Large scale spatio-temporal forcing of pelagic-coastal coupling: disentangling the effects of environmental change on intertidal invertebrate recruitment
- Authors: Muñiz, Carlota Fernández
- Date: 2019
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/75317 , vital:30400
- Description: Marine systems are driven by the relationships among organisms and environmental conditions. Anthropogenic-induced changes during the past decades have started to alter climatic drivers which have the potential to alter the physical, chemical and biological environment. In coastal systems, biogeography is influenced by the temporal variability in the conditions of the water mass. In addition, many marine benthic organisms develop in the water mass and rely on the conditions that link the pelagic and benthic systems for population maintenance. Such pelagic-coastal coupling indicates that changes in the trophic system during development can be transferred to the adult populations through changes in propagule supply. Thus, changes in environmental conditions can influence benthic populations directly (e.g. through larval advection) or indirectly, through their influence on the phytoplankton community (e.g. through the development of HABs). The South African coastline shows clear alongshore patterns of faunal biomass and species richness. On the south coast, strong longitudinal patterns of recruitment of intertidal organisms exist, with areas of particularly high recruitment. HABs of unprecedented spatio-temporal magnitude have recently developed along the south coast, including the areas where benthic recruitment is most intense. The present thesis used these blooms to study changes in intertidal recruitment directly or indirectly associated with their occurrence. Using a combination of remote sensing data to study the environmental conditions of the water mass in the innermost part of the Agulhas Bank, and estimates of mussel and barnacle recruitment rates to integrate the effects of conditions in the water mass during larval development, this thesis aimed to: (1) understand the conditions that triggered the development of an HAB of the dinoflagellate Lingulodinium polyedrum during summer of 2014, (2) determine the direct or indirect effects of that bloom on recruitment of intertidal organisms, and understand the factors that affect recruitment along the coast, (3) determine if the environmental factors during bloom development produced any carryover effects on recruit growth and mortality, and (4) determine the factors that drive changes in community biomass and composition along the south coast, the long-term trends in those factors, and possible changes experienced in recent years. Water column stability during spring, before the development of the red tide, followed by alternating periods of upwelling and relaxation during summer and autumn, seemed to promote the development and persistence of L. polyedrum. Recruitment of mussels and barnacles was estimated during the reproductive season of mussels in 2014, coinciding with the red tide, and during the following year. Alongshore patterns in recruitment were found, with higher mussel recruitment in the absence of the red tide and the opposite pattern in barnacles. Alongshore patterns in SST and chlorophyll matching those of recruitment were also found, with higher SSTs and lower chlorophyll during the red tide than the following year. Growth and mortality rates in barnacles did not differ between years during the first five months after settlement. This suggests that the factors which produced differences in recruitment between years did not produce carryover effects detectable at the temporal scales studied. Further analysis of 15 years of satellite-derived environmental data showed significant cooling trends potentially driven by a long-term seasonal acceleration of the Agulhas Current in autumn around two upwelling centres on the south coast, coinciding temporally with the reproductive period of mussels and barnacles, and spatially with the areas of highest recruitment. In addition, the comparison of SST and chl-a conditions during the first and the second half of the period of study showed that seasonality of both variables has changed in large areas over the shelf, with increasing importance of shorter-term variability, which would in turn decrease environmental predictability. Thus, the conditions observed during the present study, particularly during 2015, when upwelling seemed to be more intense, may presage the potential effects of identified long-term cooling trends at the upwelling centres. Although the general trend shows cooling around those areas, conditions can vary greatly among years, favouring different taxa. Changes in the Agulhas Current System are affected by changes in distant areas in the Indian Ocean basin. Such tele-connection is unlikely to be unique to this system and indicates the importance of understanding trends in major large scale climatic drivers and their regional effects in order to make predictions about coastal systems.
- Full Text:
- Authors: Muñiz, Carlota Fernández
- Date: 2019
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/75317 , vital:30400
- Description: Marine systems are driven by the relationships among organisms and environmental conditions. Anthropogenic-induced changes during the past decades have started to alter climatic drivers which have the potential to alter the physical, chemical and biological environment. In coastal systems, biogeography is influenced by the temporal variability in the conditions of the water mass. In addition, many marine benthic organisms develop in the water mass and rely on the conditions that link the pelagic and benthic systems for population maintenance. Such pelagic-coastal coupling indicates that changes in the trophic system during development can be transferred to the adult populations through changes in propagule supply. Thus, changes in environmental conditions can influence benthic populations directly (e.g. through larval advection) or indirectly, through their influence on the phytoplankton community (e.g. through the development of HABs). The South African coastline shows clear alongshore patterns of faunal biomass and species richness. On the south coast, strong longitudinal patterns of recruitment of intertidal organisms exist, with areas of particularly high recruitment. HABs of unprecedented spatio-temporal magnitude have recently developed along the south coast, including the areas where benthic recruitment is most intense. The present thesis used these blooms to study changes in intertidal recruitment directly or indirectly associated with their occurrence. Using a combination of remote sensing data to study the environmental conditions of the water mass in the innermost part of the Agulhas Bank, and estimates of mussel and barnacle recruitment rates to integrate the effects of conditions in the water mass during larval development, this thesis aimed to: (1) understand the conditions that triggered the development of an HAB of the dinoflagellate Lingulodinium polyedrum during summer of 2014, (2) determine the direct or indirect effects of that bloom on recruitment of intertidal organisms, and understand the factors that affect recruitment along the coast, (3) determine if the environmental factors during bloom development produced any carryover effects on recruit growth and mortality, and (4) determine the factors that drive changes in community biomass and composition along the south coast, the long-term trends in those factors, and possible changes experienced in recent years. Water column stability during spring, before the development of the red tide, followed by alternating periods of upwelling and relaxation during summer and autumn, seemed to promote the development and persistence of L. polyedrum. Recruitment of mussels and barnacles was estimated during the reproductive season of mussels in 2014, coinciding with the red tide, and during the following year. Alongshore patterns in recruitment were found, with higher mussel recruitment in the absence of the red tide and the opposite pattern in barnacles. Alongshore patterns in SST and chlorophyll matching those of recruitment were also found, with higher SSTs and lower chlorophyll during the red tide than the following year. Growth and mortality rates in barnacles did not differ between years during the first five months after settlement. This suggests that the factors which produced differences in recruitment between years did not produce carryover effects detectable at the temporal scales studied. Further analysis of 15 years of satellite-derived environmental data showed significant cooling trends potentially driven by a long-term seasonal acceleration of the Agulhas Current in autumn around two upwelling centres on the south coast, coinciding temporally with the reproductive period of mussels and barnacles, and spatially with the areas of highest recruitment. In addition, the comparison of SST and chl-a conditions during the first and the second half of the period of study showed that seasonality of both variables has changed in large areas over the shelf, with increasing importance of shorter-term variability, which would in turn decrease environmental predictability. Thus, the conditions observed during the present study, particularly during 2015, when upwelling seemed to be more intense, may presage the potential effects of identified long-term cooling trends at the upwelling centres. Although the general trend shows cooling around those areas, conditions can vary greatly among years, favouring different taxa. Changes in the Agulhas Current System are affected by changes in distant areas in the Indian Ocean basin. Such tele-connection is unlikely to be unique to this system and indicates the importance of understanding trends in major large scale climatic drivers and their regional effects in order to make predictions about coastal systems.
- Full Text:
Large scale spatio-temporal forcing of pelagic-coastal coupling: disentangling the effects of environmental change on intertidal invertebrate recruitment
- Authors: Muñiz, Carlota Fernández
- Date: 2019
- Subjects: Dinoflagellates -- South Africa , Coastal ecology -- South Africa , Climatic changes -- South Africa , Benthic ecology -- South Africa , Agulhas Current , Ocean temperature -- Agulhas Current , Ocean temperature -- Physiological effect
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/151541 , vital:39140
- Description: Marine systems are driven by the relationships among organisms and environmental conditions. Anthropogenic-induced changes during the past decades have started to alter climatic drivers which have the potential to alter the physical, chemical and biological environment. In coastal systems, biogeography is influenced by the temporal variability in the conditions of the water mass. In addition, many marine benthic organisms develop in the water mass and rely on the conditions that link the pelagic and benthic systems for population maintenance. Such pelagic-coastal coupling indicates that changes in the trophic system during development can be transferred to the adult populations through changes in propagule supply. Thus, changes in environmental conditions can influence benthic populations directly (e.g. through larval advection) or indirectly, through their influence on the phytoplankton community (e.g. through the development of HABs). The South African coastline shows clear alongshore patterns of faunal biomass and species richness. On the south coast, strong longitudinal patterns of recruitment of intertidal organisms exist, with areas of particularly high recruitment. HABs of unprecedented spatio-temporal magnitude have recently developed along the south coast, including the areas where benthic recruitment is most intense. The present thesis used these blooms to study changes in intertidal recruitment directly or indirectly associated with their occurrence. Using a combination of remote sensing data to study the environmental conditions of the water mass in the innermost part of the Agulhas Bank, and estimates of mussel and barnacle recruitment rates to integrate the effects of conditions in the water mass during larval development, this thesis aimed to: (1) understand the conditions that triggered the development of an HAB of the dinoflagellate Lingulodinium polyedrum during summer of 2014, (2) determine the direct or indirect effects of that bloom on recruitment of intertidal organisms, and understand the factors that affect recruitment along the coast, (3) determine if the environmental factors during bloom development produced any carryover effects on recruit growth and mortality, and (4) determine the factors that drive changes in community biomass and composition along the south coast, the long-term trends in those factors, and possible changes experienced in recent years. Water column stability during spring, before the development of the red tide, followed by alternating periods of upwelling and relaxation during summer and autumn, seemed to promote the development and persistence of L. polyedrum. Recruitment of mussels and barnacles was estimated during the reproductive season of mussels in 2014, coinciding with the red tide, and during the following year. Alongshore patterns in recruitment were found, with higher mussel recruitment in the absence of the red tide and the opposite pattern in barnacles. Alongshore patterns in SST and chlorophyll matching those of recruitment were also found, with higher SSTs and lower chlorophyll during the red tide than the following year. Growth and mortality rates in barnacles did not differ between years during the first five months after settlement. This suggests that the factors which produced differences in recruitment between years did not produce carryover effects detectable at the temporal scales studied. Further analysis of 15 years of satellite-derived environmental data showed significant cooling trends potentially driven by a long-term seasonal acceleration of the Agulhas Current in autumn around two upwelling centres on the south coast, coinciding temporally with the reproductive period of mussels and barnacles, and spatially with the areas of highest recruitment. In addition, the comparison of SST and chl-a conditions during the first and the second half of the period of study showed that seasonality of both variables has changed in large areas over the shelf, with increasing importance of shorter-term variability, which would in turn decrease environmental predictability. Thus, the conditions observed during the present study, particularly during 2015, when upwelling seemed to be more intense, may presage the potential effects of identified long-term cooling trends at the upwelling centres. Although the general trend shows cooling around those areas, conditions can vary greatly among years, favouring different taxa. Changes in the Agulhas Current System are affected by changes in distant areas in the Indian Ocean basin. Such tele-connection is unlikely to be unique to this system and indicates the importance of understanding trends in major large scale climatic drivers and their regional effects in order to make predictions about coastal systems.
- Full Text:
- Authors: Muñiz, Carlota Fernández
- Date: 2019
- Subjects: Dinoflagellates -- South Africa , Coastal ecology -- South Africa , Climatic changes -- South Africa , Benthic ecology -- South Africa , Agulhas Current , Ocean temperature -- Agulhas Current , Ocean temperature -- Physiological effect
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/151541 , vital:39140
- Description: Marine systems are driven by the relationships among organisms and environmental conditions. Anthropogenic-induced changes during the past decades have started to alter climatic drivers which have the potential to alter the physical, chemical and biological environment. In coastal systems, biogeography is influenced by the temporal variability in the conditions of the water mass. In addition, many marine benthic organisms develop in the water mass and rely on the conditions that link the pelagic and benthic systems for population maintenance. Such pelagic-coastal coupling indicates that changes in the trophic system during development can be transferred to the adult populations through changes in propagule supply. Thus, changes in environmental conditions can influence benthic populations directly (e.g. through larval advection) or indirectly, through their influence on the phytoplankton community (e.g. through the development of HABs). The South African coastline shows clear alongshore patterns of faunal biomass and species richness. On the south coast, strong longitudinal patterns of recruitment of intertidal organisms exist, with areas of particularly high recruitment. HABs of unprecedented spatio-temporal magnitude have recently developed along the south coast, including the areas where benthic recruitment is most intense. The present thesis used these blooms to study changes in intertidal recruitment directly or indirectly associated with their occurrence. Using a combination of remote sensing data to study the environmental conditions of the water mass in the innermost part of the Agulhas Bank, and estimates of mussel and barnacle recruitment rates to integrate the effects of conditions in the water mass during larval development, this thesis aimed to: (1) understand the conditions that triggered the development of an HAB of the dinoflagellate Lingulodinium polyedrum during summer of 2014, (2) determine the direct or indirect effects of that bloom on recruitment of intertidal organisms, and understand the factors that affect recruitment along the coast, (3) determine if the environmental factors during bloom development produced any carryover effects on recruit growth and mortality, and (4) determine the factors that drive changes in community biomass and composition along the south coast, the long-term trends in those factors, and possible changes experienced in recent years. Water column stability during spring, before the development of the red tide, followed by alternating periods of upwelling and relaxation during summer and autumn, seemed to promote the development and persistence of L. polyedrum. Recruitment of mussels and barnacles was estimated during the reproductive season of mussels in 2014, coinciding with the red tide, and during the following year. Alongshore patterns in recruitment were found, with higher mussel recruitment in the absence of the red tide and the opposite pattern in barnacles. Alongshore patterns in SST and chlorophyll matching those of recruitment were also found, with higher SSTs and lower chlorophyll during the red tide than the following year. Growth and mortality rates in barnacles did not differ between years during the first five months after settlement. This suggests that the factors which produced differences in recruitment between years did not produce carryover effects detectable at the temporal scales studied. Further analysis of 15 years of satellite-derived environmental data showed significant cooling trends potentially driven by a long-term seasonal acceleration of the Agulhas Current in autumn around two upwelling centres on the south coast, coinciding temporally with the reproductive period of mussels and barnacles, and spatially with the areas of highest recruitment. In addition, the comparison of SST and chl-a conditions during the first and the second half of the period of study showed that seasonality of both variables has changed in large areas over the shelf, with increasing importance of shorter-term variability, which would in turn decrease environmental predictability. Thus, the conditions observed during the present study, particularly during 2015, when upwelling seemed to be more intense, may presage the potential effects of identified long-term cooling trends at the upwelling centres. Although the general trend shows cooling around those areas, conditions can vary greatly among years, favouring different taxa. Changes in the Agulhas Current System are affected by changes in distant areas in the Indian Ocean basin. Such tele-connection is unlikely to be unique to this system and indicates the importance of understanding trends in major large scale climatic drivers and their regional effects in order to make predictions about coastal systems.
