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Alosa Alosa and Alosa Fallax Spp. Literature Review and Bibliography

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Alosa Alosa and Alosa Fallax Spp. Literature Review and Bibliography Alosa alosa and Alosa fallax spp. Literature Review and Bibliography R&D Technical Report W1-014/TR M. W. Aprahamian C. D. Aprahamian J. L. Baglinière R. Sabatié P. Alexandrino Research Contractors Environment Agency UMR INRA-ENSA ENSAR University of Porto University of Porto Publishing Organisation Environment Agency, Rio House,Waterside Drive, Aztec West, Almondsbury Bristol BS32 4UD Tel: 01454 624400 Fax: 01454 624409 Website: www.environment-agency.gov.uk ISBN 1 84432 109 6 © Environment Agency 2003 All rights reserved. No part of this document may be produced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise without the prior permission of the Environment Agency. The views expressed in this document are not necessarily those of the Environment Agency. Its officers, servants or agents accept no liability whatsoever for any loss or damage arising from the interpretation or use of the information, or reliance upon the views contained herein. Dissemination Status Internal: Released to Regions External: Public Domain Statement of Use The aim of this report is to provide fish biologists and fisheries managers with an overview of the biology, status and conservation options for Alosa alosa and Alosa fallax species. Research Contractor This document was produced under R&D Project W1-014 by : Aprahamian, M. W.(1), Aprahamian, C. D.(1), Baglinière, J. L(2), Sabatié, M. R.(2,3), Alexandrino, P.(4), (1) The Environment Agency P.O Box 12, Fairclough House, Knutsford Road, Warrington WA 4 1HG, UK. (2) UMR INRA-ENSA Ecobiologie et Qualité des Hydrosystèmes Continentaux, 65 rue de Saint-Brieuc, 35042 Rennes Cedex, France (3) Laboratoire d’Halieutique, ENSAR, 65 rue de Saint-Brieuc, 35042 Rennes Cedex, France. (4) Animal Genetics and Conservation Unit, Faculdade de Ciências, Universidade do Porto, Campus Agràrio de Vairão, R. Monte Castro 4480 Vila do Conde, Portugal. Environment Agency Project Manager The Environment Agency’s Project Manager for R&D Project W1-014 was Miran Aprahamian, Noth West Region. Further copies of this report are available from: Environment Agency R&D Dissemination Centre WRc, Frankland Road, Swindon, Wilts. SN5 8YF Tel: 01793 865000 Fax: 01793 514562 E-mail: [email protected] R&D TECHNICAL REPORT W1-014/TR Keywords ALOSA ALOSA; ALOSA FALLAX FALLAX; ALOSA FALLAX RHODANENSIS; ALOSA FALLAX NILOTICA; ALOSA FALLAX ALGERIENSIS; ALOSA FALLAX LACUSTRIS; ALOSA FALLAX KILLARNENSIS; NOMENCLATURE; TAXONOMY; MORPHOLOGY; DISTRIBUTION; HYBRIDIZATION; REPRODUCTION; SPAWNING; ONTOGENY; GROWTH; DIET; DISEASE; PARASITE; INTER- & INTRA-SPECIFIC INTERACTIONS; HOMING; MIGRATION; SEX RATIO; AGE STRUCTURE; RECRUITMENT; MORTALITY; POPULATION DYNAMICS; EXPLOITATION; FISHING EQUIPMENT; FISHING AREA, FISHING SEASONS; CATCHES; HABITAT PROTECTION AND MANAGEMENT; CONSERVATION STATUS; FISH PASS; CULTURE. R&D TECHNICAL REPORT W1-014/TR i PREFACE This review is for those who have and will work on Alosa, to the commercial fisherman and recreational anglers who make their living or obtain pleasure and relaxation from their