The biogeography and community structure of the fishes of South African estuaries was investigated. In all, 109 systems were examined representing two broad types: temporarily closed and permanently open estuaries. Multivariate analyses of the fish communities identified three biogeographic regions. A cool-temperate region extended along the west and southwest coasts; a warmtemperate zone stretched along the south, southeast and east coasts and a subtropical region occurred along the east coast. The boundaries of these biogeographic regions were also delineated. The general physico-chemical characteristics of the estuaries within the three biogeographic regions also reflected regional differences in climate, rainfall and ocean conditions. Estuarine temperatures followed the trend for marine coastal waters, decreasing from subtropical estuaries toward cool-temperate systems. The low rainfall and runoff in the warm-temperate region together with high evaporation rates and strong seawater input resulted in higher salinities in these estuaries. These factors also accounted for the predominantly clearer waters in warm-temperate estuaries. The estuaries in the three biogeographic regions were also shown to contain somewhat distinctive fish assemblages. Temperature and salinity appeared to be the two main factors affecting the distribution and abundance of fishes in South African estuaries. Subtropical systems were characterised by fishes mostly of tropical origin as well as certain south coast endemic species. Warm-temperate estuaries were dominated by endemic taxa with some tropical species also present. The fish fauna of cooltemperate estuaries mostly comprised south coast endemic species with cosmopolitan and temperate taxa also present. Certain functional components of the ichthyofauna also exhibited slight differences between regions. Freshwater fishes were a major component of closed subtropical estuaries while estuarine resident species were more abundant in warm-temperate estuaries. Overall, estuarine-dependent marine species dominated the fish fauna of the estuaries in all biogeographic regions, signifying that South African estuaries perform a vital nursery function for this group of fishes. Slight differences were also apparent in the trophic structure of the fishes; these were related to environmental differences between regions. Zooplanktivores and fishes that feed on aquatic macrophytes/invertebrates assumed a relatively higher importance in warm-temperate systems. Overall, detritivores dominated the estuarine fish fauna in all regions, indicating that detritus forms the main energy source in South African estuaries.

Temporarily open/closed estuaries (TOCEs) are the dominant estuary type in South Africa. These systems are often characterized by extensive beds of submerged macrophytes, which form important foraging and shelter habitats for fishes, especially for estuary-dependent fish species such as the Cape stumpnose Rhabdosargus holubi and Cape moony Monodactylus falciformis that are commonly associated with them. A loss of submerged macrophytes from an estuary has been shown to affect the fish community as well as reducing overall system productivity. The TOC East Kleinemonde Estuary, situated in the warm-temperate biogeographic region of South Africa has been subject to an ongoing long-term fish monitoring project since 1995. During the period 1995 to 2002, this estuary contained large beds of the submerged macrophytes Ruppia cirrhosa and Potamogeton pectinatus. However, subsequent to a major flood event in 2003 these macrophytes have been largely absent from this system. The effect of the loss of submerged macrophytes on the East Kleinemonde fish assemblage was investigated through an analysis of seine and gill net catch data. Seine net catches for a 12 year period, encompassing six years of macrophyte presence and six years of macrophyte senescence, revealed changes in the relative abundance of certain fish species. Vegetation-associated species such as R. holubi and M. falciformis decreased in abundance whereas sediment-associated species, especially members of the family Mugilidae, increased in abundance following loss of the macrophytes in this estuary. The critically endangered pipefish Syngnathus watermeyeri was only recorded in catches during years in which macrophyte beds were present. In addition to the analysis of catch data, the importance of macrophytes as a primary energy source for selected estuarine fishes was explored through the analysis of carbon and nitrogen stable isotopes. Prior to conducting these analyses, common methodological practices to address the presence of carbonates and lipids within isotope samples were evaluated. A subset of samples were either acid washed to remove carbonates, or lipids were removed according to the method of Bligh and Dyer (1959) as both of these compounds have been shown to affect stable carbon and nitrogen isotope ratios. The suitability of the lipid normalization models of Fry (2002) and Post et al. (2007) for samples of estuarine fish muscle were also tested. Based on this evaluation both models are suitable for use with estuarine fish muscle tissue, however since neither carbonate nor lipid content of any of the samples used in this study was high all samples were left untreated in the following analysis. Carbon isotope ratios from a wide range of fish species collected from the East Kleinemonde Estuary during the macrophyte-senescent phase were compared with individuals of the same species from the neighbouring West Kleinemonde Estuary (where extensive beds of R. cirrhosa and P. pectinatus were present) and revealed the influence of submerged macrophyte material in the diet of fishes in the latter system. However, it was apparent that these plants are not directly consumed but rather contribute to the detrital pool that forms a food source for most invertebrate and some fish species. The most significant source of carbon for East Kleinemonde fishes during the macrophyte senescent phase appeared to have a more depleted origin; probably from benthic or pelagic microalgae. In conclusion, while the importance of macrophyte beds as shelter and foraging habitats for estuarine fishes are well documented, their role in terms of the structuring and functioning of fish assemblages in TOCEs remains somewhat uncertain. The findings of this study were possibly masked by the resilience of vegetation-associated species to the loss of this habitat, as well as by life history characteristics of species such as R. holubi that allow their numerical dominance despite habitat change. Nonetheless, macrophyte senescence in the East Kleinemonde Estuary resulted in the loss of at least one species and the reduced abundance of vegetation-associated species, probably reflective of reduced food resources and/or increased vulnerability to predation. As a result, beds of submerged macrophytes are an important habitat within TOCEs.