- Full Text:
The role of upwelling in determining the composition, species distribution and genetic structure of intertidal communities in a time of climate change
- Lourenço, Carla Sofia Emídio Rodrigues
- Authors: Lourenço, Carla Sofia Emídio Rodrigues
- Date: 2018
- Subjects: Upwelling (Oceanography) , Intertidal organisms -- Morocco -- Atlantic Coast , Intertidal organisms -- Canary Current -- Effect of water currents on , Intertidal animals -- Canary Current -- Effect of water currents on , Intertidal animals -- Morocco -- Atlantic Coas , Mytilus galloprovincialis -- Morocco -- Atlantic Coast , Mytilus galloprovincialis -- Canary Current -- Effect of water currents on , Intertidal ecology -- Canary Current , Sea surface microlayer -- Morocco -- Atlantic Coast
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/61614 , vital:28042
- Description: Upwelling is an oceanographic process that strongly influences coastal species and the communities they belong to. In upwelling areas, colder, denser, nutrient-rich subsurface waters are transported to the nearshore surface, replacing warmer superficial waters that are advected offshore. Such effects influence the composition and dynamics of coastal communities, for example by affecting species abundance, recruitment, dispersal and distribution. Upwelling areas are key model regions to study the responses of coastal species to climate change because they are characterized by cooler conditions and experience lower warming rates than adjacent regions. In particular, intertidal rocky shore species are ideal coastal sentinel organisms to study distributional changes driven by climate warming because they inhabit the interface between marine and terrestrial habitats and are exposed to extremely severe environmental conditions. In fact, sharp distributional shifts have been reported for multiple intertidal species as a response to ocean warming. Although some studies have investigated the role of upwelling in influencing abundance and distribution of intertidal species, little is known about its potential as refugia against climate warming and the degree to which upwelling shapes species genetic structure is yet not fully understood. The aim of this thesis is to understand the influence of the Canary Current upwelling system on intertidal community composition, including species distribution and the genetic structure of intertidal species under current climate change. To do this, I investigated community structure of intertidal assemblages along the Atlantic shores of Morocco and Western Sahara, performed large scale surveys on species distribution, evaluated species abundance and frequency of parasitism and examined species genetic patterns. I further coupled biological data with upwelling indices, sea surface temperatures (SST) and the rate of SST warming. I demonstrate that strong upwelling influences abundance and distribution of intertidal rocky shore species and that upwelling cells can act as refugia from climate change by ameliorating thermal conditions. Upwelling cells also conserve the genetic diversity of the marine macroalga Fucus guiryi, promoting intraspecific genetic diversity by preserving unique genetic lineages. However, no evidence was found that upwelling affects the genetic structure for either F. guiryi or the brown mussel Perna perna. Instead, the genetic patterns presented in this thesis seem to result from a combination of species’ life history traits, population size and habitat suitability. My results also suggest that upwelling intensity affects the frequency of endolithic parasitism on the Mediterranean mussel Mytilus galloprovincialis. In times of climate change, upwelling events provide suitable environmental conditions for species to counter act climatic change. As upwelling is project to intensify in the future, its influence on benthic intertidal species might be greater than previously anticipated.
- Full Text:
- Authors: Lourenço, Carla Sofia Emídio Rodrigues
- Date: 2018
- Subjects: Upwelling (Oceanography) , Intertidal organisms -- Morocco -- Atlantic Coast , Intertidal organisms -- Canary Current -- Effect of water currents on , Intertidal animals -- Canary Current -- Effect of water currents on , Intertidal animals -- Morocco -- Atlantic Coas , Mytilus galloprovincialis -- Morocco -- Atlantic Coast , Mytilus galloprovincialis -- Canary Current -- Effect of water currents on , Intertidal ecology -- Canary Current , Sea surface microlayer -- Morocco -- Atlantic Coast
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/61614 , vital:28042
- Description: Upwelling is an oceanographic process that strongly influences coastal species and the communities they belong to. In upwelling areas, colder, denser, nutrient-rich subsurface waters are transported to the nearshore surface, replacing warmer superficial waters that are advected offshore. Such effects influence the composition and dynamics of coastal communities, for example by affecting species abundance, recruitment, dispersal and distribution. Upwelling areas are key model regions to study the responses of coastal species to climate change because they are characterized by cooler conditions and experience lower warming rates than adjacent regions. In particular, intertidal rocky shore species are ideal coastal sentinel organisms to study distributional changes driven by climate warming because they inhabit the interface between marine and terrestrial habitats and are exposed to extremely severe environmental conditions. In fact, sharp distributional shifts have been reported for multiple intertidal species as a response to ocean warming. Although some studies have investigated the role of upwelling in influencing abundance and distribution of intertidal species, little is known about its potential as refugia against climate warming and the degree to which upwelling shapes species genetic structure is yet not fully understood. The aim of this thesis is to understand the influence of the Canary Current upwelling system on intertidal community composition, including species distribution and the genetic structure of intertidal species under current climate change. To do this, I investigated community structure of intertidal assemblages along the Atlantic shores of Morocco and Western Sahara, performed large scale surveys on species distribution, evaluated species abundance and frequency of parasitism and examined species genetic patterns. I further coupled biological data with upwelling indices, sea surface temperatures (SST) and the rate of SST warming. I demonstrate that strong upwelling influences abundance and distribution of intertidal rocky shore species and that upwelling cells can act as refugia from climate change by ameliorating thermal conditions. Upwelling cells also conserve the genetic diversity of the marine macroalga Fucus guiryi, promoting intraspecific genetic diversity by preserving unique genetic lineages. However, no evidence was found that upwelling affects the genetic structure for either F. guiryi or the brown mussel Perna perna. Instead, the genetic patterns presented in this thesis seem to result from a combination of species’ life history traits, population size and habitat suitability. My results also suggest that upwelling intensity affects the frequency of endolithic parasitism on the Mediterranean mussel Mytilus galloprovincialis. In times of climate change, upwelling events provide suitable environmental conditions for species to counter act climatic change. As upwelling is project to intensify in the future, its influence on benthic intertidal species might be greater than previously anticipated.
- Full Text:
Assessing the impact of climate change on mangrove crabs: the role of ontogenetic macrophysiology and settlement in the persistence of central and marginal populations
- Authors: Mostert, Bruce Petrus
- Date: 2016
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/826 , vital:19994
- Description: After a brief respite in the mid to late 20th century, macro physiology has come to the fore in elucidating large scale ecological patterns and processes as physiological assumptions often form the backbone of many predictive theories associated with species distributions. Critically, macro physiological patterns are valuable in explaining physiological variation across multiple scales and provide insights into the effects of climate change on populations spanning a wide range of latitudes. This can assist in predicting possible distribution expansions, contractions or shifts in light of current climate change scenarios. From this perspective, investigating intra- and inter-specific physiological responses to environmental stress may contribute to better understanding and predicting the effects of climate change on geographical ranges. Further, investigating the physiological effects to environmental stresses across ontogenetic stages allows for the identification of weak links within the lifecycle of a species. Additionally, determining settlement characteristics along a latitudinal cline provides integrated indications of the sustainability of populations, highlighting vulnerable regions in terms of repopulation of viable habitats. In this context, the present study aimed at establishing how temperature, in a physiological context, may affect reproductive biology of two species of mangrove crab, Perisesarma guttatum and Uca urvillei at the centre (Kenya) and edge (South Africa) of their distributional range along the east coast of Africa and highlight possible consequences for range distributions. A third species, Neosarmatium africanum, only in South Africa, was included to provide additional interspecies comparisons. Furthermore, settlement characteristics of brachyuran populations at the centre and edge of their distributional range were considered in order to determine how settlement may contribute to population persistence. Physiological investigations at the centre and edge of distributional range and across ontogenetic stages (larvae, stage 2 and 4 embryos, non-gravid and gravid females) under the concept of oxygen and capacity limitation of thermal tolerance (OCLTT), revealed that, for both species, populations at the centre of their distribution (Kenya) were generally more robust to increasing temperatures and generally displayed greater physiological stability with increasing temperatures compared to their conspecifics in South Africa. Variability in physiological robustness between regions, did however, differ among ontogenetic stages and species but, overall, were evident throughout. Within and between regions, adaptation to oxygen extraction in both milieus (air or water) was displayed for present temperature conditions but aerial respiration largely alleviated increased thermal stress due to overcoming the limitations of reduced oxygen availability and diffusiveness in water for all bimodal ontogenetic stages. Brooding eggs proved to be a physiologically critical process with either heighted oxygen consumption for gravid females or collapse of physiological processes demonstrated by supressed oxygen consumption. The physiological cost of brooding eggs, referred to as maternal costs, was reflected in in both Perisesarma guttatum and Uca urvillei where, in most cases, maternal costs were negative. Again, aerial respiration was able to alleviated increased thermal stress, as shown by positive maternal costs indicating sustained maternal care, but this mechanism was species and regionally specific. Settlement patterns differed between the edge and centre of distribution of the species studied. This difference was predominantly driven by zonal preference within the mangal and/or effects of new and full moon (lunar phase). Overall, settlement dynamics were more widely variable in South Africa, both spatially and temporally, than in Kenya. Finally, empirical physiological data from ontogenetic stages present during the reproductive process (early and late stage embryos) and from non-gravid and gravid females were used in conjunction with data mined from the existing literature to parameterise an individual based model designed to simulate reproductive output at the centre and edge of distribution of Perisesarma guttatum. Physiological data indicate that, in terms of reproductive output across increasing temperatures, populations based at the centre of their distribution presently outperform their counterparts at the edge of the species’ distribution, but reproductive output stagnated as temperature rose. Edge of distribution populations consistently increased reproductive output with increasing temperatures to eventually outperform centre of distribution populations at higher simulated temperatures. Overall, results of the physiological and settlement studies suggest that with increased climate change there may be a contraction of distributional range of the study species from high latitudes to low latitudes, contrary to general poleward shifts/migrations seen in most species, with possible contractions of the entire ecosystem mirroring the disappearance of keystone mangrove macrofauna.
- Full Text:
- Authors: Mostert, Bruce Petrus
- Date: 2016
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/826 , vital:19994
- Description: After a brief respite in the mid to late 20th century, macro physiology has come to the fore in elucidating large scale ecological patterns and processes as physiological assumptions often form the backbone of many predictive theories associated with species distributions. Critically, macro physiological patterns are valuable in explaining physiological variation across multiple scales and provide insights into the effects of climate change on populations spanning a wide range of latitudes. This can assist in predicting possible distribution expansions, contractions or shifts in light of current climate change scenarios. From this perspective, investigating intra- and inter-specific physiological responses to environmental stress may contribute to better understanding and predicting the effects of climate change on geographical ranges. Further, investigating the physiological effects to environmental stresses across ontogenetic stages allows for the identification of weak links within the lifecycle of a species. Additionally, determining settlement characteristics along a latitudinal cline provides integrated indications of the sustainability of populations, highlighting vulnerable regions in terms of repopulation of viable habitats. In this context, the present study aimed at establishing how temperature, in a physiological context, may affect reproductive biology of two species of mangrove crab, Perisesarma guttatum and Uca urvillei at the centre (Kenya) and edge (South Africa) of their distributional range along the east coast of Africa and highlight possible consequences for range distributions. A third species, Neosarmatium africanum, only in South Africa, was included to provide additional interspecies comparisons. Furthermore, settlement characteristics of brachyuran populations at the centre and edge of their distributional range were considered in order to determine how settlement may contribute to population persistence. Physiological investigations at the centre and edge of distributional range and across ontogenetic stages (larvae, stage 2 and 4 embryos, non-gravid and gravid females) under the concept of oxygen and capacity limitation of thermal tolerance (OCLTT), revealed that, for both species, populations at the centre of their distribution (Kenya) were generally more robust to increasing temperatures and generally displayed greater physiological stability with increasing temperatures compared to their conspecifics in South Africa. Variability in physiological robustness between regions, did however, differ among ontogenetic stages and species but, overall, were evident throughout. Within and between regions, adaptation to oxygen extraction in both milieus (air or water) was displayed for present temperature conditions but aerial respiration largely alleviated increased thermal stress due to overcoming the limitations of reduced oxygen availability and diffusiveness in water for all bimodal ontogenetic stages. Brooding eggs proved to be a physiologically critical process with either heighted oxygen consumption for gravid females or collapse of physiological processes demonstrated by supressed oxygen consumption. The physiological cost of brooding eggs, referred to as maternal costs, was reflected in in both Perisesarma guttatum and Uca urvillei where, in most cases, maternal costs were negative. Again, aerial respiration was able to alleviated increased thermal stress, as shown by positive maternal costs indicating sustained maternal care, but this mechanism was species and regionally specific. Settlement patterns differed between the edge and centre of distribution of the species studied. This difference was predominantly driven by zonal preference within the mangal and/or effects of new and full moon (lunar phase). Overall, settlement dynamics were more widely variable in South Africa, both spatially and temporally, than in Kenya. Finally, empirical physiological data from ontogenetic stages present during the reproductive process (early and late stage embryos) and from non-gravid and gravid females were used in conjunction with data mined from the existing literature to parameterise an individual based model designed to simulate reproductive output at the centre and edge of distribution of Perisesarma guttatum. Physiological data indicate that, in terms of reproductive output across increasing temperatures, populations based at the centre of their distribution presently outperform their counterparts at the edge of the species’ distribution, but reproductive output stagnated as temperature rose. Edge of distribution populations consistently increased reproductive output with increasing temperatures to eventually outperform centre of distribution populations at higher simulated temperatures. Overall, results of the physiological and settlement studies suggest that with increased climate change there may be a contraction of distributional range of the study species from high latitudes to low latitudes, contrary to general poleward shifts/migrations seen in most species, with possible contractions of the entire ecosystem mirroring the disappearance of keystone mangrove macrofauna.
