Invasions of ecosystems by exotic species are increasing and they may often act as a significant driver of the homogenization of the Earth’s biota, resulting in global biodiversity loss. Moreover, the addition of exotic species may have dramatic effects on ecosystem structure and functioning which may result in the extirpation of indigenous species. In 2004, a large population of an unknown lymnaeid was found in the Amatikulu Hatchery, northern KwaZulu-Natal, South Africa, and was subsequently found in few garden fish ponds in Durban. In 2007, it was identified using molecular techniques as Radix rubiginosa (Michelin, 1831) – a species widespread in southeast Asia. An invasion by R. rubiginosa is however likely to go unnoticed because its shell morphology resembles some forms of the highly variable and widely distributed indigenous lymnaeid, Lymnaea natalensis Krauss, 1848. Accurate and “easy” species identifications would permit the ready assessment of introduction histories and distributions, but in the present case identification was difficult due to unclear and contradicting accounts of the indigenous L. natalensis in the literature. A redescription of L. natalensis with emphasis on conchological and anatomical characteristics was therefore presented. This will help to distinguish variation between R. rubiginosa and L. natalensis and also assist those carrying out rapid bioassessment (SASS) surveys in South African rivers in recognising R. rubiginosa should it spread. For this, shells of R. rubiginosa and L. natalensis from both the UKZN Pond and the Greyville Pond were selected into either size class 1 (shell length < 10 mm) or size class 2 (shell length ≥ 10 mm). Six shell characters, shell length (height), shell width, aperture length (height), aperture width, length of last body whorl and spire height for each specimen was measured and analysed using principal component analysis (PCA) and The invasive potential of the freshwater snail Radix rubiginosa recently introduced into South Africa discriminant functions analysis (DFA). The most useful discriminant conchological characters were shell length, length of the last body whorl and aperture width. Use of these shell characters provided simple yet effective criteria for the separation of R. rubiginosa and L. natalensis. For both size classes R. rubiginosa had larger, more broadly ovate shells with longer (higher) body whorls than either of the two populations of L. natalensis that exhibited smaller, elongated shells with shorter (lower) body whorls. Also, R. rubiginosa had a narrower aperture width compared to the larger, wider aperture of the UKZN Pond L. natalensis population. The Greyville L. natalensis population was found to have narrower apertures than both R. rubiginosa and L. natalensis (UKZN Pond). The morphology of the radula and the reproductive anatomy of R. rubiginosa and L. natalensis from both the UKZN and Greyville Ponds showed little variation. The species did however vary in the relative numbers of radula teeth in each field and this serves as an additional useful diagnostic character. Both L. natalensis populations had similar mantle pigmentation patterns but that of R. rubiginosa was different. The mantle surface of R. rubiginosa was mottled black with patches of pale white to yellow. There were also large unpigmented fields and stripes that were not observed in L. natalensis. Having found characters to conveniently separate the alien R. rubiginosa from the indigenous L. natalensis, it became increasingly important to assess the potential invasiveness of this introduced species and its likely impact. The potential invasiveness of R. rubiginosa was assessed in relation to the already invasive North American Physidae Physa acuta Draparnaud, 1805 and the indigenous L. natalensis. This was particularly important in view of the success of P. acuta as an invader in South Africa. The hatching success, frequency of egg abnormalities, embryonic development, growth, survivorship, fecundity and life history parameters (GRR, Ro, rm, T and λ) for the four snail populations were assessed at three experimental temperatures (20oC, 25oC and 30oC). The invasive potential of the freshwater snail Radix rubiginosa recently introduced into South Africa The results showed that R. rubiginosa and P. acuta had a higher growth coefficient (K), longer survivorship, higher fecundity (higher hatching success, fewer egg abnormalities, longer duration of oviposition), shorter incubation period, greater life history parameters (GRR, Ro, rm and λ) and wider temperature tolerances than the two L. natalensis populations tested. The high adaptability of P. acuta to changing environmental factors such as temperature, is in agreement with the fact that it is now more widespread in South Africa than the indigenous species L. natalensis. This has important implications for R. rubiginosa, since this species displayed reproductive attributes and a temperature tolerance that were similar and in certain cases even exceeded the performance of the invasive P. acuta. This therefore implies that R. rubiginosa has the potential to colonize a wider geographical and altitudinal range than L. natalensis, and perhaps even P. acuta. Also, the superior reproductive ability of R. rubiginosa over L. natalensis is likely to present a situation that allows for its rapid spread as well as a possible impact on the indigenous L. natalensis that might render it vulnerable. / Thesis (Ph.D.)-University of KwaZulu-Natal, Westville, 2011.

