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By David H. Cushing Duly 1973 Prepared Under RECRUITMENT AND PARENT STOCK IN FISHES By David H. Cushing WSG 73 I duly 1973 DIVISION OF MARINE RESOURCES UNIVERSITY OF WASHINGTON 98195 Prepared under the National Sea Grant Program A WASWINGTON SEA GRANT PUBLICATION RECRUITMENT AND PARENT STOCK IN FISHES by Davi d H. Cushing WSG 73-1 Published July 1973 by Division of Marine Resources UNIVERSITYOF WASHINGTONSeattle 98195 ACKNOWLEDGMENT This publication was supported by the Washington Sea Grant Program, a part of the National Sea Grant Program. The National Sea Grant Program is supported by the National Oceanic and Atmospheric Administration, U. S. Department of Commerce. Printed in the United States of America Price $7.50 Distributed by University of Washington Press, Seattle 98195 Washington residents please add sales tax! CONTENTS Chapter Page 1 The Nature of a Fish Stock Introduction 1 Early History of Racial Studies Meristic Character's 2 3 The Stocks as Defined by Fisheries Biologists The Hydrographic Containment of a Stock--Arctic Cod 48 6 Genetics in Stock Studies The Unit Stock and Recruitment 10 2 The Mechanisms of Recruitment 31 Int roduct l on ~ ~ ~ ~ ~ ~ I . ~ ~ ~ ~ 4 ~ ~ ~ ~ 31 Larval Drift 32 The Nursery Ground 35 The Variation in the Production Cycle 36 The Times of Spawning 38 The Match of Spawning to the Production Cycle 39 Recruitment and Stabilization 40 3 The Stability of Numbers 58 Introduction o ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 58 The Stability Observed . ~ ~ ~ 58 The Stabilization Mechanism ~ ~ ~ 61 Models of the Density-dependent Mechanism ~ ~ 63 Stock and Recruitment Curves 77 Introduction The Biology of the Pacific Salmon in the Development of the Ricker Curve ~ ~ ~ 77 The Ricker Curve . ~ ~ 81 The Beverton and Holt Curve ~ ~ ~ 83 Present Developments ~ ~ ~ 85 5 The Role of Fecundity 89 Introduction ~ ~ ~ ~ ~ 89 The First Method Cushing, 1970! ~ ~ ~ ~ ~ 89 The Second Method Cushing and. Harri s, in pre ss! ~ ~ 92 Conclusion ~ ~ ~ 96 iv Chapter Page 6 Recruitment Overfishing Introduction 114 The East Anglian Herring 115 The Arcto-Norwegian Cod 116 Four Pelagic Stocks 3.18 Conclusion 120 7 The Age Structure of a Population 127 Introduction 127 Density-dependent Natural Mortality 128 The Natural Mortality of the Plaice of the Southern North Sea 130 Senescent Mortality in the Plaice 132 The Age Structure in a Fish Population 134 Climate and Recruitment 144 Introduction 144 The Correlation between Year-class Strength and Climatic Change 144 The Nature of Climatic Change 146 The Biological Responseto Climatic Change 149 Conclusion 150 9 The Single Process 3,63 Introduction 163 Fecundity, Spawning, and the Density of Eggs in the Sea . 163 Egg and Larval Development 165 Feeding and Density-dependent Pr ocesses 166 Environmental Modulation 168 Stabilization 3.68 Adult Life 169 The Single Process 170 10 The Present State of the Stock and Recruitment Prob lem 3.74 Introduct ion 174 The Biology of Recruitment 174 A Provisional Yield Curve Incorporating the Dependence of Recruitment on Parent Stock 177 Conclus i on 179 Literature C'it ed,, g ~ ~,, ~ ~ ~ ~ ~ ~ ~ ~ 3.85 LIST OF FIGURES Figure Page The trend in the number of vertebrae with latitude in British Columbian herring off Vancouver Island Tester, 1935! 12 The effect of temperature upon the vertebral 0 counts of trout Taning, 1952! .............. 13 3-a The spawning ground of the Arctic cod: distr'ibution of cod eggs Hjort, 1914! . 14 The spawning ground of the Arctic cod.' distribution of echo traces in March 1955 Bostr5m, 1955!, in successive weeks, a-e.....,,............ 15 3-c The spawning ground of the Arctic cod: local re- captures from tagging on the spawning ground Hjort, 1 914 ! ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ The outward migration of the Arctic cod: recaptures in the North Norwegian coast Hjort, 1914! ........ 17 4-b The outward migration of the Arctic cod: recaptures in the Barents Sea Dannevig, 1953!............ 18 4-c The outward migration of the Arctic cod: distribution of echo traces off the North Cape in spring and autumn 1959 Hylen et al., 1961!................. 19 The return migration: recapture from eastern Barents Sea Mastov, 1944! .................... 20 The return migration: recapture from the eastern Bar ents S ea I dels on, 19 31! 21 5-c The return migration: an echo survey showing the approach of the cod to Vestfjord Saetersdal et a] 1 959! ~ ~ ~ t ~ ~ ~ ~ ~ ~ ~ ~ w ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 2 2 Migration of immature cod in the Barents Sea Mastov, 1 944 ! ~ ~ ~ s ~ ~ ~ ~ ~ ~ ~ t ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 23 An ideal picture of the migration circuit of Arctic cod . 24 The spread of tagged halibut from the point of liber- ation Thompson and Herrington, 1934! . 25 Vi Figure The transpacific migrations of the albacore Otsu > 1960! ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ t ~ 26 10 The recovery of plaice tagged on their spawning grounds one year later de Veen, 1961! .......... 