The growth and reproduction of Patella granularis (Mollusca : patellogastropoda) on the south-east coast of South Africa

Aspects of the biology and ecology of Patella granularis were investigated along a 130km stretch of the south-east coast of South Africa. Distribution, biomass, density and population structure were investigated at seven localities. In addition, a more detailed study of the growth rate and reproductive biology of populations inhabiting three different substrata (aeolianite, quartzitic sandstone and mussel shells) was conducted. The genetic relationships between these three populations was also examined, as was the foraging behaviour of the limpets inhabiting an aeolianite and a quartzitic sandstone shore. Finally, differences in food availability on the different substrata were studied. On the south-east coast, P. granularis has a wide intertidal distribution, occurring from the upper Balanoid zone through to the Cochlear zone, where it is a common inhabitant of mussel shells. The mean shell length of P. granularis was found to decrease down the shore. The largest limpets (46.6 mm shell length) were found on an offshore island in Algoa Bay. At most localities investigated, the sex ratio deviated from a 1:1 ratio with more males than females being recorded on five shores. Both limpet density and biomass were lower on the south-east coast when compared to data published for west coast populations. On the south-east coast, both density and dry biomass were highest in the lower Balanoid zone. Allozyme electrophoresis indicated that P. granularis inhabiting aeolianite, quartzitic sandstone and mussel shells are all part of a single population. Extremely high genetic identity values (0.998), low levels of heterozygosities (0.035 - 0.061), low levels of polymorphisms (25% - 31%) and low FST values (0.021) all suggest that the three populations of P. granularis form a common breeding group, despite the high levels of phenotypic plasticity observed. On all shores, P. granularis was found to grow allometrically, increasing in shell height more rapidly than shell length. Estimation of the growth rate (determined by the Von Bertalanffy growth model) of P. granularis suggested that limpets inhabiting the mussel shells grew more slowly, and attained a smaller maximum size, than those inhabiting both the aeolianite and the quartzite (K = 0.25, 0.32 and 0.33 respectively; 27.12 mm, 31.89 mm and 32.96 mm respectively). Previous work has shown that west coast P. granularis grow more quickly (K = 0.7) and reach a greater size (. 40 mm). Translocation of limpets among sites suggested that limpet size in the mussel beds was spatially constrained. Shell microgrowth bands were deposited tidally, but could not be used for aging limpets due to shell erosion. Limpets from the aeolianite had the greatest reproductive fitness, producing more eggs (.366 000/limpet) than those inhabiting quartzite (.119 500/limpet) or mussel shell limpets (.85 800/limpet). Aeolianite limpets also spawned throughout the year, whereas those from the quartzite and mussel shells spawned twice a year (once in winter and once in summer) although a great deal of interannual variability was observed. The onset of sexual maturity occurred at a similar age in all limpets (1 - 2 years) and is probably genetically entrenched. P. granularis inhabiting both an aeolianite and a quartzitic shore were active during nocturnal low tides. All limpets returned to a home scar after foraging. Whilst foraging, limpets inhabiting the aeolianite shore moved shorter distances (.17 cm) at a slower rate than those from quartzite (.30 cm). Limpets that were translocated from one substratum to the other initially moved similar distances to the source group, but after a maximum period of one week, moved distances that mirrored those moved by the resident limpets. Neither season nor tidal phase influenced the distances foraged. No directionality in foraging was found. Wear of radula teeth, particularly the pluricuspid tooth, was greater in limpets from the quartzite. It is hypothesised that the observed differences in life-history parameters and foraging behaviour of limpets both within the south-east coast and between the west and south-east coasts are related to food abundance. Chlorophyll-a, and hence microalgal biomass, was consistently higher on aeolianite (.2.5 times) than on both quartzite and mussel shells. Estimates of chlorophyll-a were higher (although not significantly) in winter. Previous studies determined that primary productivity is also higher along the west coast. Finally, the lack of evidence for migration of limpets from the low-shore to high-shore in south-east coast P. granularis is discussed. It is suggested that this species settles opportunistically within its physiological tolerances and responds morphologically to localised environmental conditions