Morphological and isozyme variations between 13 populations of the species hitherto named Patella granularis were investigated to see whether differences in shell structure between the west coast versus the south and east coasts of southern Africa are supported by other morphological features or by genetic differences. The shells showed a definite decrease in size from west to east, but this is correlated with productivity and is of no diagnostic use in distinguishing between populations. Discriminant functions analysis based on shell morphometrics failed to separate populations from the three coastal regions. Shells from the northern east coast do, however, have shell nodules with a dark pigmentation, distinctly separating them from those further south and west. No differences in radular or soft part morphology were detected between the populations, but the four northernmost populations on the east coast have a significantly shorter Z looping of the gut than the other populations along the coast. Significant microstructural differences in the sperm were also detected between these two groups of populations. Electrophoretic analysis of 16 enzyme loci failed to detect any significant differences between the west and south coast populations, but revealed a genetic identity (Nei) of 0.528 as well as four diagnostic alleles between the four northernmost populations from the east coast compared with those to the south and west. The two genetically distinct forms occurred sympatrically at one of the study sites on the east coast (Coffee Bay). It was concluded the two groupings were sufficiently different to warrant the recognition of a separate species, which is centred in KwaZulu-Natal on the east coast and extends south to Coffee Bay, from where it is replaced by P. granularis. There is, however, no evidence at all that the west coast populations are in any way separable from the remaining populations of P. granularis on the south coast.

Physiological response to environmental change is examined in species of the intertidal limpets, Patella (Prosobranchia) and Siphonaria (Pulmonata). Characteristics of heart beat were determined using impedance pneumography and these are described. Heart rate of P. granularis is related to temperature and body size, and cardiac arrest in this limpet is apparently stress-related. Siphonaria oculus may exhibit a temperature-independent, extreme, and often prolonged bradycardia (<10 beats/min). When measured shortly after aerial exposure, heart rate and oxygen consumption of the above limpet species are closely correlated. The relationships of aerial oxygen consumption with body weight and ambient temperature were determined for the above high shore species of limpet. Both have low aerial rates of oxygen consumption relative to low shore limpet species, and their QlO values decrease with increasing temperature. Diel field recordings of heart rate of S. oculus, taken during summer and winter, suggest absence of temperature acclimation, and this was also shown for oxygen consumption at high aerial temperatures (30°C) in laboratory experiments. Oxygen consumption of P. granularis is partially temperature compensated temperature acclimation). In air, even though S. oculus loses water faster, it shows greater tolerance of water loss and survives longer than P. granularis. Prolonged aerial exposure of S. oculus leads to depression of heart rate and of V02 (down to 18% of the pre-exposure rate), responses interpreted as representing adaptive metabolic rate depression. In P. granularis aerial heart rate remains constant and V02 never falls below IX 38% of pre-exposure rate. This reduction in V02 in air is considered as being stress-related, resulting from impairment of oxygen uptake. In declining oxygen tension S. capensis shows a better capacity for oxyregulation than P. granularis. On exposure to hypoxia, sand-inundation and hyposalinity, S. capensis may show typical bradycardia ( <10 beats/min), suggesting depression of aerobic metabolism, and on return to pre-exposure conditions there is no overshoot of heart rate, suggesting absence of oxygen debt. The effect of hypoxia, sand-inundation and hyposalinity on heart rate of P. granularis is variable; this becomes depressed and is often interspersed with extended cardiac arrest. When conditions are normalized, this species exhibits a clear overshoot of heart rate. The significance of differences in physiological response between patellid and siphonariid limpets is discussed with regard to their habitat segregation, particularly in the upper-shore zone (open rock and tidal pools) and sand-inundated rock substrata, where only siphonariid limpets may be found. While not previously characterized in marine gastropods, metabolic rate depression by Siphonaria, through facilitating isolation and conserving food reserves, is suggested as a key factor determining their distribution in physico-chemically more extreme and variable intertidal habitats.

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.

No description available.