Introduction. On the coast of the Bathurst Division, Eastern Cape Province, some 42 miles south- east of Grahamstown, lie two estuaries opening within a few yards of one another. These are known as the Kleinemonde. This paper embodies the results of an ecological survey of the fauna of the Larger Kleinmond Estuary. The chief concern has been the aquatic fauna but animals living along the banks have also been dealt with as far as possible, as have the birds. This is the second of South Africa's blind estuaries to be studied, the Klein River, Hermanus, having been investigated in 1947-49 by workers from the University of Cape Town. Though common in South Africa, blind estuaries are rare in parts of the world such as Europe, and present an almost entirely new field of research for the ecologist. It is hoped, therefore, that the investigation of the Kleinmond Estuary and a comparison with the results obtained from the Klein River, Hermanus, will add to our knowledge of this subject and be of service to future investigators. The history of the Kleimnond River is worthy of mention, for the lagoon was not always blind and shallow. The early maps of the 1820 settlers show the river as a navigable stream; sailing vessels anchored out at sea and sent lighters up what was then known as the Lydendoch River to collect fresh meat. The area at the mouth of the river was called Port Jessie. In these early days the region abounded with game. Hippo and elephant were common. the former inhabiting deep pools in the upper reaches of the river; pools which have since completely silted up. Constant ploughing of the slopes adjoining the river has caused vast amounts of soil to be washed by rains into the stream, and today what was within living memory a deep, clear stream is a shallow, blind and muddy river, a reflection of man's destructive methods of farming. These changes must have had a far-reaching effect on the fauna of the estuary; an effect about which we can, alas, only speculate. It is perhaps significant that though rich in numbers, the fauna is poor in species, tunicates and echinoderms being apparently entirely absent. The work involved in this survey was undertaken between September 1952 and November 1953, under the direction of Professor J . Omer-Cooper of Rhodes University, frequent and regular visits being made to the estuary. The entire month of February, 1953, was spent on the shore of the lagoon, so that animals could be observed over a continuous period. The starting date of the survey was particularly fortunate, as on September 28th, 1952, the river came down in flood and broke through the sand-bar separating the lagoon from the sea. The work thus began at a definite point in the estuarine cycle, the importance of which it is hoped to show.

