Biogeographic patterns of endolithic cyanobacteria and their negative impacts on mussels along the South African coast
- Authors: Ndhlovu, Aldwin
- Date: 2020
- Subjects: Cyanobacteria -- South Africa , Cyanobacteria -- Geographical distribution , Prokaryotes -- South Africa , Mexilhao mussel -- South Africa , Mytilus galloprovincialis -- South Africa , Coastal biology -- South Africa , Coastal ecology -- South Africa , Mussels -- Geographical distribution , Mussels -- Predators of , Mussels -- Mortality -- South Africa , Mussels -- Ecology -- South Africa , Mussels -- Growth -- South Africa , Mussels -- Fertility -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/144656 , vital:38367
- Description: Endolithic cyanobacterial species occur in a wide range of environments including cold and hot deserts as well as marine systems where they attack biological material such as corals and the shells of molluscs including limpets, mussels and abalone. Endoliths live as parasites in mussel shells, where they erode and extract calcium carbonate leading to shell weakening, creating fracture holes that lead to shell collapse and death, but they also have positive effects when they lead to discolouration of mussel shells hence giving them the ability to reduce stressful heat gain during periods of extreme heat stress. Mussels are ecological engineers on which the abundance and diversity of associated species assemblages depend. Understanding how endolithic cyanobacteria affect mussels will not only help in predicting future patterns of mussel abundances, but also future patterns of the infauna that depend on them. Firstly, I identified endolithic species infesting mussels and assessed the prevalence of endolithic parasitism in two intertidal mussel species in South Africa, the native Perna perna and the invasive Mytilus galloprovincialis. Large-scale surveys of endolithic infestation of mussels were conducted along 2500 km of the South African coast, covering three biogeographic regions: the subtropical east coast, dominated by P. perna, the warm temperate south coast where the indigenous species coexists with M. galloprovincialis, and the cool temperate west coast which is dominated by M. galloprovincialis. The prevalence of endolithic infestation was higher in the cool temperate bioregion than in the warm temperate and subtropical bioregions which did not differ and for P. perna endolithic species assemblages revealed clear groupings by bioregion. Results for endolithic induced mortality followed the same trend, with no significant difference between the two mussel species where they coexist and these results attribute biogeography of endoliths to environmental factors rather than host identity. Secondly, I assessed energy budgets of infested and clean mussels, to evaluate the energetic cost of infestation. This involved measuring energy acquisition, expenditure, calculating scope for growth and lethal temperatures (LT50s). The results revealed that endolithic cyanobacteria have a negative effect on scope for growth due to increased metabolic rates for infested mussels, with no effect of endoliths on the rates or efficiency of energy acquisition through filtration and no effect on lethal temperatures. The effects of infestation were then examined in more detail through a qualitative and quantitative analysis of mussel gonads and byssal attachment strength to the substratum. Endolithic infestation was found to affect reproduction by affecting the size (mass) of gonads, but not the density of eggs within them. Attachment strength was affected by endolithic infestation with very infested mussels requiring much less force to detach them from the substratum compared to mussels with low or no infestation. These results show that endolithic infestation affects mussel fitness by directly affecting attachment strength and by reducing their reproductive output. Thirdly, endolithic succession within mussel shells was examined by assessing endolithic species composition in different regions of the shell and as a function of time. The results on the spatial distribution of endolith species within a shell supported those for temporal succession in shells deployed in the field. Endolithic species that were early colonists of clean shells were similar to those that were found in the distal edge, the new and growing region of the shell and species that arrived late in succession were similar to endolithic species found near the umbo, the oldest region of the shell. Overall, the study shows that endolithic cyanobacteria show the effects of biogeography on species distribution and clear patterns of succession within mussel shells. Cyanobacteria affect mussels negatively; they lead to low scope for growth and hence low growth rates, low reproductive output and reduced attachment strength for infested mussels. This, in turn is expected to have indirect consequences for other species that rely on mussels as ecological engineers for their survival.