- Full Text:
Diet of coastal filter feeders : impact of factors operating at different scales
- Authors: Puccinelli, Eleonora
- Date: 2015
- Subjects: Benthic animals -- Ecology -- South Africa , Benthic animals -- Nutrition
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5929 , http://hdl.handle.net/10962/d1017807
- Description: Benthic filter feeders have a key functional role in the dynamics of coastal food web as an intermediate trophic level and bioengineers. A wide variety of factors, operating across multiple spatial scales (e.g. hydrographic regime, human activities), can affect the composition of the water column and thus the availability of food for benthic populations. Food availability in turn affects the growth, reproductive rates and survival of benthic organisms, and consequently, can influence the functioning of the entire ecosystem. This study aims to evaluate how various environmental factors may modify the diet of intertidal filter feeders living along the South African coast. Specifically, the effects of biogeography, upwelling, urbanization and freshwater input on the dietary regimes of five species of filter feeders (two mussel and three barnacle species) were investigated using fatty acid (FA) and stable isotope (SI) analyses.Strong interspecific differences were found among the five species considered. However, all species responded to factors operating at large (100s km) and meso (10s- 100s km) scales (i.e. biogeography and upwelling respectively). The barnacles exhibit habitat segregation and showed different FA and SI signatures from each other, while the two mussel species, an invasive and native species that co-occur in the same mussel beds, had partially overlapping diets. Differences in their diets were found only using FA analysis, while their SI signatures differed on only one occasion. This highlights the importance of using the appropriate tool, and ideally combined techniques, to investigate diets.FA and SI signatures of all species considered changed among the three biographical provinces (west, south and east coasts of South Africa) exhibiting similar patterns that reflect the two oceanographic regimes that characterize the coastline: the eutrophic Benguela Current on the west coast and the oligotrophic Agulhas Current on the other two coasts. Upwelling had a significant effect on FA and SI signatures, with stronger effects on the west coast than the south coast. The results indicate that benthic filter feeders at upwelling areas consumed a mix of coastal macroalgal detritus and phytoplankton, which was probably brought onshore during downwelling events. At smaller spatialscales and using repeated sampling, the influence of upwelling on the west coast was found to be pervasive, rather than discrete, so that it may be more appropriate to categorize upwelling by referring to upwelling centres and downstream areas. SI underlined a significant effect of urbanization on the diet of filter feeders with an enrichment in the δ¹⁵N being characteristic of anthropogenic effect. Although a large number of rivers characterize the South African coast, no distinct effect of freshwater input was found for either the SI or FA signatures of the filter feeders. This contrasts with earlier work on demersal species and suggests that freshwater input does not significantly affect food availability for intertidal filter feeders, and that other factors (e.g. hydrogeography) are more important in determining the diet of these populations. These results highlight that environmental and anthropogenic factors operating at different spatial and temporal scales have a profound effect on benthic ecosystems, and that they control the relationship between primary production and primary consumers in coastal areas. Above all, this work highlights the importance of understanding the spatial and temporal scales at which different factors affect feeding regimes, and their critical role in coastal food webs.
- Full Text:
- Authors: Puccinelli, Eleonora
- Date: 2015
- Subjects: Benthic animals -- Ecology -- South Africa , Benthic animals -- Nutrition
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5929 , http://hdl.handle.net/10962/d1017807
- Description: Benthic filter feeders have a key functional role in the dynamics of coastal food web as an intermediate trophic level and bioengineers. A wide variety of factors, operating across multiple spatial scales (e.g. hydrographic regime, human activities), can affect the composition of the water column and thus the availability of food for benthic populations. Food availability in turn affects the growth, reproductive rates and survival of benthic organisms, and consequently, can influence the functioning of the entire ecosystem. This study aims to evaluate how various environmental factors may modify the diet of intertidal filter feeders living along the South African coast. Specifically, the effects of biogeography, upwelling, urbanization and freshwater input on the dietary regimes of five species of filter feeders (two mussel and three barnacle species) were investigated using fatty acid (FA) and stable isotope (SI) analyses.Strong interspecific differences were found among the five species considered. However, all species responded to factors operating at large (100s km) and meso (10s- 100s km) scales (i.e. biogeography and upwelling respectively). The barnacles exhibit habitat segregation and showed different FA and SI signatures from each other, while the two mussel species, an invasive and native species that co-occur in the same mussel beds, had partially overlapping diets. Differences in their diets were found only using FA analysis, while their SI signatures differed on only one occasion. This highlights the importance of using the appropriate tool, and ideally combined techniques, to investigate diets.FA and SI signatures of all species considered changed among the three biographical provinces (west, south and east coasts of South Africa) exhibiting similar patterns that reflect the two oceanographic regimes that characterize the coastline: the eutrophic Benguela Current on the west coast and the oligotrophic Agulhas Current on the other two coasts. Upwelling had a significant effect on FA and SI signatures, with stronger effects on the west coast than the south coast. The results indicate that benthic filter feeders at upwelling areas consumed a mix of coastal macroalgal detritus and phytoplankton, which was probably brought onshore during downwelling events. At smaller spatialscales and using repeated sampling, the influence of upwelling on the west coast was found to be pervasive, rather than discrete, so that it may be more appropriate to categorize upwelling by referring to upwelling centres and downstream areas. SI underlined a significant effect of urbanization on the diet of filter feeders with an enrichment in the δ¹⁵N being characteristic of anthropogenic effect. Although a large number of rivers characterize the South African coast, no distinct effect of freshwater input was found for either the SI or FA signatures of the filter feeders. This contrasts with earlier work on demersal species and suggests that freshwater input does not significantly affect food availability for intertidal filter feeders, and that other factors (e.g. hydrogeography) are more important in determining the diet of these populations. These results highlight that environmental and anthropogenic factors operating at different spatial and temporal scales have a profound effect on benthic ecosystems, and that they control the relationship between primary production and primary consumers in coastal areas. Above all, this work highlights the importance of understanding the spatial and temporal scales at which different factors affect feeding regimes, and their critical role in coastal food webs.
- Full Text:
Fishing for resilience : herbivore and algal dynamics on coral reefs in Kenya.
- Authors: Humphries, Austin Turner
- Date: 2014
- Subjects: Coral reef conservation -- Kenya , Coral reef ecology -- Kenya , Coral reef biology -- Kenya , Coral reef fishes -- Kenya , Herbivores -- Kenya , Algae -- Control -- Kenya , Fishery management -- Kenya
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5877 , http://hdl.handle.net/10962/d1013147
- Description: Herbivory is a key process that mediates the abundance of primary producers and community composition in both terrestrial and aquatic ecosystems. On tropical coral reefs, changes in herbivory are often related to phase shifts between coral-dominance and dominance by seaweeds, or foliose macroalgae. Resilience or capacity to resist and reverse such phase shifts is, therefore, viewed as a critical function on coral reefs. This thesis used grazer exclusion and assay experiments at six sites within three different fisheries management regimes in Kenya to identify the impacts of herbivores (sea urchins and fishes) on algal dynamics in the context of coral reef resilience. First, I examined the grazing rates necessary to prevent phase shifts by quantifying consumption and algal production. Here, I found that, over a 390-day experiment, at least 50 percent of algal production must be consumed to avoid accumulation of algal biomass. Using video observations, I also showed that scraping parrotfishes remove more algae (per unit of fish biomass) than previously assumed, and that sea urchins, if released from predation, have similar impacts to fishes. Then I focused on algal succession, and found that sea urchins and fishes have different effects that are mediated by their abundances and species composition. Where sea urchins were less abundant and parrotfishes absent (e.g. young fisheries closures), progression of algae from turfs to early and then late successional macroalgae occurred rapidly and within 100 days. I then turned my focus to the removal of already established macroalgae (grown for > 1 yr in the absence of herbivores) and showed that sea urchins and browsing fishes were able to remove significant amounts of macroalgae where either herbivore was abundant. However, using multiple-choice selectivity assays and in situ video recordings, I found that browsing fishes fed very selectively with low overlap in diet among species, leading to low functional redundancy within a high diversity system. Finally, using long-term survey data (from 28 sites) to build a 43-year chronosequence, I showed that it is possible that the effects of herbivory will not be constant across transitions from open fishing to fishery closures through non-linear grazing intensity. Therefore, increases in herbivory within fisheries closures may not be immediate and may allow a window of opportunity for algae to go from turf to unpalatable macroalgae until scraping and browsing fishes fully recover from fishing (~ 20 years). The findings in this thesis are novel and raise concern over the potential implications of the slow recovery of parrotfishes or, given lower than expected functional redundancy in grazing effects, the absence of even one browsing fish species in fisheries closures. Overall, this thesis highlights the importance of herbivore community dynamics in mediating interactions among algae, and provides new insights for conservation and management actions that attempt to bolster the resilience of coral reefs.
- Full Text:
- Authors: Humphries, Austin Turner
- Date: 2014
- Subjects: Coral reef conservation -- Kenya , Coral reef ecology -- Kenya , Coral reef biology -- Kenya , Coral reef fishes -- Kenya , Herbivores -- Kenya , Algae -- Control -- Kenya , Fishery management -- Kenya
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5877 , http://hdl.handle.net/10962/d1013147
- Description: Herbivory is a key process that mediates the abundance of primary producers and community composition in both terrestrial and aquatic ecosystems. On tropical coral reefs, changes in herbivory are often related to phase shifts between coral-dominance and dominance by seaweeds, or foliose macroalgae. Resilience or capacity to resist and reverse such phase shifts is, therefore, viewed as a critical function on coral reefs. This thesis used grazer exclusion and assay experiments at six sites within three different fisheries management regimes in Kenya to identify the impacts of herbivores (sea urchins and fishes) on algal dynamics in the context of coral reef resilience. First, I examined the grazing rates necessary to prevent phase shifts by quantifying consumption and algal production. Here, I found that, over a 390-day experiment, at least 50 percent of algal production must be consumed to avoid accumulation of algal biomass. Using video observations, I also showed that scraping parrotfishes remove more algae (per unit of fish biomass) than previously assumed, and that sea urchins, if released from predation, have similar impacts to fishes. Then I focused on algal succession, and found that sea urchins and fishes have different effects that are mediated by their abundances and species composition. Where sea urchins were less abundant and parrotfishes absent (e.g. young fisheries closures), progression of algae from turfs to early and then late successional macroalgae occurred rapidly and within 100 days. I then turned my focus to the removal of already established macroalgae (grown for > 1 yr in the absence of herbivores) and showed that sea urchins and browsing fishes were able to remove significant amounts of macroalgae where either herbivore was abundant. However, using multiple-choice selectivity assays and in situ video recordings, I found that browsing fishes fed very selectively with low overlap in diet among species, leading to low functional redundancy within a high diversity system. Finally, using long-term survey data (from 28 sites) to build a 43-year chronosequence, I showed that it is possible that the effects of herbivory will not be constant across transitions from open fishing to fishery closures through non-linear grazing intensity. Therefore, increases in herbivory within fisheries closures may not be immediate and may allow a window of opportunity for algae to go from turf to unpalatable macroalgae until scraping and browsing fishes fully recover from fishing (~ 20 years). The findings in this thesis are novel and raise concern over the potential implications of the slow recovery of parrotfishes or, given lower than expected functional redundancy in grazing effects, the absence of even one browsing fish species in fisheries closures. Overall, this thesis highlights the importance of herbivore community dynamics in mediating interactions among algae, and provides new insights for conservation and management actions that attempt to bolster the resilience of coral reefs.
- Full Text:
Inter-individual variability and phenotypic plasticity : the effect of the environment on the biogeography, population structure, ecophysiology and reproduction of the sandhoppers Talorchestia capensis and Africorchestia quadrispinosa
- Authors: Baldanzi, Simone
- Date: 2014
- Subjects: Phenotypic plasticity -- Research -- Africa, Southern Talitridae -- Research -- Africa, Southern Amphipoda -- Research -- Africa, Southern Climatic changes -- Environmental aspects -- Africa, Southern Marine biology -- Africa, Southern Adaptation (Biology) -- Africa, Southern
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5846 , http://hdl.handle.net/10962/d1011447
- Description: Climatic envelope models focus on the climatic variables affecting species or species assemblages, and are important tools to investigate the effect of climate change on their geographical ranges. These models have largely been proposed in order to make successful predictions on species‘ persistence, determining which variables are likely to induce range expansion, contraction, or shifting. More recent models, including the ability and the cost for individuals to respond promptly to an environmental stimulus, have revealed that species may express phenotypic plasticity able to induce adaptation to the new environment. Consequently, understanding how species evolve to a changing climate is fundamental. From this perspective, investigating intraspecific responses to an environmental variable may contribute to better understanding and prediction of the effect of climate change on the geographical range and evolution of species, particularly in the case of widespread species. In this context, the present study aimed at establishing how environmental variables (focussing mainly on temperature) may have contributed to shape the spatial distribution, physiology, reproductive biology and connectivity of two species of Southern African sandhoppers (Talorchestia capensis and Africorchestia quadrispinosa, Amphipoda, Talitridae). Most of the work was carried out on T. capensis, due to its widespread spatial distribution. A first investigation of the biogeography of T. capensis and A. quadrispinosa, revealed that, for both species, spatial patterns of abundance, size and sex ratio were not explained by the Abundant Centre Hypothesis (greater abundance at the core of a spatial range), but rather guided by bio-physical forces. Precisely, the abundance of sandhoppers was driven by the morphodynamic state of the beach, salinity and temperatures, with strong differentiation among sites that reflected local environmental conditions. In support of these findings, strong population structure in the genetics of T. capensis was found (three main groups) when investigating its phylogeography and genetic connectivity. Although such defined structure may suggests cryptic speciation, the concomitant within-population variation in the COX1 region of mtDNA, also highlighted the importance of individual genetic variability. High individual variability was also found in the response of T. capensis to temperature, both in its physiology (thermal plasticity) and its reproductive biology (maternal effects). Since temperature is one of the main variables affecting the coastal marine systems of southern Africa and the metabolism of animals in general, its effect on the physiology and reproduction of T. capensis was therefore investigated. Thermal responses to increasing/decreasing temperatures were assessed for separated populations of T. capensis. Individual variability was reported in the oxygen consumption of T. capensis in response to temperature (high variation around the means, especially for increasing temperatures). Among population differences in thermal sensitivity were significantly correlated with air temperature variability experienced over the past 23 years, highlighting the importance of historical temperature fluctuations to the current thermal physiology of these sandhoppers. Temperature also had an important effect on the reproductive plasticity of T. capensis. Different temperatures induced mothers to adjust the size of their offspring (i.e. egg size), with larger eggs produced at lower temperatures. Interestingly, females showed strongly significant among individual variation in the size of the eggs. Given the importance of understanding rapid responses of organisms to climate change and considering the fundamental role played by phenotypic plasticity in evolution, the overall study revealed the significance of individual plasticity and variability in response to the environment and highlighted its importance. Particularly, studying the thermal physiology of separated populations and understanding within population reproductive plasticity in response to temperature, helped to clarify how differences among individual responses have important consequences at the population level, possibly explaining the widespread distribution of T. capensis.
