . 88:155-163. Hunt, R.L. 1971. Responses ofa brook trout population to habitat development in Lawrence Creek. Wise. Dept. Nat. Resour. Tech. Bull. No. 48. 35 pp. 41 ------- HW1t, R.L. 1974. Annual production by brook trout in Lawrence Creek during eleven successive years. Wise. Dept. Nat. Resour. Tech. Bull. No. 82. 28 pp. Extensive data are presented on a long-term study of the brook trout'population in this Wisconsin stream from 1960 through 1970. Production is given for the entire population and by age group for' o four stream study sections. Temporal variability in biomass is also shown by age group for April and September.' Comparison is made between production in one section that W1derwent habitat development and production in the other sections. A discussion of compensatory mechanisms in growth and survival of each age group that keep the total production at a relatively stable level is also included. ' HW1t, R.L. '1976. A long-term evaluation of trout habitat development and its relation to improving management-related research. Trans. , Amer~ Fish Soc. 105:361-364. ' HW1ter, J.G. 1959. Survival and production of pink and chum salmon in a coastal stream. J. Fish. Res. Bd. Can. 16:835-886. 'A comprehensive study of factors affecting survival and production of pink and chum salmon in Hooknose Creek, British Columbia, from 1947 through 1956. Effects of temperature, stream discharge, sex ratio, population density, and predation on these populations are analyzed. Data on the number of predators (coho salmon and sculpin) as well as weir COW1tS of pink and chum salmon fry and percent predation are given for each year of the study. 0 Hynes, H.B.N. 19:1-15. 1975. The stream and its valley. Verh. Int. Ver. Limnol. Idyll, C.1942. Food of rainbow, cutthroat, and brown trout in the Cowichan River system, B.C. J. Fish. Res~ Bd. Can. 5:448-458. Jones, A~N. 1975. A preliminary study of fish segregation in salmon spawning streams. J. Fish BioI. 7:95-104. Keeley, P.L., and T.E. Nickelson. salmonids. Ore. Wildl. Comm. 18 pp. 1974. Streamflow requirements of Ann. Prog. Rep. AFS-62-3. Keenleyside, M.H.A. 1962. Skin-diving observations of Atlantic salmon and brook trout in the Miramichi River, New BrW1swick. J. Fish. Res. Bd. Can. 19:625-634. Kennedy, M~, and P. Fitzmaurice. 197I. Salmo-trutta (L.) in Irish waters. B. (18):269-352. Growth and food of brown trout Proc. Roy~ Irish Acad. 71, Sect. 42 ------- Knight. N.J. 1980. Factors affecting the smo1t yield of coho salmon (Oncorhynchus kisutch) in three Oregon streams. M.S. thesis. Oregon State Un'iv.. Corvallis. 101 pp. Koski. K.V. 1966. The survival of coho salmon (Oncorhynchus kisutch) from egg deposition to emergence in three Oregon coastal streams. M.S. thesis. Oregon State Univ.. Corvallis. 84 pp. ' Koski. K V. 1975. The survival and fitness of two stocks of chum salmon (Oncorhynchus keta) from egg deposition to emergence in a contro11ed- stream environment at Big Beef Creek. Ph.D. thesis. Univ. of Washington. Seattle. 211 pp.. . Krohn. D.C. 1967. Production of the reticulate sculpin (Cottus perp1exus) and its predation on salmon fry in three Oregon streams. M.S. thesis. Oregon State Uni v.. Corvall is. 78 pp. , 'Lake. J.S. Austr. 1957. Trout populations and habitats in ,New South ,Wales. J~ Mar. Freshw. Res. 8:414-450. A comprehensive survey of 100 streams containing brown'and/or rainbow trout. These are divided into three general categories with' corresponding differences in mean pH. mean temperatures. bottom fauna production. and trout growth. Analysis of angling effects on the trout populations is also unde~taken. ' ' Larkin. P.A. 1971. Simulation studies of the Adams River sockeye salmon (Oncorhynchus nerka). J. Fish. Res. Bd. Can. 28:1493-1502. Latta. W.C. 1969. Some factors affecting survival of young-of-the- year brook trout. Sa1velinus fontinalis (Mitchill). in streams. pp. 229-242 in T.G. Northcote (ed.). Symposium on salmon and trout in streams; H.R:-MacMil1an Lectures in Fisheries. Univ. British Columbia. Vancouver. A study of age 0 brook trout from the Pigeon River. Michigan, in 1962. 1964. and 1965. Magnitude and periodicity of mortality are discussed in a brief review of other studies. Mortality from starvation, in relation to water temperature. is investigated in stream aquaria. Relationships between levels of groundwater and abundance of fry are also examined. The importance of grOtmdwater levels in regulating the carrying capacity of the stream is discussed. LeCren. E.D. 1969. Estimates of fish populations and production in small streams in England. Pp. 269-280 in T.G. Northcote (ed.). Symposium on salmon and trout in streams; ~R. MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. . Leonard, J.W. 1941. Some observations on the winter feeding habits of brook trout fingerlings in relation to natural food organisms present. Trans. Amer. Fish. Soc. 71:219-235. ' 43 ------- Lewis, S.L. 1969. Physical factors influencing fish populations in pools of a trout. stream. ,Trans. Amer. Fish. Soc. 98:14-19. A study of brown and rainbow trout populations in pools of Little Prickly Pear Creek, Montana, in the sunnners of 1965 and 1966. The importance of surface area, volume, depth, streamflow~ and cover in accounting for the variation in population numbers, is. examined. Combinations of these' factors are also considered as well as differential responses of ,.the two species. Cover is discussed, in terms of protection and . photonegative response; streamflow is analyzed in terms of space- food relationships. Lindroth, A. '1955. Distribution," territorial behaviour and movements of sea trout fry in the River Indalsa1ven. Inst. Freshw. Res. Drottningholm Rep. 36:l04-l19~ ' Lindsay, R.B. 1974., Distribution and survival of coho salmon fry after emigration from natal streams. M.S. thesis. Oregon State Univ., Corvallis. 41 pp. , , Lister, D.B. and H.S. Genoe. 1970. Stream habitat utilization by cohabiting underyearlings of chinook (Oncorhynchus tshawytscha) and coho (0. kisutch) salmon in the Big Qua1icum River, British Columbia. -J. Fish. Res. Bd.Can. 27: 1215-1224. Lorz, H.W. 1974. Ecology and management of brown trout in Little Deschutes River. Ore. Wildl. Connn. Fish. Res. Rep. No.8. '49 pp. Lowry, G.R. 1964. Net production, movement, and food of cutthroat trout (Salmo clarki Richardson) in three Oregon coastal streams. M.S. thesis. Oregon State Univ., Corvallis. 72 pp. A study of cutthroat trout populations in the streams of the Alsea Watershed Study in 1962 and 1963. Population size, growth rate, , and net production estimates are made. Data on monthly biomass for the 1959-1962 year classes are presented. Magnitude and timing of upstream adult and downstream kelt and smol t migrations are investigated. Trout food habits are studied during the time of emergence and early growth of coho salmon fry to determine the extent of predation. Seasonal variation in kind and quantity of food consumed is also examined. Much of the information is published in Lowry (1965 and 1966). Lowry, G.R. 1965. Movement of cutthroat trout, Salmo clarki clarki (Richardson) in three Oregon coastal streams. Trans. Amer. Fish. Soc. 94:334-338. Lowry, G.R. 1966. Production and. food of cutthroat trout in three Oregon coastal streams. J. Wild1. Mgmt. 30:754-767. 44 ------- Maciolek, J.A;" and P.R. Needham. 1952. Ecological effects of winter conditions on trout and trout foods in Convict Creek, California, 1951. Trans. Amer. Fish. Soc. 81:202-217. Maher, F.P., and P.A. Larkin. 1954. Life history of the steelhead trout 'of the Chilliwack River, British Columbia. Trans. Amer. Fish. Soc. 84:27-38. Maitland, 'P.S. 1965. The feeding relationships of salmon, trout, minnows, stone loach and three~spined sticklebacks in the River Endrick, Scotland. ' J. Anim. Ecol. 34:109-133. Major, R.L., andJ.L. Mighell. 1969. Egg-to-migrant survival of spring chinook salmon (Oncorhynchus tshawytscha) in the Yakima River, . Washington. Fish. Bull. 67:347-359. 'Manion, P.J. 1977. Downstream movement of fish in a tributary of . southern Lake Superior. Progr. Fish-Cult. 39:14-16. . Mann,R.H.K. 1971. The populations, growth 'and production of fish in four small streams in southern England. : J. Anim. Ecol. 40:155-190. Mason, J.C. 1974. Movements of fish populations in Vancouver Island: A' summary of weir operations including comments on specific life histories. 'Serv~ Tech. Rep. 483. 35 pp. ' Lynm Creek, during 1971 and 1972 Can. Fish. Mar. Mason, J.C. 1975. Seaward movement of juvenile fishes, including lunar periodicity in the movement of coho salmon (Oncorhynchus kisutch) . fry. J. Fish. Res. Bd. Can. 32:2542-2547. ' . Mason;J.C. 1976. Response of underyearling coho salmon to supplemental feeding in a natural stream. J. Wildl. Mgmt. 40:775-788. An, important study on the role of food as a limiting factor in juvenile coho salmon production in a small British Columbia stream. Supplemental feeding resulted in a six-fold increase in the stream carrying capacity. Effects of this feeding on survival, growth, and pre-winter lipid reserve are also examined. However, there were no effects on the smo1t yield the following spring. Attempts to increase winter carrying capacity by installing artificial refuges were ineffective. Complex winter behavior patterns may be responsible in limiting the population size. Mason, J.C., and D.W. Chapman. 