The primary production of Spartina maritima, the dominant macrophyte of the intertidal salt marshes of the Swartkops estuary, was determined for inclusion into the proposed ecosystem model. Several methods for measuring net production devised for North temperate Spartina species were used, as no single method was considered reliable for the local species. In addition, a new method was devised to take into account the aseasonal growth behaviour of S. maritima . Most energy for growth appears to be directed into leaf production, rather than shoot growth. Live net production values were 523-680 g m⁻²‚ yr ⁻¹, which are generally lower than estimates of N. American Spartina species. Total Live production of S. maritima for the whole Swartkops estuary was 495,6 x I0³kg yr ⁻¹, which is an estimation of potential food resources. Formation of S. maritima detritus by decomposition was significantly related to air and water temperatures. Thus detritus production showed a wide seasonal variation about a mean of 374 g m⁻² yr⁻¹. The total S. maritima detritus production of 307,5 x I0³kg for the whole Swartkops estuary was determined as an estimate of available food resources. Export of detrital material from Spartina marshes to estuarine waters was negligible. Even during a maximal Equinoctial tide, only 6,26 dry g m⁻² detritus was exported from the creek catchment area of Spartina stands. This study has shown that concepts evolved for North temperate estuaries should be carefully considered before being applied to local situations, particularly in view of the relatively low, aseasonal production of S. maritima and the negligible contribution of its detritus to estuarine waters. Furthermore, it is essential that utilisation of primary production by secondary producers be determined before the function of the estuarine ecosystem may be understood

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.

The distribution of Callianassa kraussi in southern Africa extends from Lamberts Bay on the west coast to San Martinho in Mocambique. This represents a northerly extension of the range from the previous known limit of Inhaca Island. In this area 59 localities were visited and records obtained for an additional 16. Three areas were selected for detailed study. These were the west Kleinemonde estuary, which is normally closed off from the sea by a sand bar and the open Swartkops estuary both on the southeastern Cape coast, and the Swartvlei system, which consists of a lake connected to the sea by a channel which closes intermittently, on the southern Cape coast. The nature of the burrows constructed by C. kraussi was investigated in these areas. Burrow complexes may have up to nine entrances but more commonly two to four. Laboratory studies suggest that the number of entrances does not exceed the number of prawns present and consequently burrow opening counts will give an estimate of population density. Water movement can result in the closing off of burrows while strong water and sand movement will exclude C.kraussi from an area. C.kraussi is also excluded from areas which have very coarse substrates. A three year regular sampling program showed that C.kraussi breeds mainly in winter/spring with a smaller breeding peak in summer. Egg development time at 20°C in a salinity of 35° /₀₀ is 30-33 days. There are two larval stages which last 3-5 days. Development time of the eggs is tripled at temperatures of 13-15°C while larval development time is extended to 9-14 days. Development is possibly slightly accelerated at 25 ± 3°C but the number of eggs hatching is markedly reduced. Larval development times were similar to those at 20° and 35°/₀₀. There are no planktonic larval stages. Growth was more rapid and greater size was attained in the open Swartkops estuary than in the closed Kleinemonde estuary. Prawns hatching in winter/spring breed for the first time in the following summer at an age of about 16 months and then again in the following winter/spring at an age of about 2 years. Prawns hatching in summer breed in the following winter at an age of about 18 months and then again in the immediately following summer. Longevity in both sexes is about 2 years. Dispersal is accomplished by migration of the post-larval juvenile phases at an age of 3-5 months. Non-selfmaintaining populations exist in areas where salinities are too low to permit breeding. Population densities were found to vary markedly in different areas. Investigations of osmotic and ionic regulation showed that C.kraussi is a strong hyper-regulator and thus distinct from any other known species in the genus. Volume regulatory ability is well developed and depends on variable rates of urine production. Salt loss in dilutions occurs almost equally via the gills and the urine. The general responses of C.kraussi to dilutions of sea water are discussed. Various suggestions for conservation measures based on the distribution and iife cycle of C.kraussi were made. The factors affecting the distribution of C.kraussi are discussed. The problem of the larval development in C.kraussi was discussed in relation to reviews of larval types of benthic invertebrates. The importance of the burrow, the possible route by which C.kraussi has invaded estuaries and the differences between open and closed estuaries as shown by the effects on C.kraussi were discussed.

