Crabs are conspicuous inhabitants of temperate salt marshes and tropical mangroves and interact with their environment through several processes. However, detailed information on crab community processes is absent for most South African estuaries and nearshore coastal regions. This study evaluated the primary producers supporting crab species in the salt marsh dominated Swartkops estuary and the mangrove Mngazana estuary. Various methods estimating crab abundances were also assessed in different microhabitats and the larval distribution of crabs in the coastal zone was also investigated. Various methods for estimating crab abundance have been employed in the past, each with its inherent biases. The microhabitat of a mangrove forest in Australia was structurally altered by the manipulation of the litter, pneumatophores and the associated algae. These alterations did not affect the behavioural activity or the numbers of crabs recorded in any of the experimental treatments by either visual counts or pitfall traps. However, the number of crabs caught in the pitfall traps differed between the sites. Species-specific behaviour which was not investigated in this study may bias crab abundance estimates when using pitfall traps and therefore requires further investigation. Benthic consumers inhabiting shallow coastal environments may ultimately have the origin of their nutrition in a number of possible sources. Isotopic and gut content analysis of Thalamita crenata and juvenile Scylla serrata in the Mngazana estuary in South Africa revealed that these two portunids are able to share a habitat by resource partitioning. Differences were noted for species-specific utilization of primary producers not only between seasons within a site, but also between sites. This highlighted the use of locally produced primary producers sustaining food webs in estuaries. Mangrove production in the Mngazana estuary is very important and contributes to most of the carbon in the underlying sediments in the mangrove forest. However, the relatively large number of species and biomass encountered in this estuary may also be attributed to the fact that the different species are able to exploit of a number of different resources. The variation in stable isotope analysis of the different crab species throughout the estuary indicated that these crabs able to occupy the same habitat by feeding on a number of different resources and may preferentially select for a specific primary producer. A stable isotope of crabs in the salt marsh Swartkops estuary indicated that the dominant primary producer sustaining crab communities may even take place on a relatively smallscale. Sesarma catenata found at the inner marsh site recorded more depleted carbon signatures than those encountered in the other sites approximately 100 m away, and reflected signatures similar to the locally-encountered inner marsh plants. The relatively enriched nitrogen signatures of the anthropogenically-impacted Swartkops estuary is an indication of extensive inputs due to urbanization and industrialization, in contrast to the relatively pristine Mngazana estuary which exhibited low nitrogen signatures. Emphasis has been placed on the abiotic component of the exchange of nutrients and energy, although living organisms may also be transported, both actively and passively, between ecosystems. Little variation in either species composition or abundance was found between seasons for the larval distribution of brachyuran crabs on the east coast of South Africa. Due to the lack of published larval descriptions, larvae could not be identified to species level and it was therefore not possible to identify whether the larvae were hatched or spawned in an estuary or in a marine environment, or whether the larvae originated in the northern tropical regions. Frequent wind-reversals which are common in this region may retain larvae close inshore and supply the southern temperate locations with larvae from the northern locations. In conclusion, this study has shown that in highly productive systems with a number of potential primary producers, the crabs that inhabit the estuary show a marked diversity in resource utilization which could potentially allow a number of closely related species to occupy different trophic levels. This study also highlights the importance of locally produced sources in an estuary, which may occur on very small scales and this needs to be factored in with the design of any future stable isotope studies of this nature.

