Temporarily open/closed estuaries (TOCEs) are unpredictable environments that change in response to mouth condition, which is influenced by freshwater flooding or sea storm surges. The aim of the study was to determine whether macrophyte phenology in a TOCE was event driven rather than cyclically predictable and if it differed from permanently open estuaries (POEs). Macrophyte growth and flowering phenology in response to environmental conditions was investigated in the East Kleinemonde Estuary (TOCE) and the Kowie Estuary (POE) along the eastern coastline of South Africa. The lack of freshwater flooding due to low rainfall coupled with several overwash events resulted in a prolonged period of mouth closure from September 2008 to the end of this study period in 2010. This in turn caused the inundation of the supratidal and intertidal habitats, high water level (> 1.57 m amsl) and high salinity (30 - 42 ppt) in the TOCE. Principle Components Analysis showed that high water level and reduced sediments were the most significant environmental factors affecting macrophyte phenology. Macrophyte phenology in the POE was primarily driven by temperature, sediment redox potential and salinity. The saline high water level and reduced sediment significantly reduced macrophyte cover in all habitats in the TOCE. Macrophytes in the POE maintained high cover abundance due to seasonal re-growth compared to the TOCE where cover declined over the sampling period due to the high water level. Subsequent to water level dropping by as little as 11 - 20 cm in the TOCE, the intertidal species Sarcocornia tegetaria and Salicornia meyeriana completed their life-cycles and produced viable seeds within four and three months of germinating respectively. In contrast, the Sarcocornia hybrid and S. meyeriana in the POE took longer to complete their life-cycles, namely seven and nine months respectively, while S. tegataria did not germinate in situ but reproduced vegetatively despite producing seed. In the TOCE, the submerged species Ruppia cirrhosa and Chara vulgaris completed their life-cycles within five and three months and produced a maximum of 26 242 and 196 998 seeds m-2 respectively. Due to high water level and prolonged inundation, the reproductive periods were shorter for the intertidal and reed and sedge species in the TOCE compared to the POE. Seed output during the two reproduction periods varied between the two estuaries. Sarcocornia decumbens and S. tegetaria produced a substantially higher number of seeds in the TOCE compared to the POE, namely 0 - 102 847 versus 20 661 - 48 576 seeds m-²; and 7 001 - 45 542 versus 1 587 – 16 958 seeds m-² respectively. Seed output in the TOCE was significantly higher in S. tegetaria during the second reproduction period despite the significantly lower plant cover, which may be a function of the stressful environment in the TOCE. Seed production of S. meyeriana was significantly higher in the POE compared to the TOCE, with 264 224 - 640 292 compared with 24 050 - 27 643 seeds m-², due to higher plant cover in the POE. The research suggests that macrophyte phenology in the TOCE was significantly influenced by mouth condition. Further, macrophytes were able to demonstrate considerable phenotypic plasticity in response to changing and unfavourable environmental conditions. These data can be used in mouth management plans and freshwater requirement studies in TOCEs to ensure that macrophytes can complete their life-cycles and produce viable seeds for the safeguarding of habitat persistence and ecological processes. In impacted estuaries where artificial mouth opening is practised and the macrophytes have been severely degraded or extirpated, management should ensure that the intertidal and supratidal habitats are not inundated during peak flowering and seed production periods i.e. late spring to early autumn (November to March).

