Two and a half years of data were collected from the Great Fish River and estuary using seine nets and gill nets to determine the fish species composition within these regions. The head region of the Great Fish River estuary was found to have a rich fish fauna, with the euryhaline marine fish component totally dominating the catch in all three regions. Gilchristella aesiuaria was the single most common species and several euryhaline marine species were captured in freshwater for the first time. Physical parameters were also measured in an effort to ascertain the possible factors which may affect the distribution and length frequency of the most common species within this area. Salinity was found to be the single most important factor affecting the species composition, and the sampling area was consequently divided into three regions, viz. river (< I ‰), head (1-4 ‰) and estuary (> 4 ‰). In addition, it was determined that river flow rate during the month prior to sampling also had a profound effect on species composition in all three regions. Based on the available evidence it is suggested that for most species this is related to conductivity levels rather than flow per se.

This investigation had three major objectives; firstly to document the estuarine fishes from several poorly studied Eastern Cape estuaries, secondly, to identify similarities and differences in the fish communities from permanently open and intermittently open estuaries, and thirdly, to establish which physico-chemical or environmental variables are responsible for the structuring of these communities. The study area incorporated 10 estuaries along a 70 km long stretch of the Eastern Cape coastline between the towns of Seafield (33° 32' 42" S, 27° 03' 05" E) in the south-east and Hamburg (33° 16' 45" S, 27° 29' 50" E) in the north-west. The systems investigated included two permanently open estuaries and eight intermittently open systems of varying sizes. Overall fish abundance and species richness in the study area revealed no significant seasonal variation. A contrast between open and closed estuaries was evident in the dominance of species from different estuarine dependence categories. Open estuaries comprised mainly category Ib, lla and IIb species, with a high proportion of category IV species. Closed estuaries were dominated by category IIa species, with few category IV species represented. Clear longitudinal trends in relative abundance were demonstrated for individual species. Margalefs species richness index for seine net catches, and a combination of the seine and gill net data, were significantly correlated with catchment size, mean annual runoff, estuarine area and linear length. Fish assemblages in the permanently open and intermittently open estuaries were significantly different (p<0.001) when analysed on a community and density basis. Similarly, the communities in large and small intermittently open estuaries differed significantly on a community (p=0.01) and density basis (p<0.001). These differences where caused by changes in abundance of the dominant estuarine resident and marine migrant species. The environmental variable that was dominant in accounting for these differences was the estuary mouth status. Estuarine resident species had smaller overall body lengths when compared with the marine migrant species. In addition, the estuarine resident species had smaller modal size classes in the closed estuaries when compared with the open estuaries. These differences may be linked to the foraging strategies of the species, as well as food availability and different levels of predation in the different estuary types. The marine migrant species did not show any trends in this respect although the closed systems had the largest individuals for all four dominant marine species. This finding may be linked to the inability of these species to breed in closed estuaries, thus channelling reproductive energy resources into somatic growth. In addition, prolonged periods of mouth closure prevent the emigration of large individuals back to the sea. Otter trawling was conducted in fiye estuaries and captured mostly demersal species. The otter trawl data confirmed the seine net community analysis, with significantly different communities identified in the permanently open and intermittently open estuaries (p=0.02), as well as in the small and large intermittently open estuaries (p=0.03). The environmental variables responsible for these community differences were mouth status and variables associated with estuarine size. The identification of estuary mouth status by this study as the most influential environmental factor is compounded by this "ariable affecting or being indicative of numerous other physico-chemical features. The large differences in the fish communities in different estuary types indicate the importance of each estuary type to various species. This should be taken into account before allowing any form of commercial fishing in these important nursery areas.

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.

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.