- Full Text:
- Date Issued: 2020
- Authors: Ndhlovu, Aldwin
- Date: 2020
- Subjects: Cyanobacteria -- South Africa , Cyanobacteria -- Geographical distribution , Prokaryotes -- South Africa , Mexilhao mussel -- South Africa , Mytilus galloprovincialis -- South Africa , Coastal biology -- South Africa , Coastal ecology -- South Africa , Mussels -- Geographical distribution , Mussels -- Predators of , Mussels -- Mortality -- South Africa , Mussels -- Ecology -- South Africa , Mussels -- Growth -- South Africa , Mussels -- Fertility -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/144656 , vital:38367
- Description: Endolithic cyanobacterial species occur in a wide range of environments including cold and hot deserts as well as marine systems where they attack biological material such as corals and the shells of molluscs including limpets, mussels and abalone. Endoliths live as parasites in mussel shells, where they erode and extract calcium carbonate leading to shell weakening, creating fracture holes that lead to shell collapse and death, but they also have positive effects when they lead to discolouration of mussel shells hence giving them the ability to reduce stressful heat gain during periods of extreme heat stress. Mussels are ecological engineers on which the abundance and diversity of associated species assemblages depend. Understanding how endolithic cyanobacteria affect mussels will not only help in predicting future patterns of mussel abundances, but also future patterns of the infauna that depend on them. Firstly, I identified endolithic species infesting mussels and assessed the prevalence of endolithic parasitism in two intertidal mussel species in South Africa, the native Perna perna and the invasive Mytilus galloprovincialis. Large-scale surveys of endolithic infestation of mussels were conducted along 2500 km of the South African coast, covering three biogeographic regions: the subtropical east coast, dominated by P. perna, the warm temperate south coast where the indigenous species coexists with M. galloprovincialis, and the cool temperate west coast which is dominated by M. galloprovincialis. The prevalence of endolithic infestation was higher in the cool temperate bioregion than in the warm temperate and subtropical bioregions which did not differ and for P. perna endolithic species assemblages revealed clear groupings by bioregion. Results for endolithic induced mortality followed the same trend, with no significant difference between the two mussel species where they coexist and these results attribute biogeography of endoliths to environmental factors rather than host identity. Secondly, I assessed energy budgets of infested and clean mussels, to evaluate the energetic cost of infestation. This involved measuring energy acquisition, expenditure, calculating scope for growth and lethal temperatures (LT50s). The results revealed that endolithic cyanobacteria have a negative effect on scope for growth due to increased metabolic rates for infested mussels, with no effect of endoliths on the rates or efficiency of energy acquisition through filtration and no effect on lethal temperatures. The effects of infestation were then examined in more detail through a qualitative and quantitative analysis of mussel gonads and byssal attachment strength to the substratum. Endolithic infestation was found to affect reproduction by affecting the size (mass) of gonads, but not the density of eggs within them. Attachment strength was affected by endolithic infestation with very infested mussels requiring much less force to detach them from the substratum compared to mussels with low or no infestation. These results show that endolithic infestation affects mussel fitness by directly affecting attachment strength and by reducing their reproductive output. Thirdly, endolithic succession within mussel shells was examined by assessing endolithic species composition in different regions of the shell and as a function of time. The results on the spatial distribution of endolith species within a shell supported those for temporal succession in shells deployed in the field. Endolithic species that were early colonists of clean shells were similar to those that were found in the distal edge, the new and growing region of the shell and species that arrived late in succession were similar to endolithic species found near the umbo, the oldest region of the shell. Overall, the study shows that endolithic cyanobacteria show the effects of biogeography on species distribution and clear patterns of succession within mussel shells. Cyanobacteria affect mussels negatively; they lead to low scope for growth and hence low growth rates, low reproductive output and reduced attachment strength for infested mussels. This, in turn is expected to have indirect consequences for other species that rely on mussels as ecological engineers for their survival.