River flow is important in controlling phytoplankton distribution in estuaries. Data on the effect of river inflow on phytoplankton distribution patterns in temporarily open/closed estuaries is lacking. This study investigated the influence of river inflow on size-fractionated phytoplankton biomass (Chl a), community composition and environmental parameters measured monthly over three years in two temporarily open/closed estuaries in the Eastern Cape, South Africa. A once-off primary production study over an annual cycle was completed in the Van Stadens and Maitland estuaries. The study monitored physical, chemical and biological characteristics in both estuaries to examine the effects of changes in environmental factors and river inflow. Daily sampling of physico-chemical and biological variables from river to sea was carried out in the Van Stadens to investigate short-time scale effects of changes in environmental factors and river inflow on the phytoplankton biomass. Five and three stations in the main channel of the Van Stadens and Maitland estuaries respectively were sampled at 0.5 m below the water surface and 0.5 m above the sediment surface for biological and chemical variables and at the surface, 0.25 m and every 0.5 m thereafter for physical parameters. Five stations adjacent to the main channel along the estuary were monitored for groundwater macronutrient concentrations and five additional sites located within the upper catchment of the Van Stadens River were sampled on a quarterly basis over two years. Both estuaries were characterised by distinct hydrological conditions, an overwash, an open, a closed and a semi-closed mouth phase. Flooding in the Maitland and Van Stadens estuaries in 2001 and 2002 caused sediment scour, altered channel morphology and brought about breaching of the mouth. Flood driven mouth-breaching events occurred three and four times in each of the estuaries during the study. The mouth stayed open 20 – 25 percent and was closed 60 – 65 percent of the time. In the Van Stadens the closed overwash mouth condition occurred approximately 10 – 20 percent of the time while in the Maitland it occurred less with the semi-closed mouth condition occurring 10 – 20 percent of the time. Incidents related to mouth opening not associated with strong river floods occurred approximately 10 – 15 percent of the time, although in the Maitland a semi-closed mouth state persisted more frequently than in the Van Stadens Estuary. During flooding events salinity dropped to low levels (< 5 psu) but soon recovered to brackish conditions when river flow was reduced and marine water penetrated deep upstream. Reduction in river flow combined with marine sediment deposition resulted in the closure of the mouth. During closed mouth conditions strong onshore storm surges and spring high tides introduced marine water through overwash that kept salinity high. In both estuaries salinity showed a negative correlation with rainfall (R2 = 0.12), indicative of the strong influence of marine overwash that kept salinity high thus masking the influence of freshwater. High rainfall in the Van Stadens Estuary caused high levels of turbidity that reduced light penetration at depth. Light attenuation was positively correlated with the high rainfall (R 2 = 0.26) suggesting that increased turbidity was linked to rainfall induced discharge. In contrast, in the Maitland Estuary light attenuation did not show any correlation with increased rainfall possibly because of the reduced water depth and increased euphotic zone following the floods in 2002. High river inflow introduced macronutrients in both estuaries such that dissolved inorganic phosphates (DIP) and dissolved inorganic nitrogen (DIN) concentrations in the Van Stadens Estuary were strongly correlated with rainfall (R2 = 0.78 and 0.57 respectively). In the Maitland Estuary DIP and DIN concentrations remained significantly higher (p < 0.05) compared to that in the Van Stadens suggesting that the Maitland catchment contributed greater nutrient input into the estuary and may be associated with farming activities. Phytoplankton chlorophyll a (Chl a) ranged from 0.8 – 13.9 μg L-1 in the Van Stadens and in the Maitland Estuary from 5.3 – 138 μg L-1 during the 3-year study. During the open mouth condition Chl a biomass and primary production ranged from 5.4 – 52.9 μg Chl a L-1 and 1.2 – 11.7 mg C m-2 d-1 in the Maitland and in the Van Stadens from 1.6 – 9.8 μg Chl a L-1 and 1.2 - 14 mg C m-2 d-1 respectively. Maximum annual primary production in the Maitland and Van Stadens estuaries was 8.8 and 5.1 g C m-2 y-1 respectively. When the mouth was open in the Van Stadens Estuary the microphytoplankton (> 20 μm) accounted for > 65 percent of the Chl a, whereas during closed mouth conditions they accounted for about 55 percent of the Chl a biomass. Chlorophytes became the dominant taxon in the dry summer months but were replaced by cryptophytes and dinoflagellates during the wet season. When nutrient concentrations were low during low flow conditions in the Van Stadens Estuary mixotrophic microphytoplankton became an important fraction of the water column together with phototrophic dinoflagellates and cryptophytes. In the Maitland large sized chlorophytes were the dominant taxa in late spring and summer seasons and made up more than 80 percent of the cell numbers. In the Maitland before the floods in 2002 cyanophytes were the dominant group in late spring contributing more than 75 percent in cell abundance. Data from the short-term study in the Van Stadens Estuary showed similarities and differences in the Chl a response to increased river inflow. High river inflow initially reduced Chl a biomass followed by a recovery period of a couple of days compared to a 8 – 10 week recovery period in studies monitored over seasonal and annual temporal scales. The responses may be dissimilar but help to illustrate that there are similar response patterns to environmental forcing necessary to support phytoplankton biomass at different temporal scales. This study has demonstrated that flooding events caused by strong river flow cause breaching of the mouth, a reduction in salinity and marked nutrient input. Although the causes of flooding can be similar in both estuaries the resultant effects are varied and can alter the ability of the estuary to retain water. This study was able to demonstrate that the supply of macronutrients from the catchment was strongly correlated with rainfall (R2 = 0.67) and that phytoplankton growth mainly depended on an allochthonous source of macronutrients although internal supplies could be critical at times in controlling microalgal biomass.

