Aspects of the biology of the infaunal bivalve Mollusc Solen cylindraceus (Hanley) in the Kariega estuary
- De Villiers, Casper Johannes
- Authors: De Villiers, Casper Johannes
- Date: 1990
- Subjects: Mollusks -- South Africa -- Eastern Cape Bivalves -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5639 , http://hdl.handle.net/10962/d1005321
- Description: Solen cylindraceus is an infaunal filter-feeding bivalve inhabiting the intertidal mud banks of many southern African estuaries. It is particularly abundant in the Kariega estuary (33°41'S; 26°42'E) where it reaches densities of 400m⁻² (192g shell-free dry wt. m⁻²). The Kariega is a permanently open, marine dominated estuary about 18km in length, and S. cylindraceus is most abundant in its middle and upper reaches. Some physical characteristics of the estuary (temperature, salinity, sediment and water turbidity) are described, and the possible role of these factors in determining the density and distribution of S. cylindraceus within the Kariega estuary, is discussed. The structure of the alimentary system, gills and labial palps of S. cylintfraceus is described, all of which showed no major variation from the "typical" eulamellibranchiate form. Solen cylintfraceus was found to be a euryhaline osmoconformer with a salinity tolerance range of 15-65%. When animals were removed from their burrows, osmotic equilibration of the haemolymph was rapid (1-2 hours). By contrast, in animals left undisturbed in their burrows, osmotic equilibration was retarded (72-204 hours). It is suggested that the observed decrease in the rate of change of haemolymph osmolarity for animals in their burrows is linked to the stability of the interstitial salinity. A temperature tolerance range of 5-44°C was determined for S. cylintfraceus (in situ), in which prolonged exposure to 5°C and 40-45°C (12-36 hours respectively) resulted in a decreased burrowing ability, coma and death. Animal burrowing responses were not affected by temperatures in the range 15-35°C. Field experiments were carried out over several tidal cycles, in which the measurement of crystalline style volume was used as a means of assessing extracellular digestive activity. No major variation in style volume was recorded and it appeared that S. cylindraceus did not exhibit any cyclical pattern of style dissolution and regeneration. It is suggested that S. cylindraceus feeds continuously from the water column during high tide and possibly within its burrow, at or below the water table, during low tide. At a suspensoid concentration of 5Omg l⁻¹, S. cylindraceus was found to filter water almost continuously (90-95% of the time). Time spent filtering dropped to 68% at 100mg l⁻¹ and 32% at 500mg l⁻¹. Filtration rates for summer collected animals (25°C) were 22.86 ± 4.36ml min.⁻¹, some 3ml min.⁻¹ greater than that recorded for winter (16°C) collected animals. Filtration rate may be expressed as a function of shell length by the equations: y=0.247x¹̇⁰⁶⁶ (winter) and y=0.758x⁰̇⁸²⁶ (summer). Solen cylindraceus was capable of acclimating its filtration rate to both high and low temperatures under laboratory conditions. Filtration rate exhibited a thermal optimum in the range 15-35°C, declining at higher and lower temperatures. Q₁₀ values of filtration decreased rapidly from greater than 4 to less than 2, when the thermal optimum was reached. Maximum rates generally occurred at approximately 5°C above the temperature to which the animal had been acclimated. Optimal filtration rates (19-23ml min.⁻¹) were recorded in the salinity range 15-45%. When subjected to abrupt changes in salinity, filtration rates were immediately depressed. The extent and duration of these decreased filtration rates were dependent upon the magnitude and direction of salinity change, and were always less in animals exposed to hyper- than hyposaline conditions. Animals exposed to increased temperature and simultaneous elevated or unchanged salinity, showed a slight increase in filtration rate followed by rapid acclimation. A decrease in both temperature and salinity resulted in an initial decrease in filtration rate and a longer acclimation period. The ability of S. cylindraceus to acclimate fully within a wide temperature and salinity range, and to filter maximally in hypersaline conditions may, in part, explain its unusually high abundance in the Kariega estuary, despite it being close to the southernmost limit of the animal's geographical distribution. No significant difference in flItration rate was recorded at suspensoid concentrations of 5-100mg 1⁻¹. However, at 250 and 500mg l⁻¹ filtration rates decreased significantly, and coincided with increased levels of pseudofaecal production. Solen cylindraceus retained particles down to 2.5-3.0µm with great efficiency (ca. 60-90% efficiency). Below this particle size, retention efficiency decreased rapidly and a net production