- Full Text:
- Authors: Baldanzi, Simone
- Date: 2014
- Subjects: Phenotypic plasticity -- Research -- Africa, Southern Talitridae -- Research -- Africa, Southern Amphipoda -- Research -- Africa, Southern Climatic changes -- Environmental aspects -- Africa, Southern Marine biology -- Africa, Southern Adaptation (Biology) -- Africa, Southern
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5846 , http://hdl.handle.net/10962/d1011447
- Description: Climatic envelope models focus on the climatic variables affecting species or species assemblages, and are important tools to investigate the effect of climate change on their geographical ranges. These models have largely been proposed in order to make successful predictions on species‘ persistence, determining which variables are likely to induce range expansion, contraction, or shifting. More recent models, including the ability and the cost for individuals to respond promptly to an environmental stimulus, have revealed that species may express phenotypic plasticity able to induce adaptation to the new environment. Consequently, understanding how species evolve to a changing climate is fundamental. From this perspective, investigating intraspecific responses to an environmental variable may contribute to better understanding and prediction of the effect of climate change on the geographical range and evolution of species, particularly in the case of widespread species. In this context, the present study aimed at establishing how environmental variables (focussing mainly on temperature) may have contributed to shape the spatial distribution, physiology, reproductive biology and connectivity of two species of Southern African sandhoppers (Talorchestia capensis and Africorchestia quadrispinosa, Amphipoda, Talitridae). Most of the work was carried out on T. capensis, due to its widespread spatial distribution. A first investigation of the biogeography of T. capensis and A. quadrispinosa, revealed that, for both species, spatial patterns of abundance, size and sex ratio were not explained by the Abundant Centre Hypothesis (greater abundance at the core of a spatial range), but rather guided by bio-physical forces. Precisely, the abundance of sandhoppers was driven by the morphodynamic state of the beach, salinity and temperatures, with strong differentiation among sites that reflected local environmental conditions. In support of these findings, strong population structure in the genetics of T. capensis was found (three main groups) when investigating its phylogeography and genetic connectivity. Although such defined structure may suggests cryptic speciation, the concomitant within-population variation in the COX1 region of mtDNA, also highlighted the importance of individual genetic variability. High individual variability was also found in the response of T. capensis to temperature, both in its physiology (thermal plasticity) and its reproductive biology (maternal effects). Since temperature is one of the main variables affecting the coastal marine systems of southern Africa and the metabolism of animals in general, its effect on the physiology and reproduction of T. capensis was therefore investigated. Thermal responses to increasing/decreasing temperatures were assessed for separated populations of T. capensis. Individual variability was reported in the oxygen consumption of T. capensis in response to temperature (high variation around the means, especially for increasing temperatures). Among population differences in thermal sensitivity were significantly correlated with air temperature variability experienced over the past 23 years, highlighting the importance of historical temperature fluctuations to the current thermal physiology of these sandhoppers. Temperature also had an important effect on the reproductive plasticity of T. capensis. Different temperatures induced mothers to adjust the size of their offspring (i.e. egg size), with larger eggs produced at lower temperatures. Interestingly, females showed strongly significant among individual variation in the size of the eggs. Given the importance of understanding rapid responses of organisms to climate change and considering the fundamental role played by phenotypic plasticity in evolution, the overall study revealed the significance of individual plasticity and variability in response to the environment and highlighted its importance. Particularly, studying the thermal physiology of separated populations and understanding within population reproductive plasticity in response to temperature, helped to clarify how differences among individual responses have important consequences at the population level, possibly explaining the widespread distribution of T. capensis.
- Full Text:
Molecular phylogeny, phylogeography and evolutionary adaptation of foraging behaviour amongst sympatric patellid limpets along the southern African shoreline
- Authors: Mmonwa, Kolobe Lucas
- Date: 2014
- Subjects: Limpets -- South Africa -- Adaptation , Limpets -- South Africa -- Ecology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5882 , http://hdl.handle.net/10962/d1013222
- Description: The southern African shoreline is inhabited by a great diversity of patellid limpets of which most are endemic to South Africa. These limpets have evolved foraging mechanisms that partition ecological resources and reduce interspecific competition, resulting in ecological specialists and generalists. The evolution of ecological specialization or generalization remains poorly understood and there is no agreement on how such evolutionary transitions are correlated with levels of genetic diversity. This study investigated evolutionary correlations between territoriality in foraging and genetic structure of southern African patellid limpets (Cymbula and Scutellastra spp.) using stable δ13C and δ15N ratios, mitochondrial and nuclear DNA markers. The outcomes of the study were best rationalized and understood in the context of the scale of analysis in terms of both space and taxonomy. At biogeographic scales and the interspecific level, the stable isotope signatures and genetic structure of these limpets are determined by extrinsic factors such as biogeography and oceanography. However, at the smallest site scales and the intraspecific level, the isotope signatures and genetic diversity of these limpets are significantly correlated to their foraging traits. At large scales, there are no correlations between foraging behaviour and either genetic structure or isotope signature. At smaller scales, territorial Scutellastra foragers display both isotopic enrichment and greater haplotype diversity than congeneric non-territorial foragers. Thus, the isotope signatures and genetic structure of these limpets are determined by intrinsic species-specific response linked to their foraging behaviour. However, this pattern was intriguing as differences between territorial and non-territorial limpets in both isotopic signature and genetic diversity were observed only along the south coast when the same species were compared in other biogeographic regions. The significant interaction effect between foraging behaviour and stable isotope signatures was only observed from the sites within the Agulhas Bank or which are strongly influenced by the Agulhas current. This south enrichment in isotopic ratios is due to the mixing differences between onshore and offshore waters as the Agulhas current moves from east to south. At the generic level, the correlation between foraging behaviour and isotope signatures and genetic structure were particularly profound for Scutellastra species. The molecular phylogeny revealed deep evolutionary divergence between territorial and non-territorial Scutellastra spp. This divergence was concordant with morphological differences in shell shape and radula anatomy between territorial and non-territorial species. A taxonomic review of the scutellastrid spp. is proposed, suggesting possible re-consideration of the genus as two genera characterized by either territoriality or non-territoriality. The divergence between territorial and non-territorial species in both Scutellastra and Cymbula took place approximately in the early Oligocene. Major climatic cooling and decreases in sea level occurred during the Oligocene and this probably exposed much of the lower intertidal zone, increasing new potential habitats and algal availability. The Oligocene exposure of rocky shores and algal abundance in the lower intertidal zone probably elicited resource partitioning amongst these patellid limpets and subsequently, the evolution of territorial and non-territorial species. Analyses of the demographic history of these patellid limpets revealed evidence of post-glacial spatial expansion around the Pleistocene, implying these limpets were at population equilibrium during the dramatic LGM sea temperatures. Thus, these limpets managed to expand their range during dynamic oceanographic oscillations and dramatic sea-level changes in the Pleistocene. This study highlighted the importance of applying ecological traits as a subject to investigate and comprehend the evolutionary ecology of marine herbivores. The foraging traits of these true limpets are reflected in both their stable isotope ratios and genealogy, presumably as an evolutionary consequence of competition.
- Full Text:
- Authors: Mmonwa, Kolobe Lucas
- Date: 2014
- Subjects: Limpets -- South Africa -- Adaptation , Limpets -- South Africa -- Ecology
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5882 , http://hdl.handle.net/10962/d1013222
- Description: The southern African shoreline is inhabited by a great diversity of patellid limpets of which most are endemic to South Africa. These limpets have evolved foraging mechanisms that partition ecological resources and reduce interspecific competition, resulting in ecological specialists and generalists. The evolution of ecological specialization or generalization remains poorly understood and there is no agreement on how such evolutionary transitions are correlated with levels of genetic diversity. This study investigated evolutionary correlations between territoriality in foraging and genetic structure of southern African patellid limpets (Cymbula and Scutellastra spp.) using stable δ13C and δ15N ratios, mitochondrial and nuclear DNA markers. The outcomes of the study were best rationalized and understood in the context of the scale of analysis in terms of both space and taxonomy. At biogeographic scales and the interspecific level, the stable isotope signatures and genetic structure of these limpets are determined by extrinsic factors such as biogeography and oceanography. However, at the smallest site scales and the intraspecific level, the isotope signatures and genetic diversity of these limpets are significantly correlated to their foraging traits. At large scales, there are no correlations between foraging behaviour and either genetic structure or isotope signature. At smaller scales, territorial Scutellastra foragers display both isotopic enrichment and greater haplotype diversity than congeneric non-territorial foragers. Thus, the isotope signatures and genetic structure of these limpets are determined by intrinsic species-specific response linked to their foraging behaviour. However, this pattern was intriguing as differences between territorial and non-territorial limpets in both isotopic signature and genetic diversity were observed only along the south coast when the same species were compared in other biogeographic regions. The significant interaction effect between foraging behaviour and stable isotope signatures was only observed from the sites within the Agulhas Bank or which are strongly influenced by the Agulhas current. This south enrichment in isotopic ratios is due to the mixing differences between onshore and offshore waters as the Agulhas current moves from east to south. At the generic level, the correlation between foraging behaviour and isotope signatures and genetic structure were particularly profound for Scutellastra species. The molecular phylogeny revealed deep evolutionary divergence between territorial and non-territorial Scutellastra spp. This divergence was concordant with morphological differences in shell shape and radula anatomy between territorial and non-territorial species. A taxonomic review of the scutellastrid spp. is proposed, suggesting possible re-consideration of the genus as two genera characterized by either territoriality or non-territoriality. The divergence between territorial and non-territorial species in both Scutellastra and Cymbula took place approximately in the early Oligocene. Major climatic cooling and decreases in sea level occurred during the Oligocene and this probably exposed much of the lower intertidal zone, increasing new potential habitats and algal availability. The Oligocene exposure of rocky shores and algal abundance in the lower intertidal zone probably elicited resource partitioning amongst these patellid limpets and subsequently, the evolution of territorial and non-territorial species. Analyses of the demographic history of these patellid limpets revealed evidence of post-glacial spatial expansion around the Pleistocene, implying these limpets were at population equilibrium during the dramatic LGM sea temperatures. Thus, these limpets managed to expand their range during dynamic oceanographic oscillations and dramatic sea-level changes in the Pleistocene. This study highlighted the importance of applying ecological traits as a subject to investigate and comprehend the evolutionary ecology of marine herbivores. The foraging traits of these true limpets are reflected in both their stable isotope ratios and genealogy, presumably as an evolutionary consequence of competition.
- Full Text:
Genetics and thermal biology of littorinid snails of the genera Afrolittorina, Echinolittorina and Littoraria (Gastropoda: Littorinidae) from temperate, subtropical and tropical regions
- Authors: Matumba, Tshifhiwa Given
- Date: 2013
- Subjects: Littorinidae -- South Africa Littorinidae -- Brunei Littorinidae -- Adaptation -- South Africa Littorinidae -- Adaptation -- Brunei Littorinidae -- Effect of temperature on -- South Africa Littorinidae -- Effect of temperature on -- Brunei Littorinidae -- Physiology Snails -- South Africa Snails -- Brunei Gastropoda -- South Africa Gastropoda -- Brunei Intertidal animals -- South Africa Intertidal animals -- Brunei
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5588 , http://hdl.handle.net/10962/d1001953
- Description: With the anticipated effects of climate change due to global warming, there is concern over how animals, especially ectotherms, will respond to or tolerate extreme and fluctuating environmental temperature stress. Littorinid snails are intertidal ectotherms that live high on the shore where they experience both extreme and variable conditions of temperature and desiccation stress, and are believed to live close to their tolerance limits. This study investigated the thermal biology of littorinid snails of the genera Afrolittorina, Echinolittorina and Littoraria from temperate, subtropical and tropical regions in South Africa and Brunei Darussalam using thermal tolerance, heart function, and proteome approaches. The effects of conditions, such as rate of change in temperature, acclimation, heat shock, season and starvation were also tested. In addition, the evolutionary relationships and genetic diversity between and within the South African Afrolittorina spp. were investigated using mitochondrial and nuclear markers. Genetic results confirmed that these are two distinct species, with the brown to black A. knysnaensis predominant in the cool-temperate region of South Africa and the pale blue-grey A. africana in the subtropical region. There was low genetic variation and differentiation within each species, suggesting high gene flow among populations as a result of the effects of ocean currents on the dispersal of their planktotrophic larvae. Tests using exposure to high temperatures revealed differences in the thermal tolerances, heart performance and protein profiles of species from different latitudes, regions and zones on the shore. Thermal tolerance conformed to expectations, with clear, statistically significant trends from high tolerance in subtropical species to lower tolerance in temperate species. However, for Afrolittorina spp., there were no significant differences in the thermal tolerances of conspecifics from different regions, though there was a significant difference in thermal tolerance between juveniles and adults. Overall, adults of all species showed higher thermal tolerances than juveniles. Although lethal temperatures for these species were higher in summer than winter, laboratory acclimation had no effect on heat coma temperatures. All species showed some regulation of heart rate, with a degree of independence of heart rate from temperature across mid-range temperatures. The tropical species showed quick induction and good regulation of heart rate followed by the subtropical and temperate species, which displayed mixed responses including regulation, partial regulation and lack of regulation. Overall, tropical Echinolittorina spp. showed good regulation, while the subtropical E. natalensis and Littoraria glabrata exhibited a mixture of partial regulation and regulation. The subtropical/temperate Afrolittorina spp. showed high individual variability, some animals exhibiting regulation, while others did not. These effects seem to be largely phylogenetically determined as there were no differences in the heart rate responses of Afrolittorina spp. from different regions. The temperatures at which heart rate became independent of temperature (thermoneutral zone) were within the range experienced under natural conditions. In addition, there were differences in Arrhenius breakpoint and endpoint temperatures, showing a trend from higher in tropical animals to lower for temperate animals. Conditions such as acclimation, heat shock and starvation had little or no effect on heart performance. However, a slow increase in temperature induced good regulation of heart rate with noticeable shifts of breakpoints and endpoints for Afrolittorina spp. Lastly, there were differences in the proteome responses between and within Afrolittorina spp. as a function of species, size and treatment. Although both large and small A. knysnaensis had a greater number of protein spots in their proteome than A. africana (though the difference was not significant), the later showed significantly higher differential expression of certain proteins following heat stress. In addition, juveniles of both species displayed greater numbers of protein spots in their proteome than adults. The results indicate a difference in the physiological and biochemical responses (i.e. adaptations) of these snails to temperature, and this seems to relate to differences in biogeography, phylogeny, species identity and ecology. The ability to regulate heart rate is phylogenetically determined, while thresholds and lethal limits correspond to biogeography and species ecology. The proteome seems to correspond to species ecology. The results also indicate that these littorinids can tolerate high temperature stress and in this respect they are well suited to life in the intertidal zones or habitats where temperature and other stresses or conditions are extreme and can change abruptly. However, the limited ability of these snails to acclimate to different temperatures suggests that they are already living close to their tolerance limits with small safety margins or narrow thermal windows and so may be vulnerable to small rises in substratum temperature and/or solar radiation.