Aspects of the life history of Myxus capensis which were likely to be most affected by man-induced habitat changes, were studied in the eastern Cape. Myxus capensis was found to be specialized for a catadromous life history. The fry migrate into freshwater reaches of rivers within a few months of being spawned at sea and return to an estuarine or marine environment when maximum size has been attained and large energy reserves necessary for spawning have been accumulated. Myxus capensis penetrates considerable distances up eastern Cape rivers and although present in the upper reaches of some estuaries, is only common in freshwater-dominated systems. Myxus capensis has adapted its life history style in a number of ways to overcome the constraints associated with the spatial separation of the freshwater feeding areas and the marine spawning grounds in a region prone to droughts and erratic river flow. Research on the more estuarine-dependent Mugil cephalus, which is also found in the freshwater reaches of eastern Cape rivers, enabled useful comparisons to be drawn. As the latter species is not dependent on a freshwater phase in its life cycle, it is subject to different selection pressures. This comparative approach gave insights into the adaptive significance of the life history tactics of Myxus capensis. Myxus capensis displays a number of life history tactics characteristic of migratory fish species which are thought to increase population fecundity by increasing the number of large females and hence maximizing egg production. These include a sex ratio in favour of females, faster female growth, females remaining longer in the freshwater feeding areas and reaching a larger ultimate size. Possible adaptations to the periodic isolation of the feeding and spawning areas include: (i) the development of ripe gonads only in estuarine or marine areas when access to the spawning grounds is ensured; (ii) year-round spawning and hence recruitment which enables the erratic floods to be used at any time for downriver spawning migrations and reduces the chances of total recruitment failure due to short-term adverse environmental conditions and (iii) a relatively wide range of size and age at first (and final) sexual maturity (2+ to 5+ years); this tends to ensure successful reproduction and recruitment in spite of a series of poor years, as well as dampen population fluctuations after long periods of isolation in fresh water. The dependence of Myxus capensis on the freshwater reaches of eastern Cape rivers makes this species vulnerable to man-induced habitat changes. The erection of barriers to migration has cut off large stretches of suitable habitat and almost completely eliminated Myxus capensis from certain rivers. This detrimental effect has been largely responsible for the recent inclusion of this species on the Red Data list of endangered fish species in South Africa. This study demonstrated the considerable fisheries potential of both mullet species and this should ensure that their conservation and wise utilization be given high priority. Management proposals include: (i) the netting and lifting of Myxus capensis over barriers to migration; (ii) the construction of fish ladders when feasible and (iii) the artificial propagation of this species to enable large-scale stockings of existing man-made impoundments as well as natural habitats to be carried out. It is predicted that the life history characteristics of Myxus capensis, which evolved in response to an erratic freshwater environment, should ensure that when man-induced habitat changes are rectified, this species will again flourish in eastern Cape rivers.

Early stage fishes, namely larvae and early juveniles, were collected from 12 estuaries and associated habitats in the warm temperate region of South Africa between July 1998 and December 2000. This study served to provide new information and expand on existing knowledge of early stage fish assemblages associated with various types of Eastern Cape estuaries. A total of 65 536 predominantly postflexion fish larvae were collected. Together with early juveniles, taxa comprised 72 species from 25 teleost fish families. Surf zones associated with two intermittently open estuaries showed that estuary-dependent marine species predominated in this zone. Early stage fishes responded positively during natural estuary opening events and concentrated along the estuary outflow plume, suggesting that cueing from estuary and/or river water may be taking place. A habitat study in the Swartkops Estuary, using light traps, showed that newly recruited larval fishes concentrated along the margins of the estuary and unlike their older juvenile counterparts, were poorly represented in eelgrass beds. A multi-estuary comparison, including seven permanently open and five intermittently open estuaries, indicated that early stage fish assemblages were more diverse than indicated by past investigations. Early stage fishes were also shown to concentrate in the mesohaline regions of these estuaries. Studies of estuaries with altered freshwater flow regimes were also included in the research. A regulated release of dam water in the euhaline Kromme Estuary failed to induce a cueing response from estuary-dependent marine fish larvae. This project showed that large amounts of freshwater are required to reverse the negative effects of river impoundments. The release study did show that estuary-resident fishes were sensitive to small natural pulses of freshwater and responded by spawning. However, excessive river supply through an inter-basin water transfer scheme was shown to have a flushing effect on the larvae and early juveniles of the estuary-resident Gilchristella aestuaria in the upper reaches of the Great Fish Estuary. This effect was evident in comparisons with the freshwater starved Kariega Estuary where concentrations of G. aestuaria, a species usually more abundant in freshwater rich estuaries, were considerably higher. This finding suggests that freshwater minima and maxima be considered when assessing and managing estuarine systems.