Herbivores and pathogens pose a consistent threat to plant productivity. In response, plants invest in structural and/or chemical defenses that minimize damage caused by these biotic stressors. In salt marshes along the Atlantic coast of the United States, a facultative mutualism between snails (Littoraria irrorata) and multiple species of fungi exert intense top-down control of the foundation grass species Spartina alterniflora. Since exposure to herbivores and pathogens are tightly coupled in this system, I investigated whether S. alterniflora utilizes chemical and/or structural defenses to deter both snails and fungi, and examined how plant defenses varied among S. alterniflora individuals and populations. I also assessed how other marsh plants prevent snails from establishing farms, and considered whether interspecific variation in plant chemical defenses influences marsh community structure. Initial experiments revealed that S. alterniflora chemical defenses inhibited L. irrorata and two fungi that snails commonly farm. A caging experiment determined that production of chemical defenses could not be induced in the presence of snails and fungi, nor relaxed in their absence. Through separations chemistry guided by ecological assays, I isolated two distinct classes of chemical defenses from short form S. alterniflora, one of which inhibited fungal growth and the other decreased plant palatability. In a community context, the chemical defenses produced by S. alterniflora were relatively weak compared to those of four other salt marsh plant species, which produced compounds that completely inhibited L. irrorata grazing and strongly hindered fungal growth in lab assays. Nutritional and structural differences among marsh plants did not influence feeding preferences, suggesting that plant secondary chemistry was the primary driver for food selection by snails. It appears that S. alterniflora produces weak chemical defenses that slow down or limit fungal growth and snail herbivory, and may compensate for tissue losses by producing new growth. In contrast, less abundant marsh plants express chemical defenses that completely inhibit fungal farming and deter snail grazing, but doing so may come at a cost to growth or competitive ability. As marsh dieback continues with rising herbivore densities and compounding abiotic stressors, the ecosystem services that salt marshes provide may be lost. Therefore, understanding how and under what conditions salt marsh plants resist losses to herbivores and pathogens will help predict which marsh communities are most likely to be threatened in the future. Initial experiments revealed that S. alterniflora chemical defenses inhibited L. irrorata and two fungi that snails commonly farm. A caging experiment determined that production of chemical defenses could not be induced in the presence of snails and fungi, nor relaxed in their absence. Through separations chemistry guided by ecological assays, I isolated two distinct classes of chemical defenses from short form S. alterniflora, one of which inhibited fungal growth and the other decreased plant palatability. In a community context, the chemical defenses produced by S. alterniflora were relatively weak compared to those of four other salt marsh plant species, which produced compounds that completely inhibited L. irrorata grazing and strongly hindered fungal growth in lab assays. Nutritional and structural differences among marsh plants did not influence feeding preferences, suggesting that differences in plant chemistry were the primary driver for food selection by snails. It appears that S. alterniflora produces weak chemical defenses that slow down or limit fungal growth and snail herbivory, and may compensate for tissue losses by producing new growth. In contrast, less abundant marsh plants express chemical defenses that completely inhibit fungal farming and deter snail grazing, but doing so may come at a cost to growth or competitive ability against S. alterniflora. As marsh dieback continues with rising herbivore densities and compounding abiotic stressors, the ecosystem services that salt marshes provide may be lost. Therefore, understanding how and under what conditions salt marsh plants resist losses to herbivores and pathogens will help predict which marsh communities are most likely to be threatened in the future.

The ability of prey to detect and respond appropriately to predator risk is important to overall prey fitness. Many aquatic organisms assess risk through the use of chemical cues that can change with predator diet. Two variable characteristics of diet are: 1. prey type and 2. prey mass. To assess the effect of these two characteristics on the assessment of risk by the mud crab Panopeus herbstii, I exposed mud crabs to the urine of the blue crab Callinectes sapidus fed one of 5 diet treatments: 10g of oyster shell free wet mass, 5g of oyster shell free wet mass, 10g crushed mud crabs, 5g crushed mud crabs, and a mix of 5g of oyster shell free wet mass and 5g crushed mud crab. Effects on P. herbstii foraging were tested in a previously developed bioassay by measuring shrimp consumption over a 4 hour period. I hypothesized that P. herbstii would have a larger magnitude response to urine from C. sapidus fed a diet of crushed mud crabs than to urine from C. sapidus fed a diet of oysters. I further hypothesized that P. herbstii would have a larger magnitude response to urine from C. sapidus fed a high mass diet relative to a lower mass diet. Contrary to expectations there was no observed effect of urine on P. herbstii foraging in any of the treatments. Results suggest that bioassay protocol may be unreliable suggesting further replication to determine the difference between this study and previous results. Future studies examining how P. herbstii varies with urine concentration will aid in understanding the ecological scale of this predator cue system. Determining the role of other potential cue sources will improve the predictive abilities of these studies.