1. The development of estuarine research in South Africa has been reviewed, and the present state of knowledge assessed. Little is known about many abundant estuarine species, and a previously unstudied prosobranch Nassarius kraussianus (Dunker), was selected for investigation. 2. The study area, a mudflat on the Bushmans River estuary, was divided into three habitat types: supratidal, intertidal and subtidal, each with a particular substratum and pattern of vegetation cover. The distribution of N. kraussianus is associated with the nature of the substratum and the degree of Zostera capensis Setchell cover. 3. A brief review of pertinent literature indicated the necessity of relating population parameters to environmental conditions. 4. The population biology of N. . kraussianus was investigated, and the size frequencies, sex ratios, and male/female size frequencies were recorded over three distinct periods: pre-flood, post-flood and recovery. 5. The Bushmans River flooded in July and August 1979, and the results of the flood have been described. The most immediate result was a period of low salinity, but the most persistant result was the deposition of silt, which had an important effect on the substratum and the Z. capensis beds. 6. The pre-flood population was characterised by patchy distribution, and by a large number of mature adults in relation to juveniles. Females were generally larger than males, and the sex ratio was 1:1. The presence of juveniles in consistantly low numbers indicated that reproduction may have been continuous, but that intraspecific adult/juvenile competition may have kept juvenile numbers low. 7. The post-flood period was characterised by extreme physical conditions, and the survival of a few adults. Juveniles were completely annihilated, and were absent until a few were spawned by the surviving adults towards the end of 1979 . Sex ratio was biased, and females predominated. 8. The recovery period began abruptly with the arrival of a large number of juveniles in the 2, 3 and 4 mm size classes. Recolonisation coincided with habitat recovery, and reqeneration of the Z. capensis cover. Recolonising juveniles grew quickly, reaching adult size in three months. After that, juveniles were spawned on the mudflat by the new population. During this period the sex ratio did not return to the pre-flood 1:1, but remained female biased. Males and females grew at the same rate over the period measured, but females were consistantly larger, indicating that at some stage female growth may be faster. 9. The recolonising juveniles arrived in a regenerated habitat, at a time when temperatures were high, and conditions for growth probably optimal. If intraspecific competition was a feature of the pre-flood population it would have been elimianted in the recovery phase, by the absence of adults. 10. N. kraussianus is apparently omnivorous and proabably not food limited. However, carrion, a limited resource, may be essential for growth to reproductive maturity. Adults and juveniles may compete for this resource, and any juveniles deprived of access to carrion may be stimulated to leave the parental habitat. 11. N. kraussianus is ovoviviparous, and carries up to 5 egg cases within the uterus. Each triangular egg case contains 1 egg which hatches after approximately 3 weeks, as a free swimming veliger. Yeligers metamorphose, and settle after 4-7 days, assuming a benthic mode of life. Ovoviviparity protects the developing embryo from the rigours of environmental extremes, and ensures that embryos are not swept out to sea . Additionally, veligers have a high probability of settling in the parental habitat. 12. This reproductive strategy has been compared with other prosobranchs, and the evolutionary trends within the Prosobranchia, and particularly the Nassariidae, have been related to their dispersal potential. 13. Developmental strategy is intimately related to recolonisation potential. Ovoviviparity, and the lack of pelagic larvae, did not limit the recolonisation potential of N. kraussianus, and recolonisation was probably effected by post-larval juveniles, which may disperse from neighbouring estuaries and coastal pools, by floating in coastal and tidal currents. Post-larval juveniles have two important advantages over larvae. They are less vulnerable to physical extremes, and they have a sturdy crennellated shell which probably affords them protection from predators. 14. Examples of larval recolonisation in estuarine populations have been reviewed, and the most fundamental conclusion of this work is that recolonisation is an important, and probably widespread, adaptation to estuarine conditions in South Africa.