The annual biomass of fish caught from estuaries in South Africa is currently estimated at over 24,800 tons. These estuarine fishes are caught by over 73,000 fishers, most of them recreational. Annual income derived from South Africa’s total estuarine fishery was worth approximately R430,000,000 in 1997. There is increasing concern that unless our estuarine fisheries are effectively managed, we will not be able to sustain these benefits into the future. Two factors that contribute to inadequate management of the estuarine fisheries in South Africa are a lack of data on which to base management decisions, and the lack of indicators by which to assess trends towards sustainability. The main aims of this study were to provide a description of the Kowie estuary fishery, identify suitable indicators of sustainability for this fishery, and assess its sustainability. Boat-based and shore-based roving creel surveys were carried out on the Kowie estuary between July 2000 and June 2001; 1,091 interviews were conducted with linefishers, and 277 interviews with bait collectors. In the boat-based interviews, data were collected on fisher demographics, fishing site, fishing method, choice of bait, fishing duration and catch statistics. In the shore-based surveys, additional data were collected from shore-based linefishers and bait collectors on their perceptions, attitudes, and knowledge of fishery regulations. Total annual fishing effort on the Kowie estuary was estimated at 30,952 angler hours (SD=154); 84% of it recreational, and the rest subsistence. Most fishing occurred during December and January, and decreased during winter, especially June and July. The annual yield of fish from the estuary was estimated at 16,240 fish (SD=667) or 5.99 tons (SD=0.81). By number, recreational anglers caught 69% of the annual catch. Three species dominated the catch by number: Rhabdosargus holubi (62%), Pomadasys commersonnii (17%) and Argyrosomus japonicus (7%). By mass, the dominant species caught were Argyrosomus japonicus (60%) and Pomadasys commersonnii (19%). Overall catch rate on the estuary was 0.57 fish.ang.⁻¹h⁻¹ (SD=0.24), or 0.298 kg ang.⁻¹h⁻¹ (SD=0.31). Overall catch rate by number was highest in the subsistence sector at 1.13 fish.ang.⁻¹h⁻¹(SD=0.70), while the boat-based recreational sector recorded the highest overall catch rate by mass (0.427 kg.ang.⁻¹h⁻¹, SD=0.625). Argyrosomus japonicus had the highest overall catch rate by mass on the estuary (0.496 kg ang.⁻¹h⁻¹), and Rhabdosargus holubi the highest overall catch rate by number (1.233 fish.ang.⁻¹h⁻¹). Only 19% of the catch of R. holubi was above the minimum legal size, while the estimates for P. commersonnii and A. japonicus were 21% and 25%, respectively. The annual number of bait collecting outings on the estuary was estimated at 2,889, of which 75% were subsistence. The highest numbers of bait collecting outings were recorded in December and April. The Bay of Biscay was the most popular site for bait collecting. A total of five invertebrate species were collected from the estuary to be used as bait, of which the mud prawn Upogebia africana was the dominant species. Total annual number of mud prawns collected from the estuary was estimated at 260,648; of which 41% was collected by subsistence bait collectors. Thirteen indicators were selected to assess sustainability in three fishery sectors on the Kowie estuary: namely, the shore-based recreational linefishery, the subsistence linefishery and the subsistence bait fishery. Social sustainability was evaluated on the basis of the use fishery resources to fulfil Maslow's basic human needs of food and employment, safety and security, affiliation, self-esteem and selfactualisation. Indicators of ecosystem sustainability assessed the productivity, diversity, disturbance and degree of water quality in the estuary. Institutional sustainability was assessed on the basis that management systems in the fishery should be results-oriented, consent-based, truth-seeking and adaptable. Data on indicator performance was collected during the shore-based roving creel survey, and from published literature. Arbitrarily set reference points were used to assess indicator performance, which was graded on a scale from 1 (indicating minimum probability of sustainability) to 4 (indicating maximum probability of sustainability). Sustainability was illustrated with the aid of amoeba plots. Overall sustainability was low in all three fishery sectors investigated. Nine of the 13 indicators in the shore-based recreational fishery performed poorly, while 11 of 13 in the subsistence line fishery, and 10 of 13 in the subsistence bait fishery, performed poorly. In all three fishery sectors all four selected indicators of institutional sustainability performed poorly. The probability of social sustainability was higher in the shore-based recreational line fishery, where the performance of two of the five selected indicators was very good. The probability of ecological sustainability was lowest in the shore-based recreational linefishery, while in the subsistence linefishery only one selected indicator performed very well. Recommendations made towards assessing sustainability in small-scale estuarine fisheries include the formulation of national policy for assessing sustainability in fisheries, the involvement of fishers in the assessment process, use of fisher perceptions where data gaps exist, and the use of research results to guide future management decisions. Management changes recommended for the Kowie estuary fishery include the formulation of an effective and integrated management plan, identification of the key stakeholders in the fishery, inclusion of fishers in management, the protection of the estuary’s Zostera capensis beds, and the establishment of a programme to increase research and monitoring in the fishery.

