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African Journal of Marine Science a Review and Tests of Hypotheses This article was downloaded by: On: 16 November 2010 Access details: Access Details: Free Access Publisher Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37- 41 Mortimer Street, London W1T 3JH, UK African Journal of Marine Science Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t911470580 A review and tests of hypotheses about causes of the KwaZulu-Natal sardine run P. Fréona; J. C. Coetzeeb; C. D. van der Lingenbc; A. D. Connelld; S. H. O'Donoghuee; M. J. Robertsf; H. Demarcqa; C. G. Attwoodc; S. J. Lamberthbd; L. Hutchingscf a IRD, Sète, France b Branch Fisheries, Department of Agriculture, Forestry and Fisheries, Rogge Bay, South Africa c Marine Research Institute, University of Cape Town, Rondebosch, South Africa d South African Institute for Aquatic Biodiversity, Grahamstown, South Africa e School of Biological and Conservation Sciences, Westville Campus, University of KwaZulu-Natal, Durban, South Africa f Oceans and Coasts, Department of Environmental Affairs, Rogge Bay, South Africa Online publication date: 08 November 2010 To cite this Article Fréon, P. , Coetzee, J. C. , van der Lingen, C. D. , Connell, A. D. , O'Donoghue, S. H. , Roberts, M. J. , Demarcq, H. , Attwood, C. G. , Lamberth, S. J. and Hutchings, L.(2010) 'A review and tests of hypotheses about causes of the KwaZulu-Natal sardine run', African Journal of Marine Science, 32: 2, 449 — 479 To link to this Article: DOI: 10.2989/1814232X.2010.519451 URL: http://dx.doi.org/10.2989/1814232X.2010.519451 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.informaworld.com/terms-and-conditions-of-access.pdf This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. African Journal of Marine Science 2010, 32(2): 449–479 Copyright © NISC (Pty) Ltd Printed in South Africa — All rights reserved AFRICAN JOURNAL OF MARINE SCIENCE ISSN 1814–232X EISSN 1814–2338 doi: 10.2989/1814232X.2010.519451 A review and tests of hypotheses about causes of the KwaZulu-Natal sardine run P Fréon 1*, JC Coetzee 2, CD van der Lingen 2,3 , AD Connell 4, SH O’Donoghue 5, MJ Roberts 6, H Demarcq 1, CG Attwood 3, SJ Lamberth 2,4 and L Hutchings 3,6 1 IRD, UMR 212 EME, CRHMT, avenue Jean Monnet, 34209, Sète, France 2 Branch Fisheries, Department of Agriculture, Forestry and Fisheries, Private Bag X2, Rogge Bay 8012, South Africa 3 Marine Research Institute, University of Cape Town, Private Bag X3, Rondebosch 7701, South Africa 4 South African Institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown 6140, South Africa 5 School of Biological and Conservation Sciences, Westville Campus, University of KwaZulu-Natal, Private Bag X54001, Durban 4000, South Africa 6 Oceans and Coasts, Department of Environmental Affairs, Private Bag X2, Rogge Bay 8012, South Africa * Corresponding author, e-mail : [email protected] Manuscript received June 2010; accepted August 2010 The term ‘sardine run’ is part of the cultural heritage of the South African nation and refers to a natural phenomenon that is well known to the general public but still poorly understood from an ecologi- cal perspective. This lack of understanding has stimulated numerous hypotheses, often contradic- tory, that try to explain why (ultimate factors) and how (proximate factors) the run occurs. Here, we provide a new definition of the term sardine run, review the various hypotheses about the run, and propose ways to test those hypotheses. Where possible, the results of tests that have been conducted thus far are presented and discussed. Our interpretation of the causes is that the sardine run most likely corresponds to a seasonal (early austral winter) reproductive migration of a genetically distinct subpopulation of sardine that moves along the coast from the eastern Agulhas Bank to the coast of KwaZulu-Natal (KZN) as far as Durban and sometimes beyond, in most years if not in every year. This eastward migration is constrained close to the coast by the thermal preference of sardine and the strong and warm offshore Agulhas Current. The run is facilitated by the presence of a band of cooler coastal water and by the occurrence of Natal Pulses and break-away eddies that enable sardine shoals Downloaded At: 21:50 16 November 2010 to overcome their habitat restrictions. These enabling mechanisms are most important in the area where the shelf is at its narrowest and feature most prominently off Waterfall Bluff, which has led to the coining of the ‘Waterfall Bluff gateway hypothesis’. Based on the collection of eggs off the KZN coast, sardine remain there for several months and their westward, return migration