We quantified U removal and investigated the efficacy of uranium as a quantitative tracer of groundwater discharge in a headwater salt marsh of the Okatee River, Bluffton, SC. Determining the magnitude of U removal is important for advancing U as a tracer of paleo-oceanic conditions. Since salt marsh groundwater is typically enriched in nutrients and other biologically and chemically reactive species, quantifying groundwater discharge from marshes is critical for understanding the ability of salt marshes to modify the chemistry of important species in surface waters. We hypothesized that water-column U(VI) was removed by tidally-induced advection of surface water into permeable, anoxic salt marsh sediments, a process resulting in bacterially-mediated precipitation of insoluble U(IV)O2 and/or sorption of uranium to iron-oxides at the oxic/anoxic sediment interface. Furthermore, we suggested that hydraulic pressure gradients established by marsh-surface tidal inundation and seasonally-variable rainfall promote the discharge of salt-marsh-processed, uranium-depleted groundwater to tidal creeks, producing the surface-water U-removal signal. Groundwater and surface water data revealed non-conservative uranium behavior. We documented extensive uranium removal from shallow marsh groundwater and seasonally variable uranium removal from surface waters. These observations allowed for the calculation of seasonally-dependent salt marsh uranium removal rates. On a yearly basis, our removal rate (58 to 104 mol m-2 year-1) reemphasized the importance of anoxic coastal environments for U removal. High uranium removal, high barium concentration water observed seeping from creek banks at low tide supported our hypothesis that groundwater discharge must contribute to uranium removal documented in tidal surface waters. Average site groundwater provided an analytically reasonable endmember for explaining uranium depletion in surface water. Therefore, we used three endmember mixing models for estimating the fraction of surface water with presumed a groundwater signature. Our discharge estimates of 8 to 37 L m-2 day-1 agreed closely with previously published salt marsh values. Seasonality in discharge rates can be rationalized with appeal to seasonal patterns in observed rainfall, tidal forcing, and marsh surface bioturbation. Although more work is needed, the results of this portion of the study suggest that U may be an effective quantitative tracer of groundwater discharge from salt marshes.

This collection of research examines how changes in the sensory landscape, mediated by both odor and hydrodynamic properties, impact odor-mediated predator-prey interactions in salt marsh communities. I approached this research using an interdisciplinary framework that combined field and laboratory experimentation to address issues of scale and make connections between predator behavior and patterns of predation in the field. I explored a variety of interactions mediated by changes in the sensory landscape including; indirect effects of biotic structure on associated prey, predator responses to patches of prey with differing density and distribution, and dynamic interactions between predators and prey distributions. I found that biotic structure (oyster reefs [Crassostrea virginica]) has negative indirect effects on associated hard clam prey (Mercenaria mercenaria) through the addition of oyster reef odor cues that attract predators (blue crabs [Callinectes sapidus] and knobbed whelks [Busycon carica])and increase foraging success near the structural matrix. Variation in the structure of patch-scale prey odor plumes created by multiple prey results in predator-specific patterns of predation as a function of patch density and distribution which are mediated by differences in predator sensory ability. There is a potential negative feedback loop between blue crab predators and hard clam prey distributions; clam patches assume random within-patch distributions after exposure to blue crab predators, making the detection of patches by future blue crab predators more difficult. Sensory landscapes are also mediated by water flow, which transports prey odor plumes downstream to predators. Characterization of water flow in small-scale estuary systems indicates that values of turbulent flow parameters are highly context specific and depend on both tidal type (spring, neap, normal) and site. Wind and tidal range seem to be good predictors for wave components and turbulent components of fluctuating flow parameters, respectively, although the strength of their predictive ability is dependent on time scale. Modifications of the sensory landscape through changes in structurally-induced turbulence, mixing of individual plumes from multiple prey, and bulk velocity and turbulence characteristics need to be considered when formulating predictions as to the impact of predators on naturally occurring prey populations in the field.

