Occurrence, feeding ecology, and population structure of two dolphin species, Tursiops aduncus and Delphinus delphis, off the Wild Coast of South Africa
- Authors: Caputo, Michelle Anne
- Date: 2019
- Subjects: Delphinus -- South Africa-- Kwazulu-Natal -- Ecology , Tursoops -- South Africa-- Kwazulu-Natal -- Ecology , Delphinus -- South Africa-- Kwazulu-Natal -- Feeding and feeds , Tursoops -- South Africa-- Kwazulu-Natal -- Feeding and feeds , Dolphins -- South Africa-- Kwazulu-Natal , Delphinus -- South Africa-- Kwazulu-Natal -- Behavior , Tursoops -- South Africa-- Kwazulu-Natal -- Behavior , Marine ecology -- South Africa -- Kwazulu-Natal
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/115612 , vital:34197
- Description: Dolphins are apex predators and their movements, foraging activities and population dynamics play an important role in shaping their environment. Understanding their occurrence, movement patterns, and trophic ecology is essential to their conservation, especially as they are considered sentinel species. The Wild Coast of South Africa’s Eastern Cape, is characterized by the annual sardine (Sardinops sagax) run, which serves as an important foraging event for apex predators, including Indo-Pacific bottlenose (Tursiops aduncus) and long-beaked common dolphins (Delphinus delphis). Despite the ecological significance of this event, no dedicated surveys for these species have previously been conducted in this area. The main objectives of this research were to: investigate long-term occurrence of dolphins during the sardine run to determine if there have been any changes over the past 19 years, as has been predicted from common dolphin dietary investigations and anecdotal evidence; to determine short-term occurrence and movement patterns of selected inshore delphinids within the area, which is characterized by three marine protected areas (MPAs); and to use stable isotope analyses to determine trophic relationships and population structure of bottlenose and common dolphins within the region. Long-term data consisted of 131 opportunistic aerial surveys conducted between May and July over the period 1996 to 2014. Results from these surveys indicate that common dolphins, typically associated with sardines, decreased significantly in average group size over the study period (p=0.0343). Bottlenose dolphins, demonstrated no such trend (p=0.916). For both species, greater majority (> 70% of total counts) of sightings were made inside the MPAs. Short-term boat-based surveys were conducted three times a year between June 2014 and December 2016, contributing to a total of 47 days of surveys divided into three locations: Amathole, Hluleka, and Pondoland, each containing a MPA. Density and group size data were analyzed for both species and photographic identification analysis was performed for photographs of bottlenose dolphin dorsal fins. Results indicate that animal and sighting density did not differ temporally (bottlenose dolphin: sighting density – p=0.398, individual density –p=0.781; common dolphin: sighting density –p=0.472, individual density – p=0.204). Environmental factors (sea surface temperature, depth, substrate, and distance from shore) appeared to have limited effect on individual and sighting density and group size for both species (p>0.05). Photographic identification of bottlenose dolphins resulted in 2149 individuals, with a 11.8% resighting rate, with the highest resighting rate within the Pondoland MPA (16.1%). The resighting count did not differ temporally between monthly survey based on generalized linear models (p=0.866), but did differ between study areas (p<0.0001). These results provide the first evidence of the occurrence of both species of dolphin off the Wild Coast, as they were sighted in this region in all survey months. There was no trend in density based on temporal or environmental factors, which suggests other factors are influencing their occurrence. Resightings of bottlenose dolphins within the area suggest that there is some degree of residency, though the majority of animals were only sighted on a single occasion and there was no plateau in the discovery curve. A total of 256 biopsy samples (bottlenose dolphins =128; common dolphins=128) were collected during boat-based surveys. Bottlenose dolphin samples were also collected from adjacent areas to the southwest (Algoa Bay, n=22) and northeast (KwaZulu-Natal (KZN), n=20) of the Wild Coast to investigate similarities and differences between these areas. Despite a high degree of niche overlap between the two species (41%), common dolphins fed with a broader niche (standard ellipse area probability 0.89) than bottlenose dolphins in the summer and a narrower niche in the winter (probability 0.94). There was a clear spatial variation in the diet of bottlenose dolphins along the coast, with individuals from Algoa Bay and Amathole demonstrating 0% niche overlap with individuals from KZN, but the mechanism for these differences remains unclear as other species from South African waters demonstrate a strong southwest to northeast gradient in nitrogen for the Eastern Cape coastline. This research provides valuable baseline information regarding dolphins off the Wild Coast of South Africa, which remained largely unknown. My results indicate that bottlenose dolphins may be more resident in the Wild Coast than previous predicted, and confirm that common dolphins are highly mobile in this area.
- Full Text:
- Date Issued: 2019
- Authors: Caputo, Michelle Anne
- Date: 2019
- Subjects: Delphinus -- South Africa-- Kwazulu-Natal -- Ecology , Tursoops -- South Africa-- Kwazulu-Natal -- Ecology , Delphinus -- South Africa-- Kwazulu-Natal -- Feeding and feeds , Tursoops -- South Africa-- Kwazulu-Natal -- Feeding and feeds , Dolphins -- South Africa-- Kwazulu-Natal , Delphinus -- South Africa-- Kwazulu-Natal -- Behavior , Tursoops -- South Africa-- Kwazulu-Natal -- Behavior , Marine ecology -- South Africa -- Kwazulu-Natal
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/115612 , vital:34197
- Description: Dolphins are apex predators and their movements, foraging activities and population dynamics play an important role in shaping their environment. Understanding their occurrence, movement patterns, and trophic ecology is essential to their conservation, especially as they are considered sentinel species. The Wild Coast of South Africa’s Eastern Cape, is characterized by the annual sardine (Sardinops sagax) run, which serves as an important foraging event for apex predators, including Indo-Pacific bottlenose (Tursiops aduncus) and long-beaked common dolphins (Delphinus delphis). Despite the ecological significance of this event, no dedicated surveys for these species have previously been conducted in this area. The main objectives of this research were to: investigate long-term occurrence of dolphins during the sardine run to determine if there have been any changes over the past 19 years, as has been predicted from common dolphin dietary investigations and anecdotal evidence; to determine short-term occurrence and movement patterns of selected inshore delphinids within the area, which is characterized by three marine protected areas (MPAs); and to use stable isotope analyses to determine trophic relationships and population structure of bottlenose and common dolphins within the region. Long-term data consisted of 131 opportunistic aerial surveys conducted between May and July over the period 1996 to 2014. Results from these surveys indicate that common dolphins, typically associated with sardines, decreased significantly in average group size over the study period (p=0.0343). Bottlenose dolphins, demonstrated no such trend (p=0.916). For both species, greater majority (> 70% of total counts) of sightings were made inside the MPAs. Short-term boat-based surveys were conducted three times a year between June 2014 and December 2016, contributing to a total of 47 days of surveys divided into three locations: Amathole, Hluleka, and Pondoland, each containing a MPA. Density and group size data were analyzed for both species and photographic identification analysis was performed for photographs of bottlenose dolphin dorsal fins. Results indicate that animal and sighting density did not differ temporally (bottlenose dolphin: sighting density – p=0.398, individual density –p=0.781; common dolphin: sighting density –p=0.472, individual density – p=0.204). Environmental factors (sea surface temperature, depth, substrate, and distance from shore) appeared to have limited effect on individual and sighting density and group size for both species (p>0.05). Photographic identification of bottlenose dolphins resulted in 2149 individuals, with a 11.8% resighting rate, with the highest resighting rate within the Pondoland MPA (16.1%). The resighting count did not differ temporally between monthly survey based on generalized linear models (p=0.866), but did differ between study areas (p<0.0001). These results provide the first evidence of the occurrence of both species of dolphin off the Wild Coast, as they were sighted in this region in all survey months. There was no trend in density based on temporal or environmental factors, which suggests other factors are influencing their occurrence. Resightings of bottlenose dolphins within the area suggest that there is some degree of residency, though the majority of animals were only sighted on a single occasion and there was no plateau in the discovery curve. A total of 256 biopsy samples (bottlenose dolphins =128; common dolphins=128) were collected during boat-based surveys. Bottlenose dolphin samples were also collected from adjacent areas to the southwest (Algoa Bay, n=22) and northeast (KwaZulu-Natal (KZN), n=20) of the Wild Coast to investigate similarities and differences between these areas. Despite a high degree of niche overlap between the two species (41%), common dolphins fed with a broader niche (standard ellipse area probability 0.89) than bottlenose dolphins in the summer and a narrower niche in the winter (probability 0.94). There was a clear spatial variation in the diet of bottlenose dolphins along the coast, with individuals from Algoa Bay and Amathole demonstrating 0% niche overlap with individuals from KZN, but the mechanism for these differences remains unclear as other species from South African waters demonstrate a strong southwest to northeast gradient in nitrogen for the Eastern Cape coastline. This research provides valuable baseline information regarding dolphins off the Wild Coast of South Africa, which remained largely unknown. My results indicate that bottlenose dolphins may be more resident in the Wild Coast than previous predicted, and confirm that common dolphins are highly mobile in this area.
- Full Text:
- Date Issued: 2019
Feeding ecology, residency patterns and migration dynamics of bull sharks (Carcharhinus leucas) in the southwest Indian Ocean
- Authors: Daly, Ryan
- Date: 2015
- Subjects: Bull shark -- Ecology , Bull shark -- Feeding and feeds
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5924 , http://hdl.handle.net/10962/d1017802
- Description: Bull sharks (Carcharhinus leucas) are globally distributed top predators that play an important ecological role within coastal marine communities. However, little is known about how the spatial and temporal scales of their habitat use are associated with their ecological role. In this study, a population of sub-adult and adult bull sharks were investigated within a remote subtropical marine community in the southwest Indian Ocean off the coast of southern Mozambique. The main objectives of the study were to test a minimally invasive remote biopsy sampling method; to investigate the feeding ecology of bull sharks; and to investigate the temporal and spatial scales of bull shark residency patterns and migration dynamics. Biopsy tests on free-swimming bull sharks showed that the devised sampling technique provided a minimally invasive and consistent method (biopsy retention rate = 87%) to obtain muscle tissue samples sufficiently large enough (310±78mg, mean ± SD) for stable isotope analysis. Results from the stable isotope analysis showed that adult bull sharks appeared to exhibit a shift towards consistentlyhigher trophic level prey from an expanded foraging range compared to sub-adults, possibly due to increased mobility linked with size. Additionally, bull sharks had significantly broader niche widths compared to top predatory teleost assemblages with a wide and enriched range of δ13C values relative to the local marine community, suggesting that they forage over broad spatial scales along the east coast of southern Africa. Results from the passive acoustic telemetry investigation, conducted over a period between 10 and 22 months, supported these findings showing that the majority of tagged adult sharksexhibited temporally and spatially variable residency patterns interspersed with migration events. Ten individuals undertook coastal migrations that ranged between 433 and 709km (mean = 533km) with eight of these sharks returning to the study site. During migration, individuals exhibited rates of movement between 2 and 59km.d-1 (mean = 17.58km.d-1) and were recorded travelling annual distances of between 450 and 3760km (mean = 1163km). These findings suggest that adult bull sharks are not the sedentary species once thought to be but rather, consistently move over broad spatial scales on the east coast of southern Africa and play an important predatory role shaping and linking ecological processes within the southwest Indian Ocean.
- Full Text:
- Date Issued: 2015
- Authors: Daly, Ryan
- Date: 2015
- Subjects: Bull shark -- Ecology , Bull shark -- Feeding and feeds
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5924 , http://hdl.handle.net/10962/d1017802
- Description: Bull sharks (Carcharhinus leucas) are globally distributed top predators that play an important ecological role within coastal marine communities. However, little is known about how the spatial and temporal scales of their habitat use are associated with their ecological role. In this study, a population of sub-adult and adult bull sharks were investigated within a remote subtropical marine community in the southwest Indian Ocean off the coast of southern Mozambique. The main objectives of the study were to test a minimally invasive remote biopsy sampling method; to investigate the feeding ecology of bull sharks; and to investigate the temporal and spatial scales of bull shark residency patterns and migration dynamics. Biopsy tests on free-swimming bull sharks showed that the devised sampling technique provided a minimally invasive and consistent method (biopsy retention rate = 87%) to obtain muscle tissue samples sufficiently large enough (310±78mg, mean ± SD) for stable isotope analysis. Results from the stable isotope analysis showed that adult bull sharks appeared to exhibit a shift towards consistentlyhigher trophic level prey from an expanded foraging range compared to sub-adults, possibly due to increased mobility linked with size. Additionally, bull sharks had significantly broader niche widths compared to top predatory teleost assemblages with a wide and enriched range of δ13C values relative to the local marine community, suggesting that they forage over broad spatial scales along the east coast of southern Africa. Results from the passive acoustic telemetry investigation, conducted over a period between 10 and 22 months, supported these findings showing that the majority of tagged adult sharksexhibited temporally and spatially variable residency patterns interspersed with migration events. Ten individuals undertook coastal migrations that ranged between 433 and 709km (mean = 533km) with eight of these sharks returning to the study site. During migration, individuals exhibited rates of movement between 2 and 59km.d-1 (mean = 17.58km.d-1) and were recorded travelling annual distances of between 450 and 3760km (mean = 1163km). These findings suggest that adult bull sharks are not the sedentary species once thought to be but rather, consistently move over broad spatial scales on the east coast of southern Africa and play an important predatory role shaping and linking ecological processes within the southwest Indian Ocean.
