Estuaries are intrinsically complex and dynamic ecosystems that display marked spatial and temporal variability. Because estuaries are situated at the receiving end of catchment activities, they are at particular risk of alterations to their natural complexity. The overarching objective of this study was to gain an overview of the abiotic conditions and biotic response of the Goukamma Estuary, a small temporarily open/closed estuary (TOCE) which is situated in a relatively undisturbed catchment in the Southern Cape. Physico-chemical properties drive estuarine ecology, and together with biological indicators, are commonly assessed to determine the present status of an estuary. During the study, physico-chemical parameters reflected great spatial and temporal variability in response to the mouth state over a 13 month period. Parameters ranged within expected limits, as proposed by the conceptual model for water quality of TOCE’s (Snow and Taljaard 2007). Of particular importance was the prominent occurrence of salinity stratification and hypoxic conditions (dissolved oxygen [DO] < 3 mg l-1) during both open and closed mouth states. Data indicated that in the wide and shallow lower reaches, weak stratification gradients were present and oxygenated conditions (DO > 6 mg l-1) were maintained throughout the water column mainly by wind and tidal action. However, stratification increased towards the deeper, channel-like middle and upper reaches of the estuary, followed by a significant reduction in bottom DO concentrations and development of hypoxia and anoxia. Bottom water hypoxia commonly occurs in microtidal estuaries due to the limited influence of mixing forces, mainly by wind and tidal action. The Goukamma Estuary is a channel-like microtidal estuary where stratification effectively limited oxygenation of the bottom water which resulted in frequent occurrence of bottom water hypoxia. During June 2010 when the highest local rainfall (75 mm) was recorded for the region, salinity and DO data showed that this amount of rainfall was insignificant as it did not replenish the water column of oxygen. Only the surface 0.5 m layer was fresh and oxygenated while below this, the water column was completely hypoxic. In an unimpacted state, the Goukamma Estuary is a blackwater system and is expected to be nutrient poor; however, farming activities in the catchment have resulted in elevated nutrient concentrations. This study showed that significantly higher nutrient concentrations were measured in the middle and upper reaches of the estuary, adjacent to cattle farms situated in the floodplain of these reaches. Nutrient concentrations represented mesotrophic (dissolved inorganic nitrogen [DIN] > 500 μg l-1) to eutrophic conditions (dissolved inorganic phosphorus [DIP] > 25 μg l-1). Nutrient input stimulated phytoplankton to attain a significantly high biomass, ranging between 0.3 – 112 μg l-1 (~ 7.7 ± 1.3 μg l-1; n = 128) and 0.8 – 289 μg l-1 (~ 21.1 ± 4.4 μg l-1; n = 80) during the open and closed states, respectively. High organic loads are associated with high oxygen demands which consequently result in hypoxia following decomposition. Exacerbated by natural salinity stratification which effectively limits oxygenation of the water column, unnaturally high nutrient concentrations and coinciding organic loads place the estuary at particular risk of degradation. This study captured key patterns and processes by quantifying salinity, oxygen and nutrient concentrations in addition to biological indicators (phytoplankton biomass and community composition). Considering possible budget constraints, it is recommended that monthly salinity and oxygen concentrations should be monitored as well as seasonal nutrient concentrations. It is also recommended that riparian buffer zones should be established in the middle and upper reaches of the estuary, as these vegetation buffers have been well documented to contribute to nutrient attenuation and improved water quality from agricultural run-off.

The utilisation of estuary headwater environments by young estuary- and marine-spawned fish species was investigated together with the effects of riverflow alteration, in-stream barrier effects and non-native ichthyofauna on the nursery function of these habitats. The distribution and abundance of young estuary- and marine-spawned fish were sampled using seine and fyke nets in the headwater environments of four permanently open Eastern Cape systems, namely the Great Fish, Kowie, Kariega and Sundays Estuaries. Within the suite of study systems, the first of two case studies focussed on barrier effects of in-stream structures on fish migration. This was undertaken in the Sundays River. In the second case study, predation and competition dynamics of the non-native piscivorous Micropterus salmoides on estuary-dependent fish was investigated in the estuary headwater regions of the Kowie River system. In all four estuaries, young estuary-spawned fish species dominated the ichthyofaunal community followed by marine-spawned species, despite varied freshwater inflow resulting in headwaters varying in salinity from fresh to hypersaline. Fish community structure however, differed largely between estuaries, with both freshwater abstraction and unnatural elevation of freshwater into estuaries, as a result of inter-basin transfers, affecting these communities. In-stream structures were found to effect upstream movement of fish in two ways, dependent on the type of barrier. Partial (size-dependent) and complete (species-dependent) restriction to upstream migration of fish by causeway-type instream structures were observed. Weir-type in-stream structures acted as a complete barrier to most species, regardless of fish size. Predation of estuary- and marine-spawned fish species by large sized M. salmoides was recorded, although these fish did not contribute significantly to their diet during this study. However, the main dietary components found in smaller sized M. salmoides stomachs overlap with those of juvenile estuary- and marinespawned fish species, suggesting feeding competition between the juveniles of indigenous and non-native fish species.

