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The Reproductive Biology, Condition and Feeding Ecology of the Skipjack, Katsuwonus Pelamis, in the Western Indian Ocean

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The Reproductive Biology, Condition and Feeding Ecology of the Skipjack, Katsuwonus Pelamis, in the Western Indian Ocean The reproductive biology, condition and feeding ecology of the skipjack, Katsuwonus pelamis, in the Western Indian Ocean Maitane Grande Mendizabal PhD Thesis Department of Zoology and Animal Cell Biology 2013 The reproductive biology, condition and feeding ecology of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean Tesi zuzendariak, Hilario Murua eta Nathalie Bodin Maitane Grande Mendizabal 2013ko Apirilaren 26ª TESIAREN ZUZENDARIAREN BAIMENA TESIA AURKEZTEKO Hilario Murua Aurizenea jaunak, 34108169-C I.F.Z. zenbakia duenak “The reproductive biology, condition and feeding ecology of the skipjack; Katsuwonus pelamis , in the Western Indian Ocean” izenburua duen doktorego-tesiaren zuzendari naizenak, tesia aurkezteko baimena ematen dut, defendatua izateko baldintzak betetzen dituelako. Maitane Grande Mendizabal doktoregai andreak egin du aipaturiko tesia, AZTI Tecnalia-ko Itsas-Ikerketa sailean. Pasaia, 2013(e)ko Apirilaren 23a TESIAREN ZUZENDARIA Iz.: Hilario Murua Aurizenea TESIAREN ZUZENDARIAREN BAIMENA TESIA AURKEZTEKO Nathalie Bodin andreak, pasaporte zenbakia: 07AX12437 eta 060537202886 I.F.Z . zenbakia duenak “The reproductive biology, condition and feeding ecology of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean” izenburua duen doktorego-tesiaren zuzendari naizenak, tesia aurkezteko baimena ematen dut, defendatua izateko baldintzak betetzen dituelako. Maitane Grande Mendizabal doktoregai andreak egin du aipaturiko tesia, AZTI Tecnalia-ko Itsas Ikerketa sailean. Victoria, Seychelles, 2013(e)ko Apirilaren aren 15a TESIAREN ZUZENDARIA Iz.: .Nathalie Bodin TESI-BEGIRALEAREN BAIMENA TESIA AURKEZTEKO Nik, Maren Ortiz Zarragoitia jaunak, tesi-begiralea naizen aldetik, baimena ematen dut honako tesi hau aurkeztua izateko, horretarako baldintza guztiak biltzen baititu: Tesia: “The reproductive biology, condition and feeding ecology of the skipjack; Katsuwonus pelamis , in the Western Indian Ocean” Tesia egin den saila: Zoologia eta Animali Zelulen Biologia Tesiaren egilea: Maitane Grande Mendizabal Tesiaren zuzendaria: Hilario Murua Aurizenea eta Nathalie Bodin Leioa (e)n, 2013ko Apirilaren 24 (e)an TESI-BEGIRALEA Sin.: Maren Ortiz Zaragoitia SAILAREN ADOSTASUNA Zoologia eta Animalia Zelulen Biologia Saileko Kontseiluak, 2013ko Apirilaren 10eko (e)ko bileran, “The reproductive biology, condition and feeding ecology of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean” izenburua duen doktorego-tesia aurkeztearen alde dagoela adierazi du. Hilario Murua eta Nathalie Bodin jaunaren/andrearen zuzendaritzapean egin den tesi hori Maitane Grande Mendizabal jaunak/andreak aurkeztu du sail honetan. ___________________, _______(e)ko ________________aren ______a O. E. SAILEKO ZUZENDARIA SAILEKO IDAZKARIA Iz.: __________________________ Iz.:______________________ DOKTORE GRADUAREN AKTA DOKTOREGO TESI DEFENTSAREN AKTA Doktoregai jn./and.: Maitane Grande Mendizabal TESIAREN IZENBURUA: The reproductive biology, condition and feeding ecology of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean UPV/EHUko Doktoregoko Azpibatzordeak goian adierazitako doktorego tesia kalifikatzeko aukeratutako epaimahaiak, eta adierazitako egunean bilduta, behin doktoregaiak defentsa eginda eta egin zaizkion objekzioak edota iradokizunak erantzunda, ___________________ (Aho batez edo gehiengoaz) eman du honako kalifikazioa: GAI edo EZ GAI Defentsan erabilitako hizkuntzak (hizkuntza bat baino gehiago erabili badira, zehaztu hizkuntza bakoitzean defendatutako atalak edo ehunekoak): ______________________________________________________ (e)n, 20 (e)ko aren (e)an PRESIDENTEA, IDAZKARIA Sin.: Sin.: Dk: ____________________ Dk: ______________________ 1. mahaikidea, 2. mahaikidea, 3. mahaikidea, Sin.: Sin.: Sin.: Dk: Dk: Dk: DOKTOREGAIA, Sin.: _____________________ Acknowledgement Lehen bizi eskerrak eman nahi dizkiet tesi zuzendariei. Hilariori, zuk gidatu nauzulako eta beti prest egon zarelako nire kezkak eta zalantzak entzuteko. Zure aholkuak eta animoak ezinbestekoak izan direlako. Nathalie, I would like to thank you for your dedication and patience. Your enthusiasm and passion encorauged me to continue working hard along this long project. Bekaduneei, eskerrak zuek alboan izan zaituztedala! Frikadak eta martzianadak norekin konpartittu bestela? Animo pila zuei ere! We can do it! Laborategiko neskeei, laborategiko ordu luzeak dibertigarriak egin dituzuelako. Imma, Maiteri eta Ireneri laginketa prestatzen, eskaerak egiten, eta laborategian eta almazenean ordena mantentzen laguntzeagatik. A Jefferson, Antonio y Gorka por vuestro valioso trabajo realizado en