1965. Significance of early emergence, environmental rearing capacity, and behavioral ecology of juvenile coho salmon in stream channels. J. Fish. Res. Bd. Can. 22:173-190. A study of juvenile coho salmon behavior in artificial stream . channels. Aggressive behavior is observed within 1 week of emergence and is investigated for 5 months. Territorial defense, threat, and 45 ------- chasing areexaminedo Distribution in pool. and riffle areas is studied. The effects of different times of emergence on fish size and territoriality are also examined~ Environmental rearing capacity is discussed in terms of food and stream area. McDonald,.J. 1960. The behaviour of Pacific salmon fry during their downstream migration to freshwater and saltwater nursery areas. J. Fish. Res. Bd. Can. 17:655,..676., McFadden, J.T. 1961a. An ecological comparison of six brown trout CSalmo trutta) populations. Ph.D. thesis. Pennsylvania State Univ. ,91 pp. ' McFadden, J.T. 1961bo A population study of the brook trout, Salvelinus fontinalis. Wildl. MonOgr. No.7. 73 pp. ' A comprehensive study of the brook trout population in Lawrence Creek, Wisconsin, from 1953 through 1957. Data are presented on population, estimates, lengths, age structure, density, biomass, and natural mortality in each of four study sections for each age group of trout. Length-fecundity relationships are also examined. Angling intensity is investigated by means of a complete creel census each year. Data obtained include number of fishing trips, number, sex, size, and weight of trout caught and exploitation rates. Angler mortality is compared to natural mortality of the trout, and management options are discussed as means of keeping an adequate recruitment to the population. McFadden, JoT. 1969~ DYnamics and regulation of sa1monid populations in streams. pp. 313-329 in T.G. Northcote Ced.), Symposium on salmon and trout in streams; H.~ MacMillan Lectures in Fisheries.Univ. British Columbia, Vancouver. McFadden, J,. T., G.R. Alexander, and D.S. Shetter. 1967. Numerical changes and population regulation in brook trout Salvelinus fontinalis. J. Fish. Res. Bd. Can. 24:1425-1459. McKernan, D.L., D.R. Johnson, and J.I. Hodges. 1950. Some factors influencing the trends of salmon populations in Oregon. Trans. N. Amer. Wildl. Conf. 15:427-449. One of the earlier studies attempting to explain fluctuations in salmon population levels in Oregon. Columbia River chinook salmon catch data are presented from 1866 through 1948 and coho salmon catch data from coastal rivers are presented from 1923 to 1948. -P61li.ltiOn, changes in angling regulations, hatcheries, logging, . streamflow, and salinity are analyzed as potential factors responsible for the observed fluctuations. Ih addition, the, effects of the fishery, indicated by marking experiments, fishing intensity, and economic trends, are examined.- 46 ------- McNeil, W.J. 1966. Effect of the spawning bed environment on reproduction of pink and chum salmon. Fish. Bull. 65 :495-523. . 'McNeil, W.J. 1968. Effect of streamflow on survival of pink and chum salmon in spawning beds. ?p. 96-114 in Proc. Forum on the relation between logging and salmon. Feb. 8-9:-1968. Alaska Distr., Amer. Inst. Fish. Res. BioI., Jtmeau. . McNeil, W.J. 1969. Survival of pink and chum salmon eggs and alevins. ?po 101-117 in T.G. Northcote (ed.), Symposium on salmon and trout in streams; H.R:-MacMillan Lectures in Fisheries. Univ. British Columbia. Reviews a long-term study of freshwater survival from 1940 through 1965. Also discussed are a dome-shaped production curve, density- independent mortaiity from droughts~ floods, and freezing temperatures, artd' density-dependent mortality from redd superimposition. Mense, J.B. 19700 Relation of density to brown trout movement'in a Michigan stream. Ph.D. thesis. Mich. State Univ. 82 pp. Mense, J.B. 1975. Relation 'of density to brown trout movement in a Michigan stream. Trans. Amero Fish. Soco 104:688-695. , Merrell, T.R. 1962. freshwater survival of pink salmon at Sashin Alaska. Pp. 59-72 in NoJ~ Wilimovsky (ed.), Symposium on pink H.R. MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. Creek, salmon; Presents data on number of spawners, egg deposition, number of seaward migrating fry, and freshwater survival from 1940 through 1959. Differences in magnitude and timing between odd-year runs and even-year rtmS are examined. Preliminary data in the last year of the study seem to indicate survival is related to stream gradient 0 There also seems to be a relationship between size of the rtm and section of the stream used for spawningo Miller, R.B. 1957. Permanence and size of home territory in stream-dwelling cutthroat trout. J. Fish. Res. Bd. Can. 14:687-691. Miller, W.H. 1971. Factors influencing migration of chinook salmon fry (Oncorhynchus tshawytscha) in the Salmon River, Idaho. Ph.D. thesis. Univ. Idaho, Moscow. Mills, D.H. 1964. The ecology of the young stages of the Atlantic salmon in the River Bran, Ross-Shireo Freshw. Salmon Fish. Res. Edinburgh 0 No. 32. . 58 ppo Mills, D.H. 1969. The survival of juvenile Atlantic salmon and brown trout in some Scottish streams. pp. 217-228 in T.G. Northcote (ed.), 47 ------- Symposium on salmon and trout in streams~ HoR. MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. 'Moring, J.R. 1975. The Alsea Watershed Study: Effects of logging on the aquatic resources of thr~e headwater streams of the Alsea River, Oregon. Part II -.Changes in environmental conditions. Ore. Dept. Fish Wildl., Fish. Res. Rep. No. 9~ 39 pp. Moring, J.R., and R.L. Lantz~ 1975. The Alsea Waters~ed Study: Effects' of logging on the aquatic resources of three headwater streams of the Alsea River, Oregon. Part I - Biological studies. Ore. Dept. Fish Wildl., Fish. Res. Rep. No.9. 66 pp. A summary' report, incorporating data from a number of earlier publications from the study. Three small tributaries were,studied, one clearcut down to the streambank, one clearcut with buffer strips left, and the third left uncut as a control, for 15 years. The pre-logging period was 1959 to 1965 and the post-logging period was 1967 to 1973. Biological studies concentrate mainly on coho salmon and cutthroat trout, the dominant species in the streams. ' For adults, data include numbers, timing of migration, size, sex ratio, and fecundity. Data on juveniles include emergence, growth, biomass, production, mortality, and downstream migration. Effects of logging are reported.' , Mortensen, Eo 1977. Pop~lation, survival, growth and production of trout Salmo trutta in a small Danish stream. Oikos 28:9~15. Population dynamics of brown trout in a small Danish stream'were 'examined from 1973 through 1975. Biomass of the 1971-1975 year classes is investigated. Population size, production, mortality, length-weight relationships, and smolt yield are also discussed. Mortensen, E. 1978. The population dynamics and production of trout (Salmo trutta L.) in a small Danish stream. Pp. 151-160 in J.R. ' Moring (ed.), Proc. wild trout - catchable trout symp. Ore. Dept. Fish Wildlo, Res~ Devel. Sect., Portland. Population size, survival, growth, biomass, and production of brown trout were studied in three sections of a small Danish stream from 1974 through 1976. Variability in age group composition and biomass between the three sections is shown. Density-dependent fry mortality, density-independent mortality of older fish, and variability in production-biomass ratios between age groups are also analyzed. Mundie, JoH. 1969. Ecological-J.IDplications of the diet of juvenile coho in streams. Pp. 135-152 in T.G. Northcote (ed.), Symposium on salmon and trout in streams; Ho~ MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. Mundie, J.H. 1974. Res. Bd. Can. Optimization of the salmonid nursery stream. 31:1827-1837. J. Fish. 48 ------- Murphy, M.L.1979. Predator assemblages in old-growth and logged sections of small Cascade streams. M.S. thesis. Oregon State Univ., Corvallis. 72 pp. . Murphy, M.L., and J.D. Hall. MS. Effects of clearcutting on predators and their habitat in small streams of the Cascade mountains, Oregon. Can. J. Fish. Aquat. S'ci. (in press). . Murray, A.R. J968. Numbers of Atlantic salmon and brook .trout captured and marked at the Litle Codroy River, Newfoundland, counting fence and auxiliary traps, 1954-1963. Fish. Res. Bd. Can. Tech. Rep. No. 84. . 135 pp. Magnitude and timing of upstream adult and downstream kelt and smolt migrations in this river are examined for both species. Marking experiments are used to determine movements,' utilization, and survival. Marine survival is determined from commercial catch and escapement of marked fish. Murray, A.R., and.T.J Harmon. 1969. A preliminary consideration of the factors affecting the productivity of Newfoundland streams. Fish. Res. Board Can. Tech. Rep. No. 130. 405 pp. . An extensive survey of the major streams in Newfoundland,. including. description, location, and certain morphometric, edaphic, climatic, and . biotic. characteristics. Morphometric factors examined include watershed shape, elevation, gradient, vegetational cover, runoff, and discharge. Edaphic factors include surface geology, total dissolved solids, micronutrients, acidity, conductivity, and hardness. Climatic factors examined are air temperature and precipitation. Competition, predation, fecundity, emigration, survival, and disease are the biotic factors investigated. Both fish and aquatic invertebrate populations are studied. Native salmonid species are Atlantic salmon, brook trout, and Arctic char. Pink salmon, rainbow trout, and brown trout have been introduced. In addition, the sport fishery for Atlantic salmon is examined. . Muttkowski, R.A. 1929. The ecology of trout streams in Yellowstone National Park. Roosevelt Wildl. Ann. 2:155-240. Narver, D.W. 1971. Effects of logging debris on fish production. pp. 100-111 in J.T. Krygier and J.D. Hall (eds.), A symposium: Forest land uses an~stream environment. Oregon State Univ., Corvallis. Narver, D.W., and B.C. Andersen. 