Hlatikulu Vlei, situated in the foothills of the Natal Drakensberg, is one of the priority wetlands in KwaZulu-Natal, occupying an area of 733 Hectares. The importance of Hlatikulu Vlei lies in its functions to store water, regulate stream flow and attenuate floods, as well as to provide suitable habitat for wildlife and grazing for livestock. Hlatikulu Vlei is a threatened system and has suffered the effects of human mismanagement. Forty-nine percent of the vlei has been classified as disturbed or destroyed, mainly due to the construction of two large dams and past drainage of vlei to facilitate pasture planting. The effects of grazing and fire on the plant communities has been considerably less. Vegetation communities at Hlatikulu Vlei have similarities with those at Ntabamhlope Vlei. The main plant communities present at Hlatikulu Vlei are: vlei grassland, sege-meadows, bulrushes and reedswamp. Species compositions of the mixed sedge and grass sedge-meadow community have a notably higher species diversity than similar communities sampled at Ntabamhlope Vlei and the mires at Highmoor. Soil type and moisture content are shown to be the most significant environmental factors determining the distribution of plant communities and species within the vlei. A wetland re-establishment and rehabilitation programme in the Hlatikulu Crane and Wetland Sanctuary has been effective in allowing many wetland plants to become reestablished. The sanctuary communities bear greater similarity to the sedge and rush sedge-meadow community, than the mixed sedge and grass sedge-meadow communities that were originally present. This is also reflected in the seed bank. All three Southern African crane species (Blue, Wattled and Crowned Crane) and fourteen species of waterfowl have been recorded in the Hlatikulu Crane and Wetland Sanctuary since the wetland rehabilitation programme. The waterfowl play a role in the dispersal of seeds into the sanctuary, particularly those of Schoenoplectus decipiens and Eleocharis dregeana and are in part responsible for the return of certain wetland plants to the sanctuary. The flooding of soils, the fluctuating water level and the soil type related to hummocks and to channels are shown to be responsible for the location of Cyperus denudatus, Arundinella nepalensis and Aristida junciformis in differing positions in the channels and on the hummocks and are also responsible for the maintenance and functioning of the hummocks and channels. Seed banks on the hummocks are similar to seed banks in the channels, however the extant vegetation on the hummocks is distinctly different to that in the channels. Certain species represented in the channel seed bank are being excluded from surviving to maturity. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 1996.

S'han estudiat els efectes dels factors ambientals sobre el perífiton dels sistemes lenític fluctuants del aiguamolls de l'Empordà. L'estudi s'ha realitzat als tres nivells d'integració: nivell d'ecosistema considerant el rol del perífiton envers els altres productors primaris; a nivell de comunitat, estudiant la composició específica de les diatomees i a nivell de població estudiant la plasticitat fenotípica d'una espècie de diatomea (Nitzschia frustulum).A nivell d'ecosistema s'observa que els factors que afavoreixen el predomini dels diferents tipus de productors primaris (perífiton, fitoplàncton i macròfits) són la renovació i el grau d'eutròfia de l'aigua. A nivell de comunitat els factors determinants en la composició i distribució de les espècies de diatomees són els gradients confinament-inundació així com la productivitat del sistema. En funció d'aquest factors s'han establert 5 associacions de diatomees. A nivell de població es demostra que tant la salinitat, com la relació N : P a l'aigua com el moviment de l'aigua afecten la morfologia i ultraestructura de la valva de N. frustulum. De forma interessant s'observa que la salinitat, considerada com a factor individual, afecta N. frustulum a nivell poblacional provocant-li modificacions en la morfologia de la valva, per en canvi, no afecta a nivell de comunitat, ja que totes les espècies de diatomees presents en ambients de salinitat fluctuant són eurihalines. / The effects of environmental conditions on periphytic of lentic and fluctuating waters (Empordà wetlands) at the three different levels of organisation were studied: at ecosystem level, considering the role of periphyton among the primary producers; at community level, analysing the periphytic diatom species composition; and at population level studying the phenotypic plasticity of a selected diatom species (Nitzschia frustulum). The factors favouring the predominance of different primary productors (periphyton, phytoplankton and macrophytes) were the turnover and eutrophy of the water. At the community level, we observed that the main factors affecting diatom species composition and distribution are the gradients confinement-flooding and productivity of the system. Based on these factors 5 diatom assemblages have been distinguished. At the population level, the results of the present thesis proved that the morphology and ultrastructure of N. frustulum is affected by salinity, N : P ratio and movement of the water. Interestingly the results showed that salinity, as a single environmental factor, has effect at N. frustulum population level resulting in phenotypic plasticity while there is no effect of salinity at diatom community level since in such environments with fluctuating salinity all the species present are euryhaline.