The East Kleinemonde Estuary is one of 175 temporarily open/closed estuaries (TOCEs) that represent 70 percent of estuaries in South Africa. TOCEs are small (mostly less than 100 ha), shallow estuaries (average depth < 2 m) that respond quickly to freshwater inflow events. Their connection to the sea can be highly variable resulting in considerable changes in abiotic and biotic conditions. Mouth status depends on a balance between freshwater inflow and marine influence, which in turn affects ambient abiotic conditions. The objective of the study was to identify the abiotic variables which influence macrophyte growth and habitat availability. It was hypothesised that water level and salinity were the two main drivers of macrophyte change and macrophyte habitat would respond very rapidly, in less than a month, when habitat was available. Macrophyte habitats would also have high sediment seed reserves to ensure persistence under highly variable abiotic conditions. Macrophyte cover was monitored monthly in the East Kleinemonde Estuary along three permanent transects. The dominant habitats were submerged macrophytes, intertidal salt marsh, supratidal salt marsh, reeds and sedges. The following abiotic variables; water level, water column salinity, water temperature, Secchi depth, air temperature and rainfall were also measured between March 2006 and January 2010. Time-lag responses of the macrophytes to water level and salinity changes up to four months prior to the sampling session were also assessed. The analysis of a one year dataset highlighted only water level as a driver of change in macrophyte cover, whereas the five year dataset identified salinity as an additional important abiotic driver. This is because during September 2008 to January 2010 a series of large marine overwash events maintained high salinity (> 30 ppt) and high water level (> 1.6 m amsl) in the estuary. Water level increased by up to 0.33 m due to large volumetric changes and salinity was significantly higher in the 16 month closed euhaline phase after the breach (31 ± 0.9 ppt) compared to 21.9 ± 0.9 ppt in the closed polyhaline phase before the September 2008 breach. This increase in salinity significantly reduced the cover of the submerged macrophytes Ruppia cirrhosa and Chara vulgaris. They were replaced by macroalgae during this high salinity phase. The cover of supratidal salt marsh and reed habitats was also significantly reduced during the high water level phase, which in turn would lead to the potential for bank destabilisation and erosion. Based on the average elevation above sea level position of the macrophytes in the East Kleinemonde iv Estuary, a threshold water level was identified as 1.55 amsl. This was taken to be the height above sea level at which there was a maximum cover change for each macrophyte habitat. Above this water level emergent macrophyte habitat would mainly be inundated. This, together with 30 ppt salinity, was identified as the two thresholds for macrophyte change in the East Kleinemonde Estuary. From these thresholds and the 5 year dataset four biotic states were identified as State A: open and tidal, State B: closed with a water level below 1.55 m amsl and salinity between 18 to 30 ppt, State C: closed and water level above 1.55 m amsl and salinity between 18 to 30 ppt and State D: closed and water level above 1.55 m amsl and salinity above 30 ppt. Intertidal salt marsh, reeds and sedges were dominant during the open phase. Submerged macrophytes were dominant during the closed polyhaline state and macroalgae during the closed euhaline state. The high variability of abiotic factors common in TOCEs and the response of macrophyte habitat indicated that macrophytes were resilient to changing states provided they were of relatively short (< 3 months) duration. Macrophytes in the East Kleinemonde Estuary were found to have fast growth rates and large seed reserves in the sediment. The seed banks in the East Kleinemonde, as well as the adjacent temporarily open/closed West Kleinemonde Estuary were quantified for the first time in a South African estuary. The averaged data from both estuaries showed that Charophyte öospores represented almost 72 percent of the sexual propagules in the sediment with a mean öospore density of 31 306 ± 2 293 m-2. This was despite the Charophytes being sparsely located and only representing a maximum of 32.5 percent cover in the above ground vegetation. Historically there must have been stands of Charophytes in the East Kleinemonde Estuary, such that öospores could accumulate to such high density found in this study. The second highest seed density was for the intertidal salt marsh plant Sarcocornia tegetaria (18 percent) (7 929 ± 688 seed m-2), followed by the submerged angiosperm Ruppia cirrhosa (7 percent) (2 852 ± 327 seeds m-2). Although seed density did not differ significantly with sediment depth, seeds still occurred at 20 cm below the surface of the sediment providing a regeneration source in the event of sediment scouring during a flood event. Germination studies in the greenhouse showed that most seeds were viable and Sarcocornia tegetaria began to germinate after 