1974. Fish populations of Carnation Creek and other Barkley Sound streams - 1970-1973: Data record and progress report. Fish. Res. Bd. Can.. MS Rep. 1303. 115 pp. The initial data from a long-term study of the effects of logging on the aquatic resources of a British Columbia watershed. The study' is divided into pre-logging (1970-1974), logging (1975-1979), and post-logging (after 1979) periods. In partic~lar sampling is 49 ------- conducted on sections of upper and lower Carnation Creek, "C". tributary, "1600" tributary, and Useless, Frederick, Ritherdon, and South Pachena creeks., This report contains data on population estimates, density, late summer biomass, ,growth, length-weight relationships, and condition of both resident and anadromous . salmonids. The primary species include coho and chwn .salmon, and rainbow,steelhead, and cutthroat trout. Data are updated for every year through 1977 by Andersen and Narver (1975) and Andersen (1978). Neave, F. 1947. Natural propagation of chum salmon in a coastal stream.. Fish. Res. Bd.Can. Progr. Rep., Pac~ Coast Sta. No. 70:20-21. Neave, F. 1949. Game fish populations of the Cowichan River. Res. Bd. Can. Bull. 84. 32 pp. Fish. Neave, F. 1953. Principles affecting the size of pink and chum salmon populations in British Columbia. J. Fish. Res. Bd. Can. 9:450-491. Neave, F~, and W.P. Wickett. 1953.. Factors affecting the freshwater development of Pacific salmon in British Columbia. Proc. Seventh Pac., Sci. Congr. (1949). 4:548-556. A review of some chemical, physical, and biological factors that .affect salmon populations in British Columbia streams. Chemic~l factors include dissolved oxygen and pH. Important physical factors are bottom type, particularly on spawning grounds, obstructions, streamflow, including floods, and water temperature. Predation, interspecific and intraspecific competition, and disease are important biological factors discussed. Relationships between precipitation . and adult returns two years later are also analyzed. Needham; P.R., and F.K. Cramer. 1943. Movement of trout in Convict Creek, California. J. Wild1. Mgmt. 7:142-148. Needham, P.R., and A.C. Jones. 1959. Flow, temperature, solar radiation, and ice in relation to activities of fishes in Sagehen Creek, California. Ecology 40:465-474. Needham, P.R., J.W. Moffett, and D.W. Slater. 1945. Fluctuations in wild brown trout populations in Convict Creek, California. J. Wi1dl. Mgmt. 9:9-25. A study of the brown trout populations in two sections of this stream from 1939 to 1944, one closed and the other open to angling. Reproduction, survival, seasonal weight changes, and- density are analyzed in both sections. The effects of angling on these. parameters are s~own. The possibility of stocking the stream with more trout is discussed as a method to offset angling pressure. 50 ------- Neill, R.M. 1938. The food and feeding of the brown trout (Salmo trutta L.) in relation to the organic environment. Trans. Roy. Soc. Edinburgh 59(pt. 11):481-520. Newbold, J.D., D.C. Erman, and K.B. Roby. 1980.. Logging effects on stream macroinvertebrates and the role of bufferstrips. J. Fish. Res. Bd~ Can. (In press.) . .Nickelson, T.E. 1975. Streamflow requirements of salmonids. Wildl. Comma Ann. Prog. R~p. AFS-62-4. 19 pp. Ore. Nickelson, T.E., and R.E. Hafele. 1978. Streamflow requirements of salmonids. Ore. Dept. Fish Wildl. Proj. AFS-62. 26 pp. Nickelson, T .E. ,. and R. R. Reisenbichler. 1977. Streamflow requirements of salmonids. Ore. Dept. Fish Wildl. Proj. AFS-62. 24 pp. Northcote, T.G. 1978. Migratory strategies and production in freshwater fishes. Pp. 326-~59 in S.D. Gerking (ed.), Ecology of freshwater fish production. BlackweIT Scientific Publ., Oxford. NylIian, O.L.1970. Ecological interaction of brown trout, Salmo trutta L., and brook trout, Salvelinus fontinalis (Mitchil1), in a stream. Can. Field-Nat. 84:343-350. O'Connor, J"F., and G. Power. 1976.. Production by brook trout (Salvelinus fOhtinalis) in four streams in the Matamek Watershed, Quebec. J. Fish. Res. Bd. Can. 33:6-18. Four streams were examined for brook trout populations from 1971 through 1973. Biomass of each age group each year is presented as well as total biomass in the study sections. Production and production-biomass ratios in each stream are also examinedo Differences in these parameters between streams are related to variations in stream cover, recruitment, and total biomass 0 The roles of food and cover as possible limiting factors on adult production are discussed. Orsborn, JoF., and C.H. Allman (edso). 19760 Instream flow needs: a symposium. Vol. 2. Amer. Fish. Soc. 657 pp. Osborn, C.E. 1967. A population study of the rainbow trout (Salmo gairdneri) in a central Oregon stream. M.S. thesis. Oregon State Univ., Corvallis. 65 pp. . Osterdahl, L. 1969. Thesmo1t run of a small Swedish river. pp. 205-215 in T.G. Northcote (ed.) , Symposium on salmon and trout in streams; ~R. MacMillan Lectures in Fisherieso Univ. British Columbia, Vancouver. Otto, C. 1976. Size, growth, population density and food of brown trout Sa1mo trutta L. in two sections of a south Swedish stream. J. Fish BioI. 8:477-488.. 51 ------- Overton, W.S. 1978. Study design considerations for impact assessments. pp. 24-25 in W.T. Mason, Jr. (ed.),Methods for the assessment and prediction-of mineral mining impacts on aquatic communities. U.S. Fish Wildl. Servo FWSjOBS-78j30. Pearson, L.S., K.R. Conover, and R.E. Sams. 1970. Factors affecting the natural rearing of juvenile coho salmon during the summer low flow' season. Ore. ,Fish Comm., Portland. 64 pp. (Processed.) A study of summer juvenile coho salmon populations in 19 streams in four Oregon river systems from 1962 through 1966. Relationships between minimum streamflow and coho density are examined. Mean pool velocity and fish density in pools also seem to be related. Effects of streamflow on f~sh size and incidence of disease are investigated. Fish numbers and'size are related to the rearing capacity of a stream. Shade, cover, temperature, bottom composition, stream size, comPetition, and food production can also affect coho rearing capacity. Pennak, R.W. 1971. Hydrobiologia Toward a classification of lotic habitats. 38:321-334. Phillips, R.W. 1970. Effects of sediment on the gravel environment and fish production. pp. 64-74 in J.T. Krygier and J.D. Hall (eds.), A symposium: Forest land uses and stream environment. Oregon State Uni V. , Corvallis.' . Platts, W.S. 1974. Geomorphic and aquatic conditions influencing salmonids and stream classification with application to ecosystem classification. Surface Environ. Mining, U.S. For. Serv., Billings, Montana. xiv + 199 pp. An extensive study from 1970 to 1972 on conditions influencing salmonid populations in 291 stream sites in the upper South Fork of the Salmon River watershed, Idaho. Physical and hydrologic parameters examined include climate, geological processes and types, stream channel width, depth, gradient, and order, elevation, pool- riffle ratios, and streambank vegetation and stability. Effects of these factors on the salmonid populations are analyzed. The salmonid species present include rainbow, steelhead, and brook trout, Dolly Varden, chinook salmon, and mountain whitefish. Platts, W.S., and F.E. Partridge. 1978. Rearing of chinook salmon in . tributaries of the South Fork Salmon River, Idaho.. U.S.D.A. For. Servo Res. Pap. INT-205. 12 pp. Power, G. 1973. Estimates of age, growth, standing crop and production of salmonids in ~ome north Norwegian rivers and streams. Inst. Freshw. Res. Drottningholm Rep. 53:78-111. Pritchard, A.L. 1944. Physical characteristics and behavior of pink salmon fry at McClinton Creek, B.C. J. Fish. ,Res. Bd. Can. 6:217-227. 52 ------- Raleigh, R.F. . 1978. Habitat evaluation procedure for aquatic assessments. pp. 131-137 in W.T. Mason, Jr. (ed.), Methods for the assessment and prediction of mineral mining impacts on aquatic. communities. U.S. Fish Wildl. Servo A~S/OBS-18/30. Rayner, H.J. 1937. Notes on the food of trout of.Yosemite National Park. Calif. Fish Game 23:149-156. Redmond, M.A. 1975. Natural production. Pp. 134-135 in J.R. Bohne and L. Sochasky (eds~), New England Atlantic salmon restoration conference. Int. Atlantic Salmon Spec. P~bl. Sere No.6.' Reed, R.J. 1967..0bservat:Lon of -fishes associated with spawning salmon. Trans. Amer. Fish. Soc. 96:62-67. Reimers, N. '1957. Some aspects of the relation between stream foods and trout survival. Calif. Fish Game 43:43-69. Provides data on the percentage composit~on of the major benthic fauna groups, organisms in the drift, and numbers and volume of food consumed by brown trout in Convict Creek, California. The speed and effectiveness of trout digestion at various water temperatures are also investigated. In addition, .the effects of prolonged starvation on the. condition and viability of trout in different seasons are examined. . Reimers, P.E. 1973. ,The length of residence of juvenile fall chinook salmon in Sixes River, Oregon. Ore. Fish Comm. Res. Rep. 4(2) :1-43. Reimers, P.E., and RoE. Loeffel. 1967. The length of residence of juvenile fall chinook salmon in selected Columbia River tributaries. Fish Comm. Ore. Res. Briefs 13(1):5-19. Ringstad, N.R. 1974. Food competition between freshwater sculpins (genus Cottus) and juvenile coho salmon (Oncorhynchus kisutch): experimental and ecological study in a British Columbia coastal stream. Can. Fish. Mar. Servo Tech. Rep. 457. 