Gastropods are one of the most diverse species groups in mangrove habitats, however, many of their specific roles in relation to ecological patterns and processes are currently largely unknown. The overall aim of this research project was to provide basic ecological information for key gastropod species from subtropical mangroves within a protected area. South African mangroves cover relatively small areas and are restricted to estuaries, these habitats therefore present unique opportunities and challenges to the species that occur in them. Three gastropod species, Terebralia palustris, Cerithidea decollata, and Melanoides tuberculata, all occur at their natural southernmost range limit within South Africa and were selected based on their prominence and occurrence in mangrove habitats of the iSimangaliso Wetland Park, a UNESCO World Heritage Site. Trophic linkages and resource partitioning, resource utilization rates, and ecological resilience were investigated respectively using: 1) a stable isotope (δ15N and δ13C) approach; 2) an experimental approach to quantify feeding dynamics (ingestion rate, consumption/digestion efficiency and grazing impact); and 3) a mixed-effects modelling approach to relate population responses to environmental variables. The diet of T. palustris was seasonally variable and a number of sources were incorporated by different sized snails, but their grazing impact on microphytobenthos was not significant. The results also indicated an ontogenetic shift in the dietary niche for T. palustris through robust partitioning of resources between different size classes. The diets of C. decollata and M. tuberculata were dominated by different primary resources as a function of where they occurred in the mangroves. Melanoides tuberculata consumed a wide variety of primary resources, a typical trait of an opportunistic generalist species. The ingestion rate of M. tuberculata was not dependent on the availability of microphytobenthos, and was highest when conditions were oligotrophic. The resilience of C. decollata was related to the tree-climbing behaviour of this species and its occurrence was best explained by sediment conductivity. These responses were considered in conjunction to what has previously been reported on the resilience of the mangrove trees. The results of this research project have provided new basic ecological information for all three gastropod species in this data-deficient subtropical region. This information can potentially be used in comparative studies for these species in other regions or in broader scale ecological studies. Terebralia palustris has recently experienced a range contraction along the South African coastline. This research project has shown that the diet of this species is highly variable and that food limitation and competition for resources should be considered as potential drivers of the local decline. Cerithidea decollata has in contrast expanded its distributional range in this region. This research project has shown that this species has a generalist diet and exhibits traits in relation to tolerance that are expected to have facilitated its expansion into temperate saltmarsh habitats that occur in dynamic estuaries. Melanoides tuberculata is a globally invasive species, and as South African populations are within its native range, ecological information from this region is valuable as it can be used to investigate the potential ecological effects following introduction into new habitats beyond the native range. Biological drivers have a significant impact on mangrove ecosystem functioning, particularly in relation to recycling and the retention of organic carbon generated through primary productivity. Understanding the ecological linkages that maintain ecological functioning and stability is therefore an important step towards conserving and sustainably managing threatened ecosystems such as mangrove forests.