3 days to a maximum of 82 percent after 91 days. Submerged species only germinated after 18 days with a low maximum germination of between 11 and 15 percent. This study has made an original contribution to the field of knowledge on macrophyte responses in a small TOCE as it showed that macrophyte habitats in the East Kleinemonde Estuary have a high natural variability in cover over time, they respond quickly after a disturbance event such as a mouth breach and there are large sediment seed reserves that remain viable from 2 to more than 5 years. This ensures habitat persistence even under unfavourable conditions, such as prolonged periods of mouth closure with high water level and flooding which causes loss of salt marsh species. Given this natural variability it is necessary to predict responses both spatially and temporally in order to manage and maintain ecological functioning in TOCEs. This study identified dominant macrophyte habitat for different abiotic states through the use of water level and salinity thresholds. In the determination of the freshwater requirements of any South African estuary freshwater inflow rates are provided for each estuary's past, present and possible future freshwater inflow scenarios. These flow data are generated by hydrological models and simulated monthly inflow volumes for a period of about 72 years are provided. For the East Kleinemonde freshwater requirement study for any year in that 70-odd year period, the number of high flow and low flow mouth breaches were predicted, as well as the closed state periods. The threshold water level of 1.55 m amsl was also used to filter past, present and future inflow monthly volumes to determine the frequency of the four abiotic states identified in this study. It was based on a water level/water volume equation calculation from a digital elevation model. Results showed that the total closed period in the present state was 83 percent, made up of 48 percent of the time in a polyhaline state (State C) and 35 percent in a euahaline state (State D). A second method was used to quantify available spatial habitat under different water level scenarios. A spatial model was written in Model Builder, an application in ArcGIS that allowed a series of processes to be built. A habitat map was overlaid with a bathymetric map and by selecting water level, available habitat areas were determined and empirical equations of water level versus available habitat were produced. These equations were then used to calculate the available habitat areas for monthly water level conditions from the freshwater requirement study for the past, present and two future inflow scenarios. Using both the threshold water level method and the spatial availability model method it was possible to assess the effect of the two future inflow scenarios on macrophyte habitat vi response. Scenario 1 had a 16 percent reduction in mean annual runoff (MAR) generating low flows for 88.6 percent of the time and a 3.5 percent reduction in flood events. In Scenario 2 there would be a 12 percent reduction in MAR with low flows occurring for 87.5 percent of the year, a 5.3 percent reduction in floods and an 11.5 percent reduction in the open mouth state. The model showed that Scenario 1 would have the highest submerged macrophyte area (12.56 ha versus 12.48 ha in Scenario 2), whereas Scenario 2 produced the largest mudflat and intertidal salt marsh area (7.11 ha versus 7.34 ha) due to lower water level in conjunction with the bathymetry of the estuary. A reduction in freshwater inflow to TOCEs either due to anthropogenic influences or natural precipitation cycles is one of the main threats to the optimum functioning of these estuaries. The results from this study and the two methods of assessing the effect of freshwater inflow scenarios on macrophytes in TOCEs can be integrated into the current freshwater inflow assessment methodology in South Africa, as well as adding to our understanding of the ecological functioning of these small, highly variable estuaries. The methods provide a quick assessment of macrophyte habitat associated with abiotic states under past, present and future inflow scenarios. All that is required to predict macrophyte habitat for different freshwater inflow scenarios (present, past and future) is a habitat map, a bathymetric map and the elevation range of macrophytes in the TOCE being assessed. This, together with the knowledge of response rates, provides invaluable information for the management of TOCEs to maintain their ecological functioning under altered freshwater inflow regimes.

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Thesis (Ph.D.)--Murdoch University, 2009. / Thesis submitted to the Faculty of Sustainability, Environmental and Life Sciences. Includes bibliographical references.

Thesis (M. Phil.)--University of Hong Kong, 1987.

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Thesis (Ph. D.)--Australian Catholic University, 2002. / A thesis submitted in total fulfillment of the requirements for the degree of Doctor of Philosophy. Includes bibliographical references (305-329). Also available in an electronic version via the internet.