88 pp. An Ruggles, C.P. 1959. Salmon populations and bottom fauna in the Wenatchee River, Washington. Trans. Amer. Fish. Soc. 88:186-190. Ruggles, C.P. 1966. Depth and velocity as a factor in stream rearing and production of juvenile coho salmon. Can. Fish Cult. 38:37-53. Salo, E.O.,and W.H. Bayliff. 1958. Artificial and natural production of silver salmon, Oncorhynchus kisutch, at Minter Creek, Washington. Wash. Dept. Fish. Res. Bull. 4. 76 pp. + appendix. A comprehensive long-term study of wild and hatchery coho salmon populations from 1938 to 1953. Magnitude and' timing of upstream adult and jack migrations are investigated. Numbers, lengths, and timing of both hatchery and wild smolt migrations are studied. 53 ------- For the hatchery-reared fish, fecundity, freshwater survival, marine survival, and total survival are evaluated for various rearing period durations. For wild fish, fecundity and survival are also . investigated. Comparisons of these data, as well as contributions to the commercial fishery, are made. In addition, estimates for optimum wild escapement' are presented. Salt, G.W. 1969. The role of laboratory experimentation in ecological . research. pp. 87-100 in J. Cairns, Jr. (ed.), The structure and function of fresh~water microbial communities. Virginia Polytechnic Inst. State Univ., Res. Div. Monograph 3. . Saunders, J.N., and M.W. Smith. 1~62. Physical alterat~on of stream , habitat to improve brook trout production. Trans. Amer. Fish Soc. 91:185-188. " A study of initial effects of habitat development on the brook trout population in one ,section of Hays Brook, Prince Edward Islando Data are presented on numbers of trout from 1947 through 1959 before development and in 1960 after developmento Artificial changes to the stream include 13 dams, 12 deflectors, and several covers in a 450-yd (4ll-m) study section. Changes in survival and. growth are also examined. Saunders, J.W., and M.W. Smith. 1965. Changes in a stream population of trout assoCiated with increased silt. J. Fish. Res.' Bd. Can. 22:395-404. Saunders, RoL., and J.H. Gee. 1964. Movements of young Atlantic salmon in a small stream. J. Fish. Res. Bd. Can. 21:27-36. ". Scarnecchia, DoL. 1978. Factors affecting coho salmon production in Oregon. M.S. thesis. Oregon State Univ., Corvallis. 100 pp. Schaffer, W.M., and P.F. Elson. 1975. The adaptive significance of variations in life history among local populations of Atlantic salmon in North America. Ecology 56:577-590. Schlucter, M.D., and J.A. Lichatowich. 1977. Juvenile life histories of Rogue River spring chinook salmon Oncorhynchus tshawytscha (Walbaum), as determined by scale analysis. Ore. Dept. Fish Wildl. Info. Rep. Fish. 77-5. 24 pp. Schuck, H.A. 1943. Survival, population density, growth, and movement of the wild brown trout in Crystal Creek. Trans. Amer. Fish. Soc. 73:209-230. Seegrist, D.W., and R~ Gard. 1972~ Effects of floods on trout in Sagehen Creek, California. Trans. Amer. Fisho Soc. 101:478-482. Shapovalov, L., and A.C. Taft 0 1954. The life histories of the steelhead rainbow trout (Salmo gairdneri gairdneri) and:silver salmon 54 ------- (Oncorhynchus'kisutch) with special'reference to Waddell Creek,' California, and recommendations regarding their management. Calif. Dept. Fish Game Bull. No. 98. 3?5 pp'. A comprehensive treatment of spawning migration, egg production, freshwater life, smolt migration, marine life, and survival of both species in this stream" with data from 1933 to 1944. Predators in both fresh and sal t water are als'o examined.. A review of other studies within each'topic is included as well. ' Shetter, D.S. 1961. Survival of brook trout from egg to ,fingerling stage in two Michigan trout 'streams.' Trans. Amer. Fish. Soc. 90: 252-258. ' Brook,trout populations were examined in Hunt Creek from 1943 through 1948 and in the Pigeon River from +949 through 1958. Number of spawners, fecundity, egg deposition, number of fingerlings, and survival rates are investigated. Variability in these parameters is shown through data for the entire period. Shetter, D.So, and A.So Hazzard. 1938. Species compositon and stability of fish populations in sections of three trout streams during the summer of 1937. Trans. Amer. 28l-~02. ' ' , by age groups Michigan Fish Soc. 68: , , One of the earliest studies to quantitatively show spatial variability between sections of the' same stream for brook, brown, and rainbow trout in 1935. A total of eight sections in three streams were examined for physical 'characteristics~ relative amounts of shade, trout biomass, and age group composition. Skeesick, DoG. ,1970. The fall immigration of juvenile coho salmon into a small tributary. are. Fish Comm. Res. Rep. 2(1):90-95. Slack, H.D. 1934. The winter food of brown trout (Salmo trutta L.). J. Anim. Ecol. 3 :105-108. Smith, M.W., and J.W. Saunders. 1958. Movements of brook trout, Salvelinus fontinalis (Mitchill), between and within fresh and salt water. J. Fisho Res. Bd. Can. 15:1403-1449. Smoker, W.A. 1953. Stream flow and silver salmon production in western Washington. \'lash. Dept. Fish., Fish. Res. Pap. 1(1) :5-12. Smoker, W.A. 1955. Effects of streamflow on silver salmon production in western Washington. Ph.D. thesis. Univ. Washington, Seattle. 198 pp. Solomon, D.J., and R.G. Templeton. trutta L. in a chalk stream. 1976. Movements of brown trout Salmo J. Fish BioI. 9:411-423. 55 ------- Stauffer, T.M. 1972. Age, growth, 'and downstream migration of juvenile rainbow trout in a Lake Michigan tributary. Trans. Amer. Fish. Soc. 101:18-28. ' Stein, R.A., P.E. Reimers, and J.D. Hall. juvenile coho (Oncorhynchus kisutch) ,tshaWytscha) in Sixes River, Oregon. 1737-1748. 1972. Social interaction between and fall chinook salmon (0. J. Fish. Res. Bd. Can. 29: Stewart, P.A. stream. 1970. Physical factors influencing trout density in a small Ph.D. thesis. Colorado State Univ., Ft. Collins. 78 pp. An investigation of physical variables affecting the density of brook and rainbow trout in a small Colorado stream. These variables, include mean depth, underwater overhanging rock cover, undercut banks, streamflow, and turbulence. Multiple regression analysis is, performed to determine which factors are statistically important to the density of each species. ' Straskraba,M. 1966. On the distribution of the macrofauna and fish in two streams, Lucina and Moravka. Arch. Hydrobiol. 61:515-536. Sumner, F.H. 1952. Migrations of salmonids in Sand Creek, Oregon. Amer. Fish. Soc. 82:139-150. Trans. Surber, E.W. 1951. Bottom fauna and temperature conditions in relation to trout management in St. Mary's River, Augusta County, Virginia. Virginia J. Sci. 2:190-202. Swanson, F.J., and G.W. Lienkaemper. 1978. Physical consequences of large organic debris in Pacific Northwest streams. USDA Forest Servo Gen. Tech. Rep. PNW-69. 12 pp. Symons, P.E.K. 1971. Behavioral adjustment of population density to available food by juvenile Atlantic salmon Salmo salar. J. Anim. Ecol. 40:569-587. Symons, P.E.K. 1974. Territorial behavior of juvenile Atlantic salmon reduces predation by brook trout. Can. J. Zool. 52:677-679. Symons, P.E.K., and M. Heland. 1978. Stream habitats and behavioral interactions of underyearling and yearling Atlantic salmon (Salmo salar). J. Fish. Res. Bd. Can. 35:175-183. Tagmaz'yan, Z.1. 1971. Relationship between the density of the downstream migr~tion and predation of young pink salmon [Oncorhynchus gorbuscha (Walb.)]. J. 1chthyol. 11:984-987. Tarzwell, C.M. 1938. An evaluation of the methods and results of stream improvement in the Southwest. Trans. N. Amer; Wildl. Conf. 3: 339-364. 56 ------- Tebo, L.B., Jr. 1957. Amer. For. Proc. Effects of sil tation on trout streams'. 1956:198-202. Soc. Tebo, L.B., Jr., and 1'1.1'1. Hassler. 1963. ,Food of brook, brown, and rainbow trout from streams in western North'Carolina. J. Elisha Mitchell Soc. 79:44-53. Thomas, J.D. 1962. The food and growth of brown trout (Salmo 'trutta , L.) and its feeding relationships with the salmon parr (Salmo salar L~) and the eel (Anguilla anguilla L.)in the river Teify, West Wales. J. Anim. Ecol. 31:175-205. ' Thomas, J.D. 1964. Studies on the growth of trout, Salmo trutta; from four contrasting habitats. Proc. Zool. Soc. London 142:459-509. Tomasson, T. 1978. Age and growth of cutthroat trout, Salmo clarki clarki ,(Richardson), in the Rogue River, Oregon. M~hesis. Oregon State Univ., Corvallis. 75 pp. Triska, F.J., J.R.Sedell, and S.V. Gregory. 1980. The coniferous forest stream: physical, chemical and biological interactions through , 'space and time. In R.L. Edmonds (ed.), The natural behavior and response to stress of western coniferous forests. ' Dowden, Hutchinson; and ' Ross, Inc., Stroudsburg, Pa. (In press.) , Vincent" R.E., and \'I.H. Miller. 1969. Altitudinal distribution of brown trout and other fishes in a headwater tributary of the South Platte River, Colorado. Ecology 50:464-467. Warren, C.E. 1979. Toward classification and rationale for watershed management and stream protection. U.S. Environ. Proto Agency, EPA~ 600/3-79-059. 143 pp. ' Warren, C.E., J.H. Wales, G.E. Davis, and P. Doudoroff. 1964. Trout production in an experimental stream enriched with sucrose. J. Wildl. Mgmt. 28:617-660. Waters, T.F. 1969. Invertebrate drift--ecology and significance to stream fishes. pp. 121-134 in T.G. Northcote, (ed.), Symposium on salmon and trout in streams:- H.R. MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. Watts, R.L., G.L. Trembley, and G.W. Harvey. 1942. Brook trout in Kettle Creek and tributaries. Bull. Penn. Agr. Exp. Stat. No. 437. 41 pp. Welch, H.E., P.E.K. Symons, and D.W. Narver. 1977. Some effects of potato farming and forest clearcutting on New Brunswick streams. Can. Fish. Mar. Serve Tech. Rep. 745. 13 pp. . 