Cover photograph: From the archives of the Virginia Institute of Marine Science, Gloucester Point, Virginia, United States of America

Cover photograph: From the archives of the Virginia Institute of Marine Science, Gloucester Point, Virginia, United States of America

Management techniques for elasmobranch fisheries FAO FISHERIES TECHNICAL PAPER 474 Edited by John A. Musick Virginia Institute of Marine Science Gloucester Point, Virginia United States of America and Ramón Bonfil Wildlife Conservation Society New York United States of America FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS Rome, 2005

The designations employed and the presentation of material in this information product do not imply the expression of any opinion whatsoever on the part of the Food and Agriculture Organization of the United Nations concerning the legal or development status of any country, territory, city or area or of its authorities, or concerning the delimitation of its frontiers or boundaries. ISBN 92-5-105403-7 All rights reserved. Reproduction and dissemination of material in this information product for educational or other non-commercial purposes are authorized without any prior written permission from the copyright holders provided the source is fully acknowledged. Reproduction of material in this information product for resale or other commercial purposes is prohibited without written permission of the copyright holders. Applications for such permission should be addressed to: Chief Publishing Management Service Information Division FAO Viale delle Terme di Caracalla, 00100 Rome, Italy or by e-mail to: copyright@fao.org FAO 2005

iii Foreword It is with much pleasure that the FAO has taken the opportunity to fund the printing and distribution of this manual, which is published jointly by the Fisheries Department of the FAO and the Asia-Pacific Economic Cooperation (APEC). The FAO has earlier published a guide for the implementation of its International Plan of Action - Sharks that was prepared by Dr Terry Walker of the Marine and Freshwater Resources Institute, Victoria, Australia. This publication provides a synoptic guide to the goals and requirements for fisheries authorities to start addressing the needs for the management of national elasmobranch fisheries; it has been prepared in a highly condensed style consistent with the other sister publications in the same series. The present, far more comprehensive manual, edited by John A. (Jack) Musick and Ramón Bonfil, provides detailed information for the operational fisheries manager about methods to collect and analyse data necessary to assess stocks and prepare plans to sustainably manage elasmobranch fisheries. This manual begins by establishing the objectives of fisheries management and then provides information about shark and ray identification with a key to families and a guide to regional keys. Tagging methods are then reviewed including tag data analyses. A chapter on stock structure and identification follows outlining genetic methods in particular. Two chapters respectively deal in detail with methods necessary to determine age and growth rates in sharks, and to define reproductive biology. Another chapter describes methods to estimate mortality. All of these subjects are required in order to move on to demographic modelling and stock assessments which are covered in the subsequent chapters. Data collection in the field is covered in the next two chapters, one on fishery dependent and the other on fishery independent sampling. The manual ends with chapters on management techniques, and shark utilization. Collectively, the authors have produced a manual that will be invaluable not only to those involved in management of these highly diverse and fascinating fishes and their fisheries, but also to the reader whose responsibilities extends beyond the direct involvement of resources analysis and who is involved in the management of other groups of fishes. For this, the editors and authors deserve gratitude and compliments from those who will benefit from their experience, wisdom and endeavours. R. Shotton Focal Point, FAO International Plan of Action Sharks Marine Resources Service, Fisheries Department FAO, Rome. 1 FAO. 2000. Fisheries management. 1. Conservation and management of sharks. FAO Technical Guidelines for Responsible Fisheries. No. 4, Suppl. 1. Rome, FAO. 37p.

iv Abstract This publication describes the scientific principles and techniques used for resource management of elasmobranch fisheries with emphasis on the particular context of elasmobranchs. The management characteristics of these fishes are described their common bycatch character and their biological constraints on productivity low growth rate, late maturity and low fecundity. Stock assessment of elasmobranchs is described in the context of management objectives in a wide management context. Special attention is given to accurate species identification given the prevalent aggregating of landings data across species, genera and often families in this group. Techniques and experiences for tagging elasmobranchs for population estimation are described and methods of genetic techniques for stock identification. Methods and problems involved in determining age, growth, fecundity and mortality rates are described and their use in age-structured models within the context of the reproductive biology of these fishes. Demographic models to determine the productivity of elasmobranch resources are described. Use of surveys to complement information derived from fisheries is described together with management measures. Last, practices of shark utilization are noted. Key words: Fishery management, elasmobranchs, sharks, IPOA-sharks, rays, mortality, fecundity, ageing, reproductive biology, growth, stock assessment, species identification, tagging, genetic identification, utilization. Musick, J.A.; Bonfil, R. (eds) Management techniques for elasmobranch fisheries. FAO Fisheries Technical Paper. No. 474. Rome, FAO. 2005. 251p.