- Full Text:
- Authors: Matumba, Tshifhiwa Given
- Date: 2013
- Subjects: Littorinidae -- South Africa Littorinidae -- Brunei Littorinidae -- Adaptation -- South Africa Littorinidae -- Adaptation -- Brunei Littorinidae -- Effect of temperature on -- South Africa Littorinidae -- Effect of temperature on -- Brunei Littorinidae -- Physiology Snails -- South Africa Snails -- Brunei Gastropoda -- South Africa Gastropoda -- Brunei Intertidal animals -- South Africa Intertidal animals -- Brunei
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5588 , http://hdl.handle.net/10962/d1001953
- Description: With the anticipated effects of climate change due to global warming, there is concern over how animals, especially ectotherms, will respond to or tolerate extreme and fluctuating environmental temperature stress. Littorinid snails are intertidal ectotherms that live high on the shore where they experience both extreme and variable conditions of temperature and desiccation stress, and are believed to live close to their tolerance limits. This study investigated the thermal biology of littorinid snails of the genera Afrolittorina, Echinolittorina and Littoraria from temperate, subtropical and tropical regions in South Africa and Brunei Darussalam using thermal tolerance, heart function, and proteome approaches. The effects of conditions, such as rate of change in temperature, acclimation, heat shock, season and starvation were also tested. In addition, the evolutionary relationships and genetic diversity between and within the South African Afrolittorina spp. were investigated using mitochondrial and nuclear markers. Genetic results confirmed that these are two distinct species, with the brown to black A. knysnaensis predominant in the cool-temperate region of South Africa and the pale blue-grey A. africana in the subtropical region. There was low genetic variation and differentiation within each species, suggesting high gene flow among populations as a result of the effects of ocean currents on the dispersal of their planktotrophic larvae. Tests using exposure to high temperatures revealed differences in the thermal tolerances, heart performance and protein profiles of species from different latitudes, regions and zones on the shore. Thermal tolerance conformed to expectations, with clear, statistically significant trends from high tolerance in subtropical species to lower tolerance in temperate species. However, for Afrolittorina spp., there were no significant differences in the thermal tolerances of conspecifics from different regions, though there was a significant difference in thermal tolerance between juveniles and adults. Overall, adults of all species showed higher thermal tolerances than juveniles. Although lethal temperatures for these species were higher in summer than winter, laboratory acclimation had no effect on heat coma temperatures. All species showed some regulation of heart rate, with a degree of independence of heart rate from temperature across mid-range temperatures. The tropical species showed quick induction and good regulation of heart rate followed by the subtropical and temperate species, which displayed mixed responses including regulation, partial regulation and lack of regulation. Overall, tropical Echinolittorina spp. showed good regulation, while the subtropical E. natalensis and Littoraria glabrata exhibited a mixture of partial regulation and regulation. The subtropical/temperate Afrolittorina spp. showed high individual variability, some animals exhibiting regulation, while others did not. These effects seem to be largely phylogenetically determined as there were no differences in the heart rate responses of Afrolittorina spp. from different regions. The temperatures at which heart rate became independent of temperature (thermoneutral zone) were within the range experienced under natural conditions. In addition, there were differences in Arrhenius breakpoint and endpoint temperatures, showing a trend from higher in tropical animals to lower for temperate animals. Conditions such as acclimation, heat shock and starvation had little or no effect on heart performance. However, a slow increase in temperature induced good regulation of heart rate with noticeable shifts of breakpoints and endpoints for Afrolittorina spp. Lastly, there were differences in the proteome responses between and within Afrolittorina spp. as a function of species, size and treatment. Although both large and small A. knysnaensis had a greater number of protein spots in their proteome than A. africana (though the difference was not significant), the later showed significantly higher differential expression of certain proteins following heat stress. In addition, juveniles of both species displayed greater numbers of protein spots in their proteome than adults. The results indicate a difference in the physiological and biochemical responses (i.e. adaptations) of these snails to temperature, and this seems to relate to differences in biogeography, phylogeny, species identity and ecology. The ability to regulate heart rate is phylogenetically determined, while thresholds and lethal limits correspond to biogeography and species ecology. The proteome seems to correspond to species ecology. The results also indicate that these littorinids can tolerate high temperature stress and in this respect they are well suited to life in the intertidal zones or habitats where temperature and other stresses or conditions are extreme and can change abruptly. However, the limited ability of these snails to acclimate to different temperatures suggests that they are already living close to their tolerance limits with small safety margins or narrow thermal windows and so may be vulnerable to small rises in substratum temperature and/or solar radiation.
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Intertidal patterns and processes : tracking the effects of coastline topography and settlement choice across life stages of the mussels perna perna and mytilus galloprovincialis
- Von der Meden, Charles Eric Otto
- Authors: Von der Meden, Charles Eric Otto
- Date: 2010
- Subjects: Mexilhao mussel -- Behavior -- South Africa Mytilus galloprovincialis -- Behavior -- South Africa Perna -- Behavior -- South Africa Mussels -- Ecology -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5721 , http://hdl.handle.net/10962/d1005407
- Description: Within landscapes, spatial heterogeneity is common and specific landscape features can influence propagule dispersal by wind or water, affecting population connectivity and dynamics. Coastline topographic features, such as bays and headlands, have a variety of biophysical effects on nearshore oceanography, larval transport, retention and supply, and the processes of larval settlement and recruitment. Although this has been demonstrated in several parts of the world, engendering a perception of a general ‘bay effect’, few studies have investigated this generality in a single experiment or region, by replicating at the level of ‘bay’. The Agulhas biogeographic region of the south coast of South Africa is a useful system within which to test for such generality. Using the intertidal mussels Mytilus galloprovincialis and Perna perna as model organisms, patterns of adult distribution were surveyed across four large ‘halfheart’ bays and intervening stretches of open coast, providing replication at the level of ‘bay’ and duplication of ecologically similar species. In support of a general, pervasive influence of bays on intertidal populations, mussel cover was found to be greater in bays than on the open coast for both species, although the effect was strongest for M. galloprovincialis. To explain this adult distribution, settlement, post-settlement mortality and recruitment were examined over 12mo at the same sites, with the prediction that rates of each would favour larger bay populations. Contrary to this, an interaction between month and bay-status was found, with greater settlement and recruitment on the open coast than in bays reflecting extreme settlement and recruitment events at 3 westerly open coast sites during summer. Re-analysis excluding these outliers, revealed the expected effect, of greater settlement and recruitment in bays. While this indicates the broad generality of the bay effect, it highlights exceptions and the need for replication in time and space when examining landscape effects. Measuring post-settlement mortality required testing small-scale settlement behaviour on established and newly deployed settler collectors. It was found that all settlers preferred collectors with biofilm, but that primary settlers avoided conspecific settlers, while secondary settlers were attracted to them. With discrepancies in settler attraction to new and established collectors accounted for, initial (over 2d) and longer-term (over 7d) post-settlement mortality rates were found to be substantial (ca 60 %) for both species. No topographic effect on p-s mortality was evident. Finally, recruit-settler, adult-recruit and interspecies correlations were examined at regional and local scales. Synergistic (or neutral) effects maintained the initial settlement pattern in recruit and adult populations regionally, but not at local scales; striking interspecies correlations suggested the influence of common regional transport processes. Ultimately, the results emphasize the importance of the direction of effects in different life stages and at different spatial scales, and the possibility that antagonistic effects may mask even strong patterns.
- Full Text:
- Authors: Von der Meden, Charles Eric Otto
- Date: 2010
- Subjects: Mexilhao mussel -- Behavior -- South Africa Mytilus galloprovincialis -- Behavior -- South Africa Perna -- Behavior -- South Africa Mussels -- Ecology -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5721 , http://hdl.handle.net/10962/d1005407
- Description: Within landscapes, spatial heterogeneity is common and specific landscape features can influence propagule dispersal by wind or water, affecting population connectivity and dynamics. Coastline topographic features, such as bays and headlands, have a variety of biophysical effects on nearshore oceanography, larval transport, retention and supply, and the processes of larval settlement and recruitment. Although this has been demonstrated in several parts of the world, engendering a perception of a general ‘bay effect’, few studies have investigated this generality in a single experiment or region, by replicating at the level of ‘bay’. The Agulhas biogeographic region of the south coast of South Africa is a useful system within which to test for such generality. Using the intertidal mussels Mytilus galloprovincialis and Perna perna as model organisms, patterns of adult distribution were surveyed across four large ‘halfheart’ bays and intervening stretches of open coast, providing replication at the level of ‘bay’ and duplication of ecologically similar species. In support of a general, pervasive influence of bays on intertidal populations, mussel cover was found to be greater in bays than on the open coast for both species, although the effect was strongest for M. galloprovincialis. To explain this adult distribution, settlement, post-settlement mortality and recruitment were examined over 12mo at the same sites, with the prediction that rates of each would favour larger bay populations. Contrary to this, an interaction between month and bay-status was found, with greater settlement and recruitment on the open coast than in bays reflecting extreme settlement and recruitment events at 3 westerly open coast sites during summer. Re-analysis excluding these outliers, revealed the expected effect, of greater settlement and recruitment in bays. While this indicates the broad generality of the bay effect, it highlights exceptions and the need for replication in time and space when examining landscape effects. Measuring post-settlement mortality required testing small-scale settlement behaviour on established and newly deployed settler collectors. It was found that all settlers preferred collectors with biofilm, but that primary settlers avoided conspecific settlers, while secondary settlers were attracted to them. With discrepancies in settler attraction to new and established collectors accounted for, initial (over 2d) and longer-term (over 7d) post-settlement mortality rates were found to be substantial (ca 60 %) for both species. No topographic effect on p-s mortality was evident. Finally, recruit-settler, adult-recruit and interspecies correlations were examined at regional and local scales. Synergistic (or neutral) effects maintained the initial settlement pattern in recruit and adult populations regionally, but not at local scales; striking interspecies correlations suggested the influence of common regional transport processes. Ultimately, the results emphasize the importance of the direction of effects in different life stages and at different spatial scales, and the possibility that antagonistic effects may mask even strong patterns.
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Effects of marine reserves on the biology of rocky intertidal limpets along the southern coast of South Africa
- Nakin, Motebang Dominic Vincent
- Authors: Nakin, Motebang Dominic Vincent
- Date: 2009
- Subjects: Limpets Marine parks and reserves -- South Africa Marine biology -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5680 , http://hdl.handle.net/10962/d1005365
- Description: Limpets are harvested by people in South Africa, but are selected in terms of species and size. The effects of marine reserves on the biology of commonly exploited (Helcion concolor and Scutellastra longicosta) and rarely exploited species (Cellana capensis and Scutellastra granularis) were investigated on the southeast coast of South Africa at two reserve and two non-reserve sites. For each species, a 4-way nested ANOVA was used to test the effects of month, reserve, site (nested within reserve) and area (nested within site and reserve) on population density, size structure and recruitment of these limpets. The data were collected monthly over 20 months. The overall results indicated a gradient of exploitation among species, S. longicosta was the most heavily exploited species and S. granularis the least exploited species. However, there was also a gradient of exploitation between reserves and non-reserves. Xhora was the most heavily exploited site while Nqabara was less heavily exploited. Of the two reserve sites, Cwebe had more poachers than Dwesa. In most analyses, the month x area (reserve (site)) interaction was significant. However, this was largely an artifact due to comparisons of areas in different sites and significant differences between areas within sites occurred in relatively few months. Densities were greater inside reserves for all species except C. capensis. For S. longicosta and H. concolor this was expected but not for S. granularis and the result possibly reflects its opportunistic exploitation in the absence of the preferred species or indirect effects of reserves. Commonly exploited species and the rarely exploited C. capensis clearly showed greater mean and maximum sizes in reserves but there were month/site (reserve) interactions. Months with significant differences between reserves and nonreserves in both mean and maximum sizes generally occurred more often for commonly exploited than rarely exploited species, but C. capensis showed the strongest reserve effect on maximum size. Interview surveys showed that, although not normally exploited, C. capensis is sometimes mistaken for H. concolor and this suggests that large individuals are unintentionally harvested outside reserves. There were no significant reserve effects on recruitment for any species. Although Xhora had the lowest densities and limpet sizes, it showed the highest recruitment especially for S. longicosta, suggesting that larvae can be transported far from where they are released and settle in non-reserve sites regardless of adult densities. Reserve as a main factor was not significant for the rarely exploited species, but there was a significant month x reserve interaction, with non-reserves having greater GSI values than reserves in most months. Growth rates were examined using individual tagging and cohort analysis. The two techniques gave different results, with individual tagging giving higher growth estimates than cohort analysis. Except for the territorial species S. longicosta, growth was higher in non-reserves and inversely correlated with population density. Mortality estimates using the Cormack-Jolly-Seber model indicated that the rarely exploited species had significantly greater capture probabilities in reserves while no significant reserve effects were observed for the commonly exploited species. Reserve effects on survival probability were significant only for S. longicosta, with reserves being greater than nonreserves and no significant effects for any other species. Enhanced survival in reserves was attributed to the effects of human exploitation. In theory, marine protected areas show increases in densities, sizes and reproductive output of exploited species, but the present results revealed that the efficacy of reserves depends on the status of the species, not only whether it is exploited or non-exploited, but also whether it is territorial.
- Full Text:
- Authors: Nakin, Motebang Dominic Vincent
- Date: 2009
- Subjects: Limpets Marine parks and reserves -- South Africa Marine biology -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5680 , http://hdl.handle.net/10962/d1005365
- Description: Limpets are harvested by people in South Africa, but are selected in terms of species and size. The effects of marine reserves on the biology of commonly exploited (Helcion concolor and Scutellastra longicosta) and rarely exploited species (Cellana capensis and Scutellastra granularis) were investigated on the southeast coast of South Africa at two reserve and two non-reserve sites. For each species, a 4-way nested ANOVA was used to test the effects of month, reserve, site (nested within reserve) and area (nested within site and reserve) on population density, size structure and recruitment of these limpets. The data were collected monthly over 20 months. The overall results indicated a gradient of exploitation among species, S. longicosta was the most heavily exploited species and S. granularis the least exploited species. However, there was also a gradient of exploitation between reserves and non-reserves. Xhora was the most heavily exploited site while Nqabara was less heavily exploited. Of the two reserve sites, Cwebe had more poachers than Dwesa. In most analyses, the month x area (reserve (site)) interaction was significant. However, this was largely an artifact due to comparisons of areas in different sites and significant differences between areas within sites occurred in relatively few months. Densities were greater inside reserves for all species except C. capensis. For S. longicosta and H. concolor this was expected but not for S. granularis and the result possibly reflects its opportunistic exploitation in the absence of the preferred species or indirect effects of reserves. Commonly exploited species and the rarely exploited C. capensis clearly showed greater mean and maximum sizes in reserves but there were month/site (reserve) interactions. Months with significant differences between reserves and nonreserves in both mean and maximum sizes generally occurred more often for commonly exploited than rarely exploited species, but C. capensis showed the strongest reserve effect on maximum size. Interview surveys showed that, although not normally exploited, C. capensis is sometimes mistaken for H. concolor and this suggests that large individuals are unintentionally harvested outside reserves. There were no significant reserve effects on recruitment for any species. Although Xhora had the lowest densities and limpet sizes, it showed the highest recruitment especially for S. longicosta, suggesting that larvae can be transported far from where they are released and settle in non-reserve sites regardless of adult densities. Reserve as a main factor was not significant for the rarely exploited species, but there was a significant month x reserve interaction, with non-reserves having greater GSI values than reserves in most months. Growth rates were examined using individual tagging and cohort analysis. The two techniques gave different results, with individual tagging giving higher growth estimates than cohort analysis. Except for the territorial species S. longicosta, growth was higher in non-reserves and inversely correlated with population density. Mortality estimates using the Cormack-Jolly-Seber model indicated that the rarely exploited species had significantly greater capture probabilities in reserves while no significant reserve effects were observed for the commonly exploited species. Reserve effects on survival probability were significant only for S. longicosta, with reserves being greater than nonreserves and no significant effects for any other species. Enhanced survival in reserves was attributed to the effects of human exploitation. In theory, marine protected areas show increases in densities, sizes and reproductive output of exploited species, but the present results revealed that the efficacy of reserves depends on the status of the species, not only whether it is exploited or non-exploited, but also whether it is territorial.
