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Oriented swimming and passive advection 143<br />

of critical speed (U crit) 266 with age and size. They found size to be a<br />

better predictor of speed than age. Yet, when age was expressed in terms<br />

of developmental age (i.e. days since hatching/larval duration) and<br />

when speed was normalised per species (speed equals zero at hatching<br />

and one at settlement), a unique log 10 (age) · log 10 (speed) relationship<br />

held for all ten perciform species studied by Fisher 57<br />

log 10 (U − U hatch )<br />

log 10 (U settlement − U hatch ) = log 10 (A)<br />

log 10 (A settlement )<br />

(6.8)<br />

where A is age in days post-hatch (so A settlement is what is commonly<br />

measured as pelagic larval duration from otoliths: number of daily<br />

increments between hatching and settlement marks), and U is U crit in<br />

cm s -1 (U hatch is speed at hatching, and U settlement is speed at settlement).<br />

In a more useful form, this equation provides speed in function of age<br />

post-hatch<br />

log 10 (A)<br />

log<br />

U = U hatch + 10 10 (A settlement ) log 10 (U settlement −U hatch ) (6.9)<br />

It requires U crit and age at settlement, values of which can be found in<br />

the literature, and U crit at hatching, which is more scarcely reported.<br />

Given the ranges of variation and units of age and speed (days and cm s -1<br />

respectively), this is approximately equivalent to a linear relationship,<br />

as depicted by the solid curve in Figure 6.12.<br />

To measure critical swimming speed, larvae are made to swim in<br />

a flume inside which current speed is increased every few minutes.<br />

Therefore, it measures “forced” swimming and probably overestimates<br />

speeds at which larvae would swim when not constrained. Actually,<br />

U crit was found to be approximately five times the speed at which larvae<br />

were observed to swim freely in a large tank (routine speed) 91 , and twice<br />

the speed they go in situ 25 . The development of in situ speed with size<br />

has also been described in at least two studies 190,229 . Yet, in this model,<br />

larvae are given the possibility to choose between several swimming<br />

speeds. So, what we really need is the maximum potential speed larvae<br />

could attain, discarding burst speed (burst speed is fuelled anaerobically<br />

and is only relevant to behaviours such as escape from predators 92 ).<br />

And U crit is precisely that: a measure of potential speed 25,92 .<br />

The direct effect of temperature on swimming speed is twofold: first<br />

water temperature affects muscle efficiency because fishes are ectotherms;<br />

second, water viscosity increases when temperature drops, which could<br />

slow larvae down 25 . These two effects explain in part why tropical fish<br />

larvae are in general much better swimmers than temperate ones 25<br />

(section I.5.2, page 22). Finally, temperature also affects development,<br />

and larvae develop faster — hence acquire swimming abilities faster —<br />

in warmer waters 37,267 . In a brilliant meta-analysis, O’Connor et al. 37<br />

derived a single relationship between temperature and pelagic larval<br />

duration (PLD – i.e. the duration of ontogeny), that is valid for an<br />

extremely wide range of marine taxa (crustaceans, Annelids, fishes,<br />

Critical speed is the<br />

right measure of<br />

potential speed<br />

Temperature affects the<br />

metabolism, physics,<br />

and development<br />

of swimming

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