of particles was recorded below 1.51µm. Particle retention was independent of temperature (15 and 25°C) and salinity (15 and 35%). Use was made of stable carbon isotope analyses (¹³C/¹²C ratios) in an attempt to determine the important food sources of S. cylindraceus within the Kariega estuary. The results obtained demonstrated an enrichment in δ¹³ values for S. cylindraceus from the upper (-27.9%) to the middle (-25%c) and lower (-21.6%o ) reaches of the estuary, with no seasonal variation apparent. The bivalve was substantially more depleted in ¹³C relative to the dominant aquatic macrophytes Zostera capensis (-9.1 to -15.6%o) and Spartina maritima (-12.5%o). The use of δ¹³ alone, however, to unequivocally "pin point" specific food sources of a filter feeder in a predominantly detritus based food web, is limited. It is suggested that in the Kariega estuary, riparian litter and other terrestrially derived vegetation contribute to the carbon pool. A possible contribution of ¹³C depleted food sources via chemoautotrophic and/or anaerobic pathways, to the diet of S. cylindraceus, is suggested.
- Full Text:
- Date Issued: 1990
- Authors: De Villiers, Casper Johannes
- Date: 1990
- Subjects: Mollusks -- South Africa -- Eastern Cape Bivalves -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5639 , http://hdl.handle.net/10962/d1005321
- Description: Solen cylindraceus is an infaunal filter-feeding bivalve inhabiting the intertidal mud banks of many southern African estuaries. It is particularly abundant in the Kariega estuary (33°41'S; 26°42'E) where it reaches densities of 400m⁻² (192g shell-free dry wt. m⁻²). The Kariega is a permanently open, marine dominated estuary about 18km in length, and S. cylindraceus is most abundant in its middle and upper reaches. Some physical characteristics of the estuary (temperature, salinity, sediment and water turbidity) are described, and the possible role of these factors in determining the density and distribution of S. cylindraceus within the Kariega estuary, is discussed. The structure of the alimentary system, gills and labial palps of S. cylintfraceus is described, all of which showed no major variation from the "typical" eulamellibranchiate form. Solen cylintfraceus was found to be a euryhaline osmoconformer with a salinity tolerance range of 15-65%. When animals were removed from their burrows, osmotic equilibration of the haemolymph was rapid (1-2 hours). By contrast, in animals left undisturbed in their burrows, osmotic equilibration was retarded (72-204 hours). It is suggested that the observed decrease in the rate of change of haemolymph osmolarity for animals in their burrows is linked to the stability of the interstitial salinity. A temperature tolerance range of 5-44°C was determined for S. cylintfraceus (in situ), in which prolonged exposure to 5°C and 40-45°C (12-36 hours respectively) resulted in a decreased burrowing ability, coma and death. Animal burrowing responses were not affected by temperatures in the range 15-35°C. Field experiments were carried out over several tidal cycles, in which the measurement of crystalline style volume was used as a means of assessing extracellular digestive activity. No major variation in style volume was recorded and it appeared that S. cylindraceus did not exhibit any cyclical pattern of style dissolution and regeneration. It is suggested that S. cylindraceus feeds continuously from the water column during high tide and possibly within its burrow, at or below the water table, during low tide. At a suspensoid concentration of 5Omg l⁻¹, S. cylindraceus was found to filter water almost continuously (90-95% of the time). Time spent filtering dropped to 68% at 100mg l⁻¹ and 32% at 500mg l⁻¹. Filtration rates for summer collected animals (25°C) were 22.86 ± 4.36ml min.⁻¹, some 3ml min.⁻¹ greater than that recorded for winter (16°C) collected animals. Filtration rate may be expressed as a function of shell length by the equations: y=0.247x¹̇⁰⁶⁶ (winter) and y=0.758x⁰̇⁸²⁶ (summer). Solen cylindraceus was capable of acclimating its filtration rate to both high and low temperatures under laboratory conditions. Filtration rate exhibited a thermal optimum in the range 15-35°C, declining at higher and lower temperatures. Q₁₀ values of filtration decreased rapidly from greater than 4 to less than 2, when the thermal optimum was reached. Maximum rates generally occurred at approximately 5°C above the temperature to which the animal had been acclimated. Optimal filtration rates (19-23ml min.