Globalization has seen an unprecedented dispersal of exotic and alien species worldwide resulting in worldwide homogenization and sometimes extinction of indigenous or endemic taxa. When an exotic species becomes established in a new habitat the invasive organisms are capable of having an impact on indigenous community dynamics and the overall structure and function of ecosystems. Furthermore, the impact of invasion is determined by the geographical range, abundance and the per-capita or per-biomass effect of the invader. However, the success of the introduced organisms is reliant on their ability to acclimate to the physiochemical conditions of the newly invaded environment.Freshwater ecosystems are especially vulnerable to invasions because there are numerous potential routes of introduction including intentional pathways such as stocking, and unintentional pathways such as the release of ballast water and aquarium releases. Efforts to limit the introduction of invasive species or to manage established exotic populations are often hindered by insufficient understanding of the natural history of problematic species. Relatively little is known regarding the physiological tolerances of many taxa. Knowledge about specific species ecophysiological constraints allows for the prediction of future patterns of invasion more accurately, including where an introduced organism would probably survive, thrive and disperse. Furthermore, data on the physiological tolerances of an introduced exotic organism may provide data necessary for effective management and control. This studyinvestigated three invasive species in the Nseleni River system in a protected area in KwaZulu-Natal. The species studied were, Tarebia granifera (Quilted melania – Lamarck, 1822), Pterygoplichthys disjunctivus (Suckermouth armoured catfish - Weber, 1991) and Eichhornia crassipes (water hyacinth – (Martius) Solms-Laubach,). The Nseleni River flows into Lake Nsezi which is responsible for providing potable water to the surrounding towns and industry, as well as the surrounding rural communities. The Enseleni Nature reserve has become the centre for biodiversity dispersal in the immediate area, due to the change in landscape surrounding the protected area.An important step in developing alien invasive species management strategies in protected areas is determining their extent and invasive traits. Tarebia granifera is a prosobranch gastropod originally from South-East Asia that has become invasive in several countries around the world including South Africa. Snail populations were sampled at nine sites throughout the Nseleni/Mposa river system every six weeks over a twelve month period. The snail was abundant throughout the system, especially in shallow waters of less than 1m in depth.The first positive identification the loricariid catfish Pterygoplichthys disjunctivus for the Nseleni River was in 2006. The original introduction is believed to have been via the aquarium trade. The aim of the study was to assess the usefulness of the unified framework with regard to management of fish invasions by assessing the invasion stage of the loricariid population and identifying appropriate management actions using the Blackburn et al. (2011) framework. The fish were sampled at nine different sites and three different depths over a period of twelve months, as well as when two ichthyological surveys were carried out on the Nseleni River system. This invasive fish has been located throughout the system and both male and female fish were collected. The smallest fish sampled was a fingerling of a day or two old and the smallest pregnant female was a mere 270mm TL. This is a clear indication that this fish is breeding in the river system.Although T. granifera and P. disjunctivus were abundant in the Nseleni/Mposa river system, it was not clear what their role in the system was, and in particular if they were competing with any of the indigenous species. Therefore, isotope samples were collected from numerous taxa over a two week period, with the exception of Pterygoplichthys disjunctivus samples, which were collected over 12 months. The δ13C and δ15N signatures of all samples were determined. The niche overlap between the invasive and indigenous snails was effectively zero (1.02E-13%), indicating no shared food resources. The medium ranges of dNRb (7.14) and dCRb (9.07) for the invasive fish indicate that it utilizes a wider range of food resources and trophic levels than the majority of indigenous fish. A medium CDb value (2.34) for the invasive fish species, P. disjunctivus, describes medium trophic diversity, with three indigenous species possessing higher diversity and three possessing lower diversity. Furtherresults indicated that there was no direct dietary competition between P. disjunctivus and indigenous species. Eichhornia crassipes was first recorded on the Nseleni River in 1978, and has been shown to have a significant negative impact on the biodiversity of the Nseleni/Mposa River system and therefore required a control intervention. Although biological control using the two weevil species Neochetina eichhornia (Warner) and N. bruchi (Hustache) has been credited with affecting a good level of control, the lack of a manipulated post-release evaluation experiments has undermined this statement. Five experimental plots of water hyacinth of 20m2 were sprayed with an insecticide to control weevils. After ten months the plants in the sprayed plots were significantly bigger and heavier than those in the control plots that had natural populations of the biological control agents. This study has shown unequivocally that biological control has contributed significantly to the control of water hyacinth on the Nseleni/Mposa River system.The management plan for the Enseleni Nature Reserve identifies the need to control invasive and/or exotic organisms within the boundary of the protected area. In addition, set guidelines have been implemented on how to control these organisms, so that indigenous organisms are least affected. Lack of control of exotic organisms can have serious consequences for indigenous species. It is therefore of utmost importance that the population dynamics of the invading organism be understood, what the potential impact could be and how to control them. Furthermore, it has also acknowledged the threat of possible exotic species invasions from outside of the protected area that might result in threats to the protected area and that these must be investigated, researched and managed.This thesis has identified Tarebia granifera, Pterygoplichthys disjunctivus and Eichhornia crassipes as being a threat to indigenous biodiversity within the protected area, as well as in adjacent areas to the protected area. The thesis will therefore investigate the hypothesis that both Tarebia granifera and Pterygoplichthys disjunctivus are having a direct negative effect on available food resources for indigenous species of organisms. In addition, this thesis will investigate if theNeochetina species that have previously been introduced onto E. crassipes are having any negative effect on this invasive alien aquatic plant.