The ichthyofaunal community structure, population dynamics and movement patterns in the small temporarily open/closed (TOCE) Grant’s Valley estuary, situated along the Eastern Cape coastline, were investigated over the period May 2004 to April 2005. Community structure in the littoral zone was assessed, while growth of selected ichthyofaunal species was investigated using the MULTIFAN model. Population size was assessed using mark recapture models and movement within the estuary using the Hilborn (1990) model. Total ichthyofaunal densities and biomass within the littoral zone ranged between 0.31 to 21.45 fish m⁻² and 0.20 to 4.67 g wwt m⁻², with the highest values typically recorded during the summer. Results of the study indicated that the ichthyofaunal community structure within the estuary was closely linked to the mouth phase and the establishment of a link to the marine environment via overtopping events. In the absence of any link to the sea, the ichthyofaunal community was numerically dominated by estuarine resident species, mainly Gilchristella aestuaria and to a lesser extent, by the river goby, Glossogobius callidus which collectively comprised ca. 88% of all fish sampled. The establishment of the link to the marine environment contributed to an increased contribution of marine breeding species (e.g. Rhabdosargus holubi, Myxus capensis and Atherina breviceps) to the total ichthyofaunal abundances. In contrast, total ichthyofaunal biomass was almost always dominated by marine breeding species by virtue of their larger sizes. Results of hierarchical cluster analyses did not identify any spatial patterns in the ichthyofaunal community within the littoral zone. Results of MULTIFAN analysis indicated estuarine resident fish species bred over an extended period with peaks occurring in the summer months. Conversely, marine breeding fish were shown to recruit into the estuary following overtopping and breaching events. Results of the mark-recapture experiment indicated a population of ca. 12 000 (11 219 – 13 311) individuals greater than 50mm SL. Marine-breeding species (R. holubi, Monodactylus falciformis, and two mullet species) numerically dominated the ichthyofauna, possibly as a result of their effective use of overtopping events. The two mullet species, M. capensis and Liza richardsonii, and the Cape stumpnose, R. holubi moved extensively throughout the estuary, while the remaining species exhibited restricted movement patterns possibly due to the preference for refuge and foraging areas associated with reed beds. The observed movement patterns of individual fish species appeared to be associated with both foraging behaviour and habitat selection.

Freshwater fish are the most imperilled vertebrate group with a high projected extinction rate. In general, the world's freshwater ichthyofauna is in a poor state and is threatened by habitat destruction, pollution, the effects of alien species, damming, water extraction and overfishing. Within South Africa, legislation supporting the conservation of aquatic habitats and its associated fauna is strong, but there is a lack of capacity and poor co-ordination between agencies responsible for the conservation and management. The Greater Addo Elephant National Park (GAENP) includes the upper catchments of five of the Sundays River tributaries. Since the optimal method of conserving freshwater aquatic biota is to conserve the headwaters of either the river or its tributaries, there is an opportunity to conserve the native ichthyofauna of the Sundays River system. The principal objective of this thesis was, therefore, to provide information pertinent to the conservation and management of the fishes in the Sundays River system. Sixteen species of freshwater and estuarine fish were sampled, of which eight were alien. Species composition in each of the rivers, with the exception of the Kabouga River, had changed significantly from previous studies. The distribution ranges of several indigenous species had reduced, while those of some alien species was increased. Two species not previously recorded, Labeo capensis and Gambusia affinis, were sampled. Alien species were sampled from all the tributaries. Diversity indices and community assemblage models were identified as approaches for monitoring the ichthyofauna in this species depauperate system. Diversity indices were not recommended for use as a monitoring tool, as they provided insufficient detail on community composition and structure. A modified depletion estimator was applied to multiple pass electrofishing data to determine the effects of various physico-chemical variables on the absolute abundance of Pseudobarbus afer, an endangered species. Pseudobarbus afer density was found to be positively correlated to the presence of boulders. Where P. afer were present, they were abundant. Of the total sampled catch, 21.8% was comprised of alien fishes. Clarias gariepinus was the most successful invader having penetrated all but one of the tributaries. Micropterus salmoides changed the species composition of the Wit River significantly through predation, and is potentially the most dangerous of the alien species in the Sundays River system, capable of extirpating a population of P. afer in as little as three days. Only one man-made barrier to the upstream migration of alien fishes was present on the tributaries, and several addition barriers are required to safeguard the populations of indigenous fish in the headwaters of these tributaries. Several management and conservation tools are presented and discussed, including river rehabilitation, translocation, eradication of alien species, erection of barrier weirs and captive breeding programmes. Public awareness and education is stressed because without support from the communities surrounding the GAENP, management plans for conservation of the indigenous ichthyofauna will, in all likelihood, fail.