during late winter to spring is nearly always unnoticeable because it likely occurs at depth as the fish avoid warmer surface waters. Years in which the sardine run is not detected by coastal observers could reflect either its real absence due to high water temperatures and/or other hydrographic barriers, or an eastward migration that is farther offshore and possibly deeper and is enabled by hydrographical anomalies. Keywords: migration, proximate factors, Sardinops saga x, South Africa, ultimate factors Introduction The term ‘sardine run’ (or ‘KwaZulu-Natal sardine run’) poorly understood, as evidenced by numerous hypotheses, is part of the cultural heritage of the South African nation often contradictory, that try to explain why and how the and refers to a well-known natural phenomenon off the sardine run occurs. In this paper, we first attempt to provide east coast of South Africa. The sardine run has long been a neutral and consensual meaning of the term sardine run known by the coastal population in that region because of as perceived by the general public and scientists alike. its economic importance (van der Lingen et al. 2010a), and Because of the commercial/recreational/public nature of over the last few decades has been a topic of much media the sardine run, which requires some kind of management attention and become the focus of a growing tourism industry response that will depend on how accurately the timing (Dicken 2010, Myeza et al. 2010). Despite this heightened and intensity of the ‘run’ can be predicted, our description interest, the ecological aspects of this phenomenon remain of what constitutes the run focuses on its visible aspects African Journal of Marine Science is co-published by NISC (Pty) Ltd and Taylor & Francis 450 Fréon, Coetzee, van der Lingen, Connell, O’Donoghue, Roberts, Demarcq, Attwood, Lamberth and Hutchings when it reaches the coast and elicits a human reaction. The terms ‘ultimate’ and ‘proximate’ to refer to different factors ecological meaning of the sardine run and its management influencing behaviour. Ultimate refers to the final, long-term, implications are separate issues that are also addressed in evolutionary consequences of behaviour (function), whereas this paper. proximate refers to the immediate, short-term, physiological We define the term sardine run as the visible effects of the mechanisms of behaviour (causation). Instantaneous habitat coastal, alongshore movement during early austral winter of selection is related to proximate cues and is expected to a small and variable fraction of the South African population occur on a small spatial scale (microhabitat) in response of sardine Sardinops sagax from the eastern Agulhas Bank to changes in the environment. A common example of a to the KwaZulu-Natal (KZN) coast, as far as Durban and the proximate factor, likely relevant to the sardine run, is the north coast of KZN. The sardine run is usually associated avoidance of warm waters and associated oxygen depletion. with foraging top predators including seabirds, mammals This instantaneous selection and its relationship with the (O’Donoghue et al. 2010a, 2010b), and sharks and gamefish environment are usually easy to observe and quantify. In (Dudley and Cliff 2010, Fennessy et al. 2010) that facilitate contrast, ultimate relationships may be much more difficult to its visual detection. The north-east displacement is usually resolve. Additionally, once a selective pressure has led to the initiated from late May to early June and is mostly visually evolution of a particular behaviour, that behaviour may last detected and commercially exploited (via beach-seine for epochs even when major change occurs in the environ- netting and tourism operations) when it occurs within a few ment, providing that the behaviour (termed relic behaviour) hundred metres of the coast. The duration of this coastal has at most a minimal cost regarding survival and reproduc- spectacle is highly variable, lasting until the end of June or tive success. Examples of ultimate factors are availability of even September at times (O’Donoghue et al. 2010c). Little is appropriate food, low potential predation (not instantaneous known about farther offshore movements of sardine and even predator avoidance) or favourable conditions for reproduc- less about the likely return of surviving fish to the Agulhas tive success. Bank thereafter, although eggs in plankton samples suggest An understanding of ultimate causes should also provide that sardine remain in KZN waters until as late as December insight into the long-term effects of fishing on movement (Connell 2010).
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