capture. It is also for the many people we have worked with us over the years, without whose help little would have been achieved and to the many who have freely contributed information and knowledge in the true spirit of collaboration, we hope we have not abused their generosity. We acknowledge that there still exists mistakes in this draft, especially in the references. We have supplied as much detail as we can but appreciate that in many instances it is incomplete. For this and other errors we apologise, we will, however, endeavour to correct these over time and maintain the review up to date and to help us achieve this we welcome any feedback. In the future we hope the review will be made available on the Internet. R&D TECHNICAL REPORT W1-014/TR ii R&D TECHNICAL REPORT W1-014/TR iii ACKNOWLEDGEMENTS We thank B. Steinmetz who as a Convenor of EIFAC sub-commission 1 “Biology and Management” first suggested putting together a synopsis of biological data on Alosa during a fleeting meeting at a conference in Hull (UK) over ten years ago! We also thank subsequent convenors, C. Moriarty and T. Brenner for the annual request on progress and to whom the many excuses for the lack of any development must have seemed insufferable. We also thank those who have helped us with obtaining references, translations and with the many aspects of IT, which have made this task considerably easier; We thank P. Bianco, P. Bergot, P. Boisneau, G. Boxshall, G. Casternaud, P. Caswell, F. Cassou- Leins, J. J. Cassou-Leins, S. Colclough, M. Collares-Pereira, A. Churchward, P. Economidis, E. Esteves, J. Gregory, P. Gough, B. Hardy, J. Hateley, T. Hatton-Ellis, D. Heaver, P. Henderson, M. Hendricks, J. Hillary, R. Hillman, T. Jacklin, J. King, P. Lambert, M. Le Corre, M. Larinier, S. Lester, A. Lyle, O. McCrimmon, P. Maitland, C. Mennesson-Boisneau, N. Mott, M. Nolan, N. O'Maoileideigh, S. Oesmann, A. Picken, T. Pina, E. Rochard, H. Sari, R. Seaby, M. Skora, L. Syme, C. Taverny, R. Thiel, C. Turan, V. Véron, C. Wallis, A. Walton, A. Wheeler, K. Whittaker. H. Winkler. For permission to reproduce figures we thank AULA-Verlag (GmbH Wiesbaden, Verlag für Wissenschaft und Forschung, Luisen Platz No. 2, Wiesbaden, Germany) for Figures 1.2, 1.7, 1.8, 1.11, 3.15, 3.16 and 3.17. American Society of Ichthyologists and Herpetologists for Figures 1.7, 1.8 and 1.10 Blackwell Publishing for Figures 1.15, 3.26, 3.31, 3.33 and 6.4; P. Boisneau and C. Mennesson-Boisneau for Figures 3.29 and 5.3; Bulletin Français de la Pêche et de la Pisciculture for Figures 3.20, 3.22, 3.23, 4.4, 5.12, 5.13 and 6.3; J. Dartiguelongue for Figure 3.30; By Courtesy of the Linnean Society of London Figure 1.14; Conseil Supérieur de la Pêche for Figures 3.1, 3.2, 3.22, 3.23, 4.4, 5.11, 5.12 and 6.3; FAO for Figure 1.1; INRA for Figures 3.3, 3.4 and 5.8; N. O'Maoileideigh for Figure 3.11; The Zoological Society of London for Figure 1.9. R&D TECHNICAL REPORT W1-014/TR iv CONTENTS page KEYWORDS i ACKNOWLEDGMENTS iv LIST OF FIGURES vii LIST OF TABLES xiii EXECUTIVE SUMMARY xix 1. IDENTITY 1 1.1 Nomenclature 1 1.2 Taxonomy 4 1.3 Morphology 9 2. DISTRIBUTION 39 2.1 Total Area 39 2.2 Determination of Distribution Changes 51 2.3 Hybridization 51 3. BIONOMIC AND LIFE HISTORY 52 3.1 Reproduction 52 3.2 Preadult Phase 86 3.3. Adult