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27% of their dry body weight daily. 6 cal/g dry body weight per day. This value is similar to that in Ligia which has an assimilation efficiency of 55-78% depending on the type of diet (Carefoot, 1973). 63% of the dry body weight of the animal per tidal cycle, since there are two tides per day. One obvious question is whether these responses reflect some genotypic variability in populations from the upper and lower shore. The larvae, however, are freely mixed in the plankton and settlement appears to be at random over the lower shore (Smith and Newell, 1955).

3 STRATEGIES IN THE COMPETITION FOR ENERGY IN THE INTERTIDAL ZONE One way of assessing whether an organism is likely to be able to compete successfully with its neighbours within the limits of its physiological tolerances is to study the partitioning of energy which occurs in the population. Clearly, a net energy gain will result in enhanced growth and reproduction and this provides a useful index to compare with other species occupying a similar habitat. The well-known balanced energy equation of Winberg (1956) has been extensively applied to studies on the production of 2 whole populations and all the components are expressed as kcal/m /year.

Alternatively, in those organisms which are not subject to intense desiccation stress the metabolic energy requirements may be met by aerial gas exchange through relatively unmodified gill structures. Many intertidal animals including polychaetes such as Arenicola marina, Nereis diversicolor and Thoracophelia mucronata (Wells, 1949; Dales, 1963), some bivalves such as the cockle Cardium edule (Boyden, 1972), the gastropod Nassarius (Kushins and Mangum, 1971) and intertidal fishes such as Blennius pholis (Daniel, 1971; Wallace, 1973a) are able to utilise atmospheric air at some stages of the tidal cycle as long as they are able to do so without incurring water loss.

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