- Full Text:
- Date Issued: 2015
Spatio-temporal variation in the phytobenthos and phytoplankton community structure and composition of particulate matter along a river-estuary continuum assessed using microscopic and stable isotope analyses
- Authors: Dalu, Tatenda
- Date: 2015
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/54416 , vital:26563
- Description: Phytoplankton and phytobenthos communities play an important role in lotic systems as primary producers providing essential biomolecules to higher trophic oganisms and are important indicators for environmental or ecological change. In this thesis, field studies (observational and experimental) along a river–estuary continuum were conducted to assess the spatio-temporal variation and development of phytobenthos and phytoplankton communities using a combination of stable isotope and community analyses in a temperate southern African system across four study periods: September (early spring) and November/December (late spring) 2012, and February (summer) and May/June (winter) 2013. Additionally, the sources and composition of the particulate organic matter were also analysed using stable isotope (δ15N and δ13C) analysis. The effects of substrate type and flood occurrence were assessed through experimental studies at an up- and downstream site of the river after a major flood event that occurred between October and November 2012. Common household tiles were used as artificial substrates to study the development/succession of phytobenthos communities after the flood disturbance. Distinct diatom communities were observed between upstream and downstream sites and at each site, community structure changed with time indicating succession. In addition to recording diatom characteristics on three natural substrates, namely; macrophytes, rocks and sediment, artificial substrates observations were also made on three different types of artificial substrates, namely; brick, brown clay and grey clay tiles. The natural (species richness 78) and artificial substrates (sp. richness 93) had different communities with the latter having greater species richness. Common phytobenthos taxa were not restricted to a single substrate but preference was generally high for the artificial substrates, especially brown tiles (mean sp. richness 47). Results of the redundancy analysis (RDA) analysis indicated that ammonium, conductivity, total dissolved solids, salinity, pH, oxygen reduction potential, phosphate and water depth were the major determinants of the phytobenthos composition at the two sites. The spatio–temporal variation of phytoplankton and phytobenthos communities and allochthonous organic matter along the river–estuary continuum was assessed at 8 sites using a combination of community and stable isotope analyses. A total of 178 species belonging to 78 genera were recorded with diatoms being predominant, accounting for 81.9 % of the total abundance. The total chl-a concentration along the river-estuary continuum increased from spring to a high in summer before decreasing to a low in winter. Periphyton communities were observed to be significantly different across sites (p < 0.05) in terms of species richness, abundances and isotopically The high periphytic δ15N values (range 7.9–15.2 ‰) recorded at the downstream sites compared to the pristine upstream sites (δ15N values range 4–7 ‰) suggest nutrient enrichment most likely derived from anthropogenic sources. Overall, our results reveal general patterns of periphyton communities and stable isotopes and provide improved information in the use of periphyton δ15N as an excellent indicator of anthropogenic nitrogen pollution. Ecologists are interested in the factors that control, and the variability in, the contributions of different sources to mixed organic materials traveling through lotic systems. We hypothesized that the source matter fuelling mixed organic pools in a river-estuary continuum varies over space and time, with the upper reaches of a system characterized by allochthonous-dominated material and autochthonous contributions becoming more important in the lower reaches. Samples of the mixed organic pools and allochthonous and autochthonous source materials were collected during the four study periods. The C:N ratios of suspended particulate matter (SPM) collected during summer and winter indicated that the lower reaches of the system had similar organic matter contributions from the freshwater and terrestrial sources. Stable isotope analysis in R revealed that the contributions of autochthonous organic matter were high in SPM along the entire continuum, and aquatic macrophytes were significant contributors to SPM specifically in the upper reaches. The terrestrial leaves made major contributions to the SPM in the middle regions of the system (i.e. downstream sites of the river, particularly in early and late spring). Bulk detritus had large allochthonous matter components in the lower reaches (estuary), and the contributions of aquatic macrophytes and benthic algae to bulk detritus were high (> 50 %) in the upper to middle reaches (river), but low (< 20 %) in the lower reaches (estuary). The current investigation represents the first attempt to assess the validity of the River Continuum Concept (RCC) in a southern African temperate river. The phytoplankton and phytobenthos communities, and chl-a concentration followed a trend similar to that proposed for the river continuum concept (RCC). The middle reaches based on the phytobenthos or phytoplankton communities and chl-a concentrations which were employed as proxies for primary production, were the most productive, while the upper reaches were the least primary productive. The evaluation of organic matter contributions to the SPM and detritus along the river–estuary continuum provided a baseline assessment of the nature and sources of potential food for consumers inhabiting different locations during different times of the year. Incorporating such spatio-temporal variations in SPM and detritus into food web studies will improve our understanding of the flow of carbon through aquatic systems.
- Full Text:
- Date Issued: 2015
- Authors: Dalu, Tatenda
- Date: 2015
- Language: English
- Type: text , Thesis , Doctoral , PhD
- Identifier: http://hdl.handle.net/10962/54416 , vital:26563
- Description: Phytoplankton and phytobenthos communities play an important role in lotic systems as primary producers providing essential biomolecules to higher trophic oganisms and are important indicators for environmental or ecological change. In this thesis, field studies (observational and experimental) along a river–estuary continuum were conducted to assess the spatio-temporal variation and development of phytobenthos and phytoplankton communities using a combination of stable isotope and community analyses in a temperate southern African system across four study periods: September (early spring) and November/December (late spring) 2012, and February (summer) and May/June (winter) 2013. Additionally, the sources and composition of the particulate organic matter were also analysed using stable isotope (δ15N and δ13C) analysis. The effects of substrate type and flood occurrence were assessed through experimental studies at an up- and downstream site of the river after a major flood event that occurred between October and November 2012. Common household tiles were used as artificial substrates to study the development/succession of phytobenthos communities after the flood disturbance. Distinct diatom communities were observed between upstream and downstream sites and at each site, community structure changed with time indicating succession. In addition to recording diatom characteristics on three natural substrates, namely; macrophytes, rocks and sediment, artificial substrates observations were also made on three different types of artificial substrates, namely; brick, brown clay and grey clay tiles. The natural (species richness 78) and artificial substrates (sp. richness 93) had different communities with the latter having greater species richness. Common phytobenthos taxa were not restricted to a single substrate but preference was generally high for the artificial substrates, especially brown tiles (mean sp. richness 47). Results of the redundancy analysis (RDA) analysis indicated that ammonium, conductivity, total dissolved solids, salinity, pH, oxygen reduction potential, phosphate and water depth were the major determinants of the phytobenthos composition at the two sites. The spatio–temporal variation of phytoplankton and phytobenthos communities and allochthonous organic matter along the river–estuary continuum was assessed at 8 sites using a combination of community and stable isotope analyses. A total of 178 species belonging to 78 genera were recorded with diatoms being predominant, accounting for 81.9 % of the total abundance. The total chl-a concentration along the river-estuary continuum increased from spring to a high in summer before decreasing to a low in winter. Periphyton communities were observed to be significantly different across sites (p < 0.05) in terms of species richness, abundances and isotopically The high periphytic δ15N values (range 7.9–15.2 ‰) recorded at the downstream sites compared to the pristine upstream sites (δ15N values range 4–7 ‰) suggest nutrient enrichment most likely derived from anthropogenic sources. Overall, our results reveal general patterns of periphyton communities and stable isotopes and provide improved information in the use of periphyton δ15N as an excellent indicator of anthropogenic nitrogen pollution. Ecologists are interested in the factors that control, and the variability in, the contributions of different sources to mixed organic materials traveling through lotic systems. We hypothesized that the source matter fuelling mixed organic pools in a river-estuary continuum varies over space and time, with the upper reaches of a system characterized by allochthonous-dominated material and autochthonous contributions becoming more important in the lower reaches. Samples of the mixed organic pools and allochthonous and autochthonous source materials were collected during the four study periods. The C:N ratios of suspended particulate matter (SPM) collected during summer and winter indicated that the lower reaches of the system had similar organic matter contributions from the freshwater and terrestrial sources. Stable isotope analysis in R revealed that the contributions of autochthonous organic matter were high in SPM along the entire continuum, and aquatic macrophytes were significant contributors to SPM specifically in the upper reaches. The terrestrial leaves made major contributions to the SPM in the middle regions of the system (i.e. downstream sites of the river, particularly in early and late spring). Bulk detritus had large allochthonous matter components in the lower reaches (estuary), and the contributions of aquatic macrophytes and benthic algae to bulk detritus were high (> 50 %) in the upper to middle reaches (river), but low (< 20 %) in the lower reaches (estuary). The current investigation represents the first attempt to assess the validity of the River Continuum Concept (RCC) in a southern African temperate river. The phytoplankton and phytobenthos communities, and chl-a concentration followed a trend similar to that proposed for the river continuum concept (RCC). The middle reaches based on the phytobenthos or phytoplankton communities and chl-a concentrations which were employed as proxies for primary production, were the most productive, while the upper reaches were the least primary productive. The evaluation of organic matter contributions to the SPM and detritus along the river–estuary continuum provided a baseline assessment of the nature and sources of potential food for consumers inhabiting different locations during different times of the year. Incorporating such spatio-temporal variations in SPM and detritus into food web studies will improve our understanding of the flow of carbon through aquatic systems.
- Full Text:
- Date Issued: 2015
Age, growth, reproduction and sexual dimorphism of the striped dolphin, Stenella coeruleoalba, off the south-east coast of southern Africa
- Authors: Bishop, Amy Ruth
- Date: 2014
- Subjects: Striped dolphin -- Conservation -- Africa, Southern , Striped dolphin -- Development -- Africa, Southern , Striped dolphin -- Africa, Southern -- Reproduction , Striped dolphin -- Age determination -- Africa, Southern , Dolphins -- Conservation -- Africa, Southern , Dolphins -- Development -- Africa, Southern , Dolphins -- Africa, Southern -- Reproduction , Dolphins -- Age determination -- Africa, Southern , Dimorphism (Animals)
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5848 , http://hdl.handle.net/10962/d1011600 , Striped dolphin -- Conservation -- Africa, Southern , Striped dolphin -- Development -- Africa, Southern , Striped dolphin -- Africa, Southern -- Reproduction , Striped dolphin -- Age determination -- Africa, Southern , Dolphins -- Conservation -- Africa, Southern , Dolphins -- Development -- Africa, Southern , Dolphins -- Africa, Southern -- Reproduction , Dolphins -- Age determination -- Africa, Southern , Dimorphism (Animals)
- Description: The striped dolphin, Stenella coeruleoalba, is an oceanic dolphin found worldwide in tropical to warm temperate waters. Globally populations are threatened through direct fishing, bycatch and pollution. Little is known about the life history of this species in South African waters and it is currently listed as ‘Data Deficient’ by the International Union for Conservation of Nature (IUCN). Critical to the conservation and management of this species is an understanding of basic life history parameters. For the present study, metadata and tissues from 101 stranded S. coeruleoalba along the south-east coast of southern Africa between 1969 and 2012 were analyzed to examine age and growth, male and female reproduction and sexual dimorphism. Age and growth were determined by counting the number of growth layer groups (GLG’s) present in their teeth. A von Bertalanffy growth curve gave the best fit to the data and indicated that physical maturity was reached around 21 years in males and 18 years in females. Additionally, asymptotic lengths were reached at 231cm and 224cm in males and females, respectively. The maximum ages obtained were 21 years for males and 20 years for females. Length at birth was estimated to be between 116cm and 119cm for males and between 112cm and 114cm for females. The reproductive status of the adult dolphins was assessed by histological and macroscopic analyses of gonadal tissue. Sexual maturity was attained before physical maturity in both sexes. The males reached sexual maturity between 8 and 12 years of age and between 209 and 224cm in total body length. The maximum combined testis weight comprised about 0.24% of the total body weight and sexual maturity was reached between a combined testis weight of 16 and 41g, a mean testis length of 8 and 10cm, and a mean seminiferous tubule diameter of 44 and 70μm. Females attained sexual maturity between 7 and 8 years of age and between 213 and 216 cm in total body length. The ovulation rate was 0.32 per year and the majority of ovulations occurred in the left ovary, with ovulations occurring in the right ovary only in individuals older than 10 years of age. The annual pregnancy rate was calculated to be 26%. Reproductive seasonality could not be established due to the small sample size and the absence of samples throughout the year. However, both conceptions and births appear to occur in both August and December/January. Additionally, testis weight and seminiferous tubule diameters appear to fluctuate throughout the year, which could suggest reproductive seasonality in this species. Sexual size dimorphism in mature adults was evident for a number of external measurements, however, sexual shape dimorphism was not present in this species. Mature males had significantly longer and wider pectoral fins, dorsal fins and flukes than females (p<0.05 in all cases). It was suggested that the larger appendages are required for male on male combat during intraspecific competition for females. Additionally, larger appendages aid in thermoregulation, propulsion and maneuverability. Females had a significantly greater distance between the tip upper jaw and the genital aperture and anus, which was expected due to differences in anatomy between male and female genital openings. Overall, S. coeruleoalba did not exhibit distinct sexual dimorphism and together with the relatively small testes in relation to the total body weight (0.24%) it was proposed that this species demonstrates a less extreme form of polygyny. Additionaly, the results of this study are important to the understanding of the basic biology of the species.
- Full Text:
- Date Issued: 2014
- Authors: Bishop, Amy Ruth
- Date: 2014
- Subjects: Striped dolphin -- Conservation -- Africa, Southern , Striped dolphin -- Development -- Africa, Southern , Striped dolphin -- Africa, Southern -- Reproduction , Striped dolphin -- Age determination -- Africa, Southern , Dolphins -- Conservation -- Africa, Southern , Dolphins -- Development -- Africa, Southern , Dolphins -- Africa, Southern -- Reproduction , Dolphins -- Age determination -- Africa, Southern , Dimorphism (Animals)
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5848 , http://hdl.handle.net/10962/d1011600 , Striped dolphin -- Conservation -- Africa, Southern , Striped dolphin -- Development -- Africa, Southern , Striped dolphin -- Africa, Southern -- Reproduction , Striped dolphin -- Age determination -- Africa, Southern , Dolphins -- Conservation -- Africa, Southern , Dolphins -- Development -- Africa, Southern , Dolphins -- Africa, Southern -- Reproduction , Dolphins -- Age determination -- Africa, Southern , Dimorphism (Animals)
- Description: The striped dolphin, Stenella coeruleoalba, is an oceanic dolphin found worldwide in tropical to warm temperate waters. Globally populations are threatened through direct fishing, bycatch and pollution. Little is known about the life history of this species in South African waters and it is currently listed as ‘Data Deficient’ by the International Union for Conservation of Nature (IUCN). Critical to the conservation and management of this species is an understanding of basic life history parameters. For the present study, metadata and tissues from 101 stranded S. coeruleoalba along the south-east coast of southern Africa between 1969 and 2012 were analyzed to examine age and growth, male and female reproduction and sexual dimorphism. Age and growth were determined by counting the number of growth layer groups (GLG’s) present in their teeth. A von Bertalanffy growth curve gave the best fit to the data and indicated that physical maturity was reached around 21 years in males and 18 years in females. Additionally, asymptotic lengths were reached at 231cm and 224cm in males and females, respectively. The maximum ages obtained were 21 years for males and 20 years for females. Length at birth was estimated to be between 116cm and 119cm for males and between 112cm and 114cm for females. The reproductive status of the adult dolphins was assessed by histological and macroscopic analyses of gonadal tissue. Sexual maturity was attained before physical maturity in both sexes. The males reached sexual maturity between 8 and 12 years of age and between 209 and 224cm in total body length. The maximum combined testis weight comprised about 0.24% of the total body weight and sexual maturity was reached between a combined testis weight of 16 and 41g, a mean testis length of 8 and 10cm, and a mean seminiferous tubule diameter of 44 and 70μm. Females attained sexual maturity between 7 and 8 years of age and between 213 and 216 cm in total body length. The ovulation rate was 0.32 per year and the majority of ovulations occurred in the left ovary, with ovulations occurring in the right ovary only in individuals older than 10 years of age. The annual pregnancy rate was calculated to be 26%. Reproductive seasonality could not be established due to the small sample size and the absence of samples throughout the year. However, both conceptions and births appear to occur in both August and December/January. Additionally, testis weight and seminiferous tubule diameters appear to fluctuate throughout the year, which could suggest reproductive seasonality in this species. Sexual size dimorphism in mature adults was evident for a number of external measurements, however, sexual shape dimorphism was not present in this species. Mature males had significantly longer and wider pectoral fins, dorsal fins and flukes than females (p<0.05 in all cases). It was suggested that the larger appendages are required for male on male combat during intraspecific competition for females. Additionally, larger appendages aid in thermoregulation, propulsion and maneuverability. Females had a significantly greater distance between the tip upper jaw and the genital aperture and anus, which was expected due to differences in anatomy between male and female genital openings. Overall, S. coeruleoalba did not exhibit distinct sexual dimorphism and together with the relatively small testes in relation to the total body weight (0.24%) it was proposed that this species demonstrates a less extreme form of polygyny. Additionaly, the results of this study are important to the understanding of the basic biology of the species.
- Full Text:
- Date Issued: 2014
Spatial and temporal patterns in the population and the feeding of Palaemon peringueyi (MacPherson 1990) (Decapoda, Palaemonidae) in the Kariega estuary, South Africa
- Nyalungu, Nonhlanhla Precotia Dudu
- Authors: Nyalungu, Nonhlanhla Precotia Dudu
- Date: 2013
- Subjects: Palaemonidae -- Food -- South Africa , Palaemonidae -- Behavior -- South Africa , Palaemonidae -- Habitat -- South Africa , Estuarine biology -- South Africa , Shrimp populations -- South Africa , Shrimps -- South Africa , Decapoda (Crustacea) -- South Africa
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5584 , http://hdl.handle.net/10962/d1001576 , Palaemonidae -- Food -- South Africa , Palaemonidae -- Behavior -- South Africa , Palaemonidae -- Habitat -- South Africa , Estuarine biology -- South Africa , Shrimp populations -- South Africa , Shrimps -- South Africa , Decapoda (Crustacea) -- South Africa
- Description: The spatial and temporal patterns in the population demographs and feeding ecology of the caridean shrimp, Palaemon peringueyi, was investigated over a 12 month period in the permanently open Kariega Estuary situated on the south-east coastline of South Africa. Shrimps were sampled monthly from three stations corresponding to the lower, middle and upper reaches of the Kariega Estuary. The shrimps were collected at the different stations using a pull net. Temperature, salinity, dissolved oxygen and % cover of submerged vegetation were measured at each sampling station. Finally, the feeding ecology of the shrimp was investigated employing stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes analysis. Temperature, salinity, dissolved oxygen and vegetation cover showed distinct seasonal patterns. The shrimps attained total abundances and biomass of between 0 and 245.6 individuals per m² (ind.m⁻²) and between 0 and 13.6 grams wet weight per m² (g wwt.m⁻²), respectively. Mean abundance ranged between 2.3 (± 3.0) and 88 (± 27.3) ind.m⁻² in the lower reach, between 0.9 (± 0.4) to 21.5 (± 4.5) ind.m⁻² in the middle reach and 0 to 1.6 (± 0.5) ind.m⁻² in the upper reach. Mean biomass ranged between 0.4 (± 0.5) and 4.4 (± 1.4) g wwt.m⁻² in the lower reach, between 0.1 (± 0) to 4.3 (± 0.9) g wwt.m⁻² in the middle reach and between 0 to 1.6 (± 0.5) g wwt.m⁻² in the upper reach. Greatest abundances and biomasses were recorded in winter following heavy rainfall within the catchment area. Freshwater inflow which is due to rainfall is thought to provide a cue for the recruitment of marine breeding organisms. The spatial patterns in the distribution of the shrimps showed that juveniles occupied the lower and middle reaches and adults the entire length of the estuary. There were no significant relationships between the abundance and biomass of the shrimps and the selected physico-chemical variables (p > 0.05 in all cases). Results of the stable isotope study indicated that the mean δ¹³C of the shrimps in the lower, middle and upper reaches were -12.4 (± 1.3) ‰, -15.6 (± 0.5) ‰ and -17.5 (± 0.5) ‰, respectively. The δ¹⁵N isotopes of the shrimps in the lower reach was 11.6 (± 0.5) ‰ with middle and upper reaches having similar δ¹⁵N values, 13.5 (± 0.4) ‰ and 13.4 (± 0.4) ‰, respectively. There was no significant difference in the δ¹³C and δ¹⁵N isotope (p > 0.05) between the sexes of the shrimps. Two-way ANOVA indicated a significant difference in the δ¹³C isotope of the shrimps between the sites (p < 0.05). The difference in the δ¹³C and δ¹⁵N ratios of the shrimps suggests spatial variability in the diet of P. peringueyi within the Kariega Estuary. The diet of the shrimps in the lower reach is primarily composed of Nanozostera capensis epibionts. In the middle reach the diet of the shrimps is likely comprised of the detritus of benthic algae. In the upper reach of the estuary, P. peringueyi likely derives its carbon from a combination of epibionts, benthic algae and detritus derived from Codium.
- Full Text:
- Date Issued: 2013
- Authors: Nyalungu, Nonhlanhla Precotia Dudu
- Date: 2013
- Subjects: Palaemonidae -- Food -- South Africa , Palaemonidae -- Behavior -- South Africa , Palaemonidae -- Habitat -- South Africa , Estuarine biology -- South Africa , Shrimp populations -- South Africa , Shrimps -- South Africa , Decapoda (Crustacea) -- South Africa
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5584 , http://hdl.handle.net/10962/d1001576 , Palaemonidae -- Food -- South Africa , Palaemonidae -- Behavior -- South Africa , Palaemonidae -- Habitat -- South Africa , Estuarine biology -- South Africa , Shrimp populations -- South Africa , Shrimps -- South Africa , Decapoda (Crustacea) -- South Africa
- Description: The spatial and temporal patterns in the population demographs and feeding ecology of the caridean shrimp, Palaemon peringueyi, was investigated over a 12 month period in the permanently open Kariega Estuary situated on the south-east coastline of South Africa. Shrimps were sampled monthly from three stations corresponding to the lower, middle and upper reaches of the Kariega Estuary. The shrimps were collected at the different stations using a pull net. Temperature, salinity, dissolved oxygen and % cover of submerged vegetation were measured at each sampling station. Finally, the feeding ecology of the shrimp was investigated employing stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes analysis. Temperature, salinity, dissolved oxygen and vegetation cover showed distinct seasonal patterns. The shrimps attained total abundances and biomass of between 0 and 245.6 individuals per m² (ind.m⁻²) and between 0 and 13.6 grams wet weight per m² (g wwt.m⁻²), respectively. Mean abundance ranged between 2.3 (± 3.0) and 88 (± 27.3) ind.m⁻² in the lower reach, between 0.9 (± 0.4) to 21.5 (± 4.5) ind.m⁻² in the middle reach and 0 to 1.6 (± 0.5) ind.m⁻² in the upper reach. Mean biomass ranged between 0.4 (± 0.5) and 4.4 (± 1.4) g wwt.m⁻² in the lower reach, between 0.1 (± 0) to 4.3 (± 0.9) g wwt.m⁻² in the middle reach and between 0 to 1.6 (± 0.5) g wwt.m⁻² in the upper reach. Greatest abundances and biomasses were recorded in winter following heavy rainfall within the catchment area. Freshwater inflow which is due to rainfall is thought to provide a cue for the recruitment of marine breeding organisms. The spatial patterns in the distribution of the shrimps showed that juveniles occupied the lower and middle reaches and adults the entire length of the estuary. There were no significant relationships between the abundance and biomass of the shrimps and the selected physico-chemical variables (p > 0.05 in all cases). Results of the stable isotope study indicated that the mean δ¹³C of the shrimps in the lower, middle and upper reaches were -12.4 (± 1.3) ‰, -15.6 (± 0.5) ‰ and -17.5 (± 0.5) ‰, respectively. The δ¹⁵N isotopes of the shrimps in the lower reach was 11.6 (± 0.5) ‰ with middle and upper reaches having similar δ¹⁵N values, 13.5 (± 0.4) ‰ and 13.4 (± 0.4) ‰, respectively. There was no significant difference in the δ¹³C and δ¹⁵N isotope (p > 0.05) between the sexes of the shrimps. Two-way ANOVA indicated a significant difference in the δ¹³C isotope of the shrimps between the sites (p < 0.05). The difference in the δ¹³C and δ¹⁵N ratios of the shrimps suggests spatial variability in the diet of P. peringueyi within the Kariega Estuary. The diet of the shrimps in the lower reach is primarily composed of Nanozostera capensis epibionts. In the middle reach the diet of the shrimps is likely comprised of the detritus of benthic algae. In the upper reach of the estuary, P. peringueyi likely derives its carbon from a combination of epibionts, benthic algae and detritus derived from Codium.
- Full Text:
- Date Issued: 2013
The influence of overwash and breaching events on the spatial and temporal patterns in ichthyofauna community composition in a temporarily open/closed southern African estuary
- Authors: Tweddle, Gavin Paul
- Date: 2013
- Subjects: Estuarine fishes -- Africa, Southern Marine fishes -- Africa, Southern Fishes -- Breeding -- Africa, Southern Fishes -- Effect of habitat modification on -- Africa, Southern Fishes -- Spawning -- Africa, Southern Gobiidae -- Africa, Southern Estuaries -- Africa, Southern Estuarine ecology -- Africa, Southern
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5610 , http://hdl.handle.net/10962/d1002957
- Description: This study assessed the importance of overwash and breaching events on the ichthyofaunal community structure in the medium-sized temporarily open/closed Mpekweni Estuary located on the southeast coastline of southern Africa. The fish in the littoral zone of the estuary were sampled using a 5m seine net while the channel region was sampled using two nets, a smaller meshed 30m seine net to target the estuarine spawning species and the juvenile estuarine-dependant marine spawners, and a larger meshed 50m seine net to target the larger marine and freshwater spawning species. Intensive monthly sampling over two years provided data on selected physico-chemical and biological parameters. During the two year sampling period from November 2005 to October 2007 the estuary breached in late July 2006 and remained open till the sandbar re-formed across the mouth in late December 2006. Thus, sampling encompassed three open/closed phases 1) initial closed period, 2) open period and 3) re-closed period after the berm was re-formed. The open period was divided into two phases 1) the out-flow phase and 2) the tidal phase. A total of 36 fish species representing 19 families were sampled using the various seine nets employed during the investigation. In the littoral zone, the estuarine spawners (Estuarine Utilisation Category, [EUC] I), mainly the Gobiidae, Glossogobius callidus, dominated the samples numerically and by biomass. The smaller estuarine spawning species sampled in the channel were numerically dominated by Gilchristella aestuaria in conjunction with two other EUC I species, Atherina breviceps and G. callidus. The estuarine-dependant marine spawners (EUC II), however, dominated the ichthyofaunal biomass of the channel. The abundance and biomass of the larger species targeted were dominated by estuarine-dependant marine spawning species (EUC II), principally Rhabdosargus holubi. During the closed periods of the estuary, total fish abundance and biomass displayed weak seasonal patterns. The breaching event and subsequent open period was associated with a decrease in the total abundances of fish in the littoral zone and channel of the estuary, reflecting the out-flow of estuarine biomass-rich water into the marine environment. The breaching event coincided with a shift in the community composition of the ichthyofauna, reflecting the recruitment of marine spawning species into the estuary. Numerical analysis identified two distinct spatial fish communities within the estuary, a community associated with the mouth region and one comprising the rest of the estuary. The absence of any further spatial patterns in the ichthyofaunal community structure within the Mpekweni Estuary appear to be ascribed to the virtual absence of horizontal patterns in physico-chemical and biological parameters recorded in the system. Cohort analyses were employed to determine possible recruitment events for selected estuarine and marine spawning species. The estuarine spawning species displayed continuous recruitment patterns throughout the study, which appeared to be unaffected by the breaching event. Conversely, the larger marine spawning species displayed multiple cohorts, indicating non-continuous recruitment. Subsequent retrospective analysis of cohorts for the different species identified summer recruitment peaks that coincided with the breaching event and open period. Minor recruitment of marine spawning species also occurred during overwash events. The recruitment of ichthyofauna into the estuary was quantified during three distinct hydrological events: overwash, out-flow phase immediately after breaching and tidal phase during the period when the mouth was open. Estimates of fish recruitment were highest during the outflow phase immediately after the estuary breached and declined as the estuary became tidally inundated with marine water. Although not as high as the outflow and tidal phases, recruitment was evident during overwash events. Results of the current study highlight the importance of both breaching and overwashing events in structuring the ichthyofaunal community composition in a medium-sized southern African temporarily open/closed estuary. These results are broadly in agreement with similar studies conducted both locally and in other regions of the world.
- Full Text:
- Date Issued: 2013
- Authors: Tweddle, Gavin Paul
- Date: 2013
- Subjects: Estuarine fishes -- Africa, Southern Marine fishes -- Africa, Southern Fishes -- Breeding -- Africa, Southern Fishes -- Effect of habitat modification on -- Africa, Southern Fishes -- Spawning -- Africa, Southern Gobiidae -- Africa, Southern Estuaries -- Africa, Southern Estuarine ecology -- Africa, Southern
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5610 , http://hdl.handle.net/10962/d1002957
- Description: This study assessed the importance of overwash and breaching events on the ichthyofaunal community structure in the medium-sized temporarily open/closed Mpekweni Estuary located on the southeast coastline of southern Africa. The fish in the littoral zone of the estuary were sampled using a 5m seine net while the channel region was sampled using two nets, a smaller meshed 30m seine net to target the estuarine spawning species and the juvenile estuarine-dependant marine spawners, and a larger meshed 50m seine net to target the larger marine and freshwater spawning species. Intensive monthly sampling over two years provided data on selected physico-chemical and biological parameters. During the two year sampling period from November 2005 to October 2007 the estuary breached in late July 2006 and remained open till the sandbar re-formed across the mouth in late December 2006. Thus, sampling encompassed three open/closed phases 1) initial closed period, 2) open period and 3) re-closed period after the berm was re-formed. The open period was divided into two phases 1) the out-flow phase and 2) the tidal phase. A total of 36 fish species representing 19 families were sampled using the various seine nets employed during the investigation. In the littoral zone, the estuarine spawners (Estuarine Utilisation Category, [EUC] I), mainly the Gobiidae, Glossogobius callidus, dominated the samples numerically and by biomass. The smaller estuarine spawning species sampled in the channel were numerically dominated by Gilchristella aestuaria in conjunction with two other EUC I species, Atherina breviceps and G. callidus. The estuarine-dependant marine spawners (EUC II), however, dominated the ichthyofaunal biomass of the channel. The abundance and biomass of the larger species targeted were dominated by estuarine-dependant marine spawning species (EUC II), principally Rhabdosargus holubi. During the closed periods of the estuary, total fish abundance and biomass displayed weak seasonal patterns. The breaching event and subsequent open period was associated with a decrease in the total abundances of fish in the littoral zone and channel of the estuary, reflecting the out-flow of estuarine biomass-rich water into the marine environment. The breaching event coincided with a shift in the community composition of the ichthyofauna, reflecting the recruitment of marine spawning species into the estuary. Numerical analysis identified two distinct spatial fish communities within the estuary, a community associated with the mouth region and one comprising the rest of the estuary. The absence of any further spatial patterns in the ichthyofaunal community structure within the Mpekweni Estuary appear to be ascribed to the virtual absence of horizontal patterns in physico-chemical and biological parameters recorded in the system. Cohort analyses were employed to determine possible recruitment events for selected estuarine and marine spawning species. The estuarine spawning species displayed continuous recruitment patterns throughout the study, which appeared to be unaffected by the breaching event. Conversely, the larger marine spawning species displayed multiple cohorts, indicating non-continuous recruitment. Subsequent retrospective analysis of cohorts for the different species identified summer recruitment peaks that coincided with the breaching event and open period. Minor recruitment of marine spawning species also occurred during overwash events. The recruitment of ichthyofauna into the estuary was quantified during three distinct hydrological events: overwash, out-flow phase immediately after breaching and tidal phase during the period when the mouth was open. Estimates of fish recruitment were highest during the outflow phase immediately after the estuary breached and declined as the estuary became tidally inundated with marine water. Although not as high as the outflow and tidal phases, recruitment was evident during overwash events. Results of the current study highlight the importance of both breaching and overwashing events in structuring the ichthyofaunal community composition in a medium-sized southern African temporarily open/closed estuary. These results are broadly in agreement with similar studies conducted both locally and in other regions of the world.
- Full Text:
- Date Issued: 2013
Trophodynamics of the benthic and hyperbenthic communities inhabiting the Sub-Antarctic Prince Edward Islands : stable isotope and fatty acid signatures
- Authors: Allan, Elizabeth Louise
- Date: 2011
- Subjects: Benthos -- Prince Edward Islands Benthos -- Feeding and feeds -- Prince Edward Islands Benthos -- Nutrition -- Prince Edward Islands Prince Edward Islands
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5806 , http://hdl.handle.net/10962/d1006350
- Description: The aim of this study was to investigate spatial changes in the trophic and energy pathways of the benthic community in the shallow shelf waters of the sub-Antarctic Prince Edward Islands (PEI). A combination of stable isotope and fatty acid analyses were used to provide a time-integrated view of the assimilated feeding history of selected components of the PEI benthic community. This study forms part of the larger project entitled “Variability in the Southern Ocean ecosystems” and is a contribution to the South African National Antarctic Programme (SANAP). During austral autumn 2009, benthic specimens were collected from 10 stations (from depths of 70 to 295 m) in different regions around the PEI: inter-island shelf (upstream, between and downstream of the islands) and nearshore. Historical data were combined with new data collected during 2009 to assess the long-term trends in the feeding ecology of the benthos in the region of the islands. The stable isotope and fatty acid signatures of the benthic suspension- and deposit-feeding organisms generated during this study suggested that these two communities incorporated both phytoplankton and kelp in their diets. Stable isotope, and to a lesser extent fatty acid signatures, indicated that kelp contributed more to the diets of those organisms in close proximity to the kelp beds (nearshore stations) than those from the inter-island region. Overall, however, pelagic phytoplankton was the dominant food source in the diets of all organisms, even for those living near the kelp beds. Notable exceptions were the sponges and bryozoans, in which kelp and phytoplankton contributed similar proportions to their diets, most likely resulting from a size restricted feeding mode. There were, therefore, no distinct spatial differences in the importance of the various food sources. However, fatty acid compositional data indicated increased food quality between and within the lee of the islands compared to upstream. The organisms collected upstream of the PEI had substantially lower quantities of total fatty acids (TFAs) than organisms of the same species collected from nearshore, open shelf or downstream stations. The increased food quality between and within the lee of the islands was likely a result of the “island mass effect”, which reflects increased phytoplankton concentrations at the PEI. The fatty acid profiles of hyperbenthic shrimp Nauticaris marionis, a key species in the PEI ecosystem, revealed no distinct ontogenetic or spatial patterns. This result is in contrast to the stable isotope analyses, which detected both spatial and ontogenetic differences in the diet of the shrimp. Nearshore shrimp were more ₁₃C-enriched than those from the inter-island region, suggesting increased kelp entering the food web within these regions. In addition, the shrimps demonstrated enrichment in δ₁₃C and δ₁₅N signatures with an increase in size, resulting in a relatively distinct separation of size classes, thus reflecting niche separation through their diets. The fatty acid profiles revealed that the shrimp all contained large proportions of polyunsaturated fatty acids (PUFAs) and essential fatty acids (EFAs), indicating that the quality of food consumed was similar among size class and region despite the niche separation and variation in carbon sources utilised. In addition, diatom and dinoflagellate fatty acids (20:5ω3 and 22:6ω3, respectively) occurred in the highest proportions in N. marionis, highlighting the importance of phytoplankton (indirectly) in their diet. These results represent strong evidence that phytoplankton is an importance food source in the PEI ecosystem. The temporal investigation of isotope signatures in the benthos at the PEI indicated that N. marionis demonstrated an overall depletion in δ₁₃C signatures over the period of 1984 to 2009 (nearshore: -2.55 ‰, inter-island: -2.32 ‰). Overall, the benthic community showed similar depletions in δ13C signatures (from -1.96 to -4.70 ‰), suggesting that shifts have occurred in the carbon signatures at the base of the food web. The depletion in δ₁₃C signatures of the benthos at the PEI most likely reflects increased contributions of smaller slow growing phytoplankton cells (more depleted signatures than large fast growing cells) in the diets of these organisms over time. These diet shifts, in turn, suggest a decrease in productivity has occurred at the islands, likely due to a decreased frequency and intensity of the “island mass effect”. Decreased phytoplankton productivity at the PEI likely results from the southward shift in the average position of the sub-Antarctic Front (SAF), most likely in response to climate change, which in turn promotes flow-through conditions rather than retention at the PEI.
- Full Text:
- Date Issued: 2011
- Authors: Allan, Elizabeth Louise
- Date: 2011
- Subjects: Benthos -- Prince Edward Islands Benthos -- Feeding and feeds -- Prince Edward Islands Benthos -- Nutrition -- Prince Edward Islands Prince Edward Islands
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5806 , http://hdl.handle.net/10962/d1006350
- Description: The aim of this study was to investigate spatial changes in the trophic and energy pathways of the benthic community in the shallow shelf waters of the sub-Antarctic Prince Edward Islands (PEI). A combination of stable isotope and fatty acid analyses were used to provide a time-integrated view of the assimilated feeding history of selected components of the PEI benthic community. This study forms part of the larger project entitled “Variability in the Southern Ocean ecosystems” and is a contribution to the South African National Antarctic Programme (SANAP). During austral autumn 2009, benthic specimens were collected from 10 stations (from depths of 70 to 295 m) in different regions around the PEI: inter-island shelf (upstream, between and downstream of the islands) and nearshore. Historical data were combined with new data collected during 2009 to assess the long-term trends in the feeding ecology of the benthos in the region of the islands. The stable isotope and fatty acid signatures of the benthic suspension- and deposit-feeding organisms generated during this study suggested that these two communities incorporated both phytoplankton and kelp in their diets. Stable isotope, and to a lesser extent fatty acid signatures, indicated that kelp contributed more to the diets of those organisms in close proximity to the kelp beds (nearshore stations) than those from the inter-island region. Overall, however, pelagic phytoplankton was the dominant food source in the diets of all organisms, even for those living near the kelp beds. Notable exceptions were the sponges and bryozoans, in which kelp and phytoplankton contributed similar proportions to their diets, most likely resulting from a size restricted feeding mode. There were, therefore, no distinct spatial differences in the importance of the various food sources. However, fatty acid compositional data indicated increased food quality between and within the lee of the islands compared to upstream. The organisms collected upstream of the PEI had substantially lower quantities of total fatty acids (TFAs) than organisms of the same species collected from nearshore, open shelf or downstream stations. The increased food quality between and within the lee of the islands was likely a result of the “island mass effect”, which reflects increased phytoplankton concentrations at the PEI. The fatty acid profiles of hyperbenthic shrimp Nauticaris marionis, a key species in the PEI ecosystem, revealed no distinct ontogenetic or spatial patterns. This result is in contrast to the stable isotope analyses, which detected both spatial and ontogenetic differences in the diet of the shrimp. Nearshore shrimp were more ₁₃C-enriched than those from the inter-island region, suggesting increased kelp entering the food web within these regions. In addition, the shrimps demonstrated enrichment in δ₁₃C and δ₁₅N signatures with an increase in size, resulting in a relatively distinct separation of size classes, thus reflecting niche separation through their diets. The fatty acid profiles revealed that the shrimp all contained large proportions of polyunsaturated fatty acids (PUFAs) and essential fatty acids (EFAs), indicating that the quality of food consumed was similar among size class and region despite the niche separation and variation in carbon sources utilised. In addition, diatom and dinoflagellate fatty acids (20:5ω3 and 22:6ω3, respectively) occurred in the highest proportions in N. marionis, highlighting the importance of phytoplankton (indirectly) in their diet. These results represent strong evidence that phytoplankton is an importance food source in the PEI ecosystem. The temporal investigation of isotope signatures in the benthos at the PEI indicated that N. marionis demonstrated an overall depletion in δ₁₃C signatures over the period of 1984 to 2009 (nearshore: -2.55 ‰, inter-island: -2.32 ‰). Overall, the benthic community showed similar depletions in δ13C signatures (from -1.96 to -4.70 ‰), suggesting that shifts have occurred in the carbon signatures at the base of the food web. The depletion in δ₁₃C signatures of the benthos at the PEI most likely reflects increased contributions of smaller slow growing phytoplankton cells (more depleted signatures than large fast growing cells) in the diets of these organisms over time. These diet shifts, in turn, suggest a decrease in productivity has occurred at the islands, likely due to a decreased frequency and intensity of the “island mass effect”. Decreased phytoplankton productivity at the PEI likely results from the southward shift in the average position of the sub-Antarctic Front (SAF), most likely in response to climate change, which in turn promotes flow-through conditions rather than retention at the PEI.
- Full Text:
- Date Issued: 2011
Spatial and temporal patterns in the hyperbenthic community structure in a permanently open Eastern Cape estuary, South Africa
- Authors: Heyns, Elodie R
- Date: 2010
- Subjects: Sundays Estuary -- South Africa -- Eastern Cape , Groundfishes -- South Africa -- Eatern Cape , Estuaries -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5768 , http://hdl.handle.net/10962/d1005456
- Description: The spatial and temporal (seasonal and diel) patterns in the hyperbenthic community structure (>500 μm) was investigated in the warm temperate, permanently open Kariega Estuary situated along the south-eastern coastline of South Africa. Spatial and seasonal patterns in the hyperbenthic community structure were assessed monthly at six stations along the length of the estuary over a period of twelve months. Data were collected using a modified hyperbenthic sledge, comprising two super-imposed nets. Physico-chemical data indicate the presence of a constant reverse salinity gradient, with highest salinities measured in the upper reaches and lowest at the mouth of the estuary. Strong seasonal patterns in temperature, dissolved oxygen and total chlorophyll-a (chl-a) concentrations were evident. Total average hyperbenthic densities ranged between 0.04 and 166 ind.m-3 in the lower net and between 0.12 and 225 ind.m-3 in the upper net. Hyperbenthic biomass values ranged between 0.02 and 11.9 mg.dry weight.m-3 in the lower net and between 0.02 and 17.4 mg.dry weight.m-3 in the upper net. A spatial and temporal pattern in total densities was detected with an increase in abundance over the period of September to October 2008 particularly in the middle reaches (Stations 3 and 4). Both the lower and upper nets were numerically dominated by decapods (mainly brachyuran crab zoeae) with the exception of June and July 2008 when mysids (mainly Mesopodopsis wooldridgei) dominated, making up 72.4 ± 58.14% of the total abundance in the lower net. A redundancy analysis (RDA) indicated that 99.2% of the variance in the hyperbenthic community structure could be explained by the first two canonical axes. Axis one, which accounted for 96.8% of the total variation detected in the ordination plot was highly correlated with sedimentary organic content and to a lesser extent the chl-a concentration within the Kariega Estuary. The correlations with the second canonical axis (2.4%) were less obvious, however, salinity and seston concentration were weakly correlated with this axis. Diel variability in the hyperbenthic community structure was assessed during March 2009. Samples were collected during the day and night (n = 6 for each period) using sampling gear described above. Total average hyperbenthic densities during the day (497.9 ± 254.1 ind.m-3) were significantly higher than night-time estimates (129.9 ± 38.5 ind.m-3; p<0.05). There were no significant differences in the average dayand night-time estimates of hyperbenthic biomass (p>0.05). A hierarchical cluster analysis identified two significantly distinct groupings, designated the day and night samples. Results from the SIMPER procedure indicated that the high densities of crab zoeae recorded during the day-time accounted for the majority of the dissimilarity between the day and night groupings (44.7%). In addition, it is apparent that several benthic species, especially from the cumacean and isopod orders, were absent from the hyperbenthos during the day-time and emerged into the water column at night.
- Full Text:
- Date Issued: 2010
- Authors: Heyns, Elodie R
- Date: 2010
- Subjects: Sundays Estuary -- South Africa -- Eastern Cape , Groundfishes -- South Africa -- Eatern Cape , Estuaries -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5768 , http://hdl.handle.net/10962/d1005456
- Description: The spatial and temporal (seasonal and diel) patterns in the hyperbenthic community structure (>500 μm) was investigated in the warm temperate, permanently open Kariega Estuary situated along the south-eastern coastline of South Africa. Spatial and seasonal patterns in the hyperbenthic community structure were assessed monthly at six stations along the length of the estuary over a period of twelve months. Data were collected using a modified hyperbenthic sledge, comprising two super-imposed nets. Physico-chemical data indicate the presence of a constant reverse salinity gradient, with highest salinities measured in the upper reaches and lowest at the mouth of the estuary. Strong seasonal patterns in temperature, dissolved oxygen and total chlorophyll-a (chl-a) concentrations were evident. Total average hyperbenthic densities ranged between 0.04 and 166 ind.m-3 in the lower net and between 0.12 and 225 ind.m-3 in the upper net. Hyperbenthic biomass values ranged between 0.02 and 11.9 mg.dry weight.m-3 in the lower net and between 0.02 and 17.4 mg.dry weight.m-3 in the upper net. A spatial and temporal pattern in total densities was detected with an increase in abundance over the period of September to October 2008 particularly in the middle reaches (Stations 3 and 4). Both the lower and upper nets were numerically dominated by decapods (mainly brachyuran crab zoeae) with the exception of June and July 2008 when mysids (mainly Mesopodopsis wooldridgei) dominated, making up 72.4 ± 58.14% of the total abundance in the lower net. A redundancy analysis (RDA) indicated that 99.2% of the variance in the hyperbenthic community structure could be explained by the first two canonical axes. Axis one, which accounted for 96.8% of the total variation detected in the ordination plot was highly correlated with sedimentary organic content and to a lesser extent the chl-a concentration within the Kariega Estuary. The correlations with the second canonical axis (2.4%) were less obvious, however, salinity and seston concentration were weakly correlated with this axis. Diel variability in the hyperbenthic community structure was assessed during March 2009. Samples were collected during the day and night (n = 6 for each period) using sampling gear described above. Total average hyperbenthic densities during the day (497.9 ± 254.1 ind.m-3) were significantly higher than night-time estimates (129.9 ± 38.5 ind.m-3; p<0.05). There were no significant differences in the average dayand night-time estimates of hyperbenthic biomass (p>0.05). A hierarchical cluster analysis identified two significantly distinct groupings, designated the day and night samples. Results from the SIMPER procedure indicated that the high densities of crab zoeae recorded during the day-time accounted for the majority of the dissimilarity between the day and night groupings (44.7%). In addition, it is apparent that several benthic species, especially from the cumacean and isopod orders, were absent from the hyperbenthos during the day-time and emerged into the water column at night.
- Full Text:
- Date Issued: 2010
Aspects of the ecology and biology of the isopod, Exosphaeroma hylocoetes, (Barnard, 1940) in three temporarily open/closed southern African estuaries
- Authors: Henninger, Tony Oskar
- Date: 2009
- Subjects: Estuaries -- South Africa -- Eastern Cape Estuarine ecology -- South Africa -- Eastern Cape Isopoda -- South Africa -- Eastern Cape Estuarine biology -- South Africa -- Eastern Cape Ruppia maritima
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5640 , http://hdl.handle.net/10962/d1005322
- Description: Temporarily open/closed estuaries (TOCE’s) are the numerically dominant type of estuary accounting for ≈ 70 % of all estuaries along the South African coastline. Despite their numerical dominance, aspects of the biology of organisms within these systems, particularly macrocrustacea remain poorly understood. This study firstly assessed the abundance and biomass of the most common isopod, Exosphaeroma hylocoetes, in three Eastern Cape TOCE’s, (the West Kleinemonde, East Kleinemonde and Kasouga Estuaries), and their response to mouth breaching events. This study was followed by aspects of the biology of the isopod including their utilisation of submerged macrophytes as a refuge from predation and/or possible food source, as well as the growth rates of the isopods in the laboratory under different environmental conditions. Mean isopod abundances and biomasses ranged between 0 and 4 791 ind. m⁻² and 0 and 9.65 mg dwt. m⁻² in the West Kleinemonde Estuary and between 0 and 108 ind. m⁻² and 0 and 0.318 mg dwt. m⁻² in the nearby East Kleinemonde Estuary. In the Kasouga Estuary, the values ranged between 0 and 3 650 ind. m⁻² and 0 and 5.105 mg dwt. m⁻². Temporal and spatial changes in the abundance and biomass of E. hylocoetes within the three systems was primarily linked to mouth phase, with populations declining when the mouth was open and to a lesser extent, seasonality. In all three estuaries maximum isopod abundances and biomasses were recorded in their middle and upper reaches, which could be ascribed to the presence of submerged macrophytes particularly Ruppia maritima, in two of the estuaries. Males (5.71 ± 0.41 mm) were significantly larger than females (3.99 ± 0.26 mm), but the sex ratios were skewed in favour of females, (a common feature in many isopod populations). Females were found carrying brood throughout the study, releasing offspring directly into the water column, to allow recruitment to the populations. The larger the female, the larger the brood carried (up to a maximum of 72 embryo/mancas). Results of laboratory experiments indicate that the close association of Exosphaeroma hylocoetes with submerged macrophytes is a result of the plant stands providing a refuge against predation by selected ichthyofauna. However stable carbon isotope and fatty acid analyses indicate that E. hylocoetes made use of ephiphytic algae and detritus on the stems of R. maritima, rather than the submerged macrophyte itself. There were no significant differences in the growth rates of male and female isopods at combinations of temperature (15 and 25 °C) and salinity (15 and 35 ‰). Laboratory growth studies revealed that males lived longer than females (25.77 ± 3.40 weeks vs 21.52 ± 3.00 weeks), and therefore achieved larger overall size. Females, however, reached sexual maturity (at 2.5 mm) at a faster rate (two to four weeks) than males (5.5 mm after 6.5 to 11 weeks), dependent on temperature. Exosphaeroma hylocoetes with its association with submerged macrophytes, early maturity, its growth rates and longevity, female-biased sex ratios and year-round breeding, all contribute to its success in temporary open/closed estuaries.
- Full Text:
- Date Issued: 2009
- Authors: Henninger, Tony Oskar
- Date: 2009
- Subjects: Estuaries -- South Africa -- Eastern Cape Estuarine ecology -- South Africa -- Eastern Cape Isopoda -- South Africa -- Eastern Cape Estuarine biology -- South Africa -- Eastern Cape Ruppia maritima
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5640 , http://hdl.handle.net/10962/d1005322
- Description: Temporarily open/closed estuaries (TOCE’s) are the numerically dominant type of estuary accounting for ≈ 70 % of all estuaries along the South African coastline. Despite their numerical dominance, aspects of the biology of organisms within these systems, particularly macrocrustacea remain poorly understood. This study firstly assessed the abundance and biomass of the most common isopod, Exosphaeroma hylocoetes, in three Eastern Cape TOCE’s, (the West Kleinemonde, East Kleinemonde and Kasouga Estuaries), and their response to mouth breaching events. This study was followed by aspects of the biology of the isopod including their utilisation of submerged macrophytes as a refuge from predation and/or possible food source, as well as the growth rates of the isopods in the laboratory under different environmental conditions. Mean isopod abundances and biomasses ranged between 0 and 4 791 ind. m⁻² and 0 and 9.65 mg dwt. m⁻² in the West Kleinemonde Estuary and between 0 and 108 ind. m⁻² and 0 and 0.318 mg dwt. m⁻² in the nearby East Kleinemonde Estuary. In the Kasouga Estuary, the values ranged between 0 and 3 650 ind. m⁻² and 0 and 5.105 mg dwt. m⁻². Temporal and spatial changes in the abundance and biomass of E. hylocoetes within the three systems was primarily linked to mouth phase, with populations declining when the mouth was open and to a lesser extent, seasonality. In all three estuaries maximum isopod abundances and biomasses were recorded in their middle and upper reaches, which could be ascribed to the presence of submerged macrophytes particularly Ruppia maritima, in two of the estuaries. Males (5.71 ± 0.41 mm) were significantly larger than females (3.99 ± 0.26 mm), but the sex ratios were skewed in favour of females, (a common feature in many isopod populations). Females were found carrying brood throughout the study, releasing offspring directly into the water column, to allow recruitment to the populations. The larger the female, the larger the brood carried (up to a maximum of 72 embryo/mancas). Results of laboratory experiments indicate that the close association of Exosphaeroma hylocoetes with submerged macrophytes is a result of the plant stands providing a refuge against predation by selected ichthyofauna. However stable carbon isotope and fatty acid analyses indicate that E. hylocoetes made use of ephiphytic algae and detritus on the stems of R. maritima, rather than the submerged macrophyte itself. There were no significant differences in the growth rates of male and female isopods at combinations of temperature (15 and 25 °C) and salinity (15 and 35 ‰). Laboratory growth studies revealed that males lived longer than females (25.77 ± 3.40 weeks vs 21.52 ± 3.00 weeks), and therefore achieved larger overall size. Females, however, reached sexual maturity (at 2.5 mm) at a faster rate (two to four weeks) than males (5.5 mm after 6.5 to 11 weeks), dependent on temperature. Exosphaeroma hylocoetes with its association with submerged macrophytes, early maturity, its growth rates and longevity, female-biased sex ratios and year-round breeding, all contribute to its success in temporary open/closed estuaries.
- Full Text:
- Date Issued: 2009
Trophodynamics of mesozooplankton in the the vicinity of the subtropical convergence in the Indian sector of the Southern Ocean
- Authors: Daly, Ryan
- Date: 2009
- Subjects: Zooplankton -- Antarctic Ocean , Phytoplankton -- Antarctic Ocean
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5791 , http://hdl.handle.net/10962/d1005479 , Zooplankton -- Antarctic Ocean , Phytoplankton -- Antarctic Ocean
- Description: The trophodynamics of the numerically dominant mesozooplankton (200-2000 m) in the vicinity of the Subtropical Convergence (STC) in the Indian sector of the Southern Ocean during austral autumn (April / May) 2007 were investigated as part of the Southern Ocean Ecosystem Variability Study. The survey consisted of six north-south transects each bisecting the STC between 38º to 43ºS and 38º to 41º45’E. In total, 48 stations situated at 30 nautical mile intervals were occupied over a period of ten days. Hydrographic data revealed a well defined surface and sub-surface expression of the STC, which appeared to meander considerably between 41ºS and 41º15’S. Surface chlorophyll-a (chla) concentrations were low, ranging between 0.08 and 0.68 mg chl-a.m-3 and were generally dominated by the picophytoplankton (<2 m) which made up 66.6% (SD±17.6) of the total pigment. Chl-a concentrations integrated over the top 150m of the water column ranged between 11.97 and 40.07 mg chl-a.m-2 and showed no significant spatial patterns (p>0.05). Total integrated mesozooplankton abundance and biomass during the study ranged between 3934.9 and 308521.4 ind.m-2 (mean = 47198.19; SD±62411.4 ind.m-2) and between 239.8 and 4614.3 mg Dwt.m-2 (mean = 1338.58; SD ±1060.5), respectively. Again, there were no significant spatial patterns in the total mesozooplankton abundance or biomass within the region of study (p>0.05). No significant correlations were found between biological (chlorophyll-a concentrations and zooplankton abundance) and physico-chemical variables (temperature and salinity) (p>0.05). The total mesozooplankton community was numerically dominated by copepods of the genera Pleuromamma, Calanus, Oncaea and Oithona. Other important representatives of the mesozooplankton community included the tunicate, Salpa thompsoni, and the pteropod, Limacina retroversa. At the 40% similarity level, numerical analysis identified five distinct mesozooplankton groupings within the survey area. Differences between the groupings were associated with changes in the relative contribution of numerically dominant species rather than the presence or absence of individual species. No groupings were associated with any specific feature of the front within the survey area. The feeding rates of the six most numerically abundant mesozooplankton species (Calanus simillimus, Limacina retroversa, Pleuromamma abdominalis, Clausocalanus breviceps, Oncaea conifera, Salpa thompsoni) accounting for on average 39% of the total mesozooplankton counts, were investigated using the gut fluorescence technique. For all species, the total gut pigment contents during the night time were significantly higher than the daytime values (p<0.05 for all species). The gut evacuation rates (k) for selected mesozooplankton ranged between 0.14 and 0.81 h-1. The ingestion rates ranged between 147.8 and 5495.4 ng(pigm)ind-1.day-1 which corresponded to a daily ration of between 2.4 and 10.9% body carbon. The combined grazing impact of the selected species on the daily phytoplankton standing stock was highly variable and ranged between 1.2 and 174.1% with an average of 27.3% (SD±38.78%) within the survey area. The highest grazing impact (>60%) was typically associated with those stations where the pteropod, L. retroversa, and the tunicate, S. thompsoni, contributed more than 5% of the total mesozooplankton counts. No significant differences were found in the grazing impact of any or all selected species situated either north, south or in the immediate vicinity of the front (p>0.05 in all cases). The lack of defined spatial patterns in the mesozooplankton abundance and community structure suggests that the STC did not act as a significant biogeographic barrier to the distribution of mesozooplankton during the study. It is presumed that the large scale mixing event caused by a storm prior to this study was responsible for the observed lack of elevated biological activity within the region of the STC.
- Full Text:
- Date Issued: 2009
- Authors: Daly, Ryan
- Date: 2009
- Subjects: Zooplankton -- Antarctic Ocean , Phytoplankton -- Antarctic Ocean
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5791 , http://hdl.handle.net/10962/d1005479 , Zooplankton -- Antarctic Ocean , Phytoplankton -- Antarctic Ocean
- Description: The trophodynamics of the numerically dominant mesozooplankton (200-2000 m) in the vicinity of the Subtropical Convergence (STC) in the Indian sector of the Southern Ocean during austral autumn (April / May) 2007 were investigated as part of the Southern Ocean Ecosystem Variability Study. The survey consisted of six north-south transects each bisecting the STC between 38º to 43ºS and 38º to 41º45’E. In total, 48 stations situated at 30 nautical mile intervals were occupied over a period of ten days. Hydrographic data revealed a well defined surface and sub-surface expression of the STC, which appeared to meander considerably between 41ºS and 41º15’S. Surface chlorophyll-a (chla) concentrations were low, ranging between 0.08 and 0.68 mg chl-a.m-3 and were generally dominated by the picophytoplankton (<2 m) which made up 66.6% (SD±17.6) of the total pigment. Chl-a concentrations integrated over the top 150m of the water column ranged between 11.97 and 40.07 mg chl-a.m-2 and showed no significant spatial patterns (p>0.05). Total integrated mesozooplankton abundance and biomass during the study ranged between 3934.9 and 308521.4 ind.m-2 (mean = 47198.19; SD±62411.4 ind.m-2) and between 239.8 and 4614.3 mg Dwt.m-2 (mean = 1338.58; SD ±1060.5), respectively. Again, there were no significant spatial patterns in the total mesozooplankton abundance or biomass within the region of study (p>0.05). No significant correlations were found between biological (chlorophyll-a concentrations and zooplankton abundance) and physico-chemical variables (temperature and salinity) (p>0.05). The total mesozooplankton community was numerically dominated by copepods of the genera Pleuromamma, Calanus, Oncaea and Oithona. Other important representatives of the mesozooplankton community included the tunicate, Salpa thompsoni, and the pteropod, Limacina retroversa. At the 40% similarity level, numerical analysis identified five distinct mesozooplankton groupings within the survey area. Differences between the groupings were associated with changes in the relative contribution of numerically dominant species rather than the presence or absence of individual species. No groupings were associated with any specific feature of the front within the survey area. The feeding rates of the six most numerically abundant mesozooplankton species (Calanus simillimus, Limacina retroversa, Pleuromamma abdominalis, Clausocalanus breviceps, Oncaea conifera, Salpa thompsoni) accounting for on average 39% of the total mesozooplankton counts, were investigated using the gut fluorescence technique. For all species, the total gut pigment contents during the night time were significantly higher than the daytime values (p<0.05 for all species). The gut evacuation rates (k) for selected mesozooplankton ranged between 0.14 and 0.81 h-1. The ingestion rates ranged between 147.8 and 5495.4 ng(pigm)ind-1.day-1 which corresponded to a daily ration of between 2.4 and 10.9% body carbon. The combined grazing impact of the selected species on the daily phytoplankton standing stock was highly variable and ranged between 1.2 and 174.1% with an average of 27.3% (SD±38.78%) within the survey area. The highest grazing impact (>60%) was typically associated with those stations where the pteropod, L. retroversa, and the tunicate, S. thompsoni, contributed more than 5% of the total mesozooplankton counts. No significant differences were found in the grazing impact of any or all selected species situated either north, south or in the immediate vicinity of the front (p>0.05 in all cases). The lack of defined spatial patterns in the mesozooplankton abundance and community structure suggests that the STC did not act as a significant biogeographic barrier to the distribution of mesozooplankton during the study. It is presumed that the large scale mixing event caused by a storm prior to this study was responsible for the observed lack of elevated biological activity within the region of the STC.
- Full Text:
- Date Issued: 2009
Spatial and temporal variations in metals in the sediment and water of selected Eastern Cape Estuaries, South Africa
- Authors: Orr, Kyla Kathleen
- Date: 2008
- Subjects: Marine sediments -- South Africa -- Eastern Cape , Metals -- South Africa -- Eastern Cape , Metals -- Environmental aspects , Estuaries -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape , Estuarine sediments -- South Africa -- Eastern Cape , Water quality -- South Africa -- Eastern Cape , Water pollution -- South Africa -- Eastern Cape , Environmental toxicology -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5769 , http://hdl.handle.net/10962/d1005457 , Marine sediments -- South Africa -- Eastern Cape , Metals -- South Africa -- Eastern Cape , Metals -- Environmental aspects , Estuaries -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape , Estuarine sediments -- South Africa -- Eastern Cape , Water quality -- South Africa -- Eastern Cape , Water pollution -- South Africa -- Eastern Cape , Environmental toxicology -- South Africa -- Eastern Cape
- Description: The spatio-temporal patterns in concentrations of selected metals within the sediment (Cd, Co, Cu, Fe, Pb, Ni and Zn) and water (Cd and Pb) of three permanently open estuaries (Kariega, Kowie, Great Fish) and six temporary open-closed estuaries (Mpekweni, East Kleinemonde, West Kleinemonde, Riet, Kasouga, Boknes) were investigated. The concentrations of metals were influenced by size composition and total organic content of the sediments. Enrichment factors (EFs), using Fe as a reference element, and baseline linear regression models for metals vs. Fe were calculated to assess the extent of metal enrichment in the sediments. The mean concentrations of metals in the sediments(mg·kg⁻¹) showed ranges of 0.28 – 2.31 for Cd, 1.26 – 6.24 Co, 0.69 – 6.93 for Cu, 2119 – 14912 for Fe, 2.29 –14.01 for Ni, 4.81 – 22.20 for Pb and 5.77 – 21.75 for Zn. Mean normalized enrichment factors ranged between 0.75 – 6.19 for Cd, 0.53 – 2.71 for Co, 0.22 – 0.84 for Cu, 0.30 – 1.87 for Ni, 0.99 – 3.17 for Pb and 0.14 – 0.98 for Zn. All nine estuaries had average enrichment factors of greater than 1 for Cd. In general there was no enrichment of Cu and Zn in the sediments of any of the estuaries included in this study (EFs < 1). The Kariega, East Kleinemonde, West Kleinemonde, Riet and Great Fish Estuaries showed some degree of enrichment for Co (1 < EF < 4), Ni (1 < EF < 2) and Pb (1 < EF < 4), while the Mpekweni, Kasouga, Boknes and Kowie Estuaries were unenriched with these metals (EF < 1). Enrichment factors for Cd, Co and Pb typically followed the development gradient along the estuaries, suggesting anthropogenic enrichment. The concentrations of Cd and Pb in the water of the nine estuaries were also determined. The average concentrations of Cd and Pb in the water (μg·ℓ⁻¹) ranged between 0.05 – 3.32 and 0.75 – 34.13 respectively. On average the concentrations of Cd and Pb in the water of all the estuaries were below the South African recommended water quality guidelines for coastal marine waters. Variations in metal concentrations associated with changes in hydrology (wet vs. dry season) were determined in the water and sediment of the Kariega, East Kleinemonde and Riet Estuaries. Cobalt, Pb and Ni enrichment in the Kariega Estuary sediment was significantly higher during the dry season, and the mean concentrations of Pb and Cd in the water column were 19-fold and 66-fold higher in the dry season. The elevated concentration of metals during the dry season could be related to accumulation of diffuse pollution from human activities within the catchment area. Conversely, inflow of fresh water into the estuary had the net effect of reducing the concentration and enrichment of these metals within the Kariega Estuary due to scouring and outflow of estuarine water and sediment into the marine environment. The temporal variations in metal concentrations and enrichment factors were less pronounced in the temporary open-closed estuaries than the permanently open Kariega Estuary. The observed trend can probably be related to the low anthropogenic impact within the catchment areas of these systems, and the relatively smaller size of the catchments. Significant spatial variations existed in metal enrichment in the sediment of both the East Kleinemonde and Riet estuaries, with the highest degrees of enrichment occurring in the sediments from the marine environment and lower reaches.
- Full Text:
- Date Issued: 2008
- Authors: Orr, Kyla Kathleen
- Date: 2008
- Subjects: Marine sediments -- South Africa -- Eastern Cape , Metals -- South Africa -- Eastern Cape , Metals -- Environmental aspects , Estuaries -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape , Estuarine sediments -- South Africa -- Eastern Cape , Water quality -- South Africa -- Eastern Cape , Water pollution -- South Africa -- Eastern Cape , Environmental toxicology -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5769 , http://hdl.handle.net/10962/d1005457 , Marine sediments -- South Africa -- Eastern Cape , Metals -- South Africa -- Eastern Cape , Metals -- Environmental aspects , Estuaries -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape , Estuarine sediments -- South Africa -- Eastern Cape , Water quality -- South Africa -- Eastern Cape , Water pollution -- South Africa -- Eastern Cape , Environmental toxicology -- South Africa -- Eastern Cape
- Description: The spatio-temporal patterns in concentrations of selected metals within the sediment (Cd, Co, Cu, Fe, Pb, Ni and Zn) and water (Cd and Pb) of three permanently open estuaries (Kariega, Kowie, Great Fish) and six temporary open-closed estuaries (Mpekweni, East Kleinemonde, West Kleinemonde, Riet, Kasouga, Boknes) were investigated. The concentrations of metals were influenced by size composition and total organic content of the sediments. Enrichment factors (EFs), using Fe as a reference element, and baseline linear regression models for metals vs. Fe were calculated to assess the extent of metal enrichment in the sediments. The mean concentrations of metals in the sediments(mg·kg⁻¹) showed ranges of 0.28 – 2.31 for Cd, 1.26 – 6.24 Co, 0.69 – 6.93 for Cu, 2119 – 14912 for Fe, 2.29 –14.01 for Ni, 4.81 – 22.20 for Pb and 5.77 – 21.75 for Zn. Mean normalized enrichment factors ranged between 0.75 – 6.19 for Cd, 0.53 – 2.71 for Co, 0.22 – 0.84 for Cu, 0.30 – 1.87 for Ni, 0.99 – 3.17 for Pb and 0.14 – 0.98 for Zn. All nine estuaries had average enrichment factors of greater than 1 for Cd. In general there was no enrichment of Cu and Zn in the sediments of any of the estuaries included in this study (EFs < 1). The Kariega, East Kleinemonde, West Kleinemonde, Riet and Great Fish Estuaries showed some degree of enrichment for Co (1 < EF < 4), Ni (1 < EF < 2) and Pb (1 < EF < 4), while the Mpekweni, Kasouga, Boknes and Kowie Estuaries were unenriched with these metals (EF < 1). Enrichment factors for Cd, Co and Pb typically followed the development gradient along the estuaries, suggesting anthropogenic enrichment. The concentrations of Cd and Pb in the water of the nine estuaries were also determined. The average concentrations of Cd and Pb in the water (μg·ℓ⁻¹) ranged between 0.05 – 3.32 and 0.75 – 34.13 respectively. On average the concentrations of Cd and Pb in the water of all the estuaries were below the South African recommended water quality guidelines for coastal marine waters. Variations in metal concentrations associated with changes in hydrology (wet vs. dry season) were determined in the water and sediment of the Kariega, East Kleinemonde and Riet Estuaries. Cobalt, Pb and Ni enrichment in the Kariega Estuary sediment was significantly higher during the dry season, and the mean concentrations of Pb and Cd in the water column were 19-fold and 66-fold higher in the dry season. The elevated concentration of metals during the dry season could be related to accumulation of diffuse pollution from human activities within the catchment area. Conversely, inflow of fresh water into the estuary had the net effect of reducing the concentration and enrichment of these metals within the Kariega Estuary due to scouring and outflow of estuarine water and sediment into the marine environment. The temporal variations in metal concentrations and enrichment factors were less pronounced in the temporary open-closed estuaries than the permanently open Kariega Estuary. The observed trend can probably be related to the low anthropogenic impact within the catchment areas of these systems, and the relatively smaller size of the catchments. Significant spatial variations existed in metal enrichment in the sediment of both the East Kleinemonde and Riet estuaries, with the highest degrees of enrichment occurring in the sediments from the marine environment and lower reaches.
- Full Text:
- Date Issued: 2008
The role of the euthecosome pteropod, limacina retroversa, in the polar frontal zone, Southern Ocean
- Authors: Bernard, Kim Sarah
- Date: 2007
- Subjects: Pteropoda Limacidae Zooplankton -- Antarctic Ocean Copepoda Phytoplankton
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5761 , http://hdl.handle.net/10962/d1005449
- Description: The aim of the present study was to assess the ecological role of the euthecosome pteropod, Limacina retroversa, in particular, and the mesozooplankton community, in general, in the pelagic ecosystem of the Polar Frontal Zone (PFZ), Southern Ocean. Data were collected from four oceanographic surveys to the Indian sector of the PFZ during austral autumn 2000, 2002, 2004 and 2005. Copepods, mainly Calanus simillimus, Oithona similis, Clausocalanus spp. and Ctenocalanus spp., typically dominated total mesozooplankton counts, accounting for, on average, between 75.5 % and 88.1 % (Mean = 77.4 %; SD = 13.4 %) of the total, during the present investigation. Results of the study indicate that L. retroversa may, at times, contribute substantially to total mesozooplankton abundances. During the study, L. retroversa contributed between 0.0 and 30.0 % (Mean = 5.3 %; SD = 7.1 %) to total mesozooplankton numbers. Significant small-scale variability in abundance and size structure of L. retroversa and abundance of copepods was minimal. Inter-annual variability, on the other hand, was significant between some years. Total pteropod numbers were greatest during April 2002 and 2004, while copepods exhibited greatest abundances during April 2004 only. Pearson’s Correlation analysis suggested that L. retroversa abundances were positively correlated to total surface chlorophyll-a (chl-a) concentrations. The significantly lower chl-a concentrations recorded during April 2005 may explain the reduced pteropod numbers observed during that survey. The size class structure of L. retroversa comprised mainly small and mediumsized individuals during all four surveys. This corresponds well with records from the northern hemisphere (sub-Arctic and Arctic waters) where Limacina spp. are reported to exhibit maximum spawning during mid to late-summer. Higher abundances of large individuals only occurred during April 2005, when chl-a concentrations were very low; possibly the result of delayed spawning, due to reduced food availability. Ingestion rates of the four most abundant copepods, determined using the gut fluorescence technique, ranged between 159.32 ng (pigm) ind⁻¹ day⁻¹ and 728.36 ng (pigm) ind⁻¹ day⁻¹ (Mean = 321.01 ng (pigm) ind⁻¹ day⁻¹; SD = 173.91 ng (pigm) ind⁻¹ day). Ingestion rates of L. retroversa were much higher, ranging from an average of 4 28.68 ng (pigm) ind⁻¹ day⁻¹ in April 2002 to 4 196.88 ng (pigm) ind⁻¹day⁻¹in April 2005 (Mean = 4157.36 ng (pigm) ind⁻¹ day⁻¹; SD = 35.37 ng (pigm) ind⁻¹day⁻¹). Average daily grazing rates for the pteropod varied between 0.39 mg (pigm) m⁻² day⁻¹ in April 2005 and 17.69 mg (pigm) m-2 day-1 in April 2004 (Mean = 6.13 mg (pigm) m⁻² day⁻¹; SD = 11.04 mg (pigm) m⁻² day⁻¹); corresponding average daily grazing impacts ranged between 8.4 % and 139.8 % of the phytoplankton standing stock in April 2005 and 2004, respectively (Mean = 48.5 %; SD = 84.5 %). Average daily grazing rates of the four copepods ranged from 4.58 mg (pigm) m⁻² day⁻¹ to 8.77 mg (pigm) m⁻² day⁻¹ -1, during April 2002 and 2004, respectively (Mean = 6.28 mg (pigm) m⁻² day⁻¹; SD = 5.94 mg (pigm) m⁻² day⁻¹). Collectively, the copepods removed an average of between 31.6 % and 89.8 % of the phytoplankton standing stock per day, during April 2002 and 2004, respectively (Mean = 70.8 %; SD = 86.7 %). The daily grazing impact of the copepods accounted for an average of between 40.4 % and 87.8 % of the total zooplankton grazing impact, during April 2004 and 2005, respectively (Mean = 75.0 %; SD = 65.5 %). L. retroversa was responsible for an average of 52.4 % and 59.5 % of the total zooplankton grazing impact, during April 2002 and 2004, respectively. However, during April 2005, when L. retroversa numbers were significantly lower than previous years, the pteropod contributed an average of only 7.5 % to the total zooplankton grazing impact. Thus, during the present investigation,the pteropod was responsible for removing a mean of 48.9 % of the available phytoplankton (SD = 74.9 %). The predation impact of the dominant carnivorous macrozooplankton and micronekton in the PFZ was determined during April 2004 and 2005 using daily ration estimates obtained from the literature. Additionally, gut content analysis was used to determine the contribution of L. retroversa to the diet of the dominant predators. Average predation impact ranged from 1.1 % and 5.7 % of the total mesozooplankton standing stock during April 2004 and 2005, respectively (Mean = 3.8 %; SD = 12.3 %). Chaetognaths and euphausiids dominated total carnivore numbers and made the greatest contributions to total predation impact during both years. Copepods appeared to be the main prey item of the dominant carnivorous macrozooplankton-micronekton in the region. L. retroversa was only detected in the gut contents of the amphipod, Themisto gaudichaudi, but not in either of the chaetognath species (Eukrohnia hamata and Sagitta gazellae) or the myctophid fish (Electrona spp.). The pteropod was found in 19 % of amphipod guts dissected. Pearson’s Correlation analyses showed that the four major predatory zooplankton groups found in the PFZ (chaetognaths, euphausiids, amphipods and myctophid fish) were positively correlated to abundances of L. retroversa, suggesting that the pteropod might be an important prey item for many of the carnivorous macrozooplankton/micronekton in the PFZ. To conclude, L. retroversa may play an important role in the pelagic ecosystem of the PFZ, in austral autumn. However, ocean acidification and calcium carbonate undersaturation (as a result of increased anthropogenic carbon dioxide emissions), that is predicted to occur within the next 50 – 100 years, will most likely have significant implications for the Sub-Antarctic pelagic ecosystem if L. retroversa cannot adapt quickly enough to the changes.
- Full Text:
- Date Issued: 2007
The role of the euthecosome pteropod, limacina retroversa, in the polar frontal zone, Southern Ocean
- Authors: Bernard, Kim Sarah
- Date: 2007
- Subjects: Pteropoda Limacidae Zooplankton -- Antarctic Ocean Copepoda Phytoplankton
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5761 , http://hdl.handle.net/10962/d1005449
- Description: The aim of the present study was to assess the ecological role of the euthecosome pteropod, Limacina retroversa, in particular, and the mesozooplankton community, in general, in the pelagic ecosystem of the Polar Frontal Zone (PFZ), Southern Ocean. Data were collected from four oceanographic surveys to the Indian sector of the PFZ during austral autumn 2000, 2002, 2004 and 2005. Copepods, mainly Calanus simillimus, Oithona similis, Clausocalanus spp. and Ctenocalanus spp., typically dominated total mesozooplankton counts, accounting for, on average, between 75.5 % and 88.1 % (Mean = 77.4 %; SD = 13.4 %) of the total, during the present investigation. Results of the study indicate that L. retroversa may, at times, contribute substantially to total mesozooplankton abundances. During the study, L. retroversa contributed between 0.0 and 30.0 % (Mean = 5.3 %; SD = 7.1 %) to total mesozooplankton numbers. Significant small-scale variability in abundance and size structure of L. retroversa and abundance of copepods was minimal. Inter-annual variability, on the other hand, was significant between some years. Total pteropod numbers were greatest during April 2002 and 2004, while copepods exhibited greatest abundances during April 2004 only. Pearson’s Correlation analysis suggested that L. retroversa abundances were positively correlated to total surface chlorophyll-a (chl-a) concentrations. The significantly lower chl-a concentrations recorded during April 2005 may explain the reduced pteropod numbers observed during that survey. The size class structure of L. retroversa comprised mainly small and mediumsized individuals during all four surveys. This corresponds well with records from the northern hemisphere (sub-Arctic and Arctic waters) where Limacina spp. are reported to exhibit maximum spawning during mid to late-summer. Higher abundances of large individuals only occurred during April 2005, when chl-a concentrations were very low; possibly the result of delayed spawning, due to reduced food availability. Ingestion rates of the four most abundant copepods, determined using the gut fluorescence technique, ranged between 159.32 ng (pigm) ind⁻¹ day⁻¹ and 728.36 ng (pigm) ind⁻¹ day⁻¹ (Mean = 321.01 ng (pigm) ind⁻¹ day⁻¹; SD = 173.91 ng (pigm) ind⁻¹ day). Ingestion rates of L. retroversa were much higher, ranging from an average of 4 28.68 ng (pigm) ind⁻¹ day⁻¹ in April 2002 to 4 196.88 ng (pigm) ind⁻¹day⁻¹in April 2005 (Mean = 4157.36 ng (pigm) ind⁻¹ day⁻¹; SD = 35.37 ng (pigm) ind⁻¹day⁻¹). Average daily grazing rates for the pteropod varied between 0.39 mg (pigm) m⁻² day⁻¹ in April 2005 and 17.69 mg (pigm) m-2 day-1 in April 2004 (Mean = 6.13 mg (pigm) m⁻² day⁻¹; SD = 11.04 mg (pigm) m⁻² day⁻¹); corresponding average daily grazing impacts ranged between 8.4 % and 139.8 % of the phytoplankton standing stock in April 2005 and 2004, respectively (Mean = 48.5 %; SD = 84.5 %). Average daily grazing rates of the four copepods ranged from 4.58 mg (pigm) m⁻² day⁻¹ to 8.77 mg (pigm) m⁻² day⁻¹ -1, during April 2002 and 2004, respectively (Mean = 6.28 mg (pigm) m⁻² day⁻¹; SD = 5.94 mg (pigm) m⁻² day⁻¹). Collectively, the copepods removed an average of between 31.6 % and 89.8 % of the phytoplankton standing stock per day, during April 2002 and 2004, respectively (Mean = 70.8 %; SD = 86.7 %). The daily grazing impact of the copepods accounted for an average of between 40.4 % and 87.8 % of the total zooplankton grazing impact, during April 2004 and 2005, respectively (Mean = 75.0 %; SD = 65.5 %). L. retroversa was responsible for an average of 52.4 % and 59.5 % of the total zooplankton grazing impact, during April 2002 and 2004, respectively. However, during April 2005, when L. retroversa numbers were significantly lower than previous years, the pteropod contributed an average of only 7.5 % to the total zooplankton grazing impact. Thus, during the present investigation,the pteropod was responsible for removing a mean of 48.9 % of the available phytoplankton (SD = 74.9 %). The predation impact of the dominant carnivorous macrozooplankton and micronekton in the PFZ was determined during April 2004 and 2005 using daily ration estimates obtained from the literature. Additionally, gut content analysis was used to determine the contribution of L. retroversa to the diet of the dominant predators. Average predation impact ranged from 1.1 % and 5.7 % of the total mesozooplankton standing stock during April 2004 and 2005, respectively (Mean = 3.8 %; SD = 12.3 %). Chaetognaths and euphausiids dominated total carnivore numbers and made the greatest contributions to total predation impact during both years. Copepods appeared to be the main prey item of the dominant carnivorous macrozooplankton-micronekton in the region. L. retroversa was only detected in the gut contents of the amphipod, Themisto gaudichaudi, but not in either of the chaetognath species (Eukrohnia hamata and Sagitta gazellae) or the myctophid fish (Electrona spp.). The pteropod was found in 19 % of amphipod guts dissected. Pearson’s Correlation analyses showed that the four major predatory zooplankton groups found in the PFZ (chaetognaths, euphausiids, amphipods and myctophid fish) were positively correlated to abundances of L. retroversa, suggesting that the pteropod might be an important prey item for many of the carnivorous macrozooplankton/micronekton in the PFZ. To conclude, L. retroversa may play an important role in the pelagic ecosystem of the PFZ, in austral autumn. However, ocean acidification and calcium carbonate undersaturation (as a result of increased anthropogenic carbon dioxide emissions), that is predicted to occur within the next 50 – 100 years, will most likely have significant implications for the Sub-Antarctic pelagic ecosystem if L. retroversa cannot adapt quickly enough to the changes.
- Full Text:
- Date Issued: 2007
Growth, feeding and sex change in the sequential protandric shrimp Nauticaris marionis Bate 1888 at the Prince Edward Islands (Southern Ocean)
- Authors: Vumazonke, Lukhanyiso Unam
- Date: 2005
- Subjects: Shrimps -- Prince Edward Islands , Shrimps -- Feeding and feeds , Shrimps -- Nutrition , Shrimps -- Growth
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5704 , http://hdl.handle.net/10962/d1005390 , Shrimps -- Prince Edward Islands , Shrimps -- Feeding and feeds , Shrimps -- Nutrition , Shrimps -- Growth
- Description: Demographic parameters and the general biology of the subantarctic shrimp Nauticaris marionis from the Prince Edward Islands were investigated. The carapace length is the most accurate indicator of body size and it was confirmed that N. marionis is a partially protandric hermaphrodite. Gravid females of N. marionis observed mainly in March with negligible hatching persisting until April/May. The majority of juveniles develop into males. Juveniles are characterised by an appendage on the endopodite of the first pleopod called the appendix interna or a.i.1. Juveniles that develop into males do so by growing a further appendage on the endopodite of the second pleopod. This appendage characterises males and is called the appendix masculina (or a.m.). Juveniles may develop directly into primary females through the development of an ovary and the loss of the a.i.1. The sequence differs among individuals, with some developing the ovary before losing the a.i.1 and others losing the a.i.1 first. Loss of the a.i.1 appears to be by shedding during a single moult, rather than by atrophy. Females can also develop by an alternative route as secondary females. Such animals first become males with an a.m. and then develop an ovary, thus forming an intermediate form called a tertium quid, or “third thing”. Again, there are two forms of tertium quid. Tertia quae a have the a.m. as well as the a.i.1. Tertia quae b have the a.m. in combination with an ovary but no a.i.1. Either form of tertium quid can develop into a secondary female through loss of the a.m., and in the case of the tertium quid a, loss of the a.i.1. It is unclear whether sexual differentiation occurs in the plankton or just after N. marionis settles on the benthos. The results of gut content analysis suggest that N. marionis is an opportunistic feeder, preying on a variety of prey with a preference for detritus, benthic amphipods and gastropods. Cannibalism of conspecific pleopods by large individuals of N. marionis occurred mostly in females, particularly in incubation containers. Cannibalism also occurs in males, and its occurrence depends on individual size, not gender. No diel patterns were observed in the feeding activity of N. marionis. In situ daily rations of males (carapace length <7mm) and females (>7mm) were equivalent to ≈10% and ≈5% of body dry weight, respectively. The von Bertalanffy growth curve parameters were empirically identified, by cohort analysis of data collected during 4 years, as K = 0.22239/year, L[subscript]∞ = 14.05789mm, t₀ = -0.05174, L₀ = 0.16083mm. N. marionis can survive up to seven years under natural conditions.
- Full Text:
- Date Issued: 2005
- Authors: Vumazonke, Lukhanyiso Unam
- Date: 2005
- Subjects: Shrimps -- Prince Edward Islands , Shrimps -- Feeding and feeds , Shrimps -- Nutrition , Shrimps -- Growth
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5704 , http://hdl.handle.net/10962/d1005390 , Shrimps -- Prince Edward Islands , Shrimps -- Feeding and feeds , Shrimps -- Nutrition , Shrimps -- Growth
- Description: Demographic parameters and the general biology of the subantarctic shrimp Nauticaris marionis from the Prince Edward Islands were investigated. The carapace length is the most accurate indicator of body size and it was confirmed that N. marionis is a partially protandric hermaphrodite. Gravid females of N. marionis observed mainly in March with negligible hatching persisting until April/May. The majority of juveniles develop into males. Juveniles are characterised by an appendage on the endopodite of the first pleopod called the appendix interna or a.i.1. Juveniles that develop into males do so by growing a further appendage on the endopodite of the second pleopod. This appendage characterises males and is called the appendix masculina (or a.m.). Juveniles may develop directly into primary females through the development of an ovary and the loss of the a.i.1. The sequence differs among individuals, with some developing the ovary before losing the a.i.1 and others losing the a.i.1 first. Loss of the a.i.1 appears to be by shedding during a single moult, rather than by atrophy. Females can also develop by an alternative route as secondary females. Such animals first become males with an a.m. and then develop an ovary, thus forming an intermediate form called a tertium quid, or “third thing”. Again, there are two forms of tertium quid. Tertia quae a have the a.m. as well as the a.i.1. Tertia quae b have the a.m. in combination with an ovary but no a.i.1. Either form of tertium quid can develop into a secondary female through loss of the a.m., and in the case of the tertium quid a, loss of the a.i.1. It is unclear whether sexual differentiation occurs in the plankton or just after N. marionis settles on the benthos. The results of gut content analysis suggest that N. marionis is an opportunistic feeder, preying on a variety of prey with a preference for detritus, benthic amphipods and gastropods. Cannibalism of conspecific pleopods by large individuals of N. marionis occurred mostly in females, particularly in incubation containers. Cannibalism also occurs in males, and its occurrence depends on individual size, not gender. No diel patterns were observed in the feeding activity of N. marionis. In situ daily rations of males (carapace length <7mm) and females (>7mm) were equivalent to ≈10% and ≈5% of body dry weight, respectively. The von Bertalanffy growth curve parameters were empirically identified, by cohort analysis of data collected during 4 years, as K = 0.22239/year, L[subscript]∞ = 14.05789mm, t₀ = -0.05174, L₀ = 0.16083mm. N. marionis can survive up to seven years under natural conditions.
- Full Text:
- Date Issued: 2005
Population dynamics of selected ichthyofaunal components in the temperate, temporarily open/closed Kasouga Estuary, South Africa
- Authors: Tweddle, Gavin Paul
- Date: 2005
- Subjects: Estuaries -- South Africa -- Eastern Cape , Estuarine fishes -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5742 , http://hdl.handle.net/10962/d1005428 , Estuaries -- South Africa -- Eastern Cape , Estuarine fishes -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape
- Description: The spatial and temporal pattern of ichthyofaunal community composition in relation to selected physico-chemical (temperature and salinity) and biological variables (chlorophyll-a and zooplankton) was investigated at ten stations in the temperate temporarily open/closed Kasouga estuary. In addition, the food web structure in the estuary was investigated using stable carbon isotope analysis. Results of the 5 metre seine net survey indicated that ichthyofaunal composition and biomass in the Kasouga estuary was largely determined by seasonality and mouth condition. Maximum abundance and biomass of ichthyofauna was recorded during summer or during those periods when overtopping occurred. Overtopping coincided with the recruitment of marine estuarine dependant species, which dominated the catches both numerically and in biomass. The recruitment of these species resulted in an increase in diversity of the ichthyofaunal community. There were no significant spatial patterns in the distribution of smaller ichthyofauna (<50mm SL) identified in Bray-Curtis similarity matrices using cluster analysis (Primer 5 v5.2.4). Spearman’s correlation analysis revealed that there were no significant correlations between abundance and biomass of ichthyofauna and selected physico-chemical and biological variables other than salinity (P>0.05 in all other cases). There were two distinct spatial patterns in the distribution of the larger ichthyofauna (>50mm SL). These corresponded to a grouping associated with the mouth region and a grouping associated with the remaining regions of the estuary. Stable isotope analysis indicated that the primary source of carbon utilised by the ichthyofauna of the Kasouga estuary was derived from the channel, most likely microphytobenthic algae. The contributions of the riparian and salt marsh vegetation to the total carbon flow appear to be minimal.
- Full Text:
- Date Issued: 2005
- Authors: Tweddle, Gavin Paul
- Date: 2005
- Subjects: Estuaries -- South Africa -- Eastern Cape , Estuarine fishes -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5742 , http://hdl.handle.net/10962/d1005428 , Estuaries -- South Africa -- Eastern Cape , Estuarine fishes -- South Africa -- Eastern Cape , Estuarine ecology -- South Africa -- Eastern Cape
- Description: The spatial and temporal pattern of ichthyofaunal community composition in relation to selected physico-chemical (temperature and salinity) and biological variables (chlorophyll-a and zooplankton) was investigated at ten stations in the temperate temporarily open/closed Kasouga estuary. In addition, the food web structure in the estuary was investigated using stable carbon isotope analysis. Results of the 5 metre seine net survey indicated that ichthyofaunal composition and biomass in the Kasouga estuary was largely determined by seasonality and mouth condition. Maximum abundance and biomass of ichthyofauna was recorded during summer or during those periods when overtopping occurred. Overtopping coincided with the recruitment of marine estuarine dependant species, which dominated the catches both numerically and in biomass. The recruitment of these species resulted in an increase in diversity of the ichthyofaunal community. There were no significant spatial patterns in the distribution of smaller ichthyofauna (<50mm SL) identified in Bray-Curtis similarity matrices using cluster analysis (Primer 5 v5.2.4). Spearman’s correlation analysis revealed that there were no significant correlations between abundance and biomass of ichthyofauna and selected physico-chemical and biological variables other than salinity (P>0.05 in all other cases). There were two distinct spatial patterns in the distribution of the larger ichthyofauna (>50mm SL). These corresponded to a grouping associated with the mouth region and a grouping associated with the remaining regions of the estuary. Stable isotope analysis indicated that the primary source of carbon utilised by the ichthyofauna of the Kasouga estuary was derived from the channel, most likely microphytobenthic algae. The contributions of the riparian and salt marsh vegetation to the total carbon flow appear to be minimal.
- Full Text:
- Date Issued: 2005
Population structure, physiology and feeding ecology of the mysid Mesopodopsis wooldridgei (Whittmann) in a large permanently open estuary
- Authors: Mesher, Colette Suzanne
- Date: 2005
- Subjects: Mysidacea -- Physiology , Mysidacea -- Ecology , Mysidacea -- Feeding and feeds , Zooplankton -- South Africa -- Great Fish Estuary
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5746 , http://hdl.handle.net/10962/d1005432 , Mysidacea -- Physiology , Mysidacea -- Ecology , Mysidacea -- Feeding and feeds , Zooplankton -- South Africa -- Great Fish Estuary
- Description: This project had three main aims. The first aim was to investigate the metabolic physiology of the mysid Mesopodopsis wooldridgei. Secondly, the study examined the contribution of the mysid to the zooplankton community. Finally the study attempted to clarify the primary sources of carbon utilised by the mysid using stable carbon isotope analysis. The study was conducted in the freshwater dominated permanently open Great Fish Estuary (33°30’S; 27°08’E) located along the south-eastern coastline of southern Africa. Mass specific oxygen consumption rates of Mesopodopsis wooldridgei ranged between 0.11 and 8.38 ml O2 mg ww-1 h-1 and increased with an increase in water temperature. There were no significant differences in the respiration rates between the male and female mysids (P > 0.05). At a constant temperature (20°C), the oxygen consumption rate of M. wooldridgei decreased with an increase in salinity. Salinity tolerance of the mysid was strongly modified by temperature. At 5‰ and 30°C, mysids exhibited mass mortality. The Q10 value for M. wooldridgei at 15, 25 and 35‰ was estimated at 2.34, 1.44 and 2.14, respectively. Results of the study suggest that M. wooldridgei is well adapted to surviving in environments characterised by variations in temperature and salinity. Within the Great Fish Estuary, total chl. a concentration ranged between 2.68 μg L-1 and 31.12 μg L-1 and was always dominated by large phytoplankton cells (>5 μm). Average zooplankton abundance ranged between 62 and 28 917 ind. m-3 and biomass between 10 and 203 mg Dwt m-3. The zooplankton community was numerically dominated by the calanoid copepod Pseudodiaptomus hessei, which comprised up to 100% (range between 12 and 100%) of the total zooplankton counted. Total zooplankton biomass during the day was dominated by copepods and by mysids during the nighttime. Among the mysids, M. wooldridgei was the most numerically abundant mysid and comprised <10% of the total zooplankton abundance. Numerical analysis found no seasonal pattern in the total zooplankton abundance (including mysids). The lack of any seasonality could be attributed to the continuous freshwater inflow into the estuary due to the inter-basin transfer of water from the Gariep Dam to the Fish River system. A distinct spatial pattern in the zooplankton community structure was evident with the upper stations almost entirely dominated by the copepod P. hessei, while at stations occupied in the lower reaches of the estuary, the zooplankton community comprised a mixture of freshwater, estuarine and marine breeding zooplankton species. The mysids also demonstrated a distinct spatial pattern in their distribution. Mysids were generally absent from the upper reaches, while in the middle reaches Rhopalophthalmus terranatalis and M. wooldridgei were numerically dominant. Gastrosaccus brevifissura dominated in the lower regions of the estuary. Stable isotope analysis (δ13C) indicated that the dominant source of carbon utilised by the numerically dominant copepods and juvenile M. wooldridgei within the estuary was derived from the extensive phytoplankton stocks within the system. In contrast, sub-adult and adult mysids (R. terranatalis, M. wooldridgei and G. brevifissura) appeared to consume a combination of phytoplankton and copepods. The contribution of the various sources of carbon to the total carbon intake of the mysid remains unknown.
- Full Text:
- Date Issued: 2005
- Authors: Mesher, Colette Suzanne
- Date: 2005
- Subjects: Mysidacea -- Physiology , Mysidacea -- Ecology , Mysidacea -- Feeding and feeds , Zooplankton -- South Africa -- Great Fish Estuary
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5746 , http://hdl.handle.net/10962/d1005432 , Mysidacea -- Physiology , Mysidacea -- Ecology , Mysidacea -- Feeding and feeds , Zooplankton -- South Africa -- Great Fish Estuary
- Description: This project had three main aims. The first aim was to investigate the metabolic physiology of the mysid Mesopodopsis wooldridgei. Secondly, the study examined the contribution of the mysid to the zooplankton community. Finally the study attempted to clarify the primary sources of carbon utilised by the mysid using stable carbon isotope analysis. The study was conducted in the freshwater dominated permanently open Great Fish Estuary (33°30’S; 27°08’E) located along the south-eastern coastline of southern Africa. Mass specific oxygen consumption rates of Mesopodopsis wooldridgei ranged between 0.11 and 8.38 ml O2 mg ww-1 h-1 and increased with an increase in water temperature. There were no significant differences in the respiration rates between the male and female mysids (P > 0.05). At a constant temperature (20°C), the oxygen consumption rate of M. wooldridgei decreased with an increase in salinity. Salinity tolerance of the mysid was strongly modified by temperature. At 5‰ and 30°C, mysids exhibited mass mortality. The Q10 value for M. wooldridgei at 15, 25 and 35‰ was estimated at 2.34, 1.44 and 2.14, respectively. Results of the study suggest that M. wooldridgei is well adapted to surviving in environments characterised by variations in temperature and salinity. Within the Great Fish Estuary, total chl. a concentration ranged between 2.68 μg L-1 and 31.12 μg L-1 and was always dominated by large phytoplankton cells (>5 μm). Average zooplankton abundance ranged between 62 and 28 917 ind. m-3 and biomass between 10 and 203 mg Dwt m-3. The zooplankton community was numerically dominated by the calanoid copepod Pseudodiaptomus hessei, which comprised up to 100% (range between 12 and 100%) of the total zooplankton counted. Total zooplankton biomass during the day was dominated by copepods and by mysids during the nighttime. Among the mysids, M. wooldridgei was the most numerically abundant mysid and comprised <10% of the total zooplankton abundance. Numerical analysis found no seasonal pattern in the total zooplankton abundance (including mysids). The lack of any seasonality could be attributed to the continuous freshwater inflow into the estuary due to the inter-basin transfer of water from the Gariep Dam to the Fish River system. A distinct spatial pattern in the zooplankton community structure was evident with the upper stations almost entirely dominated by the copepod P. hessei, while at stations occupied in the lower reaches of the estuary, the zooplankton community comprised a mixture of freshwater, estuarine and marine breeding zooplankton species. The mysids also demonstrated a distinct spatial pattern in their distribution. Mysids were generally absent from the upper reaches, while in the middle reaches Rhopalophthalmus terranatalis and M. wooldridgei were numerically dominant. Gastrosaccus brevifissura dominated in the lower regions of the estuary. Stable isotope analysis (δ13C) indicated that the dominant source of carbon utilised by the numerically dominant copepods and juvenile M. wooldridgei within the estuary was derived from the extensive phytoplankton stocks within the system. In contrast, sub-adult and adult mysids (R. terranatalis, M. wooldridgei and G. brevifissura) appeared to consume a combination of phytoplankton and copepods. The contribution of the various sources of carbon to the total carbon intake of the mysid remains unknown.
- Full Text:
- Date Issued: 2005
Size-fractionated phytoplankton biomass and primary production in the Southern Ocean
- Authors: Balarin, Marianne G
- Date: 1999
- Subjects: Biomass -- Analysis , Phytoplankton , Atlantic Ocean
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5765 , http://hdl.handle.net/10962/d1005453 , Biomass -- Analysis , Phytoplankton , Atlantic Ocean
- Description: The factors controlling primary production in the Southern Ocean were investigated over two years during two cruises of the South African National Antarctic Program (SANAP). The first cruise was conducted to the region of the eastern Atlantic sector of the Southern Ocean during the collaborative Scandinavian/South African Antarctic expedition conducted in austral summer (DecemberIFebruary) 1997-1998. Production studies were conducted in the vicinity of the Marginal Ice Zone (MIZ), Interfrontal Zone (IFZ) and Antarctic Polar Front(APF). The second cruise was conducted during the Third Marion Island Oceanographic Survey (MIOS III) to the region of the Sub-Antarctic Prince Edward Islands in austral autumn (April/May) 1998. Size-fractionated production rates were estimated by 14C incorporation using standard JGOFS protocols. Oceanographic data from the first cruise suggest that the three regions can be divided into two distinct regimes. Stations occupied in the vicinity of the MIZ and the APF were characterised by a shallow mixed layer depth « 40m) while at the IFZ-stations, the mixed layer depth exceeded the 1% light depth. Microphytoplankton dominated integrated chlorophyll-a biomass in the MIZ (total chlorophyll a ranged between 15.4 and 41.3 mg Chi-a. m-2) and at the APF (range between 10.7 and 31.4 mg Chi-a. m-2) , comprising > 50% of total chlorophyll-a at all these stations. Xl Within the IFZ (2 stations), nanophytoplankton dominated total integrated Chl-a biomass (range between 5.6 and 8.8 mg Chi-a. m-2) comprising, on average, 36% of the total. Picophytoplankton comprised an average of 12% of the total Chl-a biomass (range between 3.1 and 5.9 mg Chi-a. m-2) in the MIZ, 36% in the IFZ (range between 6.4 and 7.8 mg Chl-a . m-2) and 20% in the vicinity of the APF (range between 6.8 and 10.6 mg Chi-a. m-2). Total integrated primary production ranged between 316 and 729 mg C . m-2. d-1 at stations occupied in the vicinity of the MIZ, and between 292 and 317 mg C . m·2• d-l within the IFZ. At stations occupied in the region of the APF, total integrated production ranged between 708 and 926 mg C . m-2• dol. The contribution of various size fractions to total productivity generally displayed the same pattern as integrated Chl-a biomass. Microphytoplankton formed the most important contributor to total production at stations occupied in the MIZ and at the APF. Within the IFZ, nanophytoplankton dominated total daily production. Nutrient data suggest that concentrations of macro nutrients within the upper water column were above the threshold where growth would be limited. Preliminary results showed that concentrations of iron (Fe) were highest in the southern region of the MIZ and in the vicinity of the APF. During the second cruise, conducted in the vicinity of the Sub-Antarctic Front (SAF) and in the upstream, inter-island and downstream regions of the Prince Edward Islands, there was evidence of fresh water run-off from the islands, (i.e. decreased salinities and increased concentrations of ammonia and nitrate). Oceanographic data collected at the various production stations indicated that the upper water column was well mixed throughout the survey. Total integrated biomass during the study ranged between 8.5 and 20.1 mg Chi-a. m-2• No distinct patterns in total Chl-a biomass were evident. Picophytoplankton dominated total biomass comprising> 45 % of total pigment at all stations. Nanophytoplankton were the second most important contributor to total integrated biomass. Generally xu microphytoplankton contributed < 10 % of total ChI-a. Total daily integrated production was highest (442.6 mg Chi-a. m-2) at the single station occupied in the vicinity of the SAF. Outside this region, total areal production was lower, ranging from 94.5 to 353.0 mg C . m-2. d-1. With the exception of the station occupied in the vicinity of the SAF, total productivity was dominated by nanophytoplankton, which comprised between 48 and 66% of the total. Concentrations of macronutrients did not appear to be limiting to phytoplankton growth. The absence of a phytoplankton bloom in the vicinity of the islands appears to have been related to water column stability, which was influenced by the prevailing oceanographic regime during the survey. Previous studies have shown that when the SAF lies in close proximity to the islands, advecting forces prevail, resulting in the islands functioning as a flow-through system. During this study, the SAF lay immediately north of the islands. As a consequence no water was trapped in the leeward side of the islands. The results of the two cruises suggest that phytoplankton production in the four systems investigated: the Marginal Ice Zone (MIZ), Antarctic Polar Front (APF), Inter Frontal Zone (IFZ) and Prince Edward Islands (PEl), was largely controlled by water column stability. It is probable that the availability of iron, particularly in the region of the MIZ and APF, may have further contributed to the elevated production recorded in these two regions.
- Full Text:
- Date Issued: 1999
- Authors: Balarin, Marianne G
- Date: 1999
- Subjects: Biomass -- Analysis , Phytoplankton , Atlantic Ocean
- Language: English
- Type: Thesis , Masters , MSc
- Identifier: vital:5765 , http://hdl.handle.net/10962/d1005453 , Biomass -- Analysis , Phytoplankton , Atlantic Ocean
- Description: The factors controlling primary production in the Southern Ocean were investigated over two years during two cruises of the South African National Antarctic Program (SANAP). The first cruise was conducted to the region of the eastern Atlantic sector of the Southern Ocean during the collaborative Scandinavian/South African Antarctic expedition conducted in austral summer (DecemberIFebruary) 1997-1998. Production studies were conducted in the vicinity of the Marginal Ice Zone (MIZ), Interfrontal Zone (IFZ) and Antarctic Polar Front(APF). The second cruise was conducted during the Third Marion Island Oceanographic Survey (MIOS III) to the region of the Sub-Antarctic Prince Edward Islands in austral autumn (April/May) 1998. Size-fractionated production rates were estimated by 14C incorporation using standard JGOFS protocols. Oceanographic data from the first cruise suggest that the three regions can be divided into two distinct regimes. Stations occupied in the vicinity of the MIZ and the APF were characterised by a shallow mixed layer depth « 40m) while at the IFZ-stations, the mixed layer depth exceeded the 1% light depth. Microphytoplankton dominated integrated chlorophyll-a biomass in the MIZ (total chlorophyll a ranged between 15.4 and 41.3 mg Chi-a. m-2) and at the APF (range between 10.7 and 31.4 mg Chi-a. m-2) , comprising > 50% of total chlorophyll-a at all these stations. Xl Within the IFZ (2 stations), nanophytoplankton dominated total integrated Chl-a biomass (range between 5.6 and 8.8 mg Chi-a. m-2) comprising, on average, 36% of the total. Picophytoplankton comprised an average of 12% of the total Chl-a biomass (range between 3.1 and 5.9 mg Chi-a. m-2) in the MIZ, 36% in the IFZ (range between 6.4 and 7.8 mg Chl-a . m-2) and 20% in the vicinity of the APF (range between 6.8 and 10.6 mg Chi-a. m-2). Total integrated primary production ranged between 316 and 729 mg C . m-2. d-1 at stations occupied in the vicinity of the MIZ, and between 292 and 317 mg C . m·2• d-l within the IFZ. At stations occupied in the region of the APF, total integrated production ranged between 708 and 926 mg C . m-2• dol. The contribution of various size fractions to total productivity generally displayed the same pattern as integrated Chl-a biomass. Microphytoplankton formed the most important contributor to total production at stations occupied in the MIZ and at the APF. Within the IFZ, nanophytoplankton dominated total daily production. Nutrient data suggest that concentrations of macro nutrients within the upper water column were above the threshold where growth would be limited. Preliminary results showed that concentrations of iron (Fe) were highest in the southern region of the MIZ and in the vicinity of the APF. During the second cruise, conducted in the vicinity of the Sub-Antarctic Front (SAF) and in the upstream, inter-island and downstream regions of the Prince Edward Islands, there was evidence of fresh water run-off from the islands, (i.e. decreased salinities and increased concentrations of ammonia and nitrate). Oceanographic data collected at the various production stations indicated that the upper water column was well mixed throughout the survey. Total integrated biomass during the study ranged between 8.5 and 20.1 mg Chi-a. m-2• No distinct patterns in total Chl-a biomass were evident. Picophytoplankton dominated total biomass comprising> 45 % of total pigment at all stations. Nanophytoplankton were the second most important contributor to total integrated biomass. Generally xu microphytoplankton contributed < 10 % of total ChI-a. Total daily integrated production was highest (442.6 mg Chi-a. m-2) at the single station occupied in the vicinity of the SAF. Outside this region, total areal production was lower, ranging from 94.5 to 353.0 mg C . m-2. d-1. With the exception of the station occupied in the vicinity of the SAF, total productivity was dominated by nanophytoplankton, which comprised between 48 and 66% of the total. Concentrations of macronutrients did not appear to be limiting to phytoplankton growth. The absence of a phytoplankton bloom in the vicinity of the islands appears to have been related to water column stability, which was influenced by the prevailing oceanographic regime during the survey. Previous studies have shown that when the SAF lies in close proximity to the islands, advecting forces prevail, resulting in the islands functioning as a flow-through system. During this study, the SAF lay immediately north of the islands. As a consequence no water was trapped in the leeward side of the islands. The results of the two cruises suggest that phytoplankton production in the four systems investigated: the Marginal Ice Zone (MIZ), Antarctic Polar Front (APF), Inter Frontal Zone (IFZ) and Prince Edward Islands (PEl), was largely controlled by water column stability. It is probable that the availability of iron, particularly in the region of the MIZ and APF, may have further contributed to the elevated production recorded in these two regions.
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- Date Issued: 1999
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