The introduction, spread and concomitant impacts of non-native species are a global problem. Fish are among the most widely introduced vertebrate groups, with their impacts affecting multiple levels of organisation- from individuals, populations and communities, to entire ecosystems. In South Africa, the largest perceived threat to range-restricted endemic headwater stream fishes is said to be invasion by non-native fishes, however, as is the case elsewhere, invasive impacts are often a case of risk perception rather than actual risk analysis. Two range-restricted headwater species, the Eastern Cape redfin Pseudobarbus afer and the Border barb Barbus trevelyani are redlisted by the International Union for the Conservation of Nature (IUCN) as ‘Endangered’, primarily due to invasion by non-native fishes. To investigate invasions in South Africa, and provide a quantitative estimate of the impact of non-native fishes on the two imperilled endemics, P. afer and B. trevelyani, the overall aims of this thesis were to: (A) Provide a literature review on non-native fish invasions in South Africa; (B) Using two case studies on the headwaters of the perennial Keiskamma and episodic Swartkops River systems, investigate the naturalisation-invasion continuum to provide a holistic view of the invasion process in these variable environments. The specific thesis objectives were: (1) Reviewing current knowledge of invasive impacts of non-native fishes in South Africa; (2) Investigating invasibility of headwater stream environments by non-native fishes; (3) Determining the establishment success of non-native fishes, (4) Assessing the spatial and temporal impacts of invasion; (5) Understanding mechanisms responsible for non-native fish impacts; (6) Investigating the threat of non-native fish invasion on the genetic diversity of two the two headwater fishes, P. afer and B. trevelyani. Results from the literature review of fish invasions (Chapter 1) showed that South Africa has a long history of non-native fish introductions, spanning two and a half centuries. Currently, 55 species have been introduced or translocated. Many of these introduced species have become fully invasive (36%). Their impacts also span multiple levels of biological organisation. There was a general paucity of studies on fish invasions (38 studies), however, of those conducted, reviewed studies placed emphases on invasive impacts (25 studies) and the transport, introduction, establishment and spread stages of the invasion process were largely ignored. The two study systems, the Swartkops and Keiskamma Rivers, were heavily invaded and numbers of introduced species surpassed that of natives (Chapter 2, 3 and 5). Headwater streams had varying invasibility and a number of non-native species were successfully established (Chapter 2, 3, 5 and 6). The remainder of the invasions were casual incursions into headwater streams from source populations in mainstream and impoundment environments which were invasion hotspots. Irrespective of establishment, four predatory invaders (largemouth bass Micropterus salmoides, smallmouth bass M. dolomieu, brown trout Salmo trutta and rainbow trout Oncorhynchus mykiss) impacted heavily on native fish communities (Chapter 3, 4 and 5). Two broad types of invasion were documented, top down invasion by non-native O. mykiss and S. trutta and upstream invasion by M. salmoides and M. dolomieu (Chapter 3 and 5). Their impacts included changes in community structure, extirpation from invaded stream reaches resulting in contracted distribution, and isolation and fragmentation of native fish populations. The impacts of non-native predatory fishes were particularly acute for P. afer and B. trevelyani. Where non-native predatory fish occurred, P. afer and B. trevelyani had been extirpated (Chapter 3 and 5). As a result both native species exhibited contracted distributions (>20% habitat loss due to invasion). Upstream invasion by centrarchids isolated and fragmented P. afer populations into headwater refugia, while top down invasion by salmonids excluded B. trevelyani from invaded, more pristine stream reaches, by forcing the species into degraded unsuitable lower stream reaches. Predation also disrupted population processes such as adult dispersal for P. afer, and centrarchid-invaded zones acted as demographic sinks, where adults dispersing through invaded reaches were rapidly depleted. While the Mandela lineage of P. afer exhibited little within or between drainage genetic structuring, B. trevelyani was >4% divergent between drainages, and up to 2% divergent between streams within the Keiskamma River system (Chapter 7). The distribution of genetic diversity for B. trevelyani also indicated that the loss of diversity was imminent without immediate conservation interventions. This thesis has provided conclusive evidence that native fishes are vulnerable to invasion and that non-native predatory fishes have significant impacts on native fishes in Eastern Cape headwater streams. If management and conservation measures are implemented, the unwanted introduction and spread of non-native fishes may be restricted, allowing native fishes opportunities for recovery.