⁻¹) were recorded in the salinity range 15-45%. When subjected to abrupt changes in salinity, filtration rates were immediately depressed. The extent and duration of these decreased filtration rates were dependent upon the magnitude and direction of salinity change, and were always less in animals exposed to hyper- than hyposaline conditions. Animals exposed to increased temperature and simultaneous elevated or unchanged salinity, showed a slight increase in filtration rate followed by rapid acclimation. A decrease in both temperature and salinity resulted in an initial decrease in filtration rate and a longer acclimation period. The ability of S. cylindraceus to acclimate fully within a wide temperature and salinity range, and to filter maximally in hypersaline conditions may, in part, explain its unusually high abundance in the Kariega estuary, despite it being close to the southernmost limit of the animal's geographical distribution. No significant difference in flItration rate was recorded at suspensoid concentrations of 5-100mg 1⁻¹. However, at 250 and 500mg l⁻¹ filtration rates decreased significantly, and coincided with increased levels of pseudofaecal production. Solen cylindraceus retained particles down to 2.5-3.0µm with great efficiency (ca. 60-90% efficiency). Below this particle size, retention efficiency decreased rapidly and a net production of particles was recorded below 1.51µm. Particle retention was independent of temperature (15 and 25°C) and salinity (15 and 35%). Use was made of stable carbon isotope analyses (¹³C/¹²C ratios) in an attempt to determine the important food sources of S. cylindraceus within the Kariega estuary. The results obtained demonstrated an enrichment in δ¹³ values for S. cylindraceus from the upper (-27.9%) to the middle (-25%c) and lower (-21.6%o ) reaches of the estuary, with no seasonal variation apparent. The bivalve was substantially more depleted in ¹³C relative to the dominant aquatic macrophytes Zostera capensis (-9.1 to -15.6%o) and Spartina maritima (-12.5%o). The use of δ¹³ alone, however, to unequivocally "pin point" specific food sources of a filter feeder in a predominantly detritus based food web, is limited. It is suggested that in the Kariega estuary, riparian litter and other terrestrially derived vegetation contribute to the carbon pool. A possible contribution of ¹³C depleted food sources via chemoautotrophic and/or anaerobic pathways, to the diet of S. cylindraceus, is suggested.
- Full Text:
- Date Issued: 1990
Aspects of the structure and function of some gastropod columellar muscles (Mollusca)
- Authors: Frescura, Mandy
- Date: 1991
- Subjects: Gastropoda -- Research Mollusks -- Research
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5642 , http://hdl.handle.net/10962/d1005324
- Description: The columellar muscle of both limpets and coiled shell gastropods is of the paramyosin smooth type. Collagen forms an integral part of the musculature constituting about 35% of the tissue. In limpets, muscle organisation is typical of a muscular hydrostat. Tightly packed blocks of muscle, dense arrays of cross-linked collagen, large muscle cells (9 µm diameter) and thick filaments (70 nm diameter, 30 µm long) produce a tough, relatively rigid but powerful muscle. In coiled shell gastropods, muscle organisation is intermediate between a muscular and a fluid hydrostat. Finer muscle cells (6 µm diameter), thick filaments (60 nm diameter) and a loose intercellular network of collagen interspersed with fluid vesicles are features of a more pliable and extensible muscle. In addition, ultrastructural differences, such as larger numbers of mitochondria and sarcolemmal invaginations distinguish the tarsal from the columellar muscle in both limpets and coiled shell gastropods. About 25% of muscle cells in most species examined, contain a novel arrangement of thin filaments with periodic electron-dense regions. These are similar in appearance to intrafusal cells and stress-fibres of non-muscle cells. Structural analysis of isolated filaments, optical diffraction and SDS gel electrophoresis confirm the, large dimensions and the paramyosin nature of the thick filaments. Microdensitometry of the gel proteins confirms the high proportion of collagen present. No significant differences in muscle ultrastructure were found between limpets from different tidal heights. Muscle attachment areas are shown to be species-specific and positively correlated to tenacity and wave exposure. The muscle attachment mechanism is similar to that described for other molluscs. It consists of a special epithelial layer and a mucous-like material at the muscle-shell interface that possibly has an adhesive function. Although the ultrastructure of Patella is very similar to that of the anterior byssus retractor of Mytilus, its mechanical behaviour is not. The muscle has a narrow working range where maximum tensions and "catch-like" contractions develop. This narrow length range is co-incident with the in situ length at which clamping occurs. It is suggested that the large component of collagen has an important influence over the mechanical behaviour of the muscle during clamping, by cross-linking in a manner similar to that described for some echinoderm connective tissues.
