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142 Oceanography vs. behaviour<br />

Simple Euler-forward<br />

advection is appropriate<br />

the steep topography of the island or the promontory would otherwise<br />

be smoothed). The time step of the biological model is 3 hours and the<br />

output of the ROMS is stored at the same frequency so no temporal<br />

interpolation is needed.<br />

Larvae are released all around the island and the promontory, 1 km<br />

away from the 10 m isobath. The spawning strategy of adults may<br />

influence where larvae are released and how they are initially advected.<br />

For example, some tropical marine fishes were observed to spawn near<br />

the surface when tides entrain eggs toward the open ocean 185 . While<br />

potentially important, these behaviours are very location-specific and<br />

require fine scale representations of the topography and currents near<br />

the coast. This model focuses on general mesoscale features and the<br />

configuration used in the ROMS does not resolve fine scale structures,<br />

so we are only interested in what happens once the first moments of<br />

dispersal are over, 1 km away from shore. From there on, particles are<br />

advected using a simple Euler forward scheme with a 3 h time step.<br />

Particles are active in this model and de-correlate from the flow at a<br />

rate very different from that of passive particles. Therefore elaborate<br />

Lagragian advection methods, which feature a fading memory of currents<br />

in a random flight scheme or parameterised diffusion in a random<br />

walk one for example, cannot be used (see section 1.4.5, page 35). Finer<br />

advection schemes (e.g. Runge-Kutta) or shorter time steps could be<br />

used, but end positions would still have to be brought back to grid<br />

nodes. Indeed, the optimisation is performed at those points only, and<br />

optimal decisions cannot be interpolated (see Note – Computer memory<br />

and speed, page 129).<br />

6.4.2 Continuous and quantitative description of swimming<br />

behaviour<br />

In this model, at each time step, larvae can choose between several<br />

swimming speeds, oriented toward twenty-five different directions<br />

homogeneously distributed in space. In addition, instead of a three<br />

step progress, as in section 6.3, the development of swimming speed is<br />

described in a continuous fashion. To model swimming continuously<br />

in time, the ontogeny of maximum sustainable swimming speed and of<br />

swimming endurance (i.e. of the energetics of swimming) have to be<br />

described. Unfortunately, there are still very few observations of those<br />

variables throughout the larval phase.<br />

Development of swimming abilities and temperature effects<br />

Continuous, almost<br />

linear, development<br />

of swimming speed<br />

During ontogeny, maximum swimming speed increases because of<br />

allometry: even with a constant speed in body length per second (bl s -1 ),<br />

the actual speed in cm s -1 increases as larvae grow. However, on top of<br />

that, fin and muscle develop and the speed in bl s -1 actually increases<br />

during larval life 95 . A handful of studies 56,57,60 described the evolution

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