published_or_final_version / Botany / Doctoral / Doctor of Philosophy

Helcion pruinosus, the rayed limpet, is found in the midshore region of the intertidal zone, (i.e. the Balanoid zone) of rocky shores, from Saldanha Bay to Sodwana Bay along the coast of South Africa. At Gonubie (32°57'S/25°01'E) this species of limpet was found under boulders, (when not feeding), and could reach average densities of 85/ m². The limpets prefer smooth boulders, avoiding those encrusted by coralline algae or boulders with a rough texture. There was no habitat segregation between adult and juvenile H. pruinosus, i.e. there was no sign of up-shore migration by larger individuals. Males out-numbered females by, on average 1.4 : 1, which was a similar sex ratio to that of the congeneric species, H. pectunculus (Gray, 996). Unlike H. pectunculus, there were no differences in shell lengths between males and females (p = 0.946 at Gonubie, south-east coast and p = 0.961 at Kommetjie, south-west coast; t-test). Shell height could also not be used as a criterion to differentiate between the sexes. The average maximum shell length of H. pruinosus at Gonubie and Kommetjie was 25 mm, but the maximum shell lengths found were 30 mm, (one individual in each case), on both the southeast coast and west coast. Growth occurred most rapidly in the first year of life, with individuals reaching 15 mm after 1 year. In the second year limpet growth slowed to only 7 mm. Sexual maturity was reached at a shell length of between 11 to 14 mm (at an age of 8 months to 1 year). The life-span of H. pruinosus was determined at 2.55 years on the south-east coast and 2.9 years on the south-west coast. Micro-growth bands were produced tidally. Growth occurred allometrically, i.e. shell length increased at a faster rate than shell height. The limpets foraged during low tide, at night, presumably feeding on epilithic algae on the boulders. A second smaller peak of limpet activity was often recorded during the day, when low tide coincided with dawn). More limpets (up to ten times more) were active at spring tides compared to neap tides. Limpet activity was greatest during the spring tides of autumn. During all seasons limpet activity peaked 30 minutes before low water, after which numbers decreased rapidly. All limpets had retreated beneath the boulders before being covered by the flooding tide. It was concluded that the limpets were responding to both endogenous and exogenous cues, but the actual stimulus for retreat could not be determined. The limpets did not return to a fixed scar nor did they return rigidly to a home site. H. pruinosus was found to be gonochoristic. Sexes could be separated, on dissection, according to the colour of their gonads (males had white gonads, whilst that of the females was olive-green). Gonads were present throughout the year in most individuals, i.e. totally spent animals were never observed. Reproductive seasonality was similar in west coast and south-east coast animals. The gonad indices of the west coast limpets were higher, (maximum of 35%), compared to that on the southeast coast, (25%). The animals were probably trickle spawners, with some individuals liberating gametes throughout the year. There was however synchrony in gametogenesis between male and female limpets on both the west coast and south-east coast lines. Gonad indices peaked in the summer months (October to December) with a second minor peak in April/May. These peaks occurred before the release of the gametes in spawning events. The gonad index was lowest in winter, (July). The ovaries were full of mature eggs (oocytes having a diameter of 200 to 250 ~) prior to spawning. The acinal wall thickness had been decreasing prior to spawning. After spawning there was an increase in previtellogenic eggs « 100 μm) in females, and an increase in the acinal wall thickness to indicate spermatogenesis. The two populations are subjected to different environmental conditions and the actual cue for stimulating the release of gametes could not be determined. Finally, a table of differences between H pruinosus and its sister species, H pectunculus was created to show the life-style of H pruinosus.