el mar. Lagunei, zuekin barre eta negar egin dudalako eta atun mundua alde batera usten eta arnasa hartzen lagundu nauzuelako. Gora bihotzak! Ikerri, lan hau biona delako eta zurekin ibilitako bide hau errazagoa izan delako. Mil esker benetan! Xikerri, beti izan duzulako irribar bat prest opari gisa. Familiari, por vuestra ayuda y apoyo incondicional. Por los sueños oceanicos, y el azul que nos atrapa. Ondoan (nahiz eta Australian) zineten eta zaudetenoi. The work carried out in this Dr. Philos Thesis has been developed in AZTI-Tecnalia with the collaboration of IRD (Institute de Recherché pour le Developpement) and it has been financed by: • Fundación Centros Tecnológicos Iñaki Goenaga. • PEVASA - Pesquería Vasco Montañesa fishing company. • Department of Agriculture, Fisheries and Food of the Basque Government. We are also grateful for the contribution received from the EMOTION project (ANR JSV7 007 01). Part of the information contained in this Dr. Philos Thesis has been or will be published on the scientific articles: • Grande, M., Murua, H., Zudaire, I., Korta, M. 2012. Oocyte development and fecundity type of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean. J. Sea Res. 73, 117-125. • Grande, M., Murua, H., Zudaire, I., Goñi, N., Submitted. Sex ratio, reproductive timing and batch fecundity of the skipjack in the Western Indian Ocean. J. Fish Biol. • Grande, M., Murua, H., Pernet, E. J. A., Pernet, F., Zudaire, I., Bodin, N. Submitted. Lipids in female skipjack, Katsuwonus pelamis , tuna from the western Indian Ocean: Influence of the reproductive cycle and the fishing environment. Mar. Biol. • Grande, M., Murua, M., Zudaire, I., Potier, M., Ménard, F., Goñi, N., Bodin, N., Submitted. Link between the diet and reproductive activity of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean. Mar. Ecol. Prog. Ser. Contents 1. Laburpena……………………………………………………………………………1 2. Summary……………………………………………………………………………..9 3. Overview 3.1. Context and relevance of the work…………………………………………… 19 3.2. Working hypothesis………………………………………………..…………..23 3.3. Objectives.……………………………………………………………………..24 3.4. Structure of the thesis………………………………………………………….25 4. Introduction 4.1. Taxonomic classification and short description……………………………….29 4.2. Distribution, stock structure, habitat and physiological requirements………...30 4.3. Skipjack fishery………………………………………………………………..33 4.3.1. Fishery and state of the stock in the Indian Ocean……………………..36 4.3.2. Potentital effects of FADs on skipjack populations……………………39 4.4. Biology 4.4.1. Age and growth………………………………………………………...40 4.4.2. Natural mortality…………………………………………………….....44 4.4.3. Reproduction…...………………………………………………………45 4.4.4. Early life history and recruitment………………………………………52 4.4.5. Trophic ecology………………………………………………………...55 4.4.6. Life history strategy and energy allocation…………………………….57 5. Study area…………………………………………………………………………..59 6. Chapter I: Oocyte development and fecundity type of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean……………………………………………...67 7. Chapter II: Sex ratio, reproductive timing and reproductive capacity of the skipjack tuna Katsuwonus pelamis in the Western Indian Ocean……………………………87 8. Chapter III: Lipids in female skipjack tuna from the Western Indian Ocean: Influence of the reproductive cycle and the fishing environment………………...109 9. Chapter IV: Link between the reproductive activity, feeding strategy and isotopic composition of the skipjack, Katsuwonus pelamis , in the Western Indian Ocean…………………………………………………………….135 10. General Discussion………………………………………………………………..159 11. Conclusions………………………………………………………………………181 12. References..………………………………………………………………………187 13. Annex...………………………………………………………………………...…231 1. Laburpena Laburpena Egun, lanpo sabel-marradunaren ( Katsuwonus pelamis ) arrantza, munduko hiru ozeanoetako eremu tropikal eta subtropikaletan burutzen dena, arrantza jarduera ekonomiko garrantzitsuenetarikoa bilakatu da. Horren adibide urtero harrapatzen diren interes komertziala duten atunen 4.2 milioi tonetatik %58a lanpo sabel-marraduna izatea delarik (Majkowski et al. , 2011; ISSF, 2012). Indiako Ozeanoan atun espezie honen harrapaketa guztizkoaren %17a (398,000 tona) da, arrantza honek duen garrantziaren argi islatuz (Arrizabalaga et al ., 2012; ISSF, 2012). Atun arrantzaren jarduera nagusia, Indiako Ozeanoaren mendebaldean burutzen da. Espezie honek erakuzten duen produktibitate altua dela eta, Indiako Ozeanoaren Atun Batzordeak (Indian Ocean Tuna Commission IOTC ingelesez) burututako azkeneko ebaluazioak ez du lanpo sabel- marradunaren harrapaketak mugatzeko beharrik aurreikusi espeziaren egoera ona dela eta (IOTC, 2012). Hala ere, Indiako Ozeanoaren mendebaldean espezie honi dagokion ugal biologia eta ekologia ikertzen duten lanak ez dira ugariak eta informazioa ez dago eguneratuta, nahiz eta ezinbeztekoa izan arrantza kudeaketa egoki bat gauzatzeko. Hori dela
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