Behavioural energetics of the parrotfish Sparisoma viride

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1 RIJKSUNIVERSITEIT GRONINGEN Behavioural energetics of the parrotfish Sparisoma viride Flexibility in a coral reef setting PROEFSCHRIFT ter verkrijging van het doctoraat in de Wiskunde en Natuurwetenschappen aan de Rijksuniversiteit Groningen op gezag van de Rector Magnificus dr. F. van der Woude, in het openbaar te verdedigen op vrijdag 31 mei 1996 des namiddags te 4.00 uur door Julius Martinus van Rooij geboren op 27 juni 1960 te Doetinchem

2 Promotor: Co-promotor: Referent: Prof. dr. C. van den Hoek Dr. J.J. Videier Dr. A.M. Breeman

3 Stellingen behorende bij het proefschrift Behavioural energetics of the parrotfish Sparisoma viride Flexibility in a coral reef setting van Jules van Rooij I) De rol van het "small food web" op koraalriffen is tot nu toe onderschat, hetgeen echter niet betekent dat de rol van macroherbivoren overschat is. (dit proefschrift) 2) Gezien de overvloed aan zonlicht en kooldioxide is er geen enkele reden waarom op koraalriffen efficiënt met energie of koolstof omgesprongen zou moeten worden. Energiebudgetstudies zijn derhalve niet de aangewezen methode om de veronderstelde hoge oecologische efficiëntie van koraalriffen aan te tonen. 3) Tot in de jaren zestig werden initiële en terminale fase Sparisoma viride individuen als verschillende soorten aangemerkt, hetgeen verklaart waarom de soortsnaam alleen van toepassing is op de (groene) mannen. Gegeven het feit dat alle dieren het mannelijk geslacht kunnen aannemen, lijkt Sparisoma viriie een meer toepasselijke naam. 4) De rol van stochastische planktonische processen op de organisatie van koraalrif-visgemeenschappen is lang verwaarloosd maar dreigt de laatste jaren overschat te worden. (Jones 1991, contra Doherty & Williams 1988) 5) Een hoge algenbiomassa kan evenzeer duiden op een ongeschikt voedseltype als op een hoog voedselaanbod voor herbivoren. De kwaliteit van een papegaai vissen-territorium kan dan ook niet afgemeten worden aan uitsluitend de hoeveelheid hoge algenturfjes. (contra elifton 1989) 6) De spring- en doodtij cyclus loopt in een groot deel van het Canübisch gebied niet in fase met die der schijngestalten van de maan (T = 29,5 d), maar met die van de maandeclinatie CT = 27,3 d). Synchronisatie van de voortplantingscyclys van riforganismen met de schijngestalten is derhalve veelal ten onrechte in verband gebracht met de verspreiding van de larven. (contra Robertson et al. 1990) 7) De trend dat tijdschriften als gevolg van het toenemende aanbod van publikaties puur beschrijvend werk afwijzen zal averechts werken: nauwkeurige veldwaamemingen zijn vereist ten einde de relevantie te kunnen beoordelen van hypothesen die experimenteel getoetst worden. 8) Steeds krachtiger PC's en software vergemakkelijken het verwerken van grote datasets, als gevolg waarvan steeds kleinere verschillen statistisch significant gemaakt kunnen worden. Opgave van een 'relevantie-index' naast de verplichte P-waarden verdient derhalve aanbeveling. 9) Het succes van de NAVO-doctrine van de "flexible response" wordt meer bevestigd door de wijdverbreidheid van Sparisoma viride dan door het vallen van de muur. 10) Het verkorten van de aanstellingsduur van post-doe onderzoekers in verband met de wachtgeld-problematiek vormt het begin van een neerwaartse spiraal en biedt niet meer dan een zeer tijdelijke oplossing. Bij gebrek aan middelen kan beter het wachtgeld afgeschaft worden.

4 "Within the shadow of the ship I watched their rich attire: Blue, glossy green, and velvet black, They coiled and swam; and every track Was a flash of golden fire." (Uit: The rime of the ancient mariner Samuel Taylor Coleridge) Aan mijn ouders, Will en Annemarie van Rooij

5 /'l NWO Cover design: Dick Vis er Printed by Ponsen & Looijen bv, Wageningen ISBN: The re earch reported in thi thesis wa undertaken under the authority of the University of Groningen, Department of Marine Biology, P.O. Box 14, 9750 AA Haren, The Netherlands. This study was supported by the Netherlands Foundation for the Advancement of Tropical Research (WOTRO; grant no. W88-137), a subsidiary organization of the Netherlands Organization for the Advancement of Re earch (NWO).

6 Behavioural Energetics of Sparisoma viride Contents Voorwoord Overzicht & Samenvatting 3 Chapter General Introduction 13 Chapter 2 The social and mating system of the herbivorous reef fish 23 Sparisoma viride: one-male versus multi-male groups J.M. van Rooij, F.J. Kroon & J.J. Vide/er Environmental Biology of Fishes: in press Chapter 3 Resource and habitat sharing by the stoplight parrotfish 55 (Sparisoma viride), a Caribbean reef herbivore J.M. van Rooij, E. de Jong, F. Vaandrager & J.J. Vide/er Environmental Biology of Fishes: in press Chapter 4 Local variability in population structure and density of the 67 protogynous reef herbivore Sparisoma viride J.M. van Rooij, J.P. Kok & J.J. Vide/er Environmental Biology of Fishes: in press Chapter 5 Chapter 6 Chapter 7 A simple field method for stereo-photographic length 'measurement of free-swimming fish : merits and constraints J.M. van Rooij & J.J. Vide/er Journalof Experimental Marine Biology and Ecology: in press Ontogenetic, social, spatial and seasonal variations in condition of the reef herbivore Sparisoma viride J.M. van Rooij, J.H. Bruggemann, J.J. Vide/er & A.M. Breeman Marine Biology 123: , 1995 Plastic growth of the herbivorous reef fish Sparisoma viride: field evidence for a trade-off between growth and reproduction o J.M. van Rooij, J.H. Bruggemann, JoJ. Vide/er & A.M. Breeman Marine Ecology Progress Series 122: , of ~he cal lds Chapter 8 Mortality and other demographic inferences from repeated 127 visual surveys of a Sparisoma viride population. J.M. van Rooij & JoJ. Vide/er Marine Ecology Progress Series: submitted Chapter9 Estimating oxygen uptake rate from ventilation frequency in the 145 reef fish Sparisoma viride. J.M. van Rooij & J.J. Vide/er Marine Ecology Progress Series 132: 31-41, 1996

7 Contents Chapter 10 Energetics and territoriality of the reef herbivore Sparisoma viride J.M. van Rooij & J.J. Vide/er Ecological Monographs: submitled 161 Chapter 11 Summary References General Discussion

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