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SYNOPSIS of BIOLOGICAL DATA on the NORTHERN PIKE Esox Lucius Linnaeus, 1758

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SYNOPSIS of BIOLOGICAL DATA on the NORTHERN PIKE Esox Lucius Linnaeus, 1758 rAo Fisheries Synopsis No. 30 Rev. 2 SAST - Pike - 1,24(03)001,01 SYNOPSIS OF BIOLOGICAL DATA ON THE NORTHERN PIKE Esox lucius Linnaeus, 1758 FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS FIR/S30 Rev.2 FAO Fisheries Synopsis No. 30 Rev.2 SAST - Pike - 1,24(03)001,01 SYNOPSIS OF BIOLOGICAL DATA ON THE NORTHERN PIRE Esox lucius Linnaeus, 1758 Prepared by Alexander J.P. Raat Organisatie ter Verbetering van de Binnennvisserij P.O. Box 433 3430 AK Nieuwegein Netherlands FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS Rome 1988 The designations employed and the presentation of material in this publication do not imply the expression of any opinion whatsoever on the part of the Food and AgricultureOrganization of theUnited Nations concerning the legal status of any country, territory, city or area or of its authorities, or concerning the delimitation of its frontiers or boundaries. M-42 ISBN 92-5-102656-4 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means. electronic, mechanical, photocopying or otherwise, without the prior permission of the copyrfght owner. Applications for such permission, with a statement of the purpose and extent of the reproduction, should be addressed to the Director, Publications Division, Food and Agriculture Organization of the United Nations. Via delle Terme di Caracalla, 00100 Rome. Italy. © FAO 1988 FIR/S30 Rev.2 Pike PREPARATION OF THIS SYNORSIS This report was prepared as a follow-up to the offer of the Netherlands delegation to the eleventh session of EIFAC in Stavanger (FAO, 1981), to update the FAO Fisheries Synopsis of biological data on the northern pike, Esox lucius Linnaeus, 1758. In view of the enormous amount of data published on this important species since the last revision of the document (Toner and Lawler, 1969), it was necessary to rewrite it entirely. The present report is based on an extensive literature study. Sources for the literature search included the Aquatic Science Fisheries Abstracts (ASFA), the Sport Fisheries Abstracts (SFA) and the Landwirtschaftliche Zeitschrift (LWZ). Furthermore, the reference lists of the consulted publications proved to be useful tools in the collection of published and unpublished material on the species. The synopsis summarizes data on pike gathered in many countries by a large number of biologists during the last three decades, up to 1st January 1985. Its reference list only includes studies quoted in the synopsis. These papers were consulted in autopsy unless otherwise stated in the reference list. The author gratefully acknowledges the help he received from the numerous persons who have supplied him with data. It is impossible to quote all of them here, but special thanks are due to: Dr E.J. Crossman, Royal Ontario Museum, Toronto, Canada, who reviewed the chapters on taxonomy and distribution and furnished additional data on pike for North America; Dr J.D. Reist, Freshwater Institute, Winnipeg, Canada, who provided important information on the taxonomic status of the species from his unpublished Ph.D. dissertation; Dr J. Scott Campbell, Research, Branch Fisheries and Oceans, Moncton, Canada, who suppliedunpublished data on