The Great Brak temporarily open/closed estuary was subjected to a drought during the spring and summer of 2009/2010 resulting in the mouth remaining closed for a prolonged period. According to the Great Brak Estuary management programme, the mouth of the estuary had to be open for a total of 308 days during spring and summer of 2009/2010, respectively, but was closed for almost the entire two years (693 days). The aim of this study was to assess monthly changes in the abiotic characteristics (salinity, temperature, oxygen, pH and nutrients) and the biotic responses of phytoplankton and macroalgae; identify sources of nutrient input into the estuary and determine the response of the salt marsh to water level and salinity changes. The results indicated that physico-chemical parameters were similar to that previously recorded during the closed mouth condition. However mouth closure combined with elevated nutrient concentrations led to a shift from rooted submerged macrophytes to one where either microalgae or macroalgae were dominant. Soluble reactive phosphorus concentrations were significantly higher in bottom compared with surface waters. There was a significant negative correlation with SRP and dissolved oxygen for the sampling period indicating potential release of phosphorus from the sediment during closed mouth conditions. Microalgal biomass increased in response to remineralised nutrients and freshwater pulses. Flagellates were the dominant microalgal group (21718 ± 3336 cells m l-1, p < 0.05) because of their morphological ability to migrate vertically within the water column. The macroalgal cover was highest during the closed mouth state but only during winter (August 2010) when temperatures were below 20 oC. Five major point sources of nutrient input into the Great Brak Estuary were identified during rainfall periods. Point sources 4 and 5 in the upper reaches of the estuary had the highest DIN input whereas point source 3 in the middle reaches of the estuary had the highest DIP input. As a result of the drought and low water level, the salt marsh was never inundated for longer than 3 months. Die-back of Sarcocornia decumbens (r 2= -0.62, p < 0.05) was related to smothering by dead macroalgae whereas dieback of Sporobolus virginicus was related to decreasing nutrient (r2 = 0.59, p < 0.05) and salinity (r2 = 0.55, p < 0.05) levels. The physico-chemical characteristics alone did not convey the true health status of the Great Brak Estuary for the duration of the sampling (April 2010-April 2011). The study showed that microalgae and macroalgae are valuable indicators of the status of the estuary. Therefore it is suggested that bio-indicators are incorporated into the management/monitoring plan in order to assist in improving the health assessment of the Great Brak Estuary.

This thesis presents the first intensive community-level investigation of the mesozooplankton from a biogeographical transition zone estuary. The Mgazana Estuary is located along a rural, semi-undeveloped part of South Africa’s east coast, believed to represent a transition zone between the subtropical and warm-temperate biogeographical zones. The research represented in this thesis involves data collected over a five year period between 2002 and 2006, with additional data collection made in 2008. The study focussed on five areas of investigation, the first of which investigated the physico-chemical dynamics of the Mgazana Estuary. The Mgazana Estuary exhibited marked vertical, horizontal and regional structures in the hydrological environment. Prominent vertical and horizontal stratification characterised summer months. A substantial turbidity front was observed in the lower estuary during summer and winter. The upper estuary was marked by considerable variation in multiple variables but especially salinity. These structures appeared to have shifted in position over the horizontal plane, which was attributed to variation in freshwater flow. The second study focused on the spatial dynamics of mesozooplankton in the Mgazana Estuary. The zooplankton was rich and in terms of composition typical of mangrove systems. The Copepoda were dominant, numerically and in terms of taxonomic representation. Calanoids Acartiella natalensis and Pseudodiaptomus hessei characterised middle and upper reaches in summer and mostly upper reaches in winter. On community level, a highly structured assemblage arrangement was observed during summer and winter months. These trends were further scrutinised under the theoretical framework of ecological boundaries. In so doing, an agreeable spatial association emerged between specific assemblages and their environments. These trends were concluded to reflect ecoclinal as well as ecotonal properties, the latter describing interactions over narrow spatial bands of marked changes in turbidity. The third study investigated flooding events as short-lived extreme meteorological events and the influence on zooplankton. Within this regard, second-stage multivariate statistics was used to assess year-to-year variability in assemblage structures on whole-system and regional scales. The impact of two major flooding events that flushed the estuary some days prior to sampling could clearly be elucidated. Flooding emerged as a significant source of inter-annual variability in the zooplankton of the Mgazana Estuary. Second-stage