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The balance between feeding and predation 131<br />

of Acanthuridae 236 . Therefore, the larval phase is divided into three time<br />

periods, to account for the changes in swimming abilities. The limits<br />

of those three periods are defined approximately at ontogenetic shifts:<br />

end of yolk sac period and end of flexion. Swimming speed values<br />

are estimated from published records 60,236,258 . Most studies measured<br />

critical swimming speed (maximal speed of a current against which a<br />

larva can maintain its position). These speeds are probably greater than<br />

actual swimming speeds in the field 258 . Therefore, we choose lower<br />

swimming speeds for both species in each time period, while retaining<br />

the difference observed between them: 0, 13, and 36 cm s -1 for the<br />

Acanthuridae and 3, 10, and 20 cm s -1 for the Pomacentridae.<br />

During the first period, larvae extract energy from their yolk sac<br />

reserves, hence do not need to forage. In the model, their energy resources<br />

are constant and maximal. Afterward, they lose one energy<br />

unit per time step. As they have a maximum resource of five units they<br />

can only swim four time steps (24 h) before food is required. Coral<br />

reef fish larvae can swim for longer periods of time before starving (up<br />

to 194 h for Acanthuridae for example) 236 . Nevertheless, in the field,<br />

larvae are likely to avoid starvation and keep their energy resources<br />

level as high as they can. Furthermore, most studies about the swimming<br />

endurance of reef fish larvae focus on the time that a larva can<br />

swim against a current before starving or being completely exhausted,<br />

without any consideration about maintaining growth rate or integrity<br />

of metabolic pathways. However, the daily food intake of fish larvae<br />

needed to maintain their growth rate is high (50% of body weight per<br />

day is a general mean 232 ), especially for fast-growing, warm-water fish<br />

larvae 232 . Therefore, fish larvae should eat often, probably on a daily<br />

basis, during dispersal.<br />

. . . and their swimming<br />

abilities<br />

Energy intake is<br />

limiting in fish larvae<br />

Finally, the parameters for this more elaborate model can be summarised<br />

along the guidelines set by the modelling framework.<br />

Time 6 hours time step; horizon of 50 days for Acanthuridae and<br />

20 days for Pomacentridae<br />

State Energy reserves (five levels, one consumed per 6 h of swimming)<br />

and three-dimensional position in a 100 km × 50 km × 100 m<br />

domain of 720 m horizontal mesh size and 50 m vertical mesh<br />

size.<br />

Environment Spatially explicit survival and feeding probabilities; concentration<br />

around the island is described by factor f; f = 1 means<br />

homogenous repartition; as f increases, the island effect is stronger<br />

and food as well as predators are more concentrated near shore;<br />

f = 1, 1.1, 1.2 and 1.4 are tested. Static current field with incoming<br />

flow speed of 10 cm s -1 .

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