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Dissertation - HQ

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Oriented swimming and passive advection 159<br />

decisions more evenly in time. In addition, the recovery capacity of<br />

older larvae would probably be higher than that of younger stages.<br />

Therefore, swimming at maximum speed may become optimal at the<br />

end of the larval phase. Second, the relationship defining the energetic<br />

cost of swimming was identical for young and old larvae. The reasoning<br />

behind this null-hypothesis is detailed on page 146. Yet, the eventuality<br />

that swimming would be relatively more costly for young stages than<br />

for old ones — as the authors initially suggested — has to be considered.<br />

With a similar criterion of minimum energy expenditure, this should<br />

result is less swimming at the beginning of larval life. Third, early larvae<br />

may not have the sensory abilities to detect even local cues and orient in<br />

consequence. Though there is evidence that even clownfish embryos can<br />

detect sounds waves in the range of those produced by coral reef communities<br />

255 , knowledge is currently lacking in this area. Nevertheless,<br />

the two energetic constraints can be discussed. Even if early swimming<br />

is frequently observed among optimal trajectories (Figure 6.18), it does<br />

not mean that all larvae swim repeatedly for the first days after hatching.<br />

On the contrary, swimming decisions along any single optimal<br />

trajectory are quite sparse (Figure 6.17). In addition, if some larvae<br />

swim at, or close to, their maximum U crit, many also swim at speeds<br />

equivalent to only a fraction of it (Figure 6.18). Therefore, there is room<br />

for more constraints on early swimming speed and endurance before<br />

these optimal strategies become impossible energetically. Furthermore,<br />

taxa which differ in early life history (i.e. demersal vs. pelagic eggs) are<br />

usually distributed differently in space 130,162,168,274 (see also chapter 4).<br />

This supports the hypothesis that events occurring early in the larval<br />

phase have a large influence on its outcome 71 — though it might be<br />

confounded by systematic differences during the rest of the pelagic interval:<br />

species with demersal eggs are usually only passable swimmers<br />

while species with pelagic eggs are, on average, very good swimmers by<br />

the end of the larval phase 25 . Anyhow, it was already pointed out that<br />

energy not invested in swimming is available for growth, and that larval<br />

growth is particularly important for survival 48,50,170,237,238 . So a strategy<br />

minimising overall energetic expenditure, such as early swimming here,<br />

should at least be regarded as a selective advantage, if anything else.<br />

Finally, the importance of early swimming decisions is highlighted<br />

by the differences between the tropical and temperate larvae considered<br />

here. The temperate species has very weak swimming abilities,<br />

even late in larval life (U crit < 5 cm s -1 ), and, in addition, has pelagic<br />

eggs. As a consequence, in the model, temperate larvae are initially<br />

advected passively by currents and are not capable of coming back to<br />

the recruitment zone if they are entrained too far away. This explains,<br />

in part, why tropical larvae achieve self-recruitment rates of 95% and<br />

temperate larvae only 40 to 70%, and makes early retention particularly<br />

important for the latter. A natural development of this study would<br />

be to investigate the case of tropical taxa with pelagic eggs, which<br />

are passive at first, have longer pelagic lives, but are very capable<br />

. . . but is probably an<br />

evolutionary optimum

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