With the anticipated effects of climate change due to global warming, there is concern over how animals, especially ectotherms, will respond to or tolerate extreme and fluctuating environmental temperature stress. Littorinid snails are intertidal ectotherms that live high on the shore where they experience both extreme and variable conditions of temperature and desiccation stress, and are believed to live close to their tolerance limits. This study investigated the thermal biology of littorinid snails of the genera Afrolittorina, Echinolittorina and Littoraria from temperate, subtropical and tropical regions in South Africa and Brunei Darussalam using thermal tolerance, heart function, and proteome approaches. The effects of conditions, such as rate of change in temperature, acclimation, heat shock, season and starvation were also tested. In addition, the evolutionary relationships and genetic diversity between and within the South African Afrolittorina spp. were investigated using mitochondrial and nuclear markers. Genetic results confirmed that these are two distinct species, with the brown to black A. knysnaensis predominant in the cool-temperate region of South Africa and the pale blue-grey A. africana in the subtropical region. There was low genetic variation and differentiation within each species, suggesting high gene flow among populations as a result of the effects of ocean currents on the dispersal of their planktotrophic larvae. Tests using exposure to high temperatures revealed differences in the thermal tolerances, heart performance and protein profiles of species from different latitudes, regions and zones on the shore. Thermal tolerance conformed to expectations, with clear, statistically significant trends from high tolerance in subtropical species to lower tolerance in temperate species. However, for Afrolittorina spp., there were no significant differences in the thermal tolerances of conspecifics from different regions, though there was a significant difference in thermal tolerance between juveniles and adults. Overall, adults of all species showed higher thermal tolerances than juveniles. Although lethal temperatures for these species were higher in summer than winter, laboratory acclimation had no effect on heat coma temperatures. All species showed some regulation of heart rate, with a degree of independence of heart rate from temperature across mid-range temperatures. The tropical species showed quick induction and good regulation of heart rate followed by the subtropical and temperate species, which displayed mixed responses including regulation, partial regulation and lack of regulation. Overall, tropical Echinolittorina spp. showed good regulation, while the subtropical E. natalensis and Littoraria glabrata exhibited a mixture of partial regulation and regulation. The subtropical/temperate Afrolittorina spp. showed high individual variability, some animals exhibiting regulation, while others did not. These effects seem to be largely phylogenetically determined as there were no differences in the heart rate responses of Afrolittorina spp. from different regions. The temperatures at which heart rate became independent of temperature (thermoneutral zone) were within the range experienced under natural conditions. In addition, there were differences in Arrhenius breakpoint and endpoint temperatures, showing a trend from higher in tropical animals to lower for temperate animals. Conditions such as acclimation, heat shock and starvation had little or no effect on heart performance. However, a slow increase in temperature induced good regulation of heart rate with noticeable shifts of breakpoints and endpoints for Afrolittorina spp. Lastly, there were differences in the proteome responses between and within Afrolittorina spp. as a function of species, size and treatment. Although both large and small A. knysnaensis had a greater number of protein spots in their proteome than A. africana (though the difference was not significant), the later showed significantly higher differential expression of certain proteins following heat stress. In addition, juveniles of both species displayed greater numbers of protein spots in their proteome than adults. The results indicate a difference in the physiological and biochemical responses (i.e. adaptations) of these snails to temperature, and this seems to relate to differences in biogeography, phylogeny, species identity and ecology. The ability to regulate heart rate is phylogenetically determined, while thresholds and lethal limits correspond to biogeography and species ecology. The proteome seems to correspond to species ecology. The results also indicate that these littorinids can tolerate high temperature stress and in this respect they are well suited to life in the intertidal zones or habitats where temperature and other stresses or conditions are extreme and can change abruptly. However, the limited ability of these snails to acclimate to different temperatures suggests that they are already living close to their tolerance limits with small safety margins or narrow thermal windows and so may be vulnerable to small rises in substratum temperature and/or solar radiation.