The influence of desiccation and predation on vertical size gradients in populations of the gastropod Oxystele variegata (Anton) on an exposed rocky shore

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Abstract
Oxystele variegata (Anton.) exhibits a vertical size gradient contrary to the model proposed by Vermeij (1972) for low/mid intertidal species, as shell size increases in an upshore direction. Settlement occurs in the lowest zones and juveniles are restricted to the lower shore by conditions of desiccation higher up the beach. Juveniles suffer rapid water loss due to a relatively large opercular surface area and circumference and have a much lower resistance to water loss than adults. This leads to high mortality under conditions of low humidities, and juveniles caged at the top of the balanoid zone, where adults normally occur, die within a few days. As animals increase in size their resistance to desiccation rises allowing them to migrate upshore. This is a response to high rates of predation by the whelk Burnupena delalandii in the lower balanoid zone. Predation is so intense as to override the advantages of higher food availability which lead to a greater body weight for adults protected by cages on the lower shore.

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