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Hierarchical spatial structure and levels of resolution of intertidal grazing and their consequences on predictability and stability at small scales
- Authors: Diaz Diaz, Eliecer Rodrigo
- Date: 2009
- Subjects: Marine algae -- Effect of grazing on Shorelines -- South Africa Zostera marina Intertidal ecology Spatial analysis (Statistics)
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5707 , http://hdl.handle.net/10962/d1005393
- Description: The aim of this research was to assess three hierarchical aspects of alga-grazer interactions in intertidal communities on a small scale: spatial heterogeneity, grazing effects and spatial stability in grazing effects. First, using semivariograms and cross-semivariograms I observed hierarchical spatial patterns in most algal groups and in grazers. However, these patterns varied with the level on the shore and between shores, suggesting that either human exploitation or wave exposure can be a source of variability. Second, grazing effects were studied using manipulative experiments at different levels on the shore. These revealed significant effects of grazing on the low shore and in tidal pools. Additionally, using a transect of grazer exclusions across the shore, I observed unexpected hierarchical patchiness in the strength of grazing, rather than zonation in its effects. This patchiness varied in time due to different biotic and abiotic factors. In a separate experiment, the effect of mesograzers effects were studied in the upper eulittoral zone under four conditions: burnt open rock (BOR), burnt pools (Bpool), non-burnt open rock (NBOR) and non-burnt pools (NBpool). Additionally, I tested spatial stability in the effects of grazing in consecutive years, using the same plots. I observed great spatial variability in the effects of grazing, but this variability was spatially stable in Bpools and NBOR, meaning deterministic and significant grazing effects in consecutive years on the same plots. Both the significance in grazing effects and spatial stability depended on the level of resolution (species, functional, biomass) at which the algal assemblage was evaluated, suggesting hierarchical variability. In order to be able to predict spatial variability in the effects of grazers in the upper eulittoral zone using biotic and abiotic micro- and macrofactors, a conceptual model was proposed, based on data from several multiple-regressions. This linked the interactions among three elements: idiosyncratic heterogeneity, micro and macrofactors. This suggests that spatial variability can be a product of these factors, while spatial stability can be caused by the same or different combinations of factors. In conclusion, grazing and other ecological phenomena must be studied hierarchically, not only through spatiotemporal scales, but also at different levels of resolution, as these also influence our perception of patterns.
- Full Text:
- Authors: Diaz Diaz, Eliecer Rodrigo
- Date: 2009
- Subjects: Marine algae -- Effect of grazing on Shorelines -- South Africa Zostera marina Intertidal ecology Spatial analysis (Statistics)
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5707 , http://hdl.handle.net/10962/d1005393
- Description: The aim of this research was to assess three hierarchical aspects of alga-grazer interactions in intertidal communities on a small scale: spatial heterogeneity, grazing effects and spatial stability in grazing effects. First, using semivariograms and cross-semivariograms I observed hierarchical spatial patterns in most algal groups and in grazers. However, these patterns varied with the level on the shore and between shores, suggesting that either human exploitation or wave exposure can be a source of variability. Second, grazing effects were studied using manipulative experiments at different levels on the shore. These revealed significant effects of grazing on the low shore and in tidal pools. Additionally, using a transect of grazer exclusions across the shore, I observed unexpected hierarchical patchiness in the strength of grazing, rather than zonation in its effects. This patchiness varied in time due to different biotic and abiotic factors. In a separate experiment, the effect of mesograzers effects were studied in the upper eulittoral zone under four conditions: burnt open rock (BOR), burnt pools (Bpool), non-burnt open rock (NBOR) and non-burnt pools (NBpool). Additionally, I tested spatial stability in the effects of grazing in consecutive years, using the same plots. I observed great spatial variability in the effects of grazing, but this variability was spatially stable in Bpools and NBOR, meaning deterministic and significant grazing effects in consecutive years on the same plots. Both the significance in grazing effects and spatial stability depended on the level of resolution (species, functional, biomass) at which the algal assemblage was evaluated, suggesting hierarchical variability. In order to be able to predict spatial variability in the effects of grazers in the upper eulittoral zone using biotic and abiotic micro- and macrofactors, a conceptual model was proposed, based on data from several multiple-regressions. This linked the interactions among three elements: idiosyncratic heterogeneity, micro and macrofactors. This suggests that spatial variability can be a product of these factors, while spatial stability can be caused by the same or different combinations of factors. In conclusion, grazing and other ecological phenomena must be studied hierarchically, not only through spatiotemporal scales, but also at different levels of resolution, as these also influence our perception of patterns.
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A stable isotope approach to trophic ecology resolving food webs in intertidal ecosystems
- Authors: Hill, Jaclyn Marie
- Date: 2008
- Subjects: Stable isotopes Food chains (Ecology) Stable isotopes in ecological research Intertidal ecology Mussels -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5771 , http://hdl.handle.net/10962/d1005459
- Description: There are broad differences in regional oceanography and primary production around the South African coast, which we might expect to give rise to major differences in trophic pathways. δ⁻¹³C and δ⁻¹⁵N isotopic ratios of suspended particulate matter (SPM), mussels, various intertidal consumers and common macroalgae along the South African coastline were explored using stable isotope analysis to investigate biogeographic and temporal variability of isotopic signatures of marine intertidal consumers and their food sources around the coast of South Africa, with a focus on evaluating the dependence of intertidal mussels on phytoplankton and macroalgal-derived organic carbon. Isotopic equilibration rates of four mussel tissues were determined through laboratory feeding experiments, which established that adductor tissue had the slowest isotopic turnover rate, and was subsequently used as an indication of overall mussel diet. Biogeographic, temporal and nearshore/offshore trends of isotopic ratios of SPM were investigated along 10km transects perpendicular to the coast and SPM exhibited overall trends of carbon depletion when moving from west to east along the coastline and from nearshore to offshore water, in both cases suggesting a shift from macrophyte detritus to a phytoplankton signature. δ⁻¹³C signatures of SPM also revealed temporal and biogeographic variation that had strong ties to local oceanography, being closely correlated to regional hydrographic features and tidal influences. Mixing models indicated filter feeders demonstrated over 50% dependence on nearshore SPM for organic carbon and it was possible to categorize them into geographic groups based on their carbon and nitrogen signatures, suggesting biogeographic shifts in resources. Biogeographic shifts in diet were also seen in some grazers. Difficulties in relating macroalgae to mussel diet led to investigations into the isotopic changes associated with macroalgal decomposition. Variation in photosynthetic fractionation, leaching and microbial mineralization are believed to have resulted from species-specific patterns of degradation. Although the strong links between carbon signatures and local oceanography indicate that stable isotope analysis is a powerful tool for the study of water mixing and coastal hydrography in relation to food-web analyses, substantial variation in fractionation of primary consumers, along with different periods of time integration between consumers and their food sources must be considered in future studies, to resolve trophic links in marine food webs successfully.
- Full Text:
- Authors: Hill, Jaclyn Marie
- Date: 2008
- Subjects: Stable isotopes Food chains (Ecology) Stable isotopes in ecological research Intertidal ecology Mussels -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5771 , http://hdl.handle.net/10962/d1005459
- Description: There are broad differences in regional oceanography and primary production around the South African coast, which we might expect to give rise to major differences in trophic pathways. δ⁻¹³C and δ⁻¹⁵N isotopic ratios of suspended particulate matter (SPM), mussels, various intertidal consumers and common macroalgae along the South African coastline were explored using stable isotope analysis to investigate biogeographic and temporal variability of isotopic signatures of marine intertidal consumers and their food sources around the coast of South Africa, with a focus on evaluating the dependence of intertidal mussels on phytoplankton and macroalgal-derived organic carbon. Isotopic equilibration rates of four mussel tissues were determined through laboratory feeding experiments, which established that adductor tissue had the slowest isotopic turnover rate, and was subsequently used as an indication of overall mussel diet. Biogeographic, temporal and nearshore/offshore trends of isotopic ratios of SPM were investigated along 10km transects perpendicular to the coast and SPM exhibited overall trends of carbon depletion when moving from west to east along the coastline and from nearshore to offshore water, in both cases suggesting a shift from macrophyte detritus to a phytoplankton signature. δ⁻¹³C signatures of SPM also revealed temporal and biogeographic variation that had strong ties to local oceanography, being closely correlated to regional hydrographic features and tidal influences. Mixing models indicated filter feeders demonstrated over 50% dependence on nearshore SPM for organic carbon and it was possible to categorize them into geographic groups based on their carbon and nitrogen signatures, suggesting biogeographic shifts in resources. Biogeographic shifts in diet were also seen in some grazers. Difficulties in relating macroalgae to mussel diet led to investigations into the isotopic changes associated with macroalgal decomposition. Variation in photosynthetic fractionation, leaching and microbial mineralization are believed to have resulted from species-specific patterns of degradation. Although the strong links between carbon signatures and local oceanography indicate that stable isotope analysis is a powerful tool for the study of water mixing and coastal hydrography in relation to food-web analyses, substantial variation in fractionation of primary consumers, along with different periods of time integration between consumers and their food sources must be considered in future studies, to resolve trophic links in marine food webs successfully.
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A genetic and ecophysiological comparison of co-occuring indigenous (Perna perna) and invasive (Mytilus galloprovincialis) intertidal mussels
- Authors: Zardi, Gerardo Ivan
- Date: 2006
- Subjects: Perna -- South Africa Mytilus galloprovincialis -- South Africa Mussels -- South Africa Mytilidae -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5613 , http://hdl.handle.net/10962/d1003066
- Description: The Mediterranean mussel Mytilus galloprovincialis is the most successful marine invasive species in South Africa. Its presence has had significant ecological consequences on the intertidal communities of the west coast. On the south coast, M galloprovincialis co-exists and competes with the indigenous intertidal mussel Perna perna in the lower balanoid zone, where they show partial habitat segregation. The upper and the lower mussel zones are dominated by M. galloprovincialis and P. perna respectively while they co-occur in the mid zone. In this thesis M. galloprovincialis and P. perna are compared in terms of their population genetics and their ecophysiology. The success of an invader depends on its ability to react to new environmental factors, especially when compared to indigenous species. The distribution and diversity of intertidal species throughout the world are strongly influenced by periodic sand inundation and hydrodynamic stress. Occupying the lower intertidal zone, P. perna is more strongly influenced by sand (burial and sand in suspension) than M. galioprovincialis. Despite this, P. perna is more vulnerable to the effects of sand, showing higher mortality rates under experimental conditions in both the laboratory and the field. M. galioprovincialis has longer labial palps than P. perna, indicating a better ability to sort particles. This, and a higher tolerance to anoxia, explains its lower mortality rates when exposed to burial or suspended sand. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. The ability of a mussel to withstand wave-generated hydrodynamic stress depends mainly on its byssal attachment strength. The higher attachment strength of P. perna compared to M. galioprovincialis and of solitary mussels compared to mussels living within a bed (bed mussels) can be explained by more and thicker byssal threads. M galloprovincialis also has a wider shell, is subjected to higher hydrodynamic loads than P. perna and shows a higher theoretical probability of dislodgement, this is borne out under field conditions. The attachment strength of both species increased from higher to lower shore, in parallel to a gradient of a stronger wave action. Monthly measurements showed that P. perna is always more strongly attached than M. galloprovincialis and revealed seasonal fluctuations of attachment strength for both species in response to wave height. The gonad index of both species was negatively cross-correlated with attachment strength. The results are discussed in the context of the evolutionary strategy of the alien mussel, which directs most of its energy to fast growth and high reproductive output, apparently at the cost of reduced attachment strength. This raises the prediction that its invasive impact will be more pronounced at sites subjected to low or moderate wave action at heavily exposed sites. The potential of a species for invasion is also determined by the ability of the invader to disperse. Population genetics provide indirect information about dispersal through a direct measurement of gene flow. The low genetic divergence (measured as mtDNA) of M. galloprovincialis confirms its recent arrival in South Africa. In contrast, the population genetics structure of P. perna revealed strong divergence on the south-east coast, resulting in a western lineage (straddling the distributional gap of the Benguela System), and an eastern lineage, with an overlap region of the two on the south coast between Kenton-on-Sea and Haga Haga. This genetic disjunction may be caused by Agulhas Current acting as an oceanographic barrier to larval dispersal, or by different environmental selective forces acting on regional populations. Over the last ten years, M. galloprovincialis has shown a decrease or cessation of its spread to the east in exactly the region of the genetic disjunction in P. perna, again suggesting either an oceanographic barrier to larval dispersal, or increasing selection driven by sharp gradients in environmental conditions.
- Full Text:
- Authors: Zardi, Gerardo Ivan
- Date: 2006
- Subjects: Perna -- South Africa Mytilus galloprovincialis -- South Africa Mussels -- South Africa Mytilidae -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5613 , http://hdl.handle.net/10962/d1003066
- Description: The Mediterranean mussel Mytilus galloprovincialis is the most successful marine invasive species in South Africa. Its presence has had significant ecological consequences on the intertidal communities of the west coast. On the south coast, M galloprovincialis co-exists and competes with the indigenous intertidal mussel Perna perna in the lower balanoid zone, where they show partial habitat segregation. The upper and the lower mussel zones are dominated by M. galloprovincialis and P. perna respectively while they co-occur in the mid zone. In this thesis M. galloprovincialis and P. perna are compared in terms of their population genetics and their ecophysiology. The success of an invader depends on its ability to react to new environmental factors, especially when compared to indigenous species. The distribution and diversity of intertidal species throughout the world are strongly influenced by periodic sand inundation and hydrodynamic stress. Occupying the lower intertidal zone, P. perna is more strongly influenced by sand (burial and sand in suspension) than M. galioprovincialis. Despite this, P. perna is more vulnerable to the effects of sand, showing higher mortality rates under experimental conditions in both the laboratory and the field. M. galioprovincialis has longer labial palps than P. perna, indicating a better ability to sort particles. This, and a higher tolerance to anoxia, explains its lower mortality rates when exposed to burial or suspended sand. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. The ability of a mussel to withstand wave-generated hydrodynamic stress depends mainly on its byssal attachment strength. The higher attachment strength of P. perna compared to M. galioprovincialis and of solitary mussels compared to mussels living within a bed (bed mussels) can be explained by more and thicker byssal threads. M galloprovincialis also has a wider shell, is subjected to higher hydrodynamic loads than P. perna and shows a higher theoretical probability of dislodgement, this is borne out under field conditions. The attachment strength of both species increased from higher to lower shore, in parallel to a gradient of a stronger wave action. Monthly measurements showed that P. perna is always more strongly attached than M. galloprovincialis and revealed seasonal fluctuations of attachment strength for both species in response to wave height. The gonad index of both species was negatively cross-correlated with attachment strength. The results are discussed in the context of the evolutionary strategy of the alien mussel, which directs most of its energy to fast growth and high reproductive output, apparently at the cost of reduced attachment strength. This raises the prediction that its invasive impact will be more pronounced at sites subjected to low or moderate wave action at heavily exposed sites. The potential of a species for invasion is also determined by the ability of the invader to disperse. Population genetics provide indirect information about dispersal through a direct measurement of gene flow. The low genetic divergence (measured as mtDNA) of M. galloprovincialis confirms its recent arrival in South Africa. In contrast, the population genetics structure of P. perna revealed strong divergence on the south-east coast, resulting in a western lineage (straddling the distributional gap of the Benguela System), and an eastern lineage, with an overlap region of the two on the south coast between Kenton-on-Sea and Haga Haga. This genetic disjunction may be caused by Agulhas Current acting as an oceanographic barrier to larval dispersal, or by different environmental selective forces acting on regional populations. Over the last ten years, M. galloprovincialis has shown a decrease or cessation of its spread to the east in exactly the region of the genetic disjunction in P. perna, again suggesting either an oceanographic barrier to larval dispersal, or increasing selection driven by sharp gradients in environmental conditions.