Thesis (Ph. D.)--Joint Program in Oceanography (Massachusetts Institute of Technology, Dept. of Biology; and the Woods Hole Oceanographic Institution), February 2002. / Vita. / Includes bibliographical references (leaves 153-161). / I used direct measurements of nitrogen gas (N₂ fluxes and a ¹⁵N stable isotope tracer to determine the contribution of denitrification to salt marsh sediment N cycling. Denitrification in salt marsh tidal creekbottoms is a major sink for groundwater nitrate of terrestrial origin. I studied creekbottom denitrification by direct measurements of N₂ fluxes in closed chambers against a low-N₂ background. I undertook experiments and simulation modeling of sediment N₂ fluxes in closed chambers to optimize the key experimental parameters of this approach. Denitrification in these sediments was driven by the degradation of labile organic matter pools which are depleted during long incubations. Sediment thickness was the most important parameter controlling the required incubation time. Errors of up to 13% with gas headspaces and 80% with water headspaces resulted from headspace N2 accumulation and the resulting collapse of the sediment-water diffusion gradient. These errors could be eliminated by using headspaces of sufficient thickness. Headspace flushing to reduce ammonium accumulation did not affect denitrification rates, but caused transient disturbance of N₂ flux rates. Direct measurements of 0₂, C0₂, N₂, and inorganic N fluxes from the sediments of a salt marsh tidal creek were made in order to examine the interaction of denitrification with the carbon, oxygen, and N cycles. Organic carbon concentration and lability were the primary controls on metabolic rates. C0₂/N flux ratios averaged 6.1, indicating respiration driven by algal biomass. / (cont.) Allochthonous denitrification accounted for 39% of total sediment denitrification (2.7 mol N m⁻² yr⁻¹). 46% of remineralized ammonium was denitrified, while the contribution of autochthonous denitrification to 0₂ and C0₂ fluxes was 18% and 10%, respectively. A ¹⁵N-ammonium tracer was used to study competition between plants and nitrifying bacteria for remineralized ammonium. In undisturbed sediments of Spartina alterniflora, plant uptake out-competed nitrification-denitrification, with plant uptake accounting for 66% of remineralized ammonium during the growing season. Under N fertilization (15.5 mol m⁻² yr⁻¹), both plant N uptake and denitrification increased, but denitrification dominated, accounting for 72% of the available N. When plant uptake was hydrologically suppressed, nitrification-denitrification was stimulated by the excess N, shifting the competitive balance toward denitrification. / by Michael Robert Hamersley. / Ph.D.