Temporarily open/closed estuaries (TOCE’s) are the numerically dominant type of estuary accounting for ≈ 70 % of all estuaries along the South African coastline. Despite their numerical dominance, aspects of the biology of organisms within these systems, particularly macrocrustacea remain poorly understood. This study firstly assessed the abundance and biomass of the most common isopod, Exosphaeroma hylocoetes, in three Eastern Cape TOCE’s, (the West Kleinemonde, East Kleinemonde and Kasouga Estuaries), and their response to mouth breaching events. This study was followed by aspects of the biology of the isopod including their utilisation of submerged macrophytes as a refuge from predation and/or possible food source, as well as the growth rates of the isopods in the laboratory under different environmental conditions. Mean isopod abundances and biomasses ranged between 0 and 4 791 ind. m⁻² and 0 and 9.65 mg dwt. m⁻² in the West Kleinemonde Estuary and between 0 and 108 ind. m⁻² and 0 and 0.318 mg dwt. m⁻² in the nearby East Kleinemonde Estuary. In the Kasouga Estuary, the values ranged between 0 and 3 650 ind. m⁻² and 0 and 5.105 mg dwt. m⁻². Temporal and spatial changes in the abundance and biomass of E. hylocoetes within the three systems was primarily linked to mouth phase, with populations declining when the mouth was open and to a lesser extent, seasonality. In all three estuaries maximum isopod abundances and biomasses were recorded in their middle and upper reaches, which could be ascribed to the presence of submerged macrophytes particularly Ruppia maritima, in two of the estuaries. Males (5.71 ± 0.41 mm) were significantly larger than females (3.99 ± 0.26 mm), but the sex ratios were skewed in favour of females, (a common feature in many isopod populations). Females were found carrying brood throughout the study, releasing offspring directly into the water column, to allow recruitment to the populations. The larger the female, the larger the brood carried (up to a maximum of 72 embryo/mancas). Results of laboratory experiments indicate that the close association of Exosphaeroma hylocoetes with submerged macrophytes is a result of the plant stands providing a refuge against predation by selected ichthyofauna. However stable carbon isotope and fatty acid analyses indicate that E. hylocoetes made use of ephiphytic algae and detritus on the stems of R. maritima, rather than the submerged macrophyte itself. There were no significant differences in the growth rates of male and female isopods at combinations of temperature (15 and 25 °C) and salinity (15 and 35 ‰). Laboratory growth studies revealed that males lived longer than females (25.77 ± 3.40 weeks vs 21.52 ± 3.00 weeks), and therefore achieved larger overall size. Females, however, reached sexual maturity (at 2.5 mm) at a faster rate (two to four weeks) than males (5.5 mm after 6.5 to 11 weeks), dependent on temperature. Exosphaeroma hylocoetes with its association with submerged macrophytes, early maturity, its growth rates and longevity, female-biased sex ratios and year-round breeding, all contribute to its success in temporary open/closed estuaries.

