Intra-specific variability in the temporal organisation of foraging activity in the limpet Patella vulgata.

Patella vulgata is one of the most common limpet species on rocky shores throughout the north-eastern Atlantic. Foraging activity of this species has been described during various different diel and tidal phases and several factors have been suggested that might influence the timing of its activity. Among these, inclination of rock surfaces seems to have a clear role, with limpets on horizontal rocks being active during daylight at high water, while limpets on vertical rocks were active during night-time low water. However, by comparing the results of previous investigations it is difficult to separate clearly the effect of any single factor from the effects of their interactions. In the present paper we investigated the simultaneous influence of height on the shore, slope of rock surface and date of sampling (August-September and March-April) on the behaviour of a population of P. vulgata on the Isle of Man (UK). The study was carried out using modern telemetric methods, allowing the activity patterns of limpets to be recorded for long periods of time. The results showed that limpets concentrated their activity during two distinct temporal windows-daytime tide-in and night-time tide-out-while no significant activity was observed during daytime tide-out or night-time tide-in. The observed behaviour appeared to be driven by complex interactions among the three factors examined, and the influence of rock slope alone was not so strong as to override the effect of other factors completely. In addition, a key finding of this study was the conspicuous variability in the individual behaviour of P. vulgata. The greatest level of variability was observed in limpets on vertical surfaces on the upper shore, which showed an extreme switch in behavioural patterns, according to date of sampling (daylight/tide-in foraging during the spring and night-time/tide-out foraging during late summer). In addition, individuals apparently sharing the same set of conditions (inclination, height on the shore and sampling date) differed in their behaviour, and a given behavioural pattern was not always shared by all individuals from a given group. Finally, variability was also observed at the 'intra-individual' level since the same individual could be active during different tidal and diel phases. In conclusion, although the existence of endogenous rhythms has previously been demonstrated in this species, our data clearly show that P. vulgata was able to modulate its behavioural patterns in relation to a variety of exogenous determinants.