- Full Text:
- Date Issued: 2020
The epibiotic relationship between mussels and barnacles
- Authors: Bell, Caroline Margaret
- Date: 2014
- Subjects: Epibiosis , Mussels -- Ecology -- South Africa , Mussels -- Behavior -- South Africa , Mussels -- Habitat -- South Africa , Barnacles -- Ecology -- South Africa , Barnacles -- Behavior -- South Africa , Barnacles -- Habitat -- South Africa , Perna -- Behavior -- South Africa , Mytilus galloprovincialis -- Behavior -- South Africa , Marine biodiversity -- South Africa
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5847 , http://hdl.handle.net/10962/d1011460 , Epibiosis , Mussels -- Ecology -- South Africa , Mussels -- Behavior -- South Africa , Mussels -- Habitat -- South Africa , Barnacles -- Ecology -- South Africa , Barnacles -- Behavior -- South Africa , Barnacles -- Habitat -- South Africa , Perna -- Behavior -- South Africa , Mytilus galloprovincialis -- Behavior -- South Africa , Marine biodiversity -- South Africa
- Description: Epibiosis is an ecological relationship that has been described as one of the closest possible associations in marine ecosystems. In the space limited rocky intertidal, mussel beds provide important secondary space for barnacles. The epibiotic relationship between mussels and barnacles on the south-east coast of South Africa was considered at different scales, from large-scale, natural patterns of epibiosis on the rocky shore, to fine-scale settlement choices of barnacles and the effects on the condition and growth rates of individual mussels. Mussel and barnacle assemblages were generally stable over a 12-month period. The tracking of individual mussels with and without barnacle epibionts resulted in a significant increase in mortality rate of mussels with epibionts over 12 months (two-way ANOVA, p = 0.028). Barnacles on rocks, as well as on mussels, were also tracked with no significant effect of substratum on mortality of barnacles (two-way ANOVA, p = 0.119). Prevalence and intensity of barnacle infestations was also examined in relation to coastline topography on two co-occurring mussel species, the indigenous Perna perna and invasive Mytilus galloprovincialis. The results were complex, but bay status had significant effects on prevalence and intensity for both mussel species, depending on the time and zone. The effect of bay in relation to time was particularly relevant for M. galloprovincialis (four-way nested ANOVA, Season X Site(Bay): p = 0.0002), where summer prevalence was higher than that of winter in bays, regardless of zone, while in open coast sites, the effect of season was only significant in the mid zone. Patterns of intensity generally showed higher values in summer. Substratum preference by barnacles was investigated by recording settlement, survival and mortality of Chthamalus dentatus barnacles on various treatments. There was a strong preference for the rock-like plastic substratum by primary settlers (pair-wise tests of PERMANOVA: Dead < Rock mimic (p = 0.0001); Replica < Rock mimic (p = 0.019) and Live < Rock mimic (p = 0.0001)). This indicates that barnacles settle on mussel shells only as a secondary choice and that micro-topography is an important variable in barnacle settlement. The effect of barnacle epibiosis on condition index and growth of P. perna and M. galloprovincialis was also examined as a direct indication of the health of mussels subjected to the biological stress of epibiosis. Although not significant (PERMANOVA: P. perna: p(perm) = 0.890; M. galloprovincialis: p(perm) = 0.395), growth for both mussel species was slower for barnacle-infested individuals in summer, which is the main growing season for mussels in the region. Results from condition index calculations, however, showed no negative impacts of epibiotic barnacles (three-way ANCOVA: P. perna: p = 0.372; M. galloprovincialis: p = 0.762). Barnacle epibionts create a new interface between the mussel and its environment and this interaction can affect other members of the community. The possibility of the barnacle epibiont causing increased drag also needs further investigation. Biological processes operating within a wide range of physical stressors drive the interactions on the rocky shore, such as epibiosis. Overall, the results of this study suggest that the epibiotic relationship between mussels and barnacles on the south-east coast of South Africa does not significantly affect the mussel species present and that barnacles only use mussel shells as a secondary choice of substratum.