- Full Text:
Habitat segregation in competing species of intertidal mussels in South Africa
- Authors: Bownes, Sarah
- Date: 2006
- Subjects: Mussels -- South Africa Mussels -- Growth Mussels -- Habitat -- South Africa Mussels -- Larvae Perna -- South Africa Mytilus galloprovincialis -- South Africa Choromytilus meridionalis -- South Africa Mytilidae -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5706 , http://hdl.handle.net/10962/d1005392
- Description: Mytilus galloprovincialis is invasive on rocky shores on the west coast of South Africa where it has become the dominant intertidal mussel. The success of this species on the west coast and its superior competitive abilities, have led to concern that it may become invasive on the south coast at the expense of the indigenous mussel Perna perna. On shores where these species co-occur, there appears to be habitat segregation among zones occupied by mussels. M.galloprovincialis dominates the high-shore and P.perna the low-shore, with a mixed zone at mid-shore level. This study examined the factors responsible for these differences in distribution and abundance. The study was conducted in Plettenberg Bay and Tsitsikamma (70km apart) on the south coast of South Africa. Each site included two randomly selected locations (300-400m apart). A third mussel species, Choromytilus meridionalis, is found in large numbers at the sand/rock interface at one location in Plettenberg Bay. Aspects of settlement, recruitment, growth and mortality of juvenile and adult mussels were examined at different tidal heights at each site. Quantitative analysis of mussel population structure at these sites supported the initial observation of vertical habitat segregation. Post-larvae were identified to species and this was confirmed using hinge morphology and mitochondrial DNA analysis. Size at settlement was determined for each species to differentiate between primary and secondary settlement. Adult distribution of C.meridionalis was primarily determined by settlement, which was highly selective in this species. Settlement, recruitment and growth of P.perna decreased with increasing tidal height, while post-settlement mortality and adult mortality increased higher upshore. Thus all aspects of P.perna’s life history contribute to the adult distribution of this species. Presumably, the abundance of P.perna on the high-shore is initially limited by recruitment while those that survive remain prone to elimination throughout adulthood. M.galloprovincialis displayed the same patterns of settlement and recruitment as P.perna. However, post-settlement mortality in this species was consistently low in the low and high zones. Juvenile growth also decreased upshore, suggesting that M.galloprovincialis may be able to maintain high densities on the high-shore through the persistence of successive settlements of slow-growing individuals. The low cover of M.galloprovincialis on the lowshore appeared to be determined by adult interactions. M.galloprovincialis experienced significantly higher adult mortality rates than P.perna in this zone. There were seasonal variations in the competitive advantages enjoyed by each species through growth, recruitment or mortality on the low-shore. In summer, P.perna had higher recruitment rates, faster growth and lower mortality rates, while M.galloprovincialis had slightly higher recruitment rates and faster growth rates in winter. P.perna is a warm water species while M.galloprovincialis thrives on the cold-temperate west coast of South Africa. Therefore both species appear to be at the edge of their optimal temperature regimes on the south coast, which may explain the seasonal advantages of each. Nevertheless, P.perna has maintained spatial dominance on the low-shore suggesting that it may ultimately be the winner in competition between these species. M.galloprovincialis appears to have a refuge from competition with P.perna on the high-shore due to its greater tolerance of desiccation stress, while being competitively excluded from the low-shore. Warm water temperatures coupled with poor recruitment rates at most sites may limit the success of M.galloprovincialis on this coast.
- Full Text:
- Authors: Bownes, Sarah
- Date: 2006
- Subjects: Mussels -- South Africa Mussels -- Growth Mussels -- Habitat -- South Africa Mussels -- Larvae Perna -- South Africa Mytilus galloprovincialis -- South Africa Choromytilus meridionalis -- South Africa Mytilidae -- South Africa
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5706 , http://hdl.handle.net/10962/d1005392
- Description: Mytilus galloprovincialis is invasive on rocky shores on the west coast of South Africa where it has become the dominant intertidal mussel. The success of this species on the west coast and its superior competitive abilities, have led to concern that it may become invasive on the south coast at the expense of the indigenous mussel Perna perna. On shores where these species co-occur, there appears to be habitat segregation among zones occupied by mussels. M.galloprovincialis dominates the high-shore and P.perna the low-shore, with a mixed zone at mid-shore level. This study examined the factors responsible for these differences in distribution and abundance. The study was conducted in Plettenberg Bay and Tsitsikamma (70km apart) on the south coast of South Africa. Each site included two randomly selected locations (300-400m apart). A third mussel species, Choromytilus meridionalis, is found in large numbers at the sand/rock interface at one location in Plettenberg Bay. Aspects of settlement, recruitment, growth and mortality of juvenile and adult mussels were examined at different tidal heights at each site. Quantitative analysis of mussel population structure at these sites supported the initial observation of vertical habitat segregation. Post-larvae were identified to species and this was confirmed using hinge morphology and mitochondrial DNA analysis. Size at settlement was determined for each species to differentiate between primary and secondary settlement. Adult distribution of C.meridionalis was primarily determined by settlement, which was highly selective in this species. Settlement, recruitment and growth of P.perna decreased with increasing tidal height, while post-settlement mortality and adult mortality increased higher upshore. Thus all aspects of P.perna’s life history contribute to the adult distribution of this species. Presumably, the abundance of P.perna on the high-shore is initially limited by recruitment while those that survive remain prone to elimination throughout adulthood. M.galloprovincialis displayed the same patterns of settlement and recruitment as P.perna. However, post-settlement mortality in this species was consistently low in the low and high zones. Juvenile growth also decreased upshore, suggesting that M.galloprovincialis may be able to maintain high densities on the high-shore through the persistence of successive settlements of slow-growing individuals. The low cover of M.galloprovincialis on the lowshore appeared to be determined by adult interactions. M.galloprovincialis experienced significantly higher adult mortality rates than P.perna in this zone. There were seasonal variations in the competitive advantages enjoyed by each species through growth, recruitment or mortality on the low-shore. In summer, P.perna had higher recruitment rates, faster growth and lower mortality rates, while M.galloprovincialis had slightly higher recruitment rates and faster growth rates in winter. P.perna is a warm water species while M.galloprovincialis thrives on the cold-temperate west coast of South Africa. Therefore both species appear to be at the edge of their optimal temperature regimes on the south coast, which may explain the seasonal advantages of each. Nevertheless, P.perna has maintained spatial dominance on the low-shore suggesting that it may ultimately be the winner in competition between these species. M.galloprovincialis appears to have a refuge from competition with P.perna on the high-shore due to its greater tolerance of desiccation stress, while being competitively excluded from the low-shore. Warm water temperatures coupled with poor recruitment rates at most sites may limit the success of M.galloprovincialis on this coast.
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Genetic variation within two pulmonate limpet species, Siphonaria capensis and S. serrata along the South African coast
- Authors: Seaman, Jennifer Ann
- Date: 2003
- Subjects: Limpets -- South Africa Pulmonata Siphonaria Limpets -- Genetics
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5800 , http://hdl.handle.net/10962/d1006155
- Description: Dispersal rates are known to play a fundamental role in establishing the genetic structure found within a species. Dispersal in marine benthic invertebrates is largely dependent on life history strategy. The presence of pelagic larvae will potentially result in high dispersal within a species, while direct development limits dispersal. This study used two intertidal pulmonate limpet species to investigate the relationship between dispersal potential and genetic structure of populations. Siphonaria capensis produces benthic egg masses, which release pelagic larvae. These float in ocean currents before settling and metamorphosing into adults. S. serrata reproduces by direct development. The larvae hatch as completely metamorphosed juveniles after developing in an egg case not far from the parent. The high dispersal capacity of S. capensis pelagic larvae should result in high levels of within-population variation since individuals within a particular population may originate from several different populations. This should lead to high levels of gene flow along the coast and to low genetic differentiation among populations. On the other hand, the low dispersal potential of S. serrata should cause low levels of genetic variation within populations, as relatives will remain close to one another. Low levels of gene flow and high genetic differentiation among populations are expected in this species. In dealing with the relationship between life history strategy and population genetic structure in these two species, a series of indirect techniques was utilised. Multivariate analyses of the morphological variation within each species showed that variation in size was similar between the two species and largely governed by environmental factors. Variation in shell shape was largely under genetic control and supported the predictions made for each species. Variation in total proteins was relatively uninformative in terms of examining the differences between the two species. Allozyme analysis and mtDNA sequencing clearly showed differences between the species in terms of their population genetic structure. High levels of gene flow were found within S. capensis. This was strongly influenced by ocean currents, with the close inshore Agulhas current along the Transkei coast contributing to high levels of dispersal and hence gene flow. Low levels of gene flow occurred within S. serrata, resulting in low within-population variation and high among-population differentiation.
- Full Text:
- Authors: Seaman, Jennifer Ann
- Date: 2003
- Subjects: Limpets -- South Africa Pulmonata Siphonaria Limpets -- Genetics
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5800 , http://hdl.handle.net/10962/d1006155
- Description: Dispersal rates are known to play a fundamental role in establishing the genetic structure found within a species. Dispersal in marine benthic invertebrates is largely dependent on life history strategy. The presence of pelagic larvae will potentially result in high dispersal within a species, while direct development limits dispersal. This study used two intertidal pulmonate limpet species to investigate the relationship between dispersal potential and genetic structure of populations. Siphonaria capensis produces benthic egg masses, which release pelagic larvae. These float in ocean currents before settling and metamorphosing into adults. S. serrata reproduces by direct development. The larvae hatch as completely metamorphosed juveniles after developing in an egg case not far from the parent. The high dispersal capacity of S. capensis pelagic larvae should result in high levels of within-population variation since individuals within a particular population may originate from several different populations. This should lead to high levels of gene flow along the coast and to low genetic differentiation among populations. On the other hand, the low dispersal potential of S. serrata should cause low levels of genetic variation within populations, as relatives will remain close to one another. Low levels of gene flow and high genetic differentiation among populations are expected in this species. In dealing with the relationship between life history strategy and population genetic structure in these two species, a series of indirect techniques was utilised. Multivariate analyses of the morphological variation within each species showed that variation in size was similar between the two species and largely governed by environmental factors. Variation in shell shape was largely under genetic control and supported the predictions made for each species. Variation in total proteins was relatively uninformative in terms of examining the differences between the two species. Allozyme analysis and mtDNA sequencing clearly showed differences between the species in terms of their population genetic structure. High levels of gene flow were found within S. capensis. This was strongly influenced by ocean currents, with the close inshore Agulhas current along the Transkei coast contributing to high levels of dispersal and hence gene flow. Low levels of gene flow occurred within S. serrata, resulting in low within-population variation and high among-population differentiation.
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The role of microzooplankton in carbon cycling in the Southern Ocean
- Authors: Froneman, Pierre William
- Date: 1996
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:21066 , http://hdl.handle.net/10962/6234
- Description: A 3-year study was carried out on the role of microzooplankton in carbon cycling in the south Atlantic and the Atlantic sector of the Southern Ocean. Microzooplankton grazing impact on phytoplankton was estimated during austral summer and winter employing the dilution technique. Carnivory by larger zooplankton on microzooplankton during summer was estimated using in vitro incubations. Microzooplankton assemblages were always dominated by protozoans comprising ciliates and dinoflagellates. In the ( 20 um chlorophyll fraction, microzooplankton grazing was sufficient to control the growth of the nano- and picophytoplankton suggesting that, where larger microphytoplankton cells dominate, micro zooplankton maintain the background concentrations of the nano- and picophytoplankton. During winter, when small nano- and picophytoplankton cells dominate total chlorophyll concentrations, the microzooplankton grazing impact on phytoplankton is dramatically increased. Microzooplankton removed on average 37% of the initial phytoplankton stock or 70% of the daily phytoplankton production. These results suggest that in winter, micro zooplankton are the main sink for phytoplankton production. Carnivory experiments conducted with selected meso- (copepods) and macro zooplankton (euphausiids and tunicates) showed that all species examined consumed micro zooplankton in the presence of substantial chlorophyll concentrations. Microzooplankton can, therefore, be regarded as trophic intermediates between bacterioplankton, small phytoplankton cells and larger zooplankton species in the Southern Ocean. The results of this investigation suggest a spatiotemporal shift in efficiency of the biological pump mediated by changes in the size composition of the phytoplankton assemblages. South of the Antarctic Polar Front (APF) large IV microphytoplankton cells dominate the summer chlorophyll biomass, suggesting that larger zooplankton grazers represent the main sink for phytoplankton production. Under these conditions, carbon flux to the interior of the ocean will be high due to diel vertical migrations by grazers and the production of large, fast sinking faecal pellets. The sedimentation of large phytoplankton cells also contributes to flux. In the permanently open waters south of the APF and throughout the Southern Ocean during winter, small phytoplankton cells dominate total chlorophyll, resulting in the microbial loop being the main sink for phytoplankton production. The close coupling between the micro zooplankton and the microbial loop dramatically reduces the transfer of organic carbon from the surface layers to depth. Carnivory by metazoans on microzooplankton may reduce the high grazing impact of micro zooplankton and, may also represent an important source of carbon flux originating from the microbial loop.
- Full Text:
- Authors: Froneman, Pierre William
- Date: 1996
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:21066 , http://hdl.handle.net/10962/6234
- Description: A 3-year study was carried out on the role of microzooplankton in carbon cycling in the south Atlantic and the Atlantic sector of the Southern Ocean. Microzooplankton grazing impact on phytoplankton was estimated during austral summer and winter employing the dilution technique. Carnivory by larger zooplankton on microzooplankton during summer was estimated using in vitro incubations. Microzooplankton assemblages were always dominated by protozoans comprising ciliates and dinoflagellates. In the ( 20 um chlorophyll fraction, microzooplankton grazing was sufficient to control the growth of the nano- and picophytoplankton suggesting that, where larger microphytoplankton cells dominate, micro zooplankton maintain the background concentrations of the nano- and picophytoplankton. During winter, when small nano- and picophytoplankton cells dominate total chlorophyll concentrations, the microzooplankton grazing impact on phytoplankton is dramatically increased. Microzooplankton removed on average 37% of the initial phytoplankton stock or 70% of the daily phytoplankton production. These results suggest that in winter, micro zooplankton are the main sink for phytoplankton production. Carnivory experiments conducted with selected meso- (copepods) and macro zooplankton (euphausiids and tunicates) showed that all species examined consumed micro zooplankton in the presence of substantial chlorophyll concentrations. Microzooplankton can, therefore, be regarded as trophic intermediates between bacterioplankton, small phytoplankton cells and larger zooplankton species in the Southern Ocean. The results of this investigation suggest a spatiotemporal shift in efficiency of the biological pump mediated by changes in the size composition of the phytoplankton assemblages. South of the Antarctic Polar Front (APF) large IV microphytoplankton cells dominate the summer chlorophyll biomass, suggesting that larger zooplankton grazers represent the main sink for phytoplankton production. Under these conditions, carbon flux to the interior of the ocean will be high due to diel vertical migrations by grazers and the production of large, fast sinking faecal pellets. The sedimentation of large phytoplankton cells also contributes to flux. In the permanently open waters south of the APF and throughout the Southern Ocean during winter, small phytoplankton cells dominate total chlorophyll, resulting in the microbial loop being the main sink for phytoplankton production. The close coupling between the micro zooplankton and the microbial loop dramatically reduces the transfer of organic carbon from the surface layers to depth. Carnivory by metazoans on microzooplankton may reduce the high grazing impact of micro zooplankton and, may also represent an important source of carbon flux originating from the microbial loop.