57 ------- . .' W~sche,:T.A. 1973. Parametric determination of trout. Univ~ Wyoming. Water Resource Res. Ser. No. ~7. 102 pp. minimum streamflow for Inst., Water Resource , Wesche, T .A., 1976. Development and application of a trout cover rating system for IFN determinations. Pp. 224-234 in J.F. Orsborn and C.H. Allman (eds.), Instream flow needs: a-Symposium. Amer. Fish. Soc. ' White, H.C. ,1930. Some observations on the eastern brook trout (S. fontinalis) of Prince Edward Island. Trans. Amer. Fish. Soc.- 60:101-'108. White, R.J. 1975. Trout population responses to streamflow fluctuation . and habitat management in Big Roche-a-Cri Creek, Wisconsin. Verh. Int. Ver. Limnol. 19:2469-2477. White, R.J., and R.L. Hunt. 1969. Regularly occurring fluctuations in year-class strength of two brook trout populations. Trans. Wisc. Acad. Sci., Arts, and Letters. 57:135-153. Brook trout populations in Lawrence Creek and Big Roche-a-Cri Creek. were studied from 1953 through 1964. Fluctuations in population numbers of age 0 and I trout were observed in both streams, with peaks every other year. These cycles may be related to competition for food and/or space in the streams.' ' , Wickett, W.P. 1951. The coho salmon population of Nile Creek. Res. Bd. Can. Progr. Rep. (Pac.). 89:88-89. Fish. Wickett, W.P. 1958. Review of certain environmental factors affecting the production of pink and chum salmon. J. Fish. Res. Bd. Can. 15: 1103-11260 Streamflow is the primary factor investigated in relation to pink and chum salmon pr~quction in British Columbia streams. Effects of low flow on adult migration and egg deposition are discussed. Negative impacts of floods on populations are also examined. Effects of streamflow on incubation stages and the survival of deposited eggs are analyzed. Another factor discussed is spawner density, in relation to egg density and gravel permeability. Wickett, W.P. 1962. Environmental variability and reproduction potentials of pink salmon in British Columbia. Pp. 73-86 in N.J. Wilimovsky (ed.), Symposium on pink salmon; H.R. MacMillan Lectures in Fisheries. Univ. British Columbia, Vancouver. Willis, R.A. 1962. Gnat Creek weir studies. Ore. 71 pp. (Mimeo.) Final report, Fish Corom. A weir was set up on this Oregon tributary to the lower Columbia River. Counts of migrating coho and chinook salmon we~e made from 1955 58 ------- th~ough 1961. Timing of migration, adult-jack relationships" sex r~tios, fecundity, and freshwater survival of wild fish are examined. S~rvival of hatchery-released fish in this stream is also analyzed. ptel~minary data indicate that commercial gill-net catches in the l4w~~ Columbia River are related to discharge in Gnat Creek two years earlier. Wiseman, J'.S. 1951. A quantitative analysis of foods eaten' by eastern brook ,trout.' Wyoming Wild1. 15(10) :12-17.' Withler, I.L. 1966. Variability in 'life history characteristics of steelhead trout (Salmo gairdneri) along the Pacific coast of North America. J. Fish. Res; Bd. Can. 23:365-393. Wong, D.M. 1975. Aspects of the life history of 'the Paiute cutthroat trout, Salmo clarki seleniris Snyder, in North Fork Cottonwood Creek, Mono County, California, with notes on behaviour in a stream aquarium. M.A. thesis. California State Univ., Long Beach. 190 pp. Wyatt, B. 1959. Observations on the movements and reproduction of the Cascade form of cutthroat ,trout. M.S. thesis. Oregon State Univ., Corvallis. 60 pp. 59 ------- APPENDIX This appendix is a compilation of examples of the best data available on temporal and spatial variation in populations of stream salmonids . (reprinted here with permission of the copyright owners and publishers). The tables are arranged geographically--north to south, west to east. A dash in lieu of data indicates "not sampled." The tables are reprinted with permission of the following organizations and individuals: American Fisheries Society.: Tables A-9, A-11, A-13, A-22, A-23, A-28, A-31. Blackwell Scientific Publications: Table A-33. California Department of Fish and Game: Tables A-IS, A-19, A-20. Scientific Information and Publication Branch, Canada Department of Fisheries and Oceans: Tables A-3, A-29, A-30. Pacific Biological Station, Canada Department of Fisheries and Oceans, Nanaimo, British Columbia: Table A-4. . . Research and Resource Services, Canada Department of Fisheries and Oceans, St. Johns ,Newfoundland: Table A- 32. The Fisheries Society of the British Isles: Tables A-34, A-35, A-36. Fisheries Research Division, New Zealand ~Iinistry of Agricul ture and Fisheries: Table A-37. Oregon United Table A-2. Institute of Animal Resource Ecology, University of British Columbia: Table A-I. Department of Fish and Wildlife: Table A-7. '~ I States Department of Commerce, National Marine Fisheries Service: Washington State Department of Fisheries: Table A-6. The Wildlife Society: Tables A-2l, A-24. Wisconsin Academy of Science, Arts, and Letters: Table A-25. Wisconsin Department of Natural Resources: Dr. David Au: Tables A-9, A-II, A-13. Dr. Richard Gard: Tables A-16, A-17, A-18. Dr. Gordon G1ova: Table A~5. Tables A-26, A-27. Mr. Gerald Lowry: Table A-14. 60 ------- TABLE A-I. POTENTIAL EGG DEPO~:iTIONAND FRESHWATER SURVIVAL OF PINK SALMON, SASHIN CREEK, ALAS ,A, 1940-1959 (FROM MERRELL 1962). ,I' .,'. , !..,/ Brood Potential Egg .. Number of . Freshwater I' .,:. Year .Depositi.ona Mi grating Fry Survival (%) 1940 52,858,000 3,402~830 6.4 1941 88,678,000 1,024,364 1.2 1942 81,502,000 674,672 0.8 1943 14,980,000 227.,673 1.5 1944 3,904,000' 104,113 2.7 1945 5,062,000 41,900 0.8 1946 736,000 1, 168 0.2 1947 .1,330,000 26,454 2.0 1948 516,000 9,016 1.7 1949 4,800,000 176,025 3.7 1950 86,000 (50 killed) 0.1 . 1951 4,062,000 379,585 9.3 1952 run destroyed 0 1953 1, 284,000 90,219 7.0 1954 12,000 576 4.8 1955 10,286,000 1,232,872 12.2 1956 1,018,000 5,043 0.5 1957 2,587,758 588,976 22.8 1958 174,OQO 10,577 6.1 1959 40,379,327 5,332,468 13.2 a Based on 2,000 eggs per female except when actual fecundity was calculated in 1957 (1,986 eggs) and 1959 (2,021 eggs). 61 ------- T~BLE A-2. WEIR COUNTS OF COHO SALMON FRY AND SMOLTS, SASH IN CREEK, ALASKA, 1956-1968 (FROM CRONE AND BOND 1976).. . . . Year Total Count Fry Smolts t956 1957 1958 1959 1960 1961 1962 1963 1964b 1965b . 1967 1968 373 .2 ,854 218 9,923 2,699 1,209 1,236 44,023 12,000 10,000 1,665' 928 1,961 1)015 1,587 1,258 2,489 2,865 1,599a 334 1,400 1,440 . a Partial count. b Weir not functional. damaged in 1966 - no Counts are estimates "from fyke net sampl ing. Heir sampling conducted. . TABLE A-3. HEIR COUNTS OF DOWNSTREAM rUGRATING PINK AND CHUM SAU10N FRY, HOOKNOSE CREEK, BRITISH COLUMBIA, 1947-1956 (FROM HUNTER 1959). Brood Year Pink Chum Total , , 1947 33,349 108,746 142,095 1948 64,312 77 ,539 141,851 1949 54,061 44,463 98,524 1950 234,396 431,399 665,795 1951 242,993 .269,701 512,694 1952 1,227,025 182,200 1,409,225 1953 204,250 984,504 1,188,754 1954 907,458 353,761 1,261,219 1955 86,256 49,443 135,699 1956 454,148 69,830 523,978 62 ------- TIIBLE 11-4. BIOMIISS (~/i IN LATE SUI~MER) OF COIIO SilL MOil IINO RIIINBOH IItID CUTTIIROIIT TROUT III smEM1S III TliE VICINITY OF TIlE CIIIWIITrON CREEK WIlTERSII£[) , BRITISII COllJMB III , 1970-1977 (FROM NIIRVER /\NO IItlDmSON 1974; IINDERSOtl IIND NIIRVER 1975; IIND IlII0ERSON 1978). . . . ..- -.---.-------- "---_____.0____._.-- ---_.- ----.--...-- - - . ---.-- -.-- --'-'--.----"--,--" -_._-..... ---------.-.------------ -----------.----------- ------_...----- -.--.-..., , ---.-- ---"-.---.-----.-----..--.-----.-- .-. ...._---.-- ---- ------'----'-- - -. ---- ...---- 0.__..---.-- .-.-.-- .-- . ---.. .--.-- .---.---.-- .-- -_.- - --..- '4'__--"--"-.'---' - . .. ...- ---- .....- Year Lower Carnation Cr. Coho --ii" iiliiow Upper Carnation Cr. '--Cutthr-oa}--- Trib "C" CuTthroaT Trib "1600" colio"ClifiJiroai: Useless Cr. COlIn -cilTIII-I'oaT Ft"eded ck (I". . Colw" AiiTrlbow RiLherdon Cr. C"1,tTI1'rc.iiil . S. Pachena Cr. Colio- - fiii i ilho~i .-.. ..n__-.,-._-_.... _.~ ---- -'"~'---'-'_.--'-- - -.. .------...--- _.. -_._-~ --.- -.-.-.- - -.-". .~----._.__.. -- -_'____h_--____--.-.-.-.-.. .--.-..---- - -.- -- .--"'-'--.---,.-- ..- n - . _.. - -. -..-.- 1970 2.72 1. 21 1971 \. 89 0.92 3.46 1.06 0.0 5.36 1.,87 1.24 1972 1. 47 0.43 2.90 4.79 4.88 \. 93 0.84 1.f>4 1. 54 0.33 4.50 1. 31 1.11 1973 1.46 0.59 3.97 5.61\ 2.45 1. 97 0.19 1.66 3.07 0.74 0.4<' 1974 \. 59 0.49 \. 94 3.45 2.84 0.95 0.2B 2,39 0.44 0.01\ 3.31 0.59 O. HI 0\ VI 1975 \.61\ 0.44 1.66 3.71 4.19 0.39 0,70 2.01 1. 36 0.07 2.84 \.85 .0.23 1976 1. 23 0.30 1. 94 2.77 2.75 0.25 0.38 0.67 0.66 0.02 1.1\0 0.82 0.17 1977 1.62 0.32 1. 08 2.47 0.53 0.85 0.95 0.09 1. 33 1.1\0 0.37 ----.----.---. .._---._----_..~.__._._--- -- ------..-.----.---------- .-----------.-.----------------.-.- ...:.. ---..- .---. ---.-. -----.---. -.. -.- .._"- -_._~-".-..__. '---'-'---'--.--- ._----------_._~. ----.-.- ..---..----.-- ---.-.-.--------- --.--- .--- ..--.-.-.- _._---- --..----..--. --.---.--. ..-....--.- ---..--.- -- --. -----...._-- ------- T/\BLE /\-5. BIOM/\SS (20 clII/sec). - b Mainly Cottus aleuticus. -...,.--.--. ---,- ------- TABLE A-6., ESCAPE~,1ENT, POTENTIAL EGG DEPOSITION, AND FRESHHATER SURVIVAL OF WILD COHO SALMON, MINTER CREEK, WASHINGTON, 1938-1953 (FROM SALO AND BAYLIFF 1958) . Brood Females Released Egg Freshwater Year Upstream Potential Smolt Count Survival (%) 1938 967 2,657,316 35,452 1.33 1940 1,393 4,577,398 32,085 , 0.70 1942 786 ~,873,038 31,893 1. 70 1943 906 2,092,860 23,177 1.11 1944 500 1,376,500 30,408 2.21 1946 500 '1,097,000 41 ,848 3.81 1948 98 186,200. 17,839 . 9.58 1949 114 287,964 27,.781 9;65 . 1951 411 1,086,684 22,545 2.07 1952 753 . 