- Full Text:
- Date Issued: 2014
- Authors: Bell, Caroline Margaret
- Date: 2014
- Subjects: Epibiosis , Mussels -- Ecology -- South Africa , Mussels -- Behavior -- South Africa , Mussels -- Habitat -- South Africa , Barnacles -- Ecology -- South Africa , Barnacles -- Behavior -- South Africa , Barnacles -- Habitat -- South Africa , Perna -- Behavior -- South Africa , Mytilus galloprovincialis -- Behavior -- South Africa , Marine biodiversity -- South Africa
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5847 , http://hdl.handle.net/10962/d1011460 , Epibiosis , Mussels -- Ecology -- South Africa , Mussels -- Behavior -- South Africa , Mussels -- Habitat -- South Africa , Barnacles -- Ecology -- South Africa , Barnacles -- Behavior -- South Africa , Barnacles -- Habitat -- South Africa , Perna -- Behavior -- South Africa , Mytilus galloprovincialis -- Behavior -- South Africa , Marine biodiversity -- South Africa
- Description: Epibiosis is an ecological relationship that has been described as one of the closest possible associations in marine ecosystems. In the space limited rocky intertidal, mussel beds provide important secondary space for barnacles. The epibiotic relationship between mussels and barnacles on the south-east coast of South Africa was considered at different scales, from large-scale, natural patterns of epibiosis on the rocky shore, to fine-scale settlement choices of barnacles and the effects on the condition and growth rates of individual mussels. Mussel and barnacle assemblages were generally stable over a 12-month period. The tracking of individual mussels with and without barnacle epibionts resulted in a significant increase in mortality rate of mussels with epibionts over 12 months (two-way ANOVA, p = 0.028). Barnacles on rocks, as well as on mussels, were also tracked with no significant effect of substratum on mortality of barnacles (two-way ANOVA, p = 0.119). Prevalence and intensity of barnacle infestations was also examined in relation to coastline topography on two co-occurring mussel species, the indigenous Perna perna and invasive Mytilus galloprovincialis. The results were complex, but bay status had significant effects on prevalence and intensity for both mussel species, depending on the time and zone. The effect of bay in relation to time was particularly relevant for M. galloprovincialis (four-way nested ANOVA, Season X Site(Bay): p = 0.0002), where summer prevalence was higher than that of winter in bays, regardless of zone, while in open coast sites, the effect of season was only significant in the mid zone. Patterns of intensity generally showed higher values in summer. Substratum preference by barnacles was investigated by recording settlement, survival and mortality of Chthamalus dentatus barnacles on various treatments. There was a strong preference for the rock-like plastic substratum by primary settlers (pair-wise tests of PERMANOVA: Dead < Rock mimic (p = 0.0001); Replica < Rock mimic (p = 0.019) and Live < Rock mimic (p = 0.0001)). This indicates that barnacles settle on mussel shells only as a secondary choice and that micro-topography is an important variable in barnacle settlement. The effect of barnacle epibiosis on condition index and growth of P. perna and M. galloprovincialis was also examined as a direct indication of the health of mussels subjected to the biological stress of epibiosis. Although not significant (PERMANOVA: P. perna: p(perm) = 0.890; M. galloprovincialis: p(perm) = 0.395), growth for both mussel species was slower for barnacle-infested individuals in summer, which is the main growing season for mussels in the region. Results from condition index calculations, however, showed no negative impacts of epibiotic barnacles (three-way ANCOVA: P. perna: p = 0.372; M. galloprovincialis: p = 0.762). Barnacle epibionts create a new interface between the mussel and its environment and this interaction can affect other members of the community. The possibility of the barnacle epibiont causing increased drag also needs further investigation. Biological processes operating within a wide range of physical stressors drive the interactions on the rocky shore, such as epibiosis. Overall, the results of this study suggest that the epibiotic relationship between mussels and barnacles on the south-east coast of South Africa does not significantly affect the mussel species present and that barnacles only use mussel shells as a secondary choice of substratum.