Many of the larger estuaries along the South African coastline are highly degraded due to human impacts, highlighting the need to conserve pristine systems. There are approximately 100 micro-estuaries along our coastline which have not yet been identified or classified and hence their ecological functioning is unknown. The aim of this study was to identify and classify micro-estuaries along the Eastern Cape coast. Specific objectives of the study were to: 1) map the distribution of estuarine habitats in a selection of micro-estuaries and assess changes over time; 2) measure the physico-chemical characteristics of the micro-estuaries in the field; 3) investigate the distribution of vegetation, identify macrophyte species and habitat integrity and classify the micro-estuaries sampled using abiotic and botanical characteristics. A total of 30 systems were identified of which 10 were selected for sampling which took place seasonally (2015-2016). A relationship between open water surface area and short term rainfall was only evident in Cunge, Sandy Bottom and Ichabana. In the field it was observed that after heavy rainfall, open water surface area increased and mouth breaching occurred. Salinity was the only water column characteristic that showed differences; salinity increased in response to over wash events. Sand was dominant in all systems and there were no differences in sediment characteristics. A total of 8 macroalgal species, 16 estuarine macrophyte species, six freshwater species and 72 riparian plant species were found. Seed bank reserves were low in all systems despite them being vegetated. Only abiotic characteristics were used for classification using Principle Components Analysis since there were no botanical differences between systems. Micro-estuaries were larger (> 0.03 ha), had a higher salinity (oligohaline, mesohaline and polyhaline conditions) and a constant water body. Micro-outlets were smaller (< 0.03 ha), had a lower salinity (oligohaline conditions were most common) and showed periods of drying. Five micro-estuaries (Shelbertsstroom, Kwesani, Cunge, Mtwendwe and Mtendwe) and five micro-outlets (Stromatolite, Sandy Bottom, Ichabana, Palm Tree and Black Rock) were classified. This study has shown that the micro-estuaries along the Eastern Cape coast are pristine (i.e. intact riparian zone and catchment), untouched and highly dynamic systems (i.e. great variation in water surface area and mouth condition) highlighting their need for protection.

The value of mangrove habitats as fish nurseries was assessed by comparing communities of early stage and juvenile fishes between estuaries with and without mangroves. Early stage fishes were sampled using boat-based plankton towing while juveniles were sampled by seine netting. Sampling took place at five sites spaced 1 km apart starting near the estuary mouth in four estuaries along the temperate coastline of the Eastern Cape Province of South Africa. Four estuaries were selected based on shared similarities which included catchment area, estuarine area and shared habitats barring the presence of mangroves which occupied the river margins of two systems. Results revealed that early stage and juvenile fish communities (both marine- and estuary-spawned) were similar between systems with and without mangrove habitats. Differences in fish communities among estuaries were rather attributed to axial salinity gradients associated with greater freshwater input, while season and temperature produced significant variances in fish densities with Generalised Additive Models revealing responses of communities to these variables. A common estuarine-dependent fish, Rhabdosargus holubi (Family Sparidae), was further investigated to determine habitat use, residency and dietary patterns in different mangrove habitats. High habitat residency in this species was revealed during a short-term tagging study using Visible Implant Elastomer tags and long-term isotope analysis in juveniles sampled from two contrasting mangrove habitats. A wider feeding niche was observed in an eelgrass-red mangrove connected habitat when compared with more exposed white mangrove areas. Low dependence on mangrove habitats in temperate estuaries is likely due to their tidally dominated inundation and limited refuge potential due to smaller area coverage by mangroves in temperate estuaries. Relatively lower primary productivity in warm temperate mangrove areas, relative to their tropical counterparts, provides no significant feeding advantage or refuge opportunities relative to other available habitats in these estuaries. Warm temperate estuaries, which are both spatially and temporally highly variable, instead host species which are habitat generalists, able to capitalise on these highly dynamic environments.