multivariate analysis proved to be an effective analytical strategy for investigating inter-annual variability in species assemblage structures. Results from the preceding study prompted a detailed investigation into the spatio-temporal dynamics of Acartiella natalensis, the most important zooplankton species of the system aimed at elucidating flood responses. Acartiella natalensis showed indication of temporal arrest in association with flooding events in a similar fashion as is observed in seasonal variation trends in estuaries towards its geographical distribution limit south of the study area. It was concluded that A. natalensis was severely affected by floods and the possibility exist for entrainment of entire populations from the estuary during flooding events. The final study tested the hypothesis of post-flood propagation from a resting egg bank in the sediments of the Mgazana Estuary. Preliminary data revealed that numerous eggs were present in the sediments. Nauplii hatched from eggs under laboratory conditions were identified as belonging to the family Acartiidae. The preliminary mode of diapause is presented as a schematic model, emphasising the role of freshwater flow and specific environmental variables. This was the first discovery of resting eggs from a subtropical estuary from South Africa. It is hoped that the findings of this study would give rise to new research initiatives investigating the importance of resting stages in estuarine and coastal Copepoda species and the role such reproductive strategies may play in estuarine functioning.

An estuary is the last stage of a river. It is where the river meets the sea. Estuaries are one of the most significant features of the South African coastline. In recent years, South Africa has witnessed an increase in the demand for freshwater for both industrial and domestic purposes. At the same time, there has been a gradual deterioration of river systems and their catchments. To add to this, there has been a gradual reduction in the amount of recorded rainfall, which is the primary source of freshwater for rivers. This has resulted in decreased freshwater inflow into estuaries, a situation which poses a serious threat to the biological functioning of these estuaries and the services rendered to its recreational users. A deterioration of estuary services reduces the yield for subsistence households and their appeal for recration. This study uses the contingent valuation method as its primary methodology to elicit users' willingness-to-pay to reduce the negative impacts of reduced freshwater inflow into selected western and southern Cape estuaries. Eight estuaries were selected for this study; the Breede, Duiwenhoks, Great Berg, Kleinemond West, Mhlathuze, Swartvlei and Olifants estuaries. The contingent valuation (CV) method is widely used for studies of this nature because of its ability to capture active, passive and non-use values. The CV method involves directly asking people how much they would be willing to pay for specific environmental services. In this case, users were asked what they would be willing to pay to sustain freshwater inflows into selected estuaries in order to prevent the negative impacts of reduced inflows. The travel cost method (TCM) was uesed to generate an alternative comparative set of values for the purposes of convergence testing. This is because convergence testing is highly desirable as a validity test for CV estimates.

There are about 259 estuaries in South Africa that can be classified as “functioning” and their “health” status is directly related to the quantity and quality of freshwater that flows into them. Many of South Africa’s estuaries have become smaller due to a steady decrease in the amount of freshwater that flows into them. This reduction in freshwater inflows decreases their ecological functioning and undermines the recreational activities and subsistence services available from them. The National Water Act (ACT No. 36 of 1998) recognises the right of the environment to water, but a large amount of data is still needed to make management decisions on the allocation of freshwater. The objective of this research was to contribute to the management of the allocation of freshwater in the catchment areas of the Bira, Bushmans, Kasouga, Keiskamma, Kleinemond East, Nahoon and Tyolomnqa rivers by determining the recreational value of the freshwater flowing into their estuaries. This recreational value was established using the contingent valuation method. The contingent valuation method (CVM) is a technique to establish the value of a good (or service) that is not bought or sold in an actual market. This technique is frequently applied in the valuation of environmental goods, e.g. the freshwater that flows into an estuary. The CVM establishes the economic value by asking the users of an environmental good to state their willingness to pay (WTP) for a hypothetical project to prevent or bring about a change in the current condition of the environmental good. The users’ WTP is then aggregated to establish a total willingness to pay (TWTP) for the population of the users of the environmental good. The hypothetical project presented in this study is that of an increase of freshwater inflows, that would prevent (bring about) predetermined changes in environmental services provided by the selected seven estuaries.