- Full Text:
- Date Issued: 2014
Effects of coastal topography on physiology, behaviour and genetics of indigenous (Perna perna) and invasive (Mytilus galloprovincialis) mussels
- Authors: Nicastro, Katy R
- Date: 2008
- Subjects: Mussels -- Ecology -- South Africa , Perna -- Physiology -- South Africa , Perna -- Behavior -- South Africa , Mussels -- Behavior -- Environmental factors -- South Africa , Mussels -- Habitat -- South Africa , Mytilus galloprovincialis , Mytilus galloprovincialis -- Physiology -- South Africa , Mytilus galloprovincialis -- Behavior -- South Africa , Mytilus galloprovincialis -- Genetics -- South Africa , Coastal ecology -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5833 , http://hdl.handle.net/10962/d1008262
- Description: Organisms inhabit environments that have many dimensions, each of which can vary temporally and spatially. The spatial-temporal variations of environmental stressors and disturbances may have major but different effects on indigenous and invasive species, favouring either of them at different times and places. The invasive mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and, on the south coast of South Africa, it now competes and co-exists with the indigenous Perna perna in the lower eulittoral zone (referred to here as the mussel zone) The invasive and indigenous species dominate the upper and the lower mussel zones respectively, while the two co-exist in the mid-zone. My results show that intertidal mussels experience, and respond to, spatial and temporal fluctuations of several biotic and abiotic stressors. The invasive and the indigenous species adopt different strategies when reacting to environmental factors and their physiological and behavioural responses vary in time and in different habitats as different pressures become of overriding importance. Attachment strength of both species decreased in summer and increased in winter, and was higher on the open coast than in bays for both species, showing a strong positive correlation with wave force in time and space. P. perna had significantly higher attachment strength than M. galloprovincialis but, contrary to previous studies, the difference in gonad index between the two species varied according to the habitat. In bay habitats, M. galloprovincialis had a higher maximum reproductive effort than P. perna, however, on the open coast, there was no significant difference between the two species, suggesting that for the invasive species wave action is a limiting factor not only in terms of the attachment strength but also of energy availability for reproductive tissue development. Major spawning events occurred during periods of low wave action while minor spawning coincided with periods of intense hydrodynamic stress. On the open coast, gonad index was negatively correlated with attachment strength for both species while, in bays, there was no correlation between these two factors for either. The two species also showed different behaviour. In the field, M. galloprovincialis moved significantly more than P. perna over a period of six months. The higher mobility of the invasive species was also confirmed in the laboratory where, in general, M. galloprovincialis formed clumps more readily than P. perna. Taken collectively, these results suggest that channelling more energy into attachment strength limits reproductive tissue development and that, while the indigenous species invests more in byssal production, the invasive species adopts a more dynamic strategy looking for aggregation or a safer arrangement. Higher endolithic infestation and a greater expression of heat shock proteins (Hsps) in mussel populations on the open coast than in bays indicate that this habitat is a more stressful environment not only in terms of wave action. Endolith damaged mussels had significantly lower attachment strengths and condition indices than clean mussels, probably due to the need to channel energy into shell repair. The constant shell repair and expression of Hsps typical of open coast populations are energetically demanding processes. These observations suggest that on the open coast, mussels are subjected to more severe energetic constraints than in bay habitats. Wave and sand stress fluctuated seasonally with the former having a greater effect on mussel mortality on the open coast and the latter a higher impact on bay populations. Overall, mussel mortality rates were higher on the open coast than in bays. My results show that populations on the open coast had fewer private haplotypes and less genetic endemism than those inside bays. Gene flow analysis showed the relatively stable bay habitats act as source populations with greater genetic migration rates out of bays than into them. These differences in genetic structure on scales of las of kilometers show that coastal configuration strongly affects selection, larval dispersal and haplotype diversity. Environmental gradients that are key factors in species distribution over large geographical scales can also be responsible for micro-scale distributions. My results show that M. galloprovincialis colonizes the upper mussel zone where temperature is high, but is less tolerant to this stressor and has to maintain a high expression of Hsps. This suggests that temperature is probably a limiting factor in its invasion towards the sub-tropical east coast. There are inter- and intra-specific differences in responses to the environment which highlight the efforts of M. galloprovincialis and P. perna to optimize resource utilization for survival and reproduction. Determining these differences is crucial to understanding patterns of co-existence between competing indigenous and invasive species.