the HemingLake pike from Dr G.H. Lawler's files; Dr J.S. Diana, University of Michigan and Dr E.A. Huisman, Agricultural University, Wagening, Netherlands, who reviewed the chapters on metabolism; Dr L. Johnson, Freshwater Institute, Winnipeg, Canada, who supplied data from his Ph.D. dissertation on Windermere pike; Dr J. Willemsen, Ijmuiden, Netherlands, who reviewed parts of the manuscript dealing with his work on growth, culture and metabolism; Dr R.H.K. Mann, Freshwater Biological Association, East Stoke, England, who reviewed the chapters on the biology of the species. Further comments on earlier drafts of the synopsis were received from Dr H. Löffler, Langenargen, Federal Republic of Germany, Dr H.Lehtonen, Helsinki, Finland, and from the author's Dutch colleagues Dr J. Klein Breteler and Mr J. Quak, Mrs M.H.M. van Densen translated the Russian article by Shamardina (1957) on the early development of northern pike. The author has the pleasure of acknowledging the help and support of Dr M.P. Grimm of the Research Department of the Organization for Improvement of Inland Fisheries, Nieuwegein, Netherlands. His many published and unpublished data on northern pike and his invaluable knowledge and experience with freshwater fisheries have been an important source of inspiration for the author during the writing of this study. Part of the data collection was carried out by Mr J. Kampen (particularly the data on parasites) and Mr A.van Beusekon collected the data on growth. Mrs I.H. Willemsen-Vörös was extremely helpful in tracing down many publications and thus improving the bibliography of this report. Research on the biology and culture of northern pike was greatly stimulated in the Netherlands since the fifties by Mr D.E. van Drimmelen, former chairman of EIFAC and of the Organization for Improvement of Inland Fisheries. The present synopsis is therefore one of the results of his inspiring activities in the field of freshwater fisheries. Final editing of the synopsis was achieved by Ms Martina Schneider, Fishery Resources Officer, FAO Fishery Resources and Environment Division. Final revision and editing of the bibliography was carried out by Ms Gloria Soave, Research Information Unit. Distribution: For bibliographic purposes this document should be cited as follows: Author FAO Fisheries Department Paat, A.J.P., Synopsis of biological data on FAO Regional Fisheries Officers 1988 the northern pike, Esox lucius Regional Fisheries Councils and Commissions Linnaeus, 1758. FAO Fish.Synop., Selector SI and EI (E and F) (30)Rev.2:178 p. iv FIR/S30 Rev.2 Pike ABSTRACT This synopsis consolidates all the available published data on the biology, growth, migration and population dynamics of a freshwater fish species which has been exploited and cultured for food since many years in European and North American waters. FIR/S30 Rev.2 Pike CONTENTS Page TAXONOMY AND DISTRIBUTION 1.1 Nomenclature 1.1.1 Valid name 1 1.1.2 Objective synonymy 1 1.1.3 Etymology 1 1.1.4 Standard common names, vernacular names 1 1.2 Identity 1.2.1 Taxonomic status 1.2.1.1 General overview 1 1.2.1.2 Interactions between pike and muskellunge 4 1.2.2 Morphology 4 1.2.2.1 Skin, slime layer, skin patterns 4 1.2.3 Cytomorphology 5 1.2.4 Protein specificity 6 1.2.5 Blood 6 1.3 Taxonomy 7 1.3.1 Suprageneric affinities 7 1.3.2 Generic affinities 8 1.3.3 Diagnosis of the species 9 1.3.4 Key to esocids of the world 10 1.3.5 Subspecies 