A geochemical study of the composition of natural sediments and contaminating mine tailings in Hastings and Alice Arms has been carried out. Apart from the geochemical investigation on the solid fraction, pore waters from six sediment cores have been analyzed to evaluate the diagenetic processes in the sediments of these two inlets. Differences in the mineralogy as well as in the chemical composition of the sediments were used to distinguish the natural sediments from contaminated sediments and pure tailings. In Alice Arm, two types of tailings could be differentiated with these methods. The first tailings type originated from the Kitsault Mo-deposit, which was mined in the past two decades by the B.C. Molybdenum Corporation and AMAX/Canada. These tailings are characterized by a number of features which make them distinguishable from natural sediments. The K-feldspar content of the tailings is significantly increased, mainly at the expense of plagioclase, which is the main characteristic for the mineralogical identification of this tailings type. The enhanced K-feldspar content is the cause for increased K and Rb values in the Mo-mine tailings. Another indicator specific to these recent tailings is an increased Mo content. The second tailings type is derived from an older mining operation in the Kitsault Valley, most likely the Dolly Varden Silver Mine. A specific indicator for these tailings is their high Ba content which is due to large amounts of barite. This mineral phase could be identified by X-ray diffraction methods in a heavy mineral fraction of the relevant sediment layers. Both tailings types show enrichment in their Pb, Zn and S concentrations, caused by an increased amount of metal-sulphides in the ore materials. In this area, these elements can therefore serve as a general indicator for the contamination of natural sediment with either tailings type. The interstitial water analyses indicated that the sediments in both inlets become reducing at a fairly shallow depth (10-15 cm), resulting in the mobilization of Mn, Fe and Mo from the solid phase. In the deeper parts of the cores, consumption of these metals could usually be observed which is likely due to precipitation of authigenic components. The dissolved Mo concentrations in the sediments contaminated with the modern tailings were much higher than in the natural sediments. In the pore water of the pure tailings the concentration reaches some 300 times that of the overlying water, which is the highest value ever reported for saline pore waters. These high Mo concentrations must support a flux of Mo from the sediment into the overlying water, and it is shown that such a flux could increase the inventory of naturally occurring dissolved Mo in the deep waters of Alice Arm by up to 4%. / Science, Faculty of / Earth, Ocean and Atmospheric Sciences, Department of / Graduate

Pythium undulatum var. litorale Hohnk was found to infect Fucus distichus in the Squamish River estuary of southern British Columbia. This thesis adresses the questions of: 1.) whether this symbiosis can be found outside the Squamish River estuary, 2.) relationship of the infection within the estuary to the distribution of P. undulatum var. litorale in estuarine sediments, 3.) taxonomically defining those species associated with Fucus and/or in estuarine sediments, and 4.) the host parasite relationship as determined by means of histochemical and light microscope observations. Results indicated that outside the Squamish River estuary, associations between pythiaceous fungi and Fucus are uncommon in British Columbia coastal areas. Sampling of live and decaying Fucus plants from 10 field stations in British Columbia and Washington yielded only 4 species, the most common isolate being Phytophthora vesicula. Within the Squamish estuary, an association was found to exist between the distribution of P. undulatum var. 1itorale in the sediments and the distribution of infected Fucus plants. Sediment sampling from the Fraser River estuary, where Fucus does not occur, yielded P. undulatum var. litorale, suggesting that the fungus is probably indigenous to estuarine sediments. Numerous other species of Pythium were recovered from estuarine sediments, including P. butler i, P. carolinianum, P. catenulatum, P. gracile, P. torulosum , and P. volutum . Two taxa are described in detail. Pythium undulatum var. litorale was originally described by Hohnk (1953), but the varietal status was rejected by Waterhouse (1967). Arguments are presented for retention of the variety. Pythiogeten utriforme Minden is transferred to the genus Pythium and P. hohnkii is proposed as the nomen nova of this taxon. A discussion of the generic characteristics of the genus Pythiogeten is presented. In order to facilitate an understanding of the infection process by Pythium species, the anatomy and histochemistry of Fucus distichus were examined. Anatomically, F. distichus