- Full Text:
- Date Issued: 2008
- Authors: Nicastro, Katy R
- Date: 2008
- Subjects: Mussels -- Ecology -- South Africa , Perna -- Physiology -- South Africa , Perna -- Behavior -- South Africa , Mussels -- Behavior -- Environmental factors -- South Africa , Mussels -- Habitat -- South Africa , Mytilus galloprovincialis , Mytilus galloprovincialis -- Physiology -- South Africa , Mytilus galloprovincialis -- Behavior -- South Africa , Mytilus galloprovincialis -- Genetics -- South Africa , Coastal ecology -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5833 , http://hdl.handle.net/10962/d1008262
- Description: Organisms inhabit environments that have many dimensions, each of which can vary temporally and spatially. The spatial-temporal variations of environmental stressors and disturbances may have major but different effects on indigenous and invasive species, favouring either of them at different times and places. The invasive mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and, on the south coast of South Africa, it now competes and co-exists with the indigenous Perna perna in the lower eulittoral zone (referred to here as the mussel zone) The invasive and indigenous species dominate the upper and the lower mussel zones respectively, while the two co-exist in the mid-zone. My results show that intertidal mussels experience, and respond to, spatial and temporal fluctuations of several biotic and abiotic stressors. The invasive and the indigenous species adopt different strategies when reacting to environmental factors and their physiological and behavioural responses vary in time and in different habitats as different pressures become of overriding importance. Attachment strength of both species decreased in summer and increased in winter, and was higher on the open coast than in bays for both species, showing a strong positive correlation with wave force in time and space. P. perna had significantly higher attachment strength than M. galloprovincialis but, contrary to previous studies, the difference in gonad index between the two species varied according to the habitat. In bay habitats, M. galloprovincialis had a higher maximum reproductive effort than P. perna, however, on the open coast, there was no significant difference between the two species, suggesting that for the invasive species wave action is a limiting factor not only in terms of the attachment strength but also of energy availability for reproductive tissue development. Major spawning events occurred during periods of low wave action while minor spawning coincided with periods of intense hydrodynamic stress. On the open coast, gonad index was negatively correlated with attachment strength for both species while, in bays, there was no correlation between these two factors for either. The two species also showed different behaviour. In the field, M. galloprovincialis moved significantly more than P. perna over a period of six months. The higher mobility of the invasive species was also confirmed in the laboratory where, in general, M. galloprovincialis formed clumps more readily than P. perna. Taken collectively, these results suggest that channelling more energy into attachment strength limits reproductive tissue development and that, while the indigenous species invests more in byssal production, the invasive species adopts a more dynamic strategy looking for aggregation or a safer arrangement. Higher endolithic infestation and a greater expression of heat shock proteins (Hsps) in mussel populations on the open coast than in bays indicate that this habitat is a more stressful environment not only in terms of wave action. Endolith damaged mussels had significantly lower attachment strengths and condition indices than clean mussels, probably due to the need to channel energy into shell repair. The constant shell repair and expression of Hsps typical of open coast populations are energetically demanding processes. These observations suggest that on the open coast, mussels are subjected to more severe energetic constraints than in bay habitats. Wave and sand stress fluctuated seasonally with the former having a greater effect on mussel mortality on the open coast and the latter a higher impact on bay populations. Overall, mussel mortality rates were higher on the open coast than in bays. My results show that populations on the open coast had fewer private haplotypes and less genetic endemism than those inside bays. Gene flow analysis showed the relatively stable bay habitats act as source populations with greater genetic migration rates out of bays than into them. These differences in genetic structure on scales of las of kilometers show that coastal configuration strongly affects selection, larval dispersal and haplotype diversity. Environmental gradients that are key factors in species distribution over large geographical scales can also be responsible for micro-scale distributions. My results show that M. galloprovincialis colonizes the upper mussel zone where temperature is high, but is less tolerant to this stressor and has to maintain a high expression of Hsps. This suggests that temperature is probably a limiting factor in its invasion towards the sub-tropical east coast. There are inter- and intra-specific differences in responses to the environment which highlight the efforts of M. galloprovincialis and P. perna to optimize resource utilization for survival and reproduction. Determining these differences is crucial to understanding patterns of co-existence between competing indigenous and invasive species.
- Full Text:
- Date Issued: 2008
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