11 1.4 Hybrids and Mutants 11 1.4.1 Hybridization 11 1.4.1.1 General overview 11 1.4.1.2Muskellunge x northern pike 13 1.4.1.3Northern pike x Amur pike 14 1.4.1.4 Northern pike x grass pickerel 14 1.4.1.5 Chain pickerel x northern pike 14 1.4.1.6 Perca fluviatilis x northern pike 14 1.4.2 Mutants 14 1.4.3 Hardiness 15 1.4.4 Abnormalities 15 1.5 Distribution 16 1.5.1 Origin and fossil records 16 1.5.2 Present distribution 17 1.5.3 Differential distribution 18 1.5.3.1 Spawn and larvae 18 1.5.3.2Juveniles and adults 18 2. BIONOMICS AND LIFE HISTORY 19 2.1 Reproduction 19 2.1.1 Sexuality 19 vi FIR/S30 Rev.2 Pike Page 2.1.1.1 Hermaphroditism, heterosexuality 19 2.1.1.2 Sexual dimorphism 19 2.1.2 Maturity 19 2.1.3 Mating 20 2.1.4 Fertilization 20 2.1.5 Gonads 20 2.1.5.1 Testes and sperm 20 2.1.5.2 Ovaries and ova 22 2.1.5.3 Fecundity 24 2.1.6 Spawning 25 2.1.6.1 Spawning migration 25 2.1.6.2 Spawning grounds and spawning time 25 2.1.6.3 Spawning behaviour 29 2.1.7 Spawn 30 2.2 Embryonic Period 31 2.2.1 General aspects 31 2.2.2 Cleavage phase 31 2.2.3Embryonic phase 32 2.2.3.1 Effect of temperature 33 2.2.3.2 Effect of oxygen 34 2.2.3.3 Effects of other factors 35 2.2.4Hatching 35 2.2.4.1 Description 35 2.2.4.2Hatching gland cells, hatching enzyme 35 2.2.5Eleutheroembryonic phase 36 2.2.5.1 General description 36 2.2.5.2Effect of environment 36 2.3 Larval and Juvenile period 37 2.3.1 General aspects 37 2.3.2 Protopterygiolarval phase 37 2.3.3Pterygiolarval phase 38 2.3.4Feeding and feeding behaviour 40 2.3.4.1 Development of organs and feeding 40 2.3.4.2 Feeding behaviour 41 2.3.4.3 Phases in nutrition 42 2.3.4.4 Digestion and periodicity of feeding 42 2.3.4.5 Cannibalism among larvae 44 2.3.5Growth of larvae 45 2.3.6Emigration of larvae 48 2.3.7 Adult period and senescent period 51 2.4 Feeding and Food of Adult Northern Pike 51 2.4.1 Introduction 51 2.4.2Eye and vision 51 2.4.3 Lateral line and reactions to mechanical stimuli 52 2.4.4Olfactory organ and reactions to chemical stimuli 52 2.4.5Alimentary tract 52 FIR/S30 Rev.2 Pike vii Page 2.4.5.1 Mouth and dentition 52 2.4.5.2 Oesophagus and stomach 53 2.4.5.3 Intestine 53 2.4.6 Feeding behaviour 53 2.4.6.1 Behaviour of northern pike 53 2.4.6.2 Predator-prey interactions 55 2.4.7 Food selection 56 2.4.7.1 Species selection 56 2.4.7.2 Size selection 57 2.4.7.3 Seasonal food 58 2.4.8 Cannibalism 59 2.5 Growth of Northern Pike 60 2.5.1 Correlated factors 60 2.5.2 Longevity 67 2.5.3 Scales and ageing 67 2.5.4Greatest size 68 2.6Metabolism 68 2.6.1 Food conversion 68 2.6.2 Engergy deposition 69 2.6.3Energy sources 71 2.7 The Species in Relation to its Environment 71 2.7.1 Migration and local movements 71 2.7.1.1 Tagging and marking 72 2.7.2 Competitors of northern pike 73 2.7.3Predators of northern pike 74 2.7.4Vegetation 75 2.7.5Temperature 76 2.7.6 Ice 77 2.7.7 pH 77 2.7.8Oxygen 77 2.7.9Light and dark; day and night 78 2.7.10 Currents 79 2.7.11 Salinity 79 2.7.12 Pressure 79 2.7.13 Pollution 80 2.8 Parasites, Diseases 81 2.8.1 Introduction 81 2.8.2Viral diseases 81 2.8.2.1 Pike fry rhabdovirus disease 81 2.8.2.2 Egtved disease 81 2.8.2.3 Lymphocystis disease 82 2.8.2.4 Lymphosarcoma 82 2.8.3 Bacterial diseases 82 2.8.3.1 Red sore-disease 82 2.8.3.2 Black-spot disease 83 2.8.3.3 Furunculosis 83 2.8.3.4 Mycobacteriosis 83 2.8.4Parasites of northern pike 83 viii FIR/S30 Rev.2 Pike Page 3.
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