- Full Text:
- Date Issued: 1991
- Authors: Frescura, Mandy
- Date: 1991
- Subjects: Gastropoda -- Research Mollusks -- Research
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5642 , http://hdl.handle.net/10962/d1005324
- Description: The columellar muscle of both limpets and coiled shell gastropods is of the paramyosin smooth type. Collagen forms an integral part of the musculature constituting about 35% of the tissue. In limpets, muscle organisation is typical of a muscular hydrostat. Tightly packed blocks of muscle, dense arrays of cross-linked collagen, large muscle cells (9 µm diameter) and thick filaments (70 nm diameter, 30 µm long) produce a tough, relatively rigid but powerful muscle. In coiled shell gastropods, muscle organisation is intermediate between a muscular and a fluid hydrostat. Finer muscle cells (6 µm diameter), thick filaments (60 nm diameter) and a loose intercellular network of collagen interspersed with fluid vesicles are features of a more pliable and extensible muscle. In addition, ultrastructural differences, such as larger numbers of mitochondria and sarcolemmal invaginations distinguish the tarsal from the columellar muscle in both limpets and coiled shell gastropods. About 25% of muscle cells in most species examined, contain a novel arrangement of thin filaments with periodic electron-dense regions. These are similar in appearance to intrafusal cells and stress-fibres of non-muscle cells. Structural analysis of isolated filaments, optical diffraction and SDS gel electrophoresis confirm the, large dimensions and the paramyosin nature of the thick filaments. Microdensitometry of the gel proteins confirms the high proportion of collagen present. No significant differences in muscle ultrastructure were found between limpets from different tidal heights. Muscle attachment areas are shown to be species-specific and positively correlated to tenacity and wave exposure. The muscle attachment mechanism is similar to that described for other molluscs. It consists of a special epithelial layer and a mucous-like material at the muscle-shell interface that possibly has an adhesive function. Although the ultrastructure of Patella is very similar to that of the anterior byssus retractor of Mytilus, its mechanical behaviour is not. The muscle has a narrow working range where maximum tensions and "catch-like" contractions develop. This narrow length range is co-incident with the in situ length at which clamping occurs. It is suggested that the large component of collagen has an important influence over the mechanical behaviour of the muscle during clamping, by cross-linking in a manner similar to that described for some echinoderm connective tissues.
- Full Text:
- Date Issued: 1991
The growth and reproduction of Patella granularis (Mollusca : patellogastropoda) on the south-east coast of South Africa
- Authors: Vat, Laura Suzanne
- Date: 2000
- Subjects: Patellidae Limpets -- South Africa Mollusks -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5702 , http://hdl.handle.net/10962/d1005388
- Description: 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.
- Full Text:
- Date Issued: 2000
- Authors: Vat, Laura Suzanne
- Date: 2000
- Subjects: Patellidae Limpets -- South Africa Mollusks -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5702 , http://hdl.handle.net/10962/d1005388
- Description: 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.
- Full Text:
- Date Issued: 2000
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