Phase 92 3.4 Nutrition and Growth 105 3.5 Behaviour 168 4. POPULATION (SPAWNING STOCK) 179 4.1 Structure 179 4.2 Abundance and Density 189 4.3 Natality and Recruitment 190 4.4 Mortality and Morbidity 193 4.5 Dynamics of the Population 193 5. EXPLOITATION 195 5.1 Fishing Equipment 195 5.2 Fishing Areas 201 5.3 Fishing Seasons 201 5.4 Fishing Operations and Results 201 5.5 Climatic Influences 206 6. PROTECTION AND MANAGEMANT 208 6.1 Conservation Status 208 6.2 Habitat Protection and Management 212 6.3 Fish Passes 213 6.4 Screens 217 6.5 Translocation / Transplantation 218 R&D TECHNICAL REPORT W1-014/TR v 7. CULTURE 219 7.1 Collection of Spawn 219 7.2 Incubation of Eggs 220 7.3 Larval and Juvenile Rearing 220 REFERENCES (not cited in bibliography) 222 BIBLIOGRAPHY 225 APPENDIX 1 335 APPENDIX 2 342 R&D TECHNICAL REPORT W1-014/TR vi LIST OF FIGURES Figure 1.1 Alosa alosa and Alosa fallax; reproduced with permission of FAO 5 (www.fishbase.org). Insert, showing the distinct notch into which the symphysis of the lower jaw fits (Whitehead, 1985). Figure 1.2 First gill arch appearance and approximate number of gill rakers 9 for (a) A. fallax and (b) A. alosa. Figure 1.3 The relationship between number of gill rakers on the first gill 14 arch and total length for Alosa alosa (1-10, solid line) and Alosa alosa x Alosa fallax hybrids (11-14, dotted line). (1 = Rhine (Hoek, 1899); 2 = Aulne (Véron, 1999); 3 = Vilaine (Véron, 1999); 4 & 11 = Loire (Douchement, 1981); 5 = Charente (Véron, 1999); 6 = Garonne (Douchement, 1981); 7 = Adour (Douchement, 1981); 8 & 12 = Aude (Douchement, 1981); 9 & 13 = Rhône (Douchement, 1981); 10 & 14 = Sebou (Sabatié, 1993). Figure 1.4 15 The relationship between number of gill rakers on the first gill arch and total length for Alosa fallax; Anadromous populations: A. fallax fallax: 1 = Rhine (Hoek, 1899); 2 = Hass (1965); 3 = Severn (Aprahamian, 1982); 4 = Loire (Douchement, 1981); 5 = Charente (Véron, 1999); 6 = Garonne (Quignard & Kartas, 1977); 7 = Garonne (Douchement, 1981); 8 = Adour (Douchement, 1981); 9 = Loukos (Sabatié, 1993); 10 = Sebou (Sabatié, 1993); A. fallax rhodanensis: 11 = Rhône (Quignard & Kartas, 1977); 12 = Rhône (Douchement, 1981); 13 = Rhône (Le Corre, et al., 1997); 14 = Aude (Quignard & Douchement, 1991c); 15 = Aude (Douchement, 1981); 16 = Tirso (Quignard & Douchement, 1991c); A. fallax algeriensis: 17 = Miliane (Quignard & Kartas, 1977); 18 = Oubeira (Quignard & Kartas, 1977); Landlocked populations: A.fallax nilotica: 19 = Bacin (Vukovic, 1961a); 20 = Skadar (Vukovic, 1961a); A. fallax algeriensis: 21 = Ischkeul (Quignard & Kartas, 1977); A. fallax: 22 = Sidi Mohamed Ben Abdullah (Sabatié, 1993). Figure 1.5 Scale from Alosa fallax fallax.23 Figure 1.6 Skeletal features of Alosa alosa, axial view. En epineural 25 bone; ep epipleural bone; ha hemal arch; na neural arch; r rib; v vertebra; ec epicentral bone; ea epapophyses. Reproduced from Bertin (1958). Figure 1.7 The skull of the allis shad Alosa alosa.26 Figure 1.8 The skull of the twaite shad Alosa fallax fallax (a lateral view; b 26 posterior view; c dorsal view; d ventral view; e anterioventral view; als alisphenoid; boc basioccipital; bs basisphenoid; eoc exoccipital (lateral occipital); epo epiotic; eth.
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