agrees with earlier reports of other species of Fucus. The internal structure of cells was found to agree with descriptions in earlier publications, although higher physode content was noted in F. distichus. Histochemical staining suggested that cell walls of Fucus are three layered; having an outer fucan-rich layer, a middle layer composed principally of alginic acid, and an innermost layer of cellulose. Several phenolic-indicating reagents were tested on both fresh and fixed/embedded Fucus tissue, resulting in some interesting new observations of phenolics in the matrix. The host-parasite interface of P. undulatum var. 1itorale and F. distichus was also examined by use of histochemistry and the light microscope. Macroscopically, the infection of F. distichus occurs behind the most recent dichotomy, and lesions are necrotic, firm (flaccid with age), and are pink-to-red in color. Microscopically, fungal hyphae are confined to the cortical and medullary regions. Hyphae appear to penetrate host cell walls by means of an enzymatic dissolution of the alginic acid and cellulosic portions of the cell wall. Use of the Periodic Acid/Schiff's reagent shows a distinct non-staining halo at the point where hyphae cross the cell wall. Pit connections between cortical cells were observed to break down with hyphae present in only one cell, suggesting that the fungus is capable of parasitizing several cells via digestion of pits. Gemmae were observed to form in both cortical and medullary cells. The response by Fucus to infection is an active one; a hypersensitivity reaction analagous to that of higher plants is observed. Cells in advance of fungal hyphae are observed to autolyse. Normally metabolically quiescent medullary filaments are observed to have an increase in general protein levels and to have increased physode content. Physodes become polarized within the medullary cells, and coalesce to form larger units, which are then delimited from the producing cell by a cross wall. The fate of these 'giant' physodes was not observed, but it is believed that these cells autolyse and release their phenolic contents to the matrix, as levels of phenolic-reactive material were observed to increase in this region. Coupled with the buildup,of phenolics in the matrix is a decrease in the fucan component of the matrix. Stress and tear lines appear between cells, and eventually this region serves as an abscission zone by which the infected portions are dropped out of the plant. Behind the abscission zone, medullary filaments undergo transverse divisions to form irregular, cuboidal cells which function as epidermis after abscission of the lesion occurs. / Science, Faculty of / Botany, Department of / Graduate

Sharks are considered top predators in many marine ecosystems, and can play an important role in structuring those communities. As a result, it is necessary to understand the factors that influence their abundance and distribution. This is particularly important as fishery managers develop fishery management plans for sharks that identify areas that serve as essential fish habitat (EFH). This includes nursery habitat where sharks are born and juveniles spend the early part of their life. However, our understanding of shark habitat use in the northeast Florida waters is limited. The goal of this thesis was to characterize the abundance and distribution of sharks in northeast Florida estuaries, and to examine the effect of abiotic and biotic factors affecting shark habitat use. A bottom longline survey conducted from 2009 – 2011 indicated that 11 shark species use the estuarine waters of northeast Florida during summer months. Atlantic sharpnose (Rhizoprionodon terraenovae), blacktip (Carcharhinus limbatus), bonnethead (Sphyrna tiburo), and sandbar sharks (Carcharhinus plumbeus) were the most abundant species and made up 87.1% of the total catch. Month, bottom water temperature, and depth were the most important factors determining the presence and abundance of these species. This study also examined the role of prey abundance in determining the abundance of Atlantic sharpnose sharks. The probability of catching an Atlantic sharpnose shark, and the abundance of Atlantic sharpnose sharks, were most influenced by site. Neither potential prey abundance nor preferred prey abundance were not significant factors effecting Atlantic sharpnose abundance. This may be a result of prey sampling not providing an accurate measure of the true availability of prey resources. Other factors, such as predation risk, may better explain habitat use patterns of Atlantic sharpnose sharks. Continued sampling will give a better understanding of the factors influencing shark habitat use in this area.

Thesis: Sc. D., Massachusetts Institute of Technology, Department of Civil Engineering, 1975 / Vita. / Bibliography: leaves 266-271. / Sc. D. / Sc. D. Massachusetts Institute of Technology, Department of Civil Engineering