The spatial and temporal patterns in the population demographs and feeding ecology of the caridean shrimp, Palaemon peringueyi, was investigated over a 12 month period in the permanently open Kariega Estuary situated on the south-east coastline of South Africa. Shrimps were sampled monthly from three stations corresponding to the lower, middle and upper reaches of the Kariega Estuary. The shrimps were collected at the different stations using a pull net. Temperature, salinity, dissolved oxygen and % cover of submerged vegetation were measured at each sampling station. Finally, the feeding ecology of the shrimp was investigated employing stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes analysis. Temperature, salinity, dissolved oxygen and vegetation cover showed distinct seasonal patterns. The shrimps attained total abundances and biomass of between 0 and 245.6 individuals per m² (ind.m⁻²) and between 0 and 13.6 grams wet weight per m² (g wwt.m⁻²), respectively. Mean abundance ranged between 2.3 (± 3.0) and 88 (± 27.3) ind.m⁻² in the lower reach, between 0.9 (± 0.4) to 21.5 (± 4.5) ind.m⁻² in the middle reach and 0 to 1.6 (± 0.5) ind.m⁻² in the upper reach. Mean biomass ranged between 0.4 (± 0.5) and 4.4 (± 1.4) g wwt.m⁻² in the lower reach, between 0.1 (± 0) to 4.3 (± 0.9) g wwt.m⁻² in the middle reach and between 0 to 1.6 (± 0.5) g wwt.m⁻² in the upper reach. Greatest abundances and biomasses were recorded in winter following heavy rainfall within the catchment area. Freshwater inflow which is due to rainfall is thought to provide a cue for the recruitment of marine breeding organisms. The spatial patterns in the distribution of the shrimps showed that juveniles occupied the lower and middle reaches and adults the entire length of the estuary. There were no significant relationships between the abundance and biomass of the shrimps and the selected physico-chemical variables (p > 0.05 in all cases). Results of the stable isotope study indicated that the mean δ¹³C of the shrimps in the lower, middle and upper reaches were -12.4 (± 1.3) ‰, -15.6 (± 0.5) ‰ and -17.5 (± 0.5) ‰, respectively. The δ¹⁵N isotopes of the shrimps in the lower reach was 11.6 (± 0.5) ‰ with middle and upper reaches having similar δ¹⁵N values, 13.5 (± 0.4) ‰ and 13.4 (± 0.4) ‰, respectively. There was no significant difference in the δ¹³C and δ¹⁵N isotope (p > 0.05) between the sexes of the shrimps. Two-way ANOVA indicated a significant difference in the δ¹³C isotope of the shrimps between the sites (p < 0.05). The difference in the δ¹³C and δ¹⁵N ratios of the shrimps suggests spatial variability in the diet of P. peringueyi within the Kariega Estuary. The diet of the shrimps in the lower reach is primarily composed of Nanozostera capensis epibionts. In the middle reach the diet of the shrimps is likely comprised of the detritus of benthic algae. In the upper reach of the estuary, P. peringueyi likely derives its carbon from a combination of epibionts, benthic algae and detritus derived from Codium.

The role of the sandprawn, Callichirus (=Callianassa) kraussi (Stebbing), as an ecosystem engineer was assessed in the lower reach of the temporarily open/closed Kasouga Estuary situated along the Eastern Cape coastline of southern Africa over the period April 2010 to June 2011. The study comprised two distinct components, a field study and a caging experiment. The field study assessed the correlation between sand prawn densities and selected physico-chemical (organic content of the sediment and bioturbation) and biological (microphytobenthic algal concentrations and macrobenthic abundance and biomass) variables in 50 quadrants in the lower reach of the estuary. Densities of the sand prawn within the quadrants ranged from 0 to 156 ind m⁻² (mean = 37 ind m⁻²). There were no significant correlations between the densities of the sandprawn and the estimates of the organic content of the sediment and the abundance and biomass of the macrofauna (P > 0.05 in all cases). Numerical analyses failed to identify any effect of the sandprawn density on the macrofaunal community structure. The rate of bioturbation was, however, strongly correlated to the sand prawn density. Similarly, the microphytobenthic alga concentrations were significantly negatively correlated to the sand prawn densities ((P < 0.05). The absence of any distinct impact of the sandprawn on the macrobenthic community structure appeared to be related to their low densities in the lower reach of the estuary during the study. To better understand the role of the sandprawn as an ecosystem engineer, a caging experiment was conducted using inclusion and exclusion treatments (n= 5 for each treatment). Densities of the sandprawn in the inclusion treatments (80 ind m⁻²) were in the range of the natural densities within the estuary. The experiment was conducted over a period of 18 weeks in the lower reach of the estuary during summer. The presence of the sandprawn, C kraussi, contributed to a significant decrease in the microphytobenthic algal concentrations and the abundance and biomass of the macrofauna (P < 0.05 in all cases). The decrease in the microphytobenthic algal concentrations in the presence of the sandprawn appeared to be related to the res-suspension of the sediments (bioturbation) generated by the burrowing and feeding activities of the sandprawn. The observed decrease in macrofaunal abundances and biomass in the inclusion treatments appeared to be mediated by both the decreased food availability (mainly the microphytobenthic algae) and the burial of organisms within the sediments. Numerical analysis indicated that the sandprawn did, however, not contribute to a change in the species composition of the macrofauna. Results of the current study indicate that C.kraussi plays an important role in structuring the invertebrate community and energy flow within temporarily/open closed Kasouga Estuary.