Community structure and trophic relations in marine tufa stromatolite pools of the Eastern Cape

Microbialites were the dominant life-form of most shallow oceans during the Precambrian. These structures are formed by the deposition of calcium carbonate by cyanobacteria as well as the binding and trapping of sediment by these and other microalgae. In modern environments they are scarce due to several factors, including grazing pressures by metazoans, altered calcium carbonate saturation states of seawater and competition with macroalgae. The recent discovery of an extensive network of actively accreting layered microbialites (stromatolites) along the South African coastline is potentially informative from this perspective. These stromatolites form within the peritidal zone, at the interface of groundwater seepage and periodic marine incursion, forming pools trapped by the accreting fabric. The aim of this thesis was to characterise the ecosystem dynamics of a representative selection of the South African locations. During a comprehensive monthly assessment over an annual cycle, as well as for additional seasonal collections, physico-chemical measurements were monitored together with biological components such as benthic and pelagic microalgae as well as the invertebrate fauna inhabiting the stromatolite pools. These components were then assessed in terms of the potential physical and biological drivers which might explain patterns of variability. Finally, to link all of the ecosystem components, a food-web analysis was conducted, to determine the trophic linkages and, importantly, the reliance by the various consumer organisms on the stromatolite material as a food resource. Results show that the stromatolite pools are driven by a regular interplay between freshwater and marine salinity states, this being determined by tidal amplitude and ocean storm cycles. Furthermore, marine incursion represents the primary source of phosphorus for the stromatolite pools, while available nitrogen is consistently provided by the freshwater inlet stream at each site. This results in an optimum zone of primary biomass within the main stromatolite pool supported by nutrient conditions, while the shifts in salinity state occurring over a weekly tidal schedule likely exclude organisms and macrophytes that are not halotolerant. This is reflected in the benthic microalgae that form the stromatolite accretions in that they are primarily driven by salinity conditions, in addition to seasonal patterns. Interestingly, the variable nutrient conditions, both between sites and temporally, did not contribute as an important driver of the benthic microalgae but did significantly relate to the pelagic microalgae (phytoplankton). This, together with the higher biomass of benthic microalgae compared to its pelagic counterpart, suggests that the stromatolite pools are a benthic-driven system. The short duration of water retention within the stromatolite pools as a result of the constant freshwater inflow, likely also precludes nutrient build-up and favours the benthic, sessile ecosystem component, especially the stromatolite-forming microalgae. In terms of the metazoan infauna, the South African stromatolite pools support a persistent assemblage. This might be surprising given the apparently destructive influence of grazing and burrowing animals on microbial mats in terms of restricting the formation of layered accretions. However, metazoans that burrow within the stromatolite fabric were observed to coexist with clear, layered accretions. This supports the observations in some other modern microbialite habitats to suggest that metazoan disruption is clearly not the only or primary factor responsible for modern microbialite scarcity. When assessing the possible drivers of the metazoan community occupying the stromatolite matrix, both salinity patterns and resource conditions in terms of nutrient supply and macroalgal cover were consistently best related to infaunal abundance and presence/absence. This further demonstrates the role of salinity conditions in terms of providing a habitat that is restrictive to most metazoan organisms, while also suggesting that the metazoans are responding to macroalgal rather than the stromatolite microalgal conditions. To further develop this observation, the results from the stable isotope work clearly reflect a dominance of pool macroalgae in the diets of invertebrate consumers, with little to no stromatolite material consumed. This suggests that there is limited apparent destructive grazing influence by the metazoans on the stromatolite matrix, in addition to the burrowing bioturbation mentioned previously. Furthermore, the metazoan grazers may be indirectly benefitting the stromatolites by restricting macroalgal biomass, which might otherwise outcompete its microalgal counterpart. This study provides a valuable understanding of benthic-driven peritidal stromatolite ecosystems, and also, from a geological perspective of past stromatolite habitats, suggests some of the mechanisms as to why metazoans may be able to coexist with layered microbialites. Given the threats to similar habitats globally, especially in terms of water resources, management measures necessary to ensure stromatolite persistence in modern environments such as these are proposed. The possible ecological role of peritidal stromatolite habitats within the broader environment, as well as recommendations for future work, is also contextualised

Time-activity budgets of cape gannets (Morus Capensis) at Bird Island, Algoa Bay

For several decades now, seabirds have been considered to be useful indicators of the state of their prey resources because of how they reflect lower-level trophic variability through proxies such as diet or behaviour. However, collection of appropriate data is often challenging in the marine environment because of logistical or financial constraints. In this study, time-activity budgets were studied in the Cape gannet (Morus capensis), a seabird which has been advocated as a potential bioindicator for local epipelagic prey. VHF transmitters attached to PVC leg-rings were fitted to 50 adult breeding pairs during the 2011/2012 and 2012/2013 breeding seasons at Bird Island, Algoa Bay, to determine whether an automated method of continuously collecting time-activity budget data can replace conventional laborious direct observations. To validate that the foraging trip duration data generated from the automated method was a reflection of foraging effort, Time-Depth Recorders (TDRs) were simultaneously equipped to birds with VHF transmitters for three weeks. In order to assess the influence of factors other than prey availability on parent time-activity budgets, sex, chick age and body condition were measured. Additionally, chick growth and survival were recorded in order to investigate the gender-specific effects of parental time-activity budget variability on these parameters. Attachment of VHF transmitters to leg-rings of adult Cape gannets had no observable negative effects on the adults or their chicks in terms of adult body condition, nest attendance and foraging trip durations, or chick growth and survival. Furthermore, the frequency distribution of behaviour bout (foraging trip and nest attendance) durations was no different between automated and directly observed data. However, the automated method did record shorter behaviour bouts, largely attributed to the increased likelihood of direct observations missing birds returning briefly to their nests during older chick provisioning. Additionally, foraging trip duration was highly correlated to foraging effort in terms of time spent resting on the sea surface, flying and diving. The automated method therefore appears to be a good reflection of direct nest attendance observations and foraging effort. Cape gannet time-activity budgets were related to chick age and parental sex. Especially as chicks neared fledging, females spent significantly longer periods of time foraging than males, with males consequently provisioning their chicks more often. Furthermore, adults departing their nests earlier spent more time away from the nest foraging as more same-day daylight hours were available. Chick growth was a function of parent foraging trip duration and associated prey delivery rates. Chick survