An ecological reserve study by the Department of Water Affairs on the Great Brak Estuary stated that there was a need to determine how much nitrogen and phosphorus was flowing through the estuary as well as how effective the macroalgae were at removing N and P. The objective of this study was to investigate the physico-chemical characteristics in the estuary and the influence of these on the submerged macrophytes and macroalgae. A nutrient budget for the estuary was developed in order to quantify the contribution of the submerged macrophytes and macroalgae relative to other contributing sources. The Wolwedans Dam located 3 km upstream from the estuary has reduced the amount of freshwater flow to the estuary by as much as 56 percent. The estuary has been allocated 2 x 106 m3 per annum of freshwater (ecological reserve) that is used to breach the mouth once or twice a year in spring or summer. Even though this water has been made available it is not sufficient to flush the estuary. Reduced flushing has led to an accumulation of organic matter and degradation in the water quality. Physico-chemical measurements between September 2010 and July 2012 showed that dissolved oxygen values were generally below 6 mg l-1. The average NH4+ concentration in the estuary was 7 μM and increased with depth to 12 μM at 2 m depths. Concentrations >45 μM were found in February and April 2011 at the 5 m deep hole at 3.4 km upstream. Negative correlations between dissolved oxygen and NH4+ during November 2010, February 2011, April 2011 and July 2011 (r = -0.68; -0.67; -0.63; -0.96) indicated that remineralisation of organic matter had occurred. Soluble reactive phosphorus (SRP) followed a similar trend to the NH4+ and was generally below 1 μM in the water column for most months, and had peaks at 1.0 km and 3.4 km in the bottom water. The abundance of submerged macrophytes and macroalgae below the N2 bridge were mostly influenced by mouth state and river inflow. During the closed phase the dominant macroalga Cladophora glomerata had an area cover ranging from 3000 to 6000 m2 while Zostera capensis and Ruppia cirrhosa covered an area of 2000 to 3500 m2 and 1500 to 2900 m2, respectively. After an artificial breach in February 2011, water drained out of the estuary leaving the alga stranded on the marshes and as the flood tide entered the macroalga was once again redistributed in the lower reaches. The alga utilised the available nutrients in the water column and expanded its area cover from 35000 m2 in February 2011 to 64000 m2 in March 2011. However, after the floods in June 2011, Cladophora glomerata had been washed out of the system while the submerged macrophytes responded positively extending their area cover. By comparing the artificial breach with the natural breach, and the effect on the estuary, an important observation was highlighted. Increasing the current allocated ecological reserve, and using a larger volume of water to breach the mouth artificially, would result in better scouring of sediment and associated organic matter out of the estuary. This would enable better oxygenation of the water column, reduce remineralisation and minimise algal blooms.

The present study was undertaken in the temporarily open/closed Mngazi and permanently open Mngazana estuaries, located on the subtropical east coast of South Africa. The results from this research will assist decision makers in the freshwater management of these systems. Intertidal and subtidal benthic chlorophyll a concentrations, water column chlorophyll a, nutrients and several physico-chemical parameters were measured between June 2002 and November 2003. The objective of this study was to determine if the presence of freshwater in the estuaries had an effect on the microalgae of both estuaries. Five sites were sampled in the Mngazi Estuary and 14 sites were sampled in the Mngazana Estuary. The average water column chlorophyll a was significantly higher (p<0.05) in the Mngazana Estuary (surface 7.8 ± 0.7 μg.l-1, bottom 6.4 ± 0.7 μg.l-1) compared to the Mngazi Estuary (surface 4.9 ± 1.2 μg.l-1, bottom 7.3 ± 1.5 μg.l-1). There was no evidence of an REI (river-estuary interface) zone in areas where the water column chlorophyll a concentrations were high even during open mouth conditions in the Mngazi Estuary. The REI is that area where salinity is less than 10 ppt and is characterized by high water column productivity. Even though both systems received some freshwater during the summer periods, this was not enough to stimulate phytoplankton growth and nutrient availability seems to be the major factor limiting phytoplankton in these systems. Flagellates and diatoms were the dominant phytoplankton groups in both estuaries during the entire sampling session. The relative abundance of the different phytoplankton groups did not show differences between sites. The relative abundance of flagellates was in most cases greater than 60% and diatoms made up the remainder. The average benthic chlorophyll a was higher in the Mngazana Estuary (intertidal 24 ± 6 μg.g-1 subtidal 15.2 ± 3 μg.g-1) compared to the Mngazi Estuary (intertidal 15.3 ± 4.3 μg.g-1 subtidal 5.4 ± 1.6 μg.g-1). Regions with high benthic chlorophyll a concentrations had high sediment organic content. Sediment organic content was higher in the Mngazana Estuary (1 percent - 8 percent) compared to the Mngazi Estuary (4 percent – 6.8 percent). The sites situated on the Main Channel had on average significantly higher (p<0.05) benthic chlorophyll a biomass compared to Creek 1 and Creek 2 in the Mngazana Estuary. Peaks in benthic chlorophyll a concentrations occurred in the intertidal sediments in Creek 1 (50.4 ± 13.4 μg.g-1) and Creek 2 (57.4 ± 1.4 μg.g-1) in the Mngazana Estuary, the peaks occurred in winter during a period of low freshwater inflow into the estuary. Microphytobenthic biomass measured in the Mngazi Estuary is among the lowest values reported in the literature for temporarily open/closed estuaries. Statistical 4 analysis showed no significant difference between benthic chlorophyll a during the different mouth conditions and sampling sessions in the Mngazi Estuary. Microalgal responses in the Mngazana Estuary were similar to those observed in other permanently open marine dominated estuaries. In the temporarily open/closed Mngazi Estuary microalgal characteristics were different to that of other temporarily open/closed estuaries probably because the estuary was only sampled in the open and semi-closed state.

Increased abstraction of water in the catchment results in a reduced or altered pattern of river flow and this holds serious consequences for the downstream estuarine ecosystem. In South Africa this is a serious concern because freshwater is in limited supply and the demand for freshwater can be expected to increase in the future. A large multi-disciplinary consortium of South African scientists are working on projects to determine the freshwater requirements of estuarine ecosystems. As part of this, this thesis reports on research undertaken to develop mathematical models to determine the freshwater requirements of estuarine macrophytes. Three key macrophytes are selected. The macrophytes are Zostera capensis Setchell, Ruppia cirrhosa Grande, and Phragmites australis. They are common macrophytes in South African estuaries. Zostera and Ruppia are submerged macrophytes and Phragmites is an emergent macrophyte. They have different freshwater environments and therefore respond differently to alterations in freshwater flow. A first order differential equation model is used to determine the effect of different combinations of open and closed mouth conditions of the estuary on Zostera and Ruppia. The scenarios are selected to determine whether achieving a switch in states from a Zostera-dominated estuary to a Ruppia-dominated estuary is possible. To predict encroachment rates and colonisation patterns, a cellular automaton of the vegetative spread of existing Zostera beds is developed. After analysing various scenarios accounting for both an increase and a decrease in freshwater supply, the cellular automaton is extended to include interactions between Ruppia and Phragmites. The multi-species model is applied to the Kromme estuary, South Africa and the Great Brak estuary, South Africa. Various freshwater scenarios are examined from the natural runoff condition to the situation of no freshwater inflow. A sensitivity analysis of the spatial model with Zostera, Ruppia and Phragmites is conducted. / Thesis (Ph.D.)-University of Natal, Pietermaritzburg, 1998.

Increased abstraction of water in the catchment results in a reduced or altered pattern of river flow and this holds serious consequences for the downstream estuarine ecosystem. In South Africa this is a serious concern because freshwater is in limited supply and the demand for freshwater can be expected to increase in the future. A large multi-disciplinary consortium of South African scientists are working on projects to determine the freshwater requirements of estuarine ecosystems. As part of this, this thesis reports on research undertaken to develop mathematical models to determine the freshwater requirements of estuarine macrophytes. Three key macrophytes are selected. The macrophytes are Zostera capensis Setchell, Ruppia cirrhosa Grande, and Phragmites australis. They are common macrophytes in South African estuaries. Zostera and Ruppia are submerged macrophytes and Phragmites is an emergent macrophyte. They have different freshwater environments and therefore respond differently to alterations in freshwater flow. A first order differential equation model is used to determine the effect of different combinations of open and closed mouth conditions of the estuary on Zostera and Ruppia. The scenarios are selected to determine whether achieving a switch in states from a Zostera-dominated estuary to a Ruppia-dominated estuary is possible. To predict encroachment rates and colonisation patterns, a cellular automaton of the vegetative spread of existing Zostera beds is developed. After analysing various scenarios accounting for both an increase and a decrease in freshwater supply, the cellular automaton is extended to include interactions between Ruppia and Phragmites. The multi-species model is applied to the Kromme estuary, South Africa and the Great Brak estuary, South Africa. Various freshwater scenarios are examined from the natural runoff condition to the situation of no freshwater inflow. A sensitivity analysis of the spatial model with Zostera, Ruppia and Phragmites is conducted. / Thesis (Ph.D.)-University of Natal, Pietermaritzburg, 1998.

Aspects around the management of benthic soft sediment bait organisms, with special focus on the mud prawn (Upogebia africana) of eastern Cape estuaries was investigated. The recreational linefishery of the Gamtoos estuary was described, and compared to previous studies. Target fish species was identified, and a large dependency on bait sourced from estuaries needed for the capture of these species was noted, especially for spotted grunter (Pomadasys commersonnii). A comparison between bait use, success and the natural diet of target fish species was also made. The structure and distribution of sediments in the old channel mud banks was described and compared with historical data. The influence of sediments on mud prawn distribution was investigated, but no significant interactions were found at the study site. The impacts of once-off pumping and digging events, and monthly trampling on the sediments and mud prawn population was studied over a 7-month period. Initial removal rates as well as recovery time varied significantly between the two collection methods, while the largest decline in prawn numbers (to zero) with no recovery visible after seven months was caused by trampling. All disturbances caused some alteration in sediment composition, but not to such an extent that the sediments became unfavorable for mud prawns. Trampling did, however, result in the compaction of sediments to such a degree that prawns could not construct burrows. Issues around current removal quotas of bait species as well as the creation of a small-scale commercial (SSC) bait selling operation at Swartkops estuary were critically evaluated, and suggestions for the future removal rates of mud prawns based on production export calculations were made. The distribution, size, sex ratios and number of gravid females occurring along a tidal gradient as well as along horizontal gradient of the mud bank during growth (January – March) and reproductive (September – October) periods was investigated. Changes in the distribution of females between the two study periods were significant, while the distribution of reproductively active females were closely linked to the low water mark (Lower tidal levels). Females occurring in this zone were also significantly larger than females occurring towards the back of the study site. Some minor changes along the horizontal gradient were also observed. The sediment compos ition of the mud bank was found not to play a role in this distribution, leading to the suggestion that exposure to ebb and flood tide currents could influence female prawn distributions. Management recommendations based on these observations were made.

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.

The coastal transition zone (CTZ), which links the terrestrial and marine environments, has been identified as a region of high biological diversity and elevated production. Results of studies conducted in the northern hemisphere indicate that the links between estuaries and the adjacent marine environment is critical to ecological functioning within the CTZ. This study assessed the influence of selected estuaries with different hydrodynamic characteristics on the adjacent marine environment along the south-eastern coastline of southern Africa. Four estuaries were examined, including two permanently open systems, the fresh water deprived Kariega and fresh water dominated Great Fish, and two temporarily open/closed estuaries (TOCE), the Kasouga and East Kleinemonde. Results of the study indicated that outflow of estuarine water from the Great Fish Estuary contributed to a plume of less saline water being evident within the adjacent marine environment. The plume of water was associated with increased zooplankton biomass and particulate organic matter (POM) and chlorophyll-a concentrations. Adjacent to the Kariega Estuary, no evidence of fresh water outflow into the marine environment was observed. However, in the sea directly opposite the mouth of the estuary an increase in zooplankton abundance and biomass was evident. Results of numerical analyses indicated that the increase in zooplankton abundance observed adjacent to the mouth of both permanently open estuaries could not be attributed to the export of zooplankton from the estuary, but rather the accumulation of marine species within the region. The mechanisms responsible for this accumulation were not determined, but it was thought to be associated with increased food availability in the estuarine frontal zone. A similar, but less dramatic biological response was also observed in the marine environment adjacent to the two TOCEs. It is suggested that the increase in biological activity within these regions could be ascribed to seepage of estuarine or ground water through the sand bar that separates these estuaries from the sea. Results of stable carbon isotope analyses indicated that both the Great Fish and Kariega estuaries exported carbon to the nearshore marine environment. The area influenced by estuarine derived carbon was dependent on the volume of estuarine outflow to the marine environment. Adjacent to the fresh water dominated Great Fish Estuary, estuarine derived carbon was recorded up to 12km from the mouth, while adjacent to the fresh water deprived Kariega, estuarine derived carbon was only evident directly opposite the mouth. The recruitment of macrozooplankton (> 2cm) into the fresh water deprived Kariega Estuary was in the range recorded for other permanently open southern African estuaries with higher fresh water flow rates. This indicates that the mechanisms which allow estuarine dependent larvae to locate and enter estuaries are not related to fresh water inflow. In conclusion, this study has demonstrated that despite their small size relative to European and North American systems, South African permanently open and temporarily open/closed estuaries also influence biological activity within the adjacent nearshore marine environment.

Long-term interannual changes in richness, abundance, diversity and structure of the fish community in the temporarily open/closed East Kleinemonde Estuary, Eastern Cape, are described and the recruitment success of two estuary-dependent marine species assessed. In addition, laboratory experiments were conducted to assess the possible role of olfaction in the recruitment process of an estuary-dependent marine fish species. Multivariate analyses of the annual marine fish communites identified two distinct groups with more species recorded during years that succeeded spring (September to November) mouth opening events than in years following no mouth opening events in spring. Interannual community stability (IMD) and seriation (IMS) also increased from the ‘other’ to the ‘spring’ years. These results highlight the importance of the timing of mouth opening to the marine fish community in a temporarily open/closed estuary. This study also made use of long-term records of daily mouth state and linked them to the recruitment of distinct year-class cohorts in two spardis with contrasting lifehistory characteristics. Lithognathus lithognathus only recruited into the estuary in years when the mouth opened between late August and January. This was linked to the limited spawning season of this species and its inability to enter closed estuaries via wave overwash events. In contrast, recruitment by Rhabdosargus holubi juveniles appeared to be uninterrupted and was not determined by the seasonality of mouth opening. This species dominates the marine-spawning component of the East Kleinemonde Estuary and its success is attributed to an extended spawning season and its ability to recruit into estuaries during both overwash and open mouth conditions. Attraction of postflexion Rhabdosargus holubi larvae to estuary, surf zone and river water was also measured using a rectangular choice chamber. In two sets of experiments, conducted during peak recruitment periods, larvae from both the surf zone and estuary mouth region selected estuary water with a significantly higher frequency than sea water. Larvae collected in the mouth region showed a stronger preference for river water than those collected in the surf zone, thus suggesting that these fish are more attracted to freshwater influenced nursery areas once they have entered the estuary than those in the surf zone. Larvae collected in the marine environment also selected surf zone water with significantly higher frequencies than estuary water or offshore sea water, thus confirming the importance of the surf zone as an interim nursery area for postflexion R. holubi.

The growth and accumulation of macroalgae in estuaries are a common sight throughout the world (Boyer and Fong, 2005). Because macroalgae establish such complex but important links with other components within an estuary, it is essential to know which variables control and regulate their occurrences. The type of algae that will occur in an estuary depends however, on the physical and chemical features of the particular estuary (Lobban and Harrison, 1994). The focus of this study was to document and provide a listing of algae from estuaries around the Eastern Cape coast of South Africa and describe the communities which form within each estuarine habitat. The questions posed were: Do algae assemblages differ in estuaries with different environmental conditions? Are different plant communities inhabited by different algae communities? Also, are algae diverse in estuaries that are in a pristine condition? The estuaries were Mngazana, Ngqusi/Nxaxo, Mtati, Mpekweni, East Kleinemonde, West Kleinemonde, Kariega, Bushmans, Kowie and the Sundays estuary. Percentage cover surveys were conducted and environmental parameters recorded to determine community structure of algae and their environmental tolerances. A total of 49 different macroalgal species were identified from 11 estuaries, which were dominated by Chlorophytes (23 species) and Rhodophytes (17 species). The Ochrophytes consisted of 10 species. The highest number of species recorded was 29 at the Bushmans estuary (March 2009), while the lowest number (4 species) was recorded at Mtati (October 2008). Common Chlorophytes found included Ulva prolifera, Chaetomorpha linum and Codium tenue. In the Division Ochrophyta, species from the Genus Ectocarpus were the most common. Of the red algae filamentous reds, Polysiphonia incompta and Polysiphonia kowiensis were the common algae found at estuaries. Salinity (p= 0.029861; n= 49) and temperature (p= 0.037641; n= 49) were the driving environmental parameters of estuaries and significantly influenced richness of species. In ordination analyses, patterns of biogeographic distribution were noticeable and were described by salinity. Four gradients caused by salinity were therefore found – which is inversely correlated to water clarity. Moreover, a clear split between temporarily open/closed and the permanently open estuaries was observed. Samples grouped into four communities presumably based on the similarity in conditions of estuaries. Within each estuarine community and major type of estuary there was a range of environmental conditions. However, the macroalgae within each community may not have been colonists of or found in greatest abundances in the estuaries of these communities per se, but may have grouped together in this manner based on the optimum growing conditions for them. Also, specific algal groups were not directly determined by the type of estuary they were found in; they were driven by environmental ranges rather. The results therefore support hypothesis one which states species distribution and diversity differ and are influenced by environmental gradients of the estuary. Although close species and habitat relationships were apparent from the ordination analyses, providing support for hypothesis two which states “Different plant communities in estuaries will be inhabited by different macroalgae communities”, the evidence was not substantial enough to provide sufficient support for the hypothesis. Lastly, the hypothesis “Macroalgae are diverse in estuaries that are in a pristine condition” was not accepted as highest species numbers were found in estuaries of greatest anthropogenic disturbance (e.g. Bushmans estuary) and not in estuaries which where ecologically more natural (e.g. Mtati estuary). By doing monthly macroalgal sampling over a period of one year for example one would obtain greater information of biodiversity, and patterns of macroalgal seasonality may be uncovered as well. This is an area of reasearch which needs to be investigated in the future as it would greatly add to the understanding of the role macroalge have in an estuarine environment.

The role of the sandprawn, Callichirus (=Callianassa) kraussi (Stebbing), as an ecosystem engineer was assessed in the lower reach of the temporarily open/closed Kasouga Estuary situated along the Eastern Cape coastline of southern Africa over the period April 2010 to June 2011. The study comprised two distinct components, a field study and a caging experiment. The field study assessed the correlation between sand prawn densities and selected physico-chemical (organic content of the sediment and bioturbation) and biological (microphytobenthic algal concentrations and macrobenthic abundance and biomass) variables in 50 quadrants in the lower reach of the estuary. Densities of the sand prawn within the quadrants ranged from 0 to 156 ind m⁻² (mean = 37 ind m⁻²). There were no significant correlations between the densities of the sandprawn and the estimates of the organic content of the sediment and the abundance and biomass of the macrofauna (P > 0.05 in all cases). Numerical analyses failed to identify any effect of the sandprawn density on the macrofaunal community structure. The rate of bioturbation was, however, strongly correlated to the sand prawn density. Similarly, the microphytobenthic alga concentrations were significantly negatively correlated to the sand prawn densities ((P < 0.05). The absence of any distinct impact of the sandprawn on the macrobenthic community structure appeared to be related to their low densities in the lower reach of the estuary during the study. To better understand the role of the sandprawn as an ecosystem engineer, a caging experiment was conducted using inclusion and exclusion treatments (n= 5 for each treatment). Densities of the sandprawn in the inclusion treatments (80 ind m⁻²) were in the range of the natural densities within the estuary. The experiment was conducted over a period of 18 weeks in the lower reach of the estuary during summer. The presence of the sandprawn, C kraussi, contributed to a significant decrease in the microphytobenthic algal concentrations and the abundance and biomass of the macrofauna (P < 0.05 in all cases). The decrease in the microphytobenthic algal concentrations in the presence of the sandprawn appeared to be related to the res-suspension of the sediments (bioturbation) generated by the burrowing and feeding activities of the sandprawn. The observed decrease in macrofaunal abundances and biomass in the inclusion treatments appeared to be mediated by both the decreased food availability (mainly the microphytobenthic algae) and the burial of organisms within the sediments. Numerical analysis indicated that the sandprawn did, however, not contribute to a change in the species composition of the macrofauna. Results of the current study indicate that C.kraussi plays an important role in structuring the invertebrate community and energy flow within temporarily/open closed Kasouga Estuary.