Extending Darwin's investigations on the barnacle life-history

Abstract

Darwin's magnificent study of the stalked, sessile and fossil barnacles, perhaps his greatest work, was started not many years after the systematic position of cirripedes within the class Crustacea had been accepted. It was completed at a time when histology and microtomy were not developed and when living specimens could only be seen on occasional visits to the seaside. Yet 130 years later it remains the standard text. It was on a visit to Tenby that he observed that barnacles were sensitive to vibration and, in seeking an acoustic organ, he mistakenly seized upon the oviducal gland. This led to an ever increasing series of misinterpretations of the female generative mechanism. Associated with this error was the belief that the ovary and cement glands were homologous in both cyprid and adult. He later confessed to “having blundered terribly over the cement glands”, but it was probably his search for examples of organs changing their function during evolution that led to this blunder. Similarly, he convinced himself mistakenly that the ovigerous fræna, which hold fast the egg masses to the mantle lining in stalked barnacles, became modified to gills in sessile barnacles. Darwin's interest in barnacle reproduction led to his discovery of complemental males attached to hermaphrodite individuals. He proved that they were dwarf cirripedes and not parasites. His knowledge that male and hermaphrodite flowers could co-exist helped him to accept that these apparently unnecessary beings could be present also in barnacles. We now know that complemental males are not confined to stalked barnacles but are also present in certain balanids. Though Darwin did not include the dioecious parasitic group, the Rhizocephala, in his studies, he recognized the wide repertoire of sexual arrangements from hermaphroditism to dioecy which now provides challenging material to our further understanding of sex allocation, control and evolution. Since Darwin dealt mainly with dried or preserved specimens he did not describe any of the details of the liberation of nauplii or the searching behaviour of the cyprid. Current investigators have shown that the former involves a prostaglandin which activates the embryos. Searching cyprids have been shown to be able to recognize their own and other species. Charles would have seen in this phenomenon a relevance both to survival and to the evolution of epizoic barnacles. Despite Charles' belief that his barnacle work would be “for ever unapplied” it has in fact been the foundation on which all studies of commercially important barnacle fouling have been based.