Previous research into bottom-up processes on saltmarshes has mainly focused on the influence of plant succession on herbivores. This study will present original research exploring the influence of bottom-up processes in a saltmarsh ecosystem between three trophic levels: Orchestia, redshanks, and sparrowhawks. Density dependence, may be the dominant top-down effect when higher numbers of sparrowhawks and redshanks are present, and may mask top-down and bottom-up trait effects which are constant. Bottom-up effects begin to emerge when cold conditions force redshanks from muddy creeks onto the saltmarsh to forage for Orchestia, because their primary prey, Corophium become less available. Larger flocks form and feeding on Orchestia requires them to balance a need to profit from the best available feeding patches and to be vigilant to sparrowhawk attack. Redshank vulnerability is compounded, because Orchestia hide in cold temperatures, so probing in the soil with their heads down makes them more vulnerable to sparrowhawk attack. Larger flocks may be able to exploit areas closer to sparrowhawk-concealing cover at the terrestrial boundary because they feel safer in greater numbers. Warmer temperatures make Orchestia more active which attracts redshanks, which can simultaneously feed and be vigilant because they peck and catch crawling and jumping Orchestia with their heads up. Consequently, increased flock size may temporarily depress Orchestia abundance, so that redshanks become spaced, leaving isolated individuals more vulnerable to attack. Therefore, it is a temperature-dependent bottom-up process which impacts upon both Orchestia and redshank behaviour, which then may influence the hunting success of sparrowhawks. Whether the characteristics of this saltmarsh ecosystem and the trophic dynamics can be compared to other examples is questionable. Saltmarshes probably differ in their topography and the way in which environmental conditions affect them that then defines which species are present and how these species interact.

Climate change and increasing sea level elevation are predicted to increase salinity in estuarine tidal wetlands in the Southeastern United States. Since much of the ecosystem function in these areas is predicated upon salinity regimes, many fundamental changes are likely to occur as a result. The influence of salinity and SO4 2- reduction on PO4 3- release from tidal wetland soils was evaluated along a salinity gradient at three sites in The St. John’s River, Florida using both field and laboratorybased methods. Porewater was sampled over the course of 10 months to determine ambient levels of SO4 2- and PO4 3-. Lab-based experiments, soils samples were subjected to seawater and SO4 2- treatments in an attempt to induce PO4 3- release. Salinity was lowest at Sixmile Creek (0.45 ± 0.1 g kg-1) and Goodby’s Creek (2.05 ± 2.3 g kg-1) and much higher at Sister’s Creek (27.81 ± 3.1 g kg-1). The organic content of soils was highest (82.35% ± 5.11) at Sixmile Creek, intermediate at Goodby’s Creek (64.45% ± 7.02) and lowest at Sister’s Creek (32.11% ± 9.61). Total soil P was highest at the freshwater Sixmile Creek (1101.64 ± 220.2 μg g-1), intermediate at the brackish Goodby’s Creek (719.61 ± 114.3 μg g-1) and lowest at the Sister’s Creek saltmarsh (475.85 ± 110.9 μg g-1). Porewater PO4 3- was higher at Sixmile and Goodby’s Creek sites (9.44 ± 15.6, 8.99 ± 14.7 !g L-1, respectively) compared to Sister’s Creek (0.6 ± 3.1 !g L-1). Porewater SO4 2- was lower at Sixmile (70.73± 57.58 !g L-1) and Goodby’s Creeks (124.35 ± 152.5 !g L-1) compared to Sister’s Creek (1931.41 ± 557.82 !g L-1). Temporal and spatial trends indicated that SO4 2- and PO4 3- in porewater was likely due to floodwater content and that direct reaction between analytes in soils was unlikely. The addition of aerated seawater failed to cause PO4 3- release from any sites. The incubation of soils under anaerobic conditions, in the presence of Na2SO4 induced SO4 2- reduction, but inhibited PO4 3- flux from both Sixmile and Goodby’s Creek, which is attributed here to likely S- toxicity (Roychoudhury et al., 1999). PO4 3- flux from Sister’s Creek increased in association with Na2SO4 concentration, likely due to more Fe availability to mitigate Stoxicity. Ambient seawater additions to soils under anaerobic conditions followed a similar trend, but the results were not statistically conclusive. Overall, both field and labbased data indicated that Tidal wetland porewater PO4 3- likely originates from floodwaters and that increased salinity and SO4 2- reduction did not directly enhance soil PO4 3- fluxes.