1,929,186 31,363 L63 1953 491 1,150,413 18,620 . 1.62 TABLE A'-7.. COUNTS OF SPAWNI.NG COHO SALMON AND SMOLTS AT DOWNSTREAM I~EIR ON GNAT CREEK, OREGON, 1954-1959 (FROM WILLIS 1962). . Brood Year Female Spawners Smolt Count 1955 26 2,996 1956 29 1 ,847 1957 67 . 1,013 1958 40 1,061 1959 45 3,226 65 ------- TABLE A-8. ESCAPEMENT, POTENTIAL EGG DEPOSITION, AND FRESHWATER SURVIVAL OF COHO SALMON. DEER CREEK, OREGON, 1959~1971 (FROM KNIGHT 1980). Brood. Female Egg Smolt . Freshwater Year Escapement Potentiala Count Survival (%) 1959 21 43,197 ' 1,917 4.44 1960 19 44,156 2,210 5.00 1961 28 67,620 2,775 4.10 1962 18 ,'42,030 2,082 4.95 1963 27 62,964, 2,368 3.76 1964 44 104,940 1,836 1. 75 1965 24 55,176 2,245 4.07 196,6 56 141,798 2,461 1. 74 1967 23 52,815 2,160 4.09 .1968 39 . 80,301 1,484 1.85 1969 8 15,484 738 ' 4.77 1970 10 22,119 1,072 4.85 1971 36 73,134 1,923 2.63 a Calculated from reg~ession equation (Koski 1966), Y = -3,184 + 7.81 X, where X = average length in mm (from unpublished data) and Y = individual fecundity. Total fecundity equals Y times the number of female spawners. TABLE A-9. ' ESTIMATED BIOMASS (g/m2) OF, JUVENILE COHO SALMON, DEER CREEK, OREGON, 1959-1968 (FROM CHAPMAN 1965 AND AU 1972). DATA ARE INTERPOLATED FOR THE BEGINNING OF EACH MONTH INDICATED, FROM POPULATION ESTIMATES MADE LESS FREQUENTLY THROUGHOUT THE YEAR. 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968 June 4.8 2.7 2.1 5.9' 4.2 2.1 4.8 5.0 8.7 2.3 July 4.0 2.5 2.5 3.6 3.7 2.1 5.9 4.3 7.2 3.0 Aug 3.1 2.3 3.0 3.0 3.4 2.1 4.3 3.8 6.2 3.7 Sept 2.9 2.3 3.2 4.0 3.1 2.6 3.2 3.6 5.9 4.4 Oct 3.1 2.5 3.4 4.7 2.5 3.1 2.9 3.8 6.1 5.0 . Nov 3.6 2.7 3.4 5.1 2.4 3.5 2.8 4.0 6.3 5.3 Dec 3.6 2.0 3.2 4.0 2.4 3.7 2.7 4.1 6.1 5.1 Jan 3.8 2.0 3.8 4.5 2.5 3.6 3.0 4.2 4.4 4.1 Feb 4.0 2.0 1.7 4.5 2.5 2.8 2.9 4.2 3..5 2.9 Mar 4.2 2.0' 1.7 3.6 2.4 1.9 2.4 3.7 3.2 2.2 Apr 1.8 1.3 1.3 2.1 1.6 1.5 0.8 1.9 1.9 1.4 May 0.5 0.4 0.6 0.5 66 ------- TABLE A-10. ESCAPEMENT, POTENTIAL EGG DEPOSITION, AND FRESHWATER SURVIVAL OF COHO SALMON, FLYNN CREEK, OREGON, 1959-1971 (FROM KNIGHT 1980). Brood Female Egg Smolt Freshwater Year Escapement Potentiala Count Survival (%) 1959 8 17 ,368 875 5.04 1960 26 66,742 776 1.16 1961 51 131,427 " 1,354 1.03 1962 2 4,644 565 12.17 1963 20 44,220 736 1.66 1964 . 10 . 24,020 663 2.76 . 1965. 11 26,565 968 3.64 1966 55 138,050 616 . 0.45 1967 10 23,130 430 1.86 1968 19 38,931 207 0.53 1969 5 9,625 140' 1.45 1970 "5 13,745 330 2.40. 1971 18 37,404 404 1.08 a Calculated from regression equation (Koski 1966), Y = -3,184 + 7.81 X, where X = average length in mm (from unpublished data) and Y = average individual fecundity. Total fecundity equalsY times the number of female spawners. TABLE A-II. ESTIMATED BIOMASS (g/m2) OF JUVENILE. COHO SALMON, FLYNN CREEK, OREGON, 1959-1968 (FROM CHAPMAN 1965 AND AU 1972). DATA ARE INTERPOLATED FOR THE BEGINNING OF EACH MONTH INDICATED, FROM POPULATION ESTIMATES MADE LESS FREQUENTLY THROUGHOUT THE~YEAR. 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968 June 4.1 2.9 2.1 8.3 1.3 2.2 4.1 4.0 6.0 1.1 July 3.3 2.8 2.0 3.0 1.7 1.7 1.7 2.8 3.9 1.3 Aug 3.0 2.7 2.1 2.3 2.0 1.5 1.4 2.6 2.9 1.8 Sept 3.1 2.6 2.3 2.6 2.2 1.7 1.6 2.9 2.1 2.3 Oct 2.9 2.5 2.5 3.4 2.4 1.8 1.7 2.8 1.8 2.5 Nov 2.9 2.5 2.5 3.8 '2.2 2.0 1.7 2.6 1.7 2.3 Dec 2.4 2.0 3.5 4.5 2.0 1.9 1.7 2.4 1.7 1.8 Jan 2.2 1.9 2.1 4.1 1.8 1.9 1.7 2.5 1.8 1.1 Feb 2.2 1. 7- 2.0 4.1 1.6 1.7 1.6 2.5 1.7 0.9 Mar 2.6 1.8 1.8 3.8 1.4 1.7 1.1 2.1 1.6 . 0.8 Apr 1.5 1.7 1.6 2.6 1.1 1.3 0.4 0.9 1.4 0.5 May 0.5 1.3 0.7 1.4 67 ------- TABLE A-12. ESCAPEMENT, POTENTIAL EGG DEPOS ITION, AND FRESHWATER SURVIVAL OF COHO SALMON, NEEDLE BRANCH, OREGON, .1959-1971 (FROM KNIGHT 1980). Brood Female Egg a Smolt Freshwater Year Escapement Potential Count Survival. (%) 1959 2b 4,471 b 462 10.3 1960 2 4,192 . 223 5.32 1961 15 33,135 470 1.42 1962 4 . 9,632d 314 3.26 1963 15c 33,530d 160 0.477 1964 25c 55,884d 286 0.512 1965 28c 62,590 333 0.532 1966 19 46,664 277 0~594 1967 15 40,460 421 1.04 1968 17 35,088 194 0.55 1969 1 2,666 76 2.85 1970 2 5,386 113 2.10 1971 18 35,604 369 1.04 " a Calculated from regression equation (Koski 1966)~ Y = -3,184 + 7.81 X,. . where X = average length in mm (from unpublished data) and Y = average individual fecundity. Total fecundity equals Y times the number of female sp~wners. . . b Estimated equivalents from 1,627 planted fry. c Estimated from redd surveys. d Estimated from mean female length (693.9 mm) from the other years of the study. ~ 68 ------- TABLE A-13.. ESTIMATED BIOMASS (g/m2) OF .JUVENILE COHO SALMON, NEEDLE BRANCH, OREGON, 1959-1968 (FROM CHAPMAN 1965 AND AU 1972). DATA ARE INTERPOLATED FOR THE BEGINNING .OF EACH MONTH INDICATED, FROM POPULATION ESTIMATES MADE LESS FREQUENTLY THROUGHOUT THE YEAR. 1959 1960 1961 1962 1963 1964 1965 1966 1967 1968 June 2.3 1.4 2.1 7.2 3.5 4.0 5.0 7.6 6.9 3.1 July 1.6 1.7 1.9 3.9 3.0 2.4 3.9 4.0 9.0 3.4 Aug 1.8 2.0 1.8 3..2 .2.9 1.5 3.5 3.1 7.8 3.8 Sept 1.9 2.0 1.8 2.8 2.7 1.3 3.4 3.7 6.5 4.2 Oet 2.2 2.2 1.7 2.8 2.8 1.2 3.3 '4.3 6.2 4..6 Nov 2.4 2.4. 1.7 2.9 2.7 1.0 2.9 4.1 3.2 4.4 Dee 2.0 2.2 1.8 2.8 2.6 0.7 2.2 4.4 3.1. 3.7 Jan 1.9 2.2 1.8 3.3 2.3 0.8 1.9 4.0 3.2 2.4 Feb 1.9 2.4 1.7 3.1 2.3 0.9 1.3 3.1 3.9 1.7 Mar 2.4 1.5 1.4 3.1 1.5 1.0 0.8 1.5 2.6 0.4 Apr 2.4' 1.1 1.8 2.1. 1.0 0.8 0.4 0.4 0.2 0.1 May . 0.9 0.3 0.7 0.9 TABLE A-14. BIOMASS (g/m2 IN SEPTEMBER) OF CUTTHROAT TROUT, ALSEA l,/ATERSHED . STUDY, 1962-1973 (~ROM LOWRY 1964 AND UNPUBLISHED DATA). .. Year Deer Cr. Flynn Cr. Needle Br. 1962 5.07 . 5.82. 3.89 1963 .2.93 3.54 3.41 1964 . 1.90 4.04 3.16 1965 2.93 2.72 2.97 1966 . 2.05 2.73 1.09 .1967 3.29 4.26 0.68 1968 2.15 2.71 1.65 1969 2.80 3.70 1.46 1970 3.83 4.01 1.14 1971 4.20 4.27 1.32 1972 4.03 4.13 1.39 1973 3.79 1.53 69 ------- TABLE A-15. BIOMASS (g/m2) OF SALt10NID SPECIES IN THREE NORTHERN CALIFORNIA STREAMS; 1967-1969 (FROM BURNS 1971). . No. Fork, Casper Cr. Coho Steel head S. Fork Yager Cr. Steel head Godwood Cr. a Coho Trout June 1967 0.18 1.09 July 1967 Aug 1967 Oct 1967 0.15 1.46 June 1968 0.13 1.16 July 1968 Aug 1968 Oct 1968 0.19 1.44 June 1969 0.61 0.98 . Jul y 1969. Aug 1969 Oct 1969 0.81 1.13 a 1 Stee head and cutthroat. 1.09 0.57 3.22 0.76 0.49 4.21 0.34 0.51 2.94 TABLE A-16. BIOMASS OF BROOK TROUT (g/m2 IN MID-AUGUST) IN 10 SECTIONS OF SAGEHEN CREEK, CALIFORNIA, 1952-1961 (FROM R. GARD PERS. COMM.). SECTION I IS UPSTREAM. I I! II! IV V VI VII VIII IX X 1952 5.15 13.90 3.75 1.48 0.39 0.80 2.01 1.06 0.24 0.01 1953 4.71 14.24 4.74 1. 38 0.50 3.25 1.46 0.67 0.07 0 1954 4.85 10.12 2.17 1.28 0 1.29 1.01 1.60 0 0 1955 4.48 6.87 2.45 0.41 0.01 0.74 0.06 1.31 0 0 1956 2.47 6.65 1.80 0.96 0.18 0.55 0.41 0.13 0.01 0 1957 4.56 3.67 1.88 1.49 0.25 0.80 0.18 0.19 0 0 1958 2.24 2.91 1.23 0.25 0.53 1.14 0.96 1.84 0 0 1959 7.40 2.31 0.85 0.35 1. 78 0.24 0.19 0.01 0 1960 4.63 2.32 1.95 0.19 0.54 1.64 0.86 0.75 0 0 1961 3.36 2.50 0.86 1.12 0.55 2.85 1.56 0.40 0 0 70 ------- I TABLE A-17. BIOMASS OF BRO~1N TROUT (g/m2 IN MID-AUGUST) IN' 10 SE~TIOt-lS OF ' SAGEHEN CREEK, CALIFORNIA, 1952-1961 (FROM R. GARD PERS. COMM.). SECTION I IS UPSTREAM. I II' III .IV V VI VII VIII IX X 1952 0' 0' 0 0 0 0 0.63 ' 8.56 1.99 0.40 1953 0 0 0 0 0 0.08 6.34 1.18 1.18 0.78' ,1954 0 0 0 0, 0 0 1.69 '1.47 2.03 0.67 1955. 0 0 0 0 0 0.01 3.96 1.66 4.18 0.45 1956 0 0 0 0 a 0 1.48 1.06 2.54 0.11 1957 0 0 0 0 0 0.44 0.91 0 2.95 0.01 1958 0 0 0.54 0 0 0.08 0 0.13' 4.11 0.16 1959 0 0 0 0 0.02 1.30 1. 97 2.82 0 ,1960 0 0 0 0 0.06 0 1.30 0.37 2.93 0 1961 0 0 0 0 0 0.02 1. 51 0.16 1. 74 0.01 TABLE A-18. . BIOMASS OF RAINBOW TROUT (g/m2 IN MID-'AUGUST) IN 10 SECTIONS OF . SAGEHEN CREEK, CALIFORNIA, 1952-1961 (FROM R. GARD PERS. COMM.). SECTION I IS UPSTREAM'. I II III IV V VI ' VII VIII IX X 1952 0 1.12 1.45 4.69 0.75 .1. 01 0.99 0.54 0.21 0.07 1953 0 1.27 4.30 4.78 0.77 1.20 ,1.46 0.99 0.28 0.03 1954 0 1.42 2.31 3.89 0.37 1. 73 2.42 0.45 0.01 (j 1955 0 1.04 2.82 4.07 0.31 2.26 1.43 0.77 0 0.12 1956 0 0.41 1. 30 2.21 0.45 1. 75 0.41 0.06 0 0 1957 0 1.09 1.23 3.45 0.44 2.93 1.82 0.64 0.15 0 1958 0 1.12 0.91 2.35 1.43 3.55 1. 79 0.75 0.59 0.07 1959' 1.04 0.54 4.02 0.37 5.30 2.50 0.91 0 0 1960 0 0.37 0.83 4.60, 0.83 4.29 1.54 0.11 0.49 0 1961 0 0.41 0.36 5.91 0.89 6.67 1.64 0.96 0 0 71 ------- ikd TABLE A-19. ESCAPEMENT, POTENTIAL EGG DEPOSITION, AND FRESHWATER SURVIVAL OF COHO SALMON, WADDELL CREEK, CALIFORNIA, 1933-1940 (FROM SHAPOVALOV AMD TAFT 1954) Brood Year 1933 1934 1935 1936 1937 1938 1939 1940 Femal e Escapement 222 309 59 157 37 56 150 115 Egg Potential 560,690 725,014 141,233 377,352 91 ,728 130,074 396,321 257,886 Smolt Count 3,573 4,911 1,067 1,926 852 1,740 152 711 Freshwater Survival (%) 0.64 0.68 0.76 0.51 0.93 1.34 0.038 0.28 TABLE A-20. DOWNSTREAM TRAP COUNTS OF STEELHEAD TROUT BY AGE GROUP, WADDELL CREEK, CALIFORNIA, 1933-1942 (FROM SHAPOVALOV AND TAFT 1954). 0 I II III Year Number Percent Number Percent Number Percent Number Percent 1933-34 1934-35 1935-36 1936-37 -1937-38 1938-39 1939-40 1940-41 1941-42 604 699 1,365 1,875 1,946 691 2,239 3,306 2,009 19 39 35 53 57 11 64 59 35 741 578 1,655 1,191 1,015 3,699 945 2,049 . 2,834 24 32 42 34 30 60 27 36 50 1,657 484 830 451 410 1,720 . 292 251 843 53 27 21 13 12 28 8 4 15 112 28 90 11 19 77 7 9 33 4 2 2 a 1 1 a a 1 a < 1 percent. 72 ------- TIIOLE A-21. BIOHASS (g/m2) OF BROOK, RAINBOII, AND BROI-IN TROUT, TROllr CREEK, MONTAflA, 1950-1951 (FR0I1110LTON 1953). SECTION I IS UPSTREAM. Sec ti on 2 -----------.----- ------ --------------- -'--'--'----'------------'.-_n______...--- --.--..-.--.----.-----------.------.. 4 ---..-- -----.-.-. _._- . -----.--.,.- -'.'--'-'-----'----'--'-----'- -.----.+---- ----...------..-- -- ..-_.._--_.-..-. ._-. ---- "..-- ----- ,.