- Full Text:
Dispersal, settlement and recruitment : their influence on the population dynamics of intertidal mussels
- Authors: Phillips, Tracey Elizabeth
- Date: 1995
- Subjects: Perna -- South Africa -- Eastern Cape -- Algoa bay Mytilus galloprovincialis -- South Africa -- Eastern Cape -- Algoa bay Mytilidae -- South Africa -- Eastern Cape -- Algoa bay
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5663 , http://hdl.handle.net/10962/d1005347
- Description: Recruitment of planktonic larvae into sedentary benthic populations regulates the population dynamics of marine invertebrates. The processes controlling recruitment, however, are poorly understood, and recruitment remains largely unpredictable, which complicates management of exploited shellfish resources. The mussels Perna perna, Choromytilus meridionalis and Mytilus galloprovincialis, found on the south coast of southern Africa, have planktonic larvae and sedentary adult stages. This thesis examines dispersal, settlement and early post-settlement growth and mortality, and their effect on recruitment and demography of intertidal mussel populations in the region of Algoa Bay on the south coast of southern Africa. Temporal and spatial variation in the body mass, density and size structure of mussels, the distribution of bivalve larvae on plankton grids in the nearshore zone and the distribution of a recently introduced invasive mussel, Mytilus galloprovincialis, were examined between 1989 and 1992. Furthermore, data on hourly or daily changes in wind strength and direction, air and sea surface temperatures and low and high tide levels in the study region, were obtained. There were 3-4 peaks in spawning (characterised by an abrupt decline in weight) and settlement activity annually. These peaks varied in exact timing, intensity and duration between sites and over time. However, at a site, spawning was followed by settlement 4-8 weeks later, and there was a significant (P < 0.05) direct correlation between spawning intensity prior to the appearance of a new cohort and the cohort density (settlement intensity). The stochastic spatial and temporal variation in breeding activity was superimposed on a more general pattern of a higher intensity of spawning and settlement in Algoa Bay than on the open coast, and a higher settlement intensity on coastal sandstone shores than on dune rock shores. Spawning was more frequent in winter and spring, and the probability of spawning and settlement peaked around the spring and autumn equinox, if temperature and wind conditions were suitable. Larval behaviour had little effect on their dispersal in the well-mixed nearshore region. Larvae were passively dispersed by currents, and their dispersal range and direction depended on prevailing winds and local topography. The sharp decline in density of recruit and adult M. galloprovincialis with increasing distance from the point of introduction, showed that some larvae were carried by wind generated currents over moderately long distances (-100 km). However, since most (76 %) M. galloprovincialis recruited within 4 km of the parent population, it is possible that larvae become trapped in small gullies and crevices around rocky shores, and have a limited dispersal range. This could explain the link between local patterns of spawning and settlement. The distribution and abundance of settlers on the shore was influenced by larval behaviour and the availability of settlement, substrata. Larvae preferred to settle primarily on foliose coralline algae and migrate to the adult mussel bed when they were larger (0.60-7 mm), but larvae also settled directly on adult mussels, possibly because the amount of coralline algae was limited. Both direct and secondary settlement were considered to be important in maintaining mussel populations since the rate of settlement was low(generally < 60 000.m-2). Cohort analyses showed that prior to maturity post-settlement growth (- 30 mm in 10 months) and mortality rates (60-100%) were high, but varied. When settlement intensity was low this variability uncoupled the relationship between spawning and recruitment intensity. Multiple regression analysis showed that together reproductive effort (gamete output), settlement intensity, growth and mortality prior to maturity, accounted for 76 % of the variance in recruitment into mature adult populations. The low settlement rate coupled with the short life span of mussels « 3 years), meant that populations underwent marked spatial and temporal variations in structure and abundance as settlement intensity varied, but there were consistent general differences between mussel populations on dune rock and sandstone shores in Algoa Bay and on the open coast. It was concluded from these results that, spawning intensity and post-settlement growth and mortality, rather than dispersal, regulated recruitment and the structure and abundance of intertidal P. perna and C. meridionalis populations along the south coast of southern Africa. On the basis of these results it is recommended that species with limited dispersal, variable recruitment and high natural mortality, such as P. perna, should be conserved by protecting a small part of the population in reserves, and controlling utilisation outside reserves to minimize disturbance to local brood stocks. Furthermore, since the potential for reseeding adjacent exploited areas is limited, several small reserves placed at regular intervals along the coast would be more effective than a single large reserve.
- Full Text:
- Authors: Phillips, Tracey Elizabeth
- Date: 1995
- Subjects: Perna -- South Africa -- Eastern Cape -- Algoa bay Mytilus galloprovincialis -- South Africa -- Eastern Cape -- Algoa bay Mytilidae -- South Africa -- Eastern Cape -- Algoa bay
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5663 , http://hdl.handle.net/10962/d1005347
- Description: Recruitment of planktonic larvae into sedentary benthic populations regulates the population dynamics of marine invertebrates. The processes controlling recruitment, however, are poorly understood, and recruitment remains largely unpredictable, which complicates management of exploited shellfish resources. The mussels Perna perna, Choromytilus meridionalis and Mytilus galloprovincialis, found on the south coast of southern Africa, have planktonic larvae and sedentary adult stages. This thesis examines dispersal, settlement and early post-settlement growth and mortality, and their effect on recruitment and demography of intertidal mussel populations in the region of Algoa Bay on the south coast of southern Africa. Temporal and spatial variation in the body mass, density and size structure of mussels, the distribution of bivalve larvae on plankton grids in the nearshore zone and the distribution of a recently introduced invasive mussel, Mytilus galloprovincialis, were examined between 1989 and 1992. Furthermore, data on hourly or daily changes in wind strength and direction, air and sea surface temperatures and low and high tide levels in the study region, were obtained. There were 3-4 peaks in spawning (characterised by an abrupt decline in weight) and settlement activity annually. These peaks varied in exact timing, intensity and duration between sites and over time. However, at a site, spawning was followed by settlement 4-8 weeks later, and there was a significant (P < 0.05) direct correlation between spawning intensity prior to the appearance of a new cohort and the cohort density (settlement intensity). The stochastic spatial and temporal variation in breeding activity was superimposed on a more general pattern of a higher intensity of spawning and settlement in Algoa Bay than on the open coast, and a higher settlement intensity on coastal sandstone shores than on dune rock shores. Spawning was more frequent in winter and spring, and the probability of spawning and settlement peaked around the spring and autumn equinox, if temperature and wind conditions were suitable. Larval behaviour had little effect on their dispersal in the well-mixed nearshore region. Larvae were passively dispersed by currents, and their dispersal range and direction depended on prevailing winds and local topography. The sharp decline in density of recruit and adult M. galloprovincialis with increasing distance from the point of introduction, showed that some larvae were carried by wind generated currents over moderately long distances (-100 km). However, since most (76 %) M. galloprovincialis recruited within 4 km of the parent population, it is possible that larvae become trapped in small gullies and crevices around rocky shores, and have a limited dispersal range. This could explain the link between local patterns of spawning and settlement. The distribution and abundance of settlers on the shore was influenced by larval behaviour and the availability of settlement, substrata. Larvae preferred to settle primarily on foliose coralline algae and migrate to the adult mussel bed when they were larger (0.60-7 mm), but larvae also settled directly on adult mussels, possibly because the amount of coralline algae was limited. Both direct and secondary settlement were considered to be important in maintaining mussel populations since the rate of settlement was low(generally < 60 000.m-2). Cohort analyses showed that prior to maturity post-settlement growth (- 30 mm in 10 months) and mortality rates (60-100%) were high, but varied. When settlement intensity was low this variability uncoupled the relationship between spawning and recruitment intensity. Multiple regression analysis showed that together reproductive effort (gamete output), settlement intensity, growth and mortality prior to maturity, accounted for 76 % of the variance in recruitment into mature adult populations. The low settlement rate coupled with the short life span of mussels « 3 years), meant that populations underwent marked spatial and temporal variations in structure and abundance as settlement intensity varied, but there were consistent general differences between mussel populations on dune rock and sandstone shores in Algoa Bay and on the open coast. It was concluded from these results that, spawning intensity and post-settlement growth and mortality, rather than dispersal, regulated recruitment and the structure and abundance of intertidal P. perna and C. meridionalis populations along the south coast of southern Africa. On the basis of these results it is recommended that species with limited dispersal, variable recruitment and high natural mortality, such as P. perna, should be conserved by protecting a small part of the population in reserves, and controlling utilisation outside reserves to minimize disturbance to local brood stocks. Furthermore, since the potential for reseeding adjacent exploited areas is limited, several small reserves placed at regular intervals along the coast would be more effective than a single large reserve.
- Full Text:
The conflict between adaptation and constraint : the case of the Siphonariid limpets
- Authors: Chambers, Richard Jonathan
- Date: 1995
- Subjects: Siphonaria -- Adaptation Limpets -- Adaptation Pulmonata
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5657 , http://hdl.handle.net/10962/d1005340
- Description: The reproductive strategies of marine invertebrates have been related to various aspects of both their ecology, and their phylogenetic history. It has been the purpose of this thesis to try and separate these components among Siphonaria, a group of marine pulmonates. The taxonomy of these species is revised and I conclude that nine species are valid. All species deposit benthic egg masses and development may be either direct (S. anneae, S. compressa, S. dayi, S. nigerrima, S. serrata and S. tenuicostulata) or planktonic (S. capensis, S.concinna and S. oculus). Data on distribution and life-history relating to mode of larval development is then presented for 26 species of Siphonaria worldwide. Fifteen species are direct developers, nine are planktonic developers and a further two appear to have a dual developmental capacity, retaining both the velar swimming apparatus of a planktonic developer and the crawling foot of a direct developer. Direct developing species hatch from larger egg capsules, and generally occur higher on the shore than planktonic developers. Worldwide, planktonic developers are more widespread than direct developers, and individual planktonic species may have a greater latitudinal range. In most S~honaria subgenera, mode of larval development appears to be constant, although two subgenera (Patel/opsis and Sacculosiphonaria) include both developmental types. Locally, the intertidal zonation of three sympatric species (S. capensis, S. concinna and S. serrata) does not support a model which predicts direct development on the high shore and planktonic development on the low shore. However, distributions do correspond to particular intertidal microhabitats, and while there may be no, direct relationship between mode of larval development and intertidal height, the physical structure of egg masses, and the microhabitats used for spawning appear adaptive with regards to desiccation in the intertidal. S. concinna (planktonic development) and S. serrata (direct development) occur in similar microhabitats and are likely to be under similar selection pressures. In having different modes of larval development, there appears to be more than one optimal solution in a particular selective regime. In addition, both species seem to apportion similar amounts of energy to reproduction for each spawning episode, and also annually suggesting an optimum allocation of resources to reproduction. Genetic investigations using PolyAcrylamide Gel Electrophoresis (PAGE) confirm the status of the southern African species initially described, and indicate greater genetic variability associated with planktonic developing species than direct developing species. The systematic relationships revealed by DNA fingerprinting support the current classification systems, and also have implications with regards the evolution of larval development: direct development may be the plesiomorphic condition in, and among, some Siphonaria groups. There are both phylogenetic and adaptive explanations for the distribution of reproductive mode among benthic marine invertebrates. An evolutionary question, however, is not just a matter of either adaptation or constraint, it is a combination of these. Both contribute to the distribution of developmental mode among Siphonaria.
- Full Text:
- Authors: Chambers, Richard Jonathan
- Date: 1995
- Subjects: Siphonaria -- Adaptation Limpets -- Adaptation Pulmonata
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5657 , http://hdl.handle.net/10962/d1005340
- Description: The reproductive strategies of marine invertebrates have been related to various aspects of both their ecology, and their phylogenetic history. It has been the purpose of this thesis to try and separate these components among Siphonaria, a group of marine pulmonates. The taxonomy of these species is revised and I conclude that nine species are valid. All species deposit benthic egg masses and development may be either direct (S. anneae, S. compressa, S. dayi, S. nigerrima, S. serrata and S. tenuicostulata) or planktonic (S. capensis, S.concinna and S. oculus). Data on distribution and life-history relating to mode of larval development is then presented for 26 species of Siphonaria worldwide. Fifteen species are direct developers, nine are planktonic developers and a further two appear to have a dual developmental capacity, retaining both the velar swimming apparatus of a planktonic developer and the crawling foot of a direct developer. Direct developing species hatch from larger egg capsules, and generally occur higher on the shore than planktonic developers. Worldwide, planktonic developers are more widespread than direct developers, and individual planktonic species may have a greater latitudinal range. In most S~honaria subgenera, mode of larval development appears to be constant, although two subgenera (Patel/opsis and Sacculosiphonaria) include both developmental types. Locally, the intertidal zonation of three sympatric species (S. capensis, S. concinna and S. serrata) does not support a model which predicts direct development on the high shore and planktonic development on the low shore. However, distributions do correspond to particular intertidal microhabitats, and while there may be no, direct relationship between mode of larval development and intertidal height, the physical structure of egg masses, and the microhabitats used for spawning appear adaptive with regards to desiccation in the intertidal. S. concinna (planktonic development) and S. serrata (direct development) occur in similar microhabitats and are likely to be under similar selection pressures. In having different modes of larval development, there appears to be more than one optimal solution in a particular selective regime. In addition, both species seem to apportion similar amounts of energy to reproduction for each spawning episode, and also annually suggesting an optimum allocation of resources to reproduction. Genetic investigations using PolyAcrylamide Gel Electrophoresis (PAGE) confirm the status of the southern African species initially described, and indicate greater genetic variability associated with planktonic developing species than direct developing species. The systematic relationships revealed by DNA fingerprinting support the current classification systems, and also have implications with regards the evolution of larval development: direct development may be the plesiomorphic condition in, and among, some Siphonaria groups. There are both phylogenetic and adaptive explanations for the distribution of reproductive mode among benthic marine invertebrates. An evolutionary question, however, is not just a matter of either adaptation or constraint, it is a combination of these. Both contribute to the distribution of developmental mode among Siphonaria.
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Environmental physiology of the intertidal limpets Patella (Prosobranchia) and Siphonaria (Pulmonata)
- Authors: Marshall, David John
- Date: 1992
- Subjects: Prosobranchia Pulmonata Patellidae Siphonaria Limpets
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5686 , http://hdl.handle.net/10962/d1005372
- Description: 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.
- Full Text:
- Authors: Marshall, David John
- Date: 1992
- Subjects: Prosobranchia Pulmonata Patellidae Siphonaria Limpets
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5686 , http://hdl.handle.net/10962/d1005372
- Description: 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.
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