v Contents Foreword Abstract Contributors Acknowledgements Dedication iii iv vi vii vii 1. INTRODUCTION: MANAGEMENT OF SHARKS AND THEIR RELATIVES (ELASMOBRANCHII) 1 John A. Musick 2. THE PURPOSE OF STOCK ASSESSMENT AND THE OBJECTIVES OF FISHERIES MANAGEMENT 6 Ramón Bonfil 3. TAXONOMY AND FIELD TECHNIQUES FOR IDENTIFICATION AND AVAILABLE REGIONAL GUIDES 15 John D. Stevens 4. TAGGING METHODS AND ASSOCIATED DATA ANALYSIS 45 Robert J. Latour 5. GENETICS: STOCK IDENTIFICATION 62 Edward J. Heist 6. AGE AND GROWTH OF ELASMOBRANCH FISHES 76 Kenneth J. Goldman 7. REPRODUCTIVE BIOLOGY 103 Christina L. Conrath 8. MORTALITY ESTIMATION 127 Colin A. Simpfendorfer, Ramón Bonfil and Robert J. Latour 9. DEMOGRAPHIC MODELS: LIFE TABLES, MATRIX MODELS AND REBOUND POTENTIAL 143 Colin A. Simpfendorfer 10. FISHERY STOCK ASSESSMENT MODELS AND THEIR APPLICATION TO SHARKS 154 Ramón Bonfil 11. FISHERY-DEPENDENT SAMPLING: TOTAL CATCH, EFFORT AND CATCH COMPOSITION 182 Alexia C. Morgan and George H. Burgess 12. FISHERY INDEPENDENT SAMPLING: SURVEY TECHNIQUES AND DATA ANALYSES 201 Paul J. Rago 13. MANAGEMENT MEASURES 216 Terence I. Walker 14. SHARK UTILIZATION 243 John A. Musick

vi Contributors Ramón Bonfil, Wildlife Conservation Society, 2300 Southern Blvd., Bronx, NY, 10460, USA. E-mail: rbonfil@wcs.org George H. Burgess, Florida Museum of Natural History, Division of Fishes, University of Florida, Gainsville, Florida, USA. E-mail: gburgess@flmnh.ufl.edu Christina L. Conrath, Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, Virginia 23062, USA. E-mail: conrath@vims.edu Kenneth J. Goldman, Alaska Department of Fish and Game, 3298 Douglas Place, Homer, AK 99603, USA. E-mail: ken_goldman@fishgame.state.ak.us Edward J. Heist, Fisheries and Illinois Aquaculture Center, Southern Illinois University at Carbondale, 1125 Lincoln Drive, Carbondale, IL 62901-6511, Carbondale, Illinois, USA E-mail: edheist@siu.edu Robert J. Latour, Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, Virginia, 23062, USA. E-mail: latour@vims.edu Alexia C. Morgan, Florida Museum of Natural History, Division of Fishes, University of Florida, Gainsville, Florida, USA. John A. Musick, Virginia Institute of Marine Science, College of William and Mary, PO Box 1346, Gloucester Point, Virginia 23062, USA. E-mail: jmusick@vims.edu Paul Rago, Population Dynamics Branch, National Marine Fisheries Service, 166 Water St., Woods Hole, MA 02543, USA. E-mail: paul.rago@noaa.gov Colin A. Simpfendorfer, Center for Shark Research, Mote Marine Laboratory, 1600 Ken Thompson Parkway, Sarasota, FL 34236, USA. E-mail: colins@mote.org John D. Stevens, Marine Research, Commonwealth Scientific and Industrial Research Organization (CSIRO), Hobart, Tasmania, Australia. E-mail: john.d.stevens@csiro.au Terence I. Walker, Marine and Freshwater Resources Institute, PO Box 114, Queenscliff, Victoria 3225, Australia. E-mail: Terry.Walker@dpi.vic.gov.au

vii Acknowledgements Thanks are due to Malanie Harbin whose efforts in editorial management of this project were invaluable and to Ruth Hershner who was responsible for the layout and draft production of this publication. In addition, we thank Enric Cortes, John Graves, William Hamlett, Rob Latour and Colin Simpfendorfer for their efforts in providing scientific reviews of manuscripts. Lastly, thanks are due to Sonja Fordham, Stetson Tinkham, Colin McIff and the APEC Fisheries Working Group without whom this project would not be possible. This is a contribution from the IUCN Shark Specialists Group, the National Shark Research Consortium and also Virginia Institute of Marine Science Contribution No. 2562. Dedication This book is dedicated to the shark fishers and fishery managers. May they have the wisdom and will to achieve and maintain sustainable shark fisheries.