The spatial and temporal pattern of ichthyofaunal community composition in relation to selected physico-chemical (temperature and salinity) and biological variables (chlorophyll-a and zooplankton) was investigated at ten stations in the temperate temporarily open/closed Kasouga estuary. In addition, the food web structure in the estuary was investigated using stable carbon isotope analysis. Results of the 5 metre seine net survey indicated that ichthyofaunal composition and biomass in the Kasouga estuary was largely determined by seasonality and mouth condition. Maximum abundance and biomass of ichthyofauna was recorded during summer or during those periods when overtopping occurred. Overtopping coincided with the recruitment of marine estuarine dependant species, which dominated the catches both numerically and in biomass. The recruitment of these species resulted in an increase in diversity of the ichthyofaunal community. There were no significant spatial patterns in the distribution of smaller ichthyofauna (<50mm SL) identified in Bray-Curtis similarity matrices using cluster analysis (Primer 5 v5.2.4). Spearman’s correlation analysis revealed that there were no significant correlations between abundance and biomass of ichthyofauna and selected physico-chemical and biological variables other than salinity (P>0.05 in all other cases). There were two distinct spatial patterns in the distribution of the larger ichthyofauna (>50mm SL). These corresponded to a grouping associated with the mouth region and a grouping associated with the remaining regions of the estuary. Stable isotope analysis indicated that the primary source of carbon utilised by the ichthyofauna of the Kasouga estuary was derived from the channel, most likely microphytobenthic algae. The contributions of the riparian and salt marsh vegetation to the total carbon flow appear to be minimal.

The Mngazana and Mngazi estuaries are situated just south of Port St. Johns close to the biogeographical boundary between the warm temperate and subtropical zones along the east coast of Southern Africa. The Mngazana estuary is a permanently open estuary with three species of mangrove. The Mngazi estuary is a temporarily open/closed system. The ichthyofauna was sampled with seine nets and gillnets during June 2005, August 2005, November 2005, February 2006 and April 2006. The fish community structure was significantly different between the two estuaries with the differences being especially noticeable in the larger fish sampled with gillnets. There were significant differences in the fish community sampled with the seine net on both spatial and temporal scales in both of the estuaries. Certain warm temperate species were less abundant and certain subtropical species were more abundant at the Mngazana estuary than they were during a study conducted in 1975. Stomach content analysis revealed that most species had similar diets in the two estuaries with the only differences occurring among benthic feeding species. The diet of sub-adult piscivores comprised of a large proportion of penaeid prawns while the juveniles feed mainly on mysids. The zooplanktivorous fish feed mainly on copepods and mysids with the proportions varying between the species. Benthic feeding fishes showed the greatest variability in diet with plant material, detritus, crabs and copepods being the main food sources. Stable isotopes revealed that particulate organic matter and microphytobenthos were the most important carbon sources to fish in the Mngazi estuary. In the Mngazana estuary a combination of mangrove material, detritus, particulate organic matter and Zostera material were the most important carbon sources. The importance of each of these sources did, however, vary greatly between the different trophic guilds of the fish in the Mngazana estuary. The distribution of certain fish species was investigated in relation to physical parameters (temperature, salinity, turbidity and depth) and biological parameters (prey and predators) in the environment. It was found that most small fish (< 100 mm SL) were strongly correlated with turbidity where-as they showed weak correlations with their prey. Sub-adult piscivores fell into two groups, i.e. those that could tolerate turbid water and those that were only found in clear water. The group that also occurred in turbid water showed very strong correlations with their prey but the clear water group did not. It would appear that the small and thus vulnerable fish seemed to occur in areas that offered them protection rather than areas where their food supply is most abundant, where-as the piscivores were more influenced by prey distribution, as long as the physical conditions were tolerable to them.

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.