Estuaries require sufficient quantity, quality and appropriate timing of freshwater inflow, referred to as environmental flow requirements, to ensure adequate health and functioning. In South Africa the environmental flow requirements of estuaries is determined using Resource Directed Measures (RDM). The present health and importance of an estuary must be considered when determining the ecological flow required to maintain an estuary in its desired state. An Estuarine Health Index that quantifies changes in abiotic and biotic components from natural conditions to present day is used. Health of biotic components is assessed according to changes in species richness, abundance and community composition. There has, however, been debate regarding the calculation of these attributes. In particular, for macrophytes, the inclusion of all habitat within the Estuarine Functional Zone (i.e. 5 m topographical contour) would affect the health and changes over time determined in past assessments. This is due to different areas being included as different areas would be included The aim of this study was to test the validity and suggest improvements to South Africa’s RDM macrophyte health score determination. The health of macrophytes were assessed at varying levels of intensity from desktop studies to thorough field studies. Rapid field studies provided a visual estimate of macrophyte health; while the intermediate and comprehensive assessments quantified change by mapping the distribution of macrophyte habitats from aerial photographs. These approaches were applied to the 64 estuaries within the Mvoti-Mzimkulu Water Management Area (WMA 11), situated in KwaZulu-Natal (KZN) on the subtropical east coast. Historically, these estuaries have supported restricted macrophyte habitats. Consequently, estuaries that presently or historically supported range limited habitats such as mangroves and swamp forest are considered important. Submerged macrophytes are scarce in KZN estuaries due to siltation and low turbidity and thus estuaries supporting this macrophyte habitat are also important. Based on available literature important estuaries were highlighted and selected for field studies. Transects spanning from the estuary water channel to the boundary of the EFZ provided a generalised distribution of KZN vegetation along an elevation gradient. The transition from estuarine to terrestrial vegetation can be used to improve the current EFZ boundaries, as estuarine area has been found to occur outside of the 5 m contour in some estuaries. Results from the desktop assessment indicated that most of the estuaries are moderately modified. There was a 50 % similarity in the macrophyte health scores determined by the desktop assessment to the 2011 National Biodiversity Assessment (NBA). Field studies mostly confirmed the desktop assessment aside from seven estuaries that had different scores. There has been a significant loss of macrophyte habitat with 100% loss of certain habitats from 9 of the 22 estuaries for which there were field assessments. Submerged macrophytes were not found at any of the estuaries that were assessed in the field. Black mangroves, Bruguiera gymnorrhiza (L.) Lam., were rediscovered at Ngane Estuary, however the few individuals could not be considered a ‘mappable’ community (i.e. <0.5 ha). Mtamvuna and Mkomazi estuaries both supported small stands of mangroves that, compared to data from 2006, appeared healthy. Macrophyte habitats and surrounding coastal forest matched the species composition described in the Vegetation Map of South Africa. The freshwater mangrove or Powder puff tree, Barringtonia racemosa (L.) Roxb, was not abundant in the estuaries. It was only found at four (Damba, Fafa, Little Amanzimtoti and Mvoti) of the estuaries that were assessed in the field. Important estuaries, from a botanical perspective, included some of the larger estuaries such as Mgeni, Durban Bay and Sipingo. Smaller, healthier estuaries that presently support mangrove and swamp forest habitat were also identified as important from a botanical perspective. Macrophyte habitats have mainly been lost due to non-flow related pressures. Sugarcane cultivation occurred within the floodplain of 25% of the estuaries. Nutrient enrichment was evident for 12.5% of the estuaries and reed encroachment was evident for 27% of estuaries for which there were field assessments. Development, aside from the N2 road bridges, was evident for 40% of the estuaries assessed in the field. Invasive plant species, including aquatic invasive plants, were present in all estuaries assessed in the field. The extent of invasive plant species was related to the degree of disturbance and surrounding land use pressures. The Mkomazi and Mvoti estuaries that were assessed at a comprehensive level, as they have earmarked for further water abstraction, had the poorest macrophyte health scores. The macrophyte health of these estuaries was much lower than previously determined by the NBA. The decline in health was attributed to the removal of macrophyte habitat for sugarcane cultivation and development as well as displacement by invasive plant species. The areas covered by the macrophyte habitats also differed from the NBA highlighting the importance of updated mapping and ground truthing. Largely differing macrophyte health scores were produced when attributes were calculated using different combinations of macrophyte habitats. Results indicate that only macrophyte habitats and no other floodplain vegetation should be included when calculating abundance. All macrophyte habitats, physical habitats and floodplain should be incorporated when calculating community composition. The minimum of these attributes is used as the overall macrophyte health score as a precautionary approach is followed. Scoring of health is subjective and the only benchmark for determining the most appropriate method is comparion with previous RDM studies. This study illustrates the need for a standardised RDM scoring method that is presented in a manner that ensures the same results irregardless of the speciliast conducting the study. The updated estuary health and importance scores for WMA 11 are necessary to inform management, particularly as few of the estuaries receive formal protection. In conclusion, this study contributed to the limited knowledge of the estuaries of WMA 11 and assisted in determining appropriate methods for assessing the health and importance of estuary macrophytes.

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).