--. ----------- --_.__..--- ---- -'-- - ---.- "-"--'----,-,----- -_._----~-- ..-.-......-- .'-__'_h. ---_.. Date Auy. Nov. I-lay Aug. 24 18 20 12 1950 1950 1951 1951 July Sept. flov. !Iay Aug.' 25 14 19 30 4 1950 1950 1950 1951 1951 _..----..-. --" ----------.-'-----"'--- ---- --... .'-.;---.-.-.---. _._---_._.~- Br'ook 13.0 34.9 12.4 10.B 3.B B.O 3.7 2.1 5.3 3.6, 6.5 5.0 6.6 11. 1 0.4 0.0 4.6 6.7 13.4 RainlJow 2.0 3.0 0.9 D.9 o Brown o o o ~---------- -...-.-----------.------------.---.----------.-.--------------.---.------------.---- 3 ---------- ------------.--.-..------- -_-4____..- '.'--- .. ..__.--_.~-..- Aug. Sept. !Iov. June I\ug. 9 14 4 3 11 1950 1950 1950 1951 1951 Aug. Oct. May Aug. 1 20 17' 5 1950 1950 1951 1951 ----- --------.-------- --------- . -------.----.-.- --'.-. --- --- 2.2 2.1 :3.4 1.3 2.1 4.9 5.0 5.0 6.9 5.7 0.5 0.5 10.6 0.1 0.3 2.1 1.4 1.1 0.2 3.9 6.4 3.0 6.2 0.6 9.7 4.1 0.7 .----.---..-.-----.--- ---.------.------------.------..----.---.----------------- --------- -.-- -~-_._--- ----.--_.- --- -- -.-------..-.-- -. ~n TABLE A-2? BIOMASS (y/l) OF BROHII, RAIUBOH. AND BROOK TI!OUT HI 11 SECTIONS or LITTLE PRICKLY PEAI{ CREEK; MONTANA, SUMMER 1966 (FIWH ELSER 1960). SECTION 1 IS UPSTREAM . See ti on 3 4<1 -------------.---------.---.------.-----.-..---- - ---.---------.-.--.---------- ---_._.~----_. --- --.-.-------.--. -.-- .---.--- 9 lOa ..... VI -.--.---..----------.--.-------------------------.-----------------.----.----..-----------..--.-------------------.-..---..- ..-- lA 2 5 6<1 Oa 7 --~--_."._"-------_._-_._._.._---------_._-,--_._---------------------.---------.------------------.--.-.----------.-----'-.-..---- 2 2308 1862 2266 29% 2630 3157 1255 3213 Area (III ) 1093 21'1S 17'10 Mean width (III) 5.5 2.7 5.5 7.3 7.6 8.2 B.O 7.3 6.'1 7.6 13.4 ,.Brown l. 57 3.14 12.9 16.7 3.36 17.6 4.37 17. 1 10.'1 11. l' 1. 57. Rainbow 4.70 5.0'1 5.60 5.72 0.90 6.39 2.02 7.85 1. 91 5.9'1 '1.04 Brook 1. 01 10.8 3.81 2.02 0.22 0.'15 0.11 0.3'1 0.67 0.22 0.11 Total trout 7.23 19.0 22.3 24.4 4.'18 2'1.'1 6.50 25.3 13.0 17.3 5.72 a Altered sections. .- ---.--.- ---.- - --.-- -.-.-------.- -.-----..- --~- ---.---- ----.---'--.-----.--.-.-.-----. .---' -------- -- -----..---.--------.-----.--. --..._~ --- --r---------...--- ----._------- ------.-.-.. --..-- ----.. --,---- _.~_._.- -- -~.. ..- .-.- .--....----- ---.--.-- ~ ---.-.--..---.- -.----.----..--.-- ---.._-----.~_.- -'-----'--------.-. .--.. --_u- ..----.------.--.-.--. ------- TABLE A-23. NUMBER OF CUTTIIROAT TROUT TRAPPED, AND IIUMBER REMAINING IN STREAM AFTER TRAP WAS REMOVED,a ARNICA CREEK, YELlOWSTOtIE PARK, WYOMING, 1950-1958 (FROM BENSON 1960), --~_._------ -------..-.------.-----..-----------.----------------------'-----'-------'-----.--'-----------.-----'- .._-----_._-------_._---------------------~-------------------.-.------------.----------.--------.---------.- , . Year. Oil te trap removed _~~9~~.!.!r.~!lJI- Q.---- Trapped In Stream .._.A9~.:.9EP.!!P.-l.__-- TI'apped, In Stream fi~ap~~a---9!'.9-f!H{ ream Tota 1 frappe-a---'n Stream Grand To ta 1 . --'-'-_._-----'--'--'-'--'-'----~--------------------'-------'---------'--'-----'-------'----.---------.-..---. 1950 Sept. 28 9,556 300 95 9,951 9,951 1951 Oct. 6 5,240 365 39 5,641 5,644 1952 Sept. 21 502 792 242 1.536 1,536 1953 Sept. 28 1,332 708 763 407 82 67 2,177 1,182 3,359 1954 Sept. 14 4,151 943 47 5,141 5,141 1955 Sept. 24 4,182 244 340 15 7 0 4,529 259 4,788 1956 Sept. 13 4,268 612 386 56 132 5 4,786 673 5,459 1957 Sept. 25 2,850 405 121 270 5 3 . 2,976 678 3,654 --..j 1958 Aug. 31 36 1,950 9 30 15 46 1,995 2,041 .j:>. --------------- ..--------.------......-..----.------------------------'--'----'-------------'--.-'---'-'--'-.--.---.--'--'---.--.----- . . ---'--.---------'------'--'-'--------------,---------,-----.".-'------'--"-------'--.-------,.-----,,- a Data not available on number of fish in stream after dismantling of trap for 1950, 1951, 1952, and 1954. ------- TABLE A-24. BIOMASS (g/m2) OF BROOK TROUT, LAWRENCE CREEK, \.:IISCONSIN, 1953- 1957 (FROM McFADDEN 1961b). SECTION A is UPSTREAM Section A B C.. D Total Mean. Sept. 1953 10.30 9.81 8.40 5.52 34.03 8.51 Sept. 1954 14.53 12.36 8.81 6.31 42.01 10.50 April 1955 11. 18 9.64 6.82 6.60 34.24 8.56 . Sept. 1955 7.11 8.25 6..10 3.02 24.48 6.12 April 1956 4.41 4.2"7 3.89 2.45 15.02 3.75 Sept. 1956 10.76. 6.01 5.50 1. 30 23.57 5.89 April 1957 14.48. 6.93 5.35 3.37 30.13 7.53 Sept. 1957 . 26.18 10.74 6.37 4.20 47.49 11.87 TABLE A-25. SEPTEMBER POPULATION ESTIMATES OF AGES 0 AND I BROOK TROUT IN LAWRENCE CREEK AND ~IG ROCHE-A-CRI CREEK, WiSCONSIN, 1953-1964 (FROM WHITE AND HUNT 1969). . . Lawrence Creek Year. 0 I 1953 10,113 2,040 1954 13,523 2,749 1955 5,720 . 2,754 1956 10,853 816 1957 13 ,258 3,370 1958 4,166 4,393 1959 22,646 1,044 1960 8,507 3,324 1961 14,313 2,360 1962 7,611 4,523 1963 10,367 2,388 1964 9,680 4,382 Big Roche-a-Cri Creek o ..1 2,012 6,229 2,637 9,915 4.,361 5,632 4,964 7,420 1,135 . 474. 1,817 1,257 2,630 1,609 1,623 1,072 75 ------- ll\(}l[ 11-26. BIOr1J\SS (g/n?)a OF BROOK TROUT BY IIGE GROUP III IIPRll 11110 SEPHMBER, llll/REtICE CHEEK, IIISCONSIN, 1960-1970 (FHOM ItUNT 1974).. . -.----. -.- -'-'------.-------.---..-- ----,------ '--___"0 --- ---- ----- .-- ------ --...-..--.-.-- --.--.-- -"---.,----, - .-..--.--. ----....----. ------- -..----....-.-.----..-- -"----'-'-"--' . .--..-.-...--.--... - --~-----_. -.__'__.0_._- ----'--- -- ------"----------".----'----- ---------"- -- -..---.. ---- -.. --'''-------'----'---.--. ---.-.---."---.---.. -.------... - ------- ----- -.-.."0_....- .-.---..-- Year .-.----..--.-.---- .____~I_i.l. ..---- ... .--..---....-.-....-..- .--- I II III IVt Total "---.--..-.-----.-.- .S_e.P. ~e.!!!.~E___.__---_._-- .:..----.-- ___h_- o [ II III IV+ Total -- - -------------...- ----------.-----------..---- --------'------'-'----- ---_.,-~---_.__._---.-_._._---_._._..._--_._-----_._----..-.-------. 1960 4.00 0.375 .0.542 0.0245 4.94 2.01 4.75 0.105 0.0417 0 6.91 1961 l. 97 1.48 0.0245 0.0294 3.50 3.83 4.39 0.922 0.0172 0 9.16 1962 5.48 2.30 0.412 0.0049 8.20 1. 70 6.0B 0.507 0.0930 0 8.38 1963 2.71 4.62 0.333 0.035B 7.75 2.80 3.39 1. 78 0.137 0.0392 11.15 1964 5.20 2.72 1.20 0.108 9.23 2.51 5.56 0.995 0.387 0.0319 9.48 1965 2.43 3.30 0.517 0.174 6.47 2.13 4.30 1.13 0.167 0.0294 . 7.81 1966 3.96 3.38 0.934 0.130 8.40 2.54 6.19. 1.64 0.299 0.0392 10.7 1967 3.15 5.65 1.04 0.427 10.3 2.14 4.03 2.02 0.332 0.0613 8.63 " 1968 3.35 4.43 l. 62 0.287 9.69 2.85 3.40 0.BG3 0.280 0.0686 7.46 0\ . 1969 5.25 3.75 0.699 0.194 9.B9 3.10 4.51 1. 02 0.177 0.0809 8.89 1970 3.88 5.17 0.701 0.142 9.89 3.32 3.78 0.9110 0.145 0;0466 8.27 r1ean 3.77 3.38 0.729 0.146 8.02 2.63 4.58 1. 09 U.194 0.0361 8.53 . --.- -.. '.-.----."--'---------'------"-'----'-'--'- .--------.------.----- --.-----..-.--.-------.-.-.---------.-----....-------..-----.-- ._- --- _. .------------------.------.-.----- ...- -.- .....__.-- -.---.-.- --------_.._---_. ------------.-- _._----- ....---.-- ---- --------.--.-.-.----.-. a Biomass f.'om IIppendix Table 1 has been divided by stream of 4.0fi ha. area ------- TIIRlE 11-27. ANNUlll pHODUCTI orl (9/nl /yr) OF OIWOK THOUT OY SECT ION Arm AGE GHOUl', lllHHENCE CHEEK, IH SCONS IN, 1960-1970 (FROM HUNT 1974). SECTION II IS UPSTREN1. " -----------------.---------.----------- --------------.-.---.---.. ---.---------.-----------.-----------.-.--.---. -_._----._-_._--------~----_.__._---_. --- " . -.----- Year Section" -A----Ir------c~n_____----o- ----_._------~~J1.r:.()~-~._- --.---..----. o I' II . III IVI Stream Total _._----~-------------._----------- .,-----_.._---------_._-_._---_.__.__._--_._-----_.~_...--'--- 1960 13.0 10.1 13.4 14.0 4.1 7.7 0.3 0.4 <0.1 12.5 1961 17.2 13.5 10.2 8.0 6.8 3.9 1.1 <0.1 0.1 11.9 1962 14.0 10.0 11. 1 IO.IJ 3.8 6.2 0.9 0.2 <0.1 1l.2 1963 16.5 12.0 12.9 11. 5 6.4 4.0 2.2 0.2 <0.1 12.9 1964 19.8 12.8 9.8 1J.6 5.2 5.2 1.2 0.5 <0. I 12.2 1965 19.5 9.8 11.0 5.4 4.3 4.2 1.9 0.2 <0.1 10.6 1966 15.2 12.6 9.8 6.3 3.1 5.7 1.4 0.4 <0.1 10.6. 1967 21.7 9.4 10.9 6.7 3.8 4.5 2.4 0.5 <0. I 11.2 -...j 19GIJ 21.3 12.0 1J.9 5.1 4.5 "4.0 1.9 0.6 O. I II. 1 -...j 1969 25.8 12.0 7.9 6.6 4.7 5.3 1.6 0.3 0.1 12.0 1970 20.5 13.2 ro.1 7.8 5.3 4.6 2.2 0.2 <,), 1 12.3 Mean 18.8 11.6 10.6 8.2 4.3 5.0 1.6 .0.3 <0. I 11. 7 -----------------------.---..--------.-------.-.---------.--.---.---.---- --------------------------.--......---.-------------..-. ------- TADLE A-28. PIIYSICAL CllIIRIlCTERIST/CS ANIIIIIOMASS (g/II?) OF BROOK, DROWN, AND RAINOOW TROUT III SECT/OtiS OF TlmEE MICIIIGAN STREAMS, 1937 (FROt1 SIiElTER AND IIAZZIIRD 1931:). -----.-.- "..-- -------.---------..----.-.--..--.---------.------.-----.--------. ----.--'------------.------------- -----.----.---.-----------------------..------ .--- ------ -~ -------.-.- -.-.---- - ---1"'----------'-' .---. .--. -'--'----.----.---.-...-----.. ----._--. Stream Section Length (m) Mean Hidth (m) Mean Depth (elll) Velocity (em/see) Re 1 a t i ve Shade Oiomass ifi:Ook'------ ifrown---lfiilnljpw-'--T 0 taT . -----_._~-_.._-----------~---_._----_._---_.- ---.---------------.-.----..--------..---.----- South 8raneh, Upper 29.5 7.6 24.9 22.9 Pa rt 1 y 0.95 0 0.35 1. 30 Pine River Middle 29.0 5.4 27.7 26.2 Densely. 0.87 0 0.53 1.40 Lower 31.5 6.7 34.8 20.7 Partly 2.40 U.27 1. 82 4.49 . Little Manistee Upper 32.5 8.5 43.7 4t.!! Partly 0.21 1. 92 1. 75 3.88 River Middle 46.9 11.2 44.2 Exposed 0.12 0.73 2.54 3.39 North IIraneh, Upper 42.6 7.5 25.7 94.5. Partly 0.028 0.84 0 0.87 Ooardman River Middle 29.0 8.5 29.0 46.3 Exposed 0.041 0.13 0 0.17 "-I Lower 37.0 9.1 23.1 51.5 Partly 0.048 0.10 0 0.15 'JO --.---.------------------------ -------- ---.