27 ll-a The distribution of isozyme. in the North At.lantic cod: a cline in hemoglobin in cod along the coast of Norway de Ligny, 1969! 28 11 b The distribution of isozymes in the North Atlantic cod distzibution of the hemoglobin HbI' in the North Atlantic de Ligny, 1969! 29 11-c The distribution of isozymes in the North Atlantic c cod: distribution of the transferrin, Tf in the Nozth Atlantic de Ligny, 1969! 30 12 The center of the spawning ground of plaice in the Southern Bight after Simpson, 1959!, the larval drift in the water of maximum salinity and. minimum turbidity and the Texel gate to the inshore region Dietrich, 1 954! ~ ~ ~ ~ ~ ~ ~ ~ e ~ ~ ~ t ~ t ~ ~ ~ ~ ~ ~ t a ~ 42 13-a Compensation depths in the Southern Bight of the North Sea from January to May in coastal water types 1, 3, 5, 7, and 9 43 13-b Critical depths in the Southern Bight of the North Sea from January to May in coastal water types 1, 3, 5, 7~ and 9 s ~ ~ ~ ~ ~ ~ e e ~ ~ ~ ~ ~ ~ ~ o ~ ~ 43 14 The distribution of 0-group plaice on the Dutch coast particularly in the Waddensee Zijlstra, 1972! ...... 44 Catch curve for larval winter flounder 45 Catch curve for juvenile flounder of age-groups 0 and I of the 1958 and 1959 year-classes in the Mystic River Estuary 46 Numbers of 0-group plaice of the 1964 and 1965 year-classes caught per standard tow at five positions in Red Wharf Bay, Anglesey, U.K. ~ ~ ~ ~ ~..... ~ ~ ~ 47 Size distribution of 1965 year class of plaice, by months in Firemore Bay, Loch Ewe, Scotland 48 vr1 Figure Page 19 The dependence of growth rate of 0-group plaice upon their mortality rate in Firemore Bay, Loch Ewe, S cotland ~ ~ ~ ~ ~ ~ ~ ~ ~ I ~ ~ ~ ~ ~ ~ ~ t ~ ~ ~ 1 1 ~ 49 20 The three types of production cycle in phytoplankton color: A! central North Sea and Iceland!, B! coastal Atlantic, and C! oceanic Atlantic . 50 21- a Time series of peak spawning dates for sockeye stocklets in the Fraser River system Anonymous, 1950-1966; Cushing, 1970! 52 2 1-b Distribution of days of spawning after the date of first spawning of the Norwegian herring, estimated from the age of spawn in a grab sample Runnstrqhn, 1941!; the numbers in the upper table give the number of positive samples on a given day 53 21-c Mean dates of peak catches of cod in the Lofoten fishery, 1894-1967 Anonymous, 1894-1968!; the trend line x- -x! i s f it ted by moving means 22 Survival curves for plaice eggs and larvae in the Great Orme spawning area of the Irish Sea in 1955 and the Southern Bight of the North Sea in 1962, 1963, and 1968 Harding and Talbot, in press! 55 23 The importance of the 1963 year-class of plaice in the Southern North Sea D. W. Harding and R. C. A. Bannister, unpublished manuscript! . 56 24 The three sources of variance in recruitment shown on stock and recruitment curve Cushing, 1968! 57 25 Herring and salmon recruitments 68 26 Gadoid recruitments 69 27 Flatfish recruitments, North Sea sole Anonymous, 1970! .. 70 28 Variation in numbers and in timing of the outburst of Asfemonella fossa in Windermere Lund, 1964!...... 71 Ricker's stock and recruitment curve Ricker, 1958! 72 30 The decline in survival as recruits/stock! with increasing stock in the southern North Sea plaice Beverton, 1962! 73 Figure Page 31-a Results of model investigations: dependence of mortality on density 74 31-b Results of model investigations: dependence of final weight on food 75 31 c Results of model investigations. dependence of mortality on food..................... 76 32 Ricker's 958! family of stock/recruitment curves .... 98 33 The index of density-dependence for four stocks of pink salmon 99 34 The index of density-dependence in eight red salmon stocks and one chum salmon stock ............. 100 35 The index of density-dependence in four Atlantic herring stocks 101 The index of density-dependence in four Pacific herring stocks 102 37 The index of density-dependence for the California sardine and the yellowfin tuna ~ . 103 38 The index of density-dependence in four stocks of flatfish ~ ~ ~ s 104 39 The index of density-dependence in four stocks of g adoids 105 40 The indices of density-dependence and their errors at twice the standard error! arranged in order of fecundity, lowest at the top . , . ~ . ~ 106 41 Relation between the index of density-dependence and the cube root of the fecundity ............ 107 42 Stock/recruitment curves for three stocks of red salmon .. 108 Stock/recruitment curves for two Atlantic herring stocks . 109 Stock/recruitment curves for four Pacific herring stocks . 110 45 Stock/recruitment curves for four flatfish stocks . 111 Stock/recruitment curves for four gadoid stocks . 112 Figure Page 47 The maxima of four stock/recruitment curves with errors ., 113 48 The dependence of recruitment upon parent stock for the Downs her'ring Cushing, 1968a! ............ 122 49 Catches of the Hokkaido herring and the fishing effort exerted on the stock Cushing, 1971!...
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