--------.--------- .------,..--- ------- TABLE A-29. NUMBER OF BROOK TROUT PRESENT IN SEPTEMBER IN HUNT CREEK, MICHIGAN BY AGE~GROUP (FROM MCFADDEN ET AL. 1967). Year 0 I II III IV Tota 1 1949 4,471 2,036 287 14 0 " 6,808 1950 3,941 2,013 304 13 0 6,271 1951 4,287 1,851 265 16 1 6,820 1952 5,033 1,763 261 16 0 7;073 1953 5,387 1,637 175 13 0 7,212 1954 6,325 2,035 234 13 0 8,607 1955 4,235 2 ,325 383 " 24 0 6,947 1956" 4,949 1,612 392 51 1 7,005 1957 6,703 1,796 309 . 33 1 " 8,842 1958 5,097 2,653 355 26 2 8 , 133 1959 4,038 2,395 685 68 0 7,186 1960 5,057 2,217 473 47 1 7,795 1961 2,809 2,017 409 23 0 5,258 1962 . 5,052 1,589 448 52 2 7,143 79 ------- TABLE 11-30. MEAN ANNUAL BIOMASS (g/m2) OF BROOK THOUT IN STREAMS IN MATAMEK ~'ATERSItEO. QU[(JEC. 1971-1973 (FROM O'CONNOR 111m POWER 1976). -.-.- .---.- ------~. ---"------'------------------ -'-~'- _....--_....._----- --_.__._-~------_._---_.~...~,._-_._-_._._----- '-------'--'--"--'-'---.------'- --- -'---'----- _. ----------_. ----'.'--'-"-'.----..' ---....---.- ---. ---........---.------ -------.-.--.----..-..-------- ...---.--------- --_.-.----_._--,- ..+ Stream See ti 0/1 Length (m) Average Hidth (m) Year . Biomass u Dt"u --------l+--------~-----------T~------ - --- -;n--- ------5+------ ---6:j-- - ----To-taT Kaikhosru 6.1 -- -- ._----------- --.'-'-"---'.----' .,-- ---'-'-'--' --.--- -.._--- -----'--'- ----- ---- '-----------'--'--~- -.- ,...- -."-.. .'--'--'- ..-.. -- .--- --.--.. -------'------'--.---..-.-.-.'-- 355 Ga 11 i enne 330 6.6 Tehillie3man 620 15. 1 00 Q Sherry 215 4.2 1971 1972 1971 1972 1971 1977 1971 1972 1973 0.35 0.35 0.08 0.08 0.23 0.28 0.28 1. 93 0.33 0.54 0.05 0.17 0 4.17 2.23 0.66 0.44 0.22 0.06 0 3.96 2.40 2.12 0.57 0.14 0.10 0 5.33 2.83 1. 74 0.49 0.00 0.14 0 5.28 0.51 0.26 0.25 0.05 0.01 0.06 1. 21 0.42 0.48 0.32 0.07 0.01 0.03 1.42 1. 25 0.87 1.13 0.25 0 0 3.78 0.79 0.51 0.30 n.15 0 0 2.03 0.57 .0.2B 0.16 0.17 0 0 1. 4 7 .._-~---_._------_._---------------_._--------- -_.~-----_._._------_.._---_._.__._---_._---_._-- ----.-.---.-----.---------.------------..--------.------.-.-.---...--------------.------.--.------- --- ------- TABLE A-31. NUMBERS OF BROOK TROUT HI A 411-m SECTION OF HAYES BROOK, PRINCE Em~ARD ISLAND, 1947-1960 (FROM,SAUNDERS,.lXND Sf1ITH 1962). Year, Age 0 Age I + Total 1947 588 351 939 1948 729 342 1,071 1949 539 279 818 1950 321 223 544 1951 166 418 584 1952 611 372 983 1953 308 362 670 1954 468 294 762 1955 758 383 1,141 1956 580 467 1,047 1957 350 363 713 1958 4$1 314 795 .1959a 371 352 723 1960 526 611 1,137 a Afte; habitat,development. TABLE A.,.32. COUNTS OF ATLANTIC SAU10N SMOL TS AND SEAl-lARD MIGRATING BROOK TROUT, LITTLE CODROY RIVER, NE~/FOUNDLAND, 1954-1963 (FROM ~1URRAY 1968)., ' Year Sa 1 man Trout 1954 12,210 1955 11 ,248 1956 14,772 706 1957 8,900 1,067 1958 9 , 341 889 1959 12,099 1,074 1960 7,829 457 1961 8,058 312 1962 8,193 698 1963 7,326 485 81 ------- TABLE A-33. BIOI-lASS (g/m2) OF BROUN TROUT IN TRIBUTARIES AND TilE MAIN STEM OF TIlE UPPEH RIVER lEES SYSTEM, EIiGlAND, ]967-]970 (FROM CRISP [T Al. ]974). . "---'---'--------"'----------'- ..---- ----.--..-.--------.-.. "-'."-- --'----'.'----- - .---.- --- -- -------- ---._u_-.---.-----.---- -... ".. -..--.----.---.- -".'.------.--..-.--..-... --"------ - --------- --------"---_. .--..--.- -.---.-. ..-..------------ .----.- --.-.-. - - ...-----.- - ---- -_.._..~-_...__._--...._.._._. -.-- .-. .--- .-.- ..----.---------.. .-- -..-.--.-. -.- - -'. Mai ze Beck River Tees below Cauldron Snout River Tees above the Weel Wee1head SHe Dubby Sike Mattergi11 Sike lod!1eg i 11 Sike -.-------------- ---------------.---------.---.--------.----------.----"--- ----'--------.-'."--.'---------.-.------. --.--.- Section length (lit) 70.9 30.5 23,4 .97.5 40.8 47.2 45.7 Mean Width (Ill) 9.94 6.91 ]1.20 1. ]6 1. 54 3.26 3.0!i Section Area 2 705 211 262 ]13 63 ]54 139 (m ) -'--'-"-_._~-"-'--'-------_.,--,-,-------,----,-,-------.-------.----.-- -_._._._--_.._-~ -----------.'--------.'-' -_u_.---------- --._--- -'------'- August 1967 3.63 3.65 3.2] 5. ]4 Uc tober 1967 1,43 2.16 0.08 ]6.63 6.85 5.84 3.34 May ]968 0.81 1. 28 2.81 2.55 .1.03 0.32 July ]968 0.30 1.28 0.37 3,43 4.69 1.72 0.60 00 2.24 0.01 4.73 3.20 ] 1. 76 5.25 N October 1968 0.92 May ]969 0.48 1.11 1. 96 2,45 1.18 1. 87 August 1969 0.42 1.2] 4.44 4.09 2.68 1. 21 October 1969 1.18 0.96 5.45 3.]4 4.87 1. 26 May 1970 1.05 2.05 1. 56 1.66 1.63 0.57 "."..-..._- -'--"--'------------------------------ '-------'----.. -'---~---'-'-'--'-'-'.'-----'----------,-----,-,-- _....~-------_._._-- ---.---------- ---~--"-'-------'----'-'---'-----. ------ --.-------------.---.-----.--..----- --'-.---"---------'-'--'- - ---. ....- ------- TABLE A-34. MEAN (1963-1972) BIOMIISS (gll) OF BROWN TROUT IN FivE TlUBIITI\RIES OF TIlE RIVER TEES SYSTEM, ENGLAND, IN MAY, AUGUST. Atm OCTOBER (FROM CRISP ET AI.. 1975), . . '~----'----"- -.----.. -.-.-.----------. --_.- --- --------.---...---. _..~------_.. "---,.-. -." -----.------....--------------------.-.- ---'---'--'--'--- -.'-.-- .- --.--.--.-.---.. -.-- '-'--"-----'---'-" - '--------- - - --- _no _.. ------------ --- --. ----- -.--. ------ - ------ --.---.--..-.--..-----.-.-.---.. ..- . -- .--.---- .-...----.. ~loss Burn Nether lIearth Sike Trout Bed Great Dodgen Pot Sike 'A' Great Dodgen Pot Sike 'B' -- -- ------------- -..---".-- ------ "--- .--.- - --_.... -...-- ..-.. .'--- -------- ------.----" --- -'---'--'-'-'- -.-----... .-... "--- . - -...,.---..-- .--- .-.---- --.--.-..-- ---- Meall IHdth (m)a 1.9 2 a 254 Surface Area (m ) Minimum 0.8 May t1ean 1. 19 Maximum 1.6 Minill1U111 1.4 August Mean 2.00 00 Max imlun 2.9 VI. Minimum 1.0 Oc toher tlean 1. 37 Maximum 2.2 4.1 5.6 0.9 1.6 209 205 34 115 0.7 1.0 3.0 1.1 1. 14 1. 45 4.55 1.85 1.5 1.8 6.8 2.7 1.3 2.4 3.5 1.6 2.62 3.97 5.54 3.77 3.3 6.2 10.1 7. I 1.2 1.0 5.3 2.9 2.02 1.47 5.41 3.8n 2.8 1.9 8.1 5.1 . a Based 011 measurements made in tlay 1968. ..-..-------.--.---.------.-------..------------.---------.--------. -----.__.__._----------~._-----_._,-_._-_._._-------_.- -------------------.------------.----.----------------.---.----.--.--.-.-------.--------------- --- --'------.--.----.-.--...--- ------- TABLE A-35. PRODUCTION (g/nh OF ATLANTIC SALMON MID BROWN mOUT IN TltREE SECTIONS OF SIIELlIGlI1I BURN, SCOTlAND, 1966-196f1 (FROM EGGLISIIAW 1970). ---- .-.- -----.-.---...---.----.-- --.---- --_.- -'-------'-".'-'--- .-..-.--.---- -- ..--- "- .-- -----------..----.--------- -----'--'-----'--------'---,- . , . . ---- -.- '-----.-,--..---------------.----.---.------ --". ---.---..------ --'-'~----'- -.------.-.----- --------------....----..--------- --'---_.,----_._~_._- , . . Year,Class 1963----1964-----1965-- ---1%6------1 967 -------T~68 Annual , To ta 1 s Sec ti on Length (III) Mean Width (III) Section Area (1112) -,. .-------------.----.---- -.-----------------.------------.. --------- ---'------------------------'--------'-_'_"_0. --..----..-.----.-.---. - -,------ ..-. - . Sec ti on I-Ups trealll 30.8 4.10 126.0 1966 Salmon 0.23 1. 57 1. 25 4.47 7.52 Trout 0.65 1. 47 3.42 4.10 9.64 1967 Salilion 0 0.11 0.63 2.22 7.38 10.84 T rou t 0 0.52 0.77 5.05 5:82 12.16 1968 Salilion 0 0 0.16 0.64 2.53 8.54 11.87 Trout 0 0 0 0.94 3. JO 2.67 6.71 Sec ti on 2 27.9 3.07 86.0 1966 Salmon 0 1. 12 1.07 3.52 5.71 Trou t 0.51 2.49 4.69 2.80 1O.62a '00 1967 Salmon 0 0.03 O. HJ 1. 92 6.59 87.2 -too frout 0 0 2.07 5.95 3.98 12.00 1968 Salmon 0 0 0 0.43 2.51 7.28 10.22 Trout 0 0 0 1. 91 4.40 2.39 8.70 Sect ion 3-Downs tream 27.5 3.43 9'1.0 1966 Salmon 0 0.51 1.03 4.69 6.23 Trout 0.41 1. 96 '1.44 3.95 10..76 1967 Salmon 0 0.30 0.52 2.87 8.59 12.28 Trout 0 0.46 2.02 5.68 4.54 12.70 1968 Sa lilion 0 0 0 0.72 2.56 7.89 11.17 Tt'out 0 0 0 1.45 3.92 2.114 7.81 --.-- ----- .._--_._----------_.~--------,-.- ------.-.---..-...--- ".---'-------'- -+-----'-----'_.-__h.'__,.,. -..,.+.-. -----"---'-~---'-'----'-- ----'- -- ---. - -. ---.-..- -----'--~'---'- --- . ."-".-'.' '-----'.-. --_. - _.__.m___.__.----- .. --------.--... -',-----,-,_u.,--".,. - - "-- "---.. __n___.,..--..----.---. - - ---'----'----'-'- -'---'--'--.- - - -'----____-.n.- ----------- .-..., -.-- .-..--------.- a hlcludes 0.13 9/1112 production of the 1962 yea,. class. ------- TABLE A-36. BIOMASS (g/m2) OF ATLANTIC SAU1or: AND BRO\'m TROUT AT THE END OF THE GROWING SEASON, SHELLIGAN BURN, SCOTLAND, 1966-1975 (FROM EGGLISHAt'J Arm SHACKLEY 1977). TOTAL BIOMASS INCLUDES :1: STANDARD ERROR. Salmon Trout Total . Year 0+ 1+ '2+ All 0+ 1+ 2+ All Biomass 1966 3.5 1.0 1.5 6 . 0:1: 1. 9 2.5 4.0 3.2 9.8:t1.1 15.8:1:0.9 1967 2.7 1.7 0.2 4.6:1: 1. 2 3.6 6.1 1.8 11.4:1:1.5 16.1:1:2.6 1968 2.5 1.7 0.7 4.9:1:0.9 1.9 3.2 0.9 6 . 0:1: 1. 1 10.9:1:2.1 1969 4.4 2.2 0.7 7 . 3:1: 1. 9 5.0 3.8 2.9 11.6:1:2.1 18.9:1:2.8 1970 1.9 . 2.5 0.3 4.7:1:2.3 1.9 6.0 2.1 10.1:1:2.5 14. .8:1:3.9 1971 4.7 2.4 0.5 7.6:1: 1. 9 3.8 4.9 1.4 10.1:t2.9 17~7:t4.2 1972 2.6 4.6 0.3 7.5:1:2.8 3.2 6.9 0.8 10.9:1:3.0 18.4:1:4.2 1973 3.7 3.8 0.2' 7.6:1:1.4 4.0 5.4 1.2 10.7:1:2.5 18.3:1:3.2 1974 3.2 3.4 .0.0 6.6:t1.2 2.5 4.5 0.4 7.4:t2.2 14.0:1:3.2 1975 6.1 2.9 0.2 9.2:1:2.6 1.9 4.3 0.8 7. l:t2 . 1 16.3:1:4.0 Me.an . 3.5 2.6 0.5 6.6:1:1.5 3.0 4.9 1.6 . 9.5:1:2.0 16.l:t2.4 TABLE A-37. BIOMASS (g/m2) OF BROWN TROUT IN SIX SECTIONS OF HOROKIHISTREAM, NEW ZEALAND, 1940-1941 (FROM ALLEN 1951). SECTION I IS DOWNSTREAM. . Zone I IIM IIR III IV V Length (m) 3,167 2,035 1 ,918. 2,719 1,602 527 Total July 1940 24.1 31.5 11. 7 24.9 41.0 18.8 25.6 Oct. 1940 28.0 36.1 11. 7 25.7 46.4 24.7 28.6 Jan. 1941 34.1 32.7 11.0 16.4 29.8 21.9 26.5 May 1941 22.8 42.8 14.9 21. 3 26.0 24.4 25.8 Oct. 1941 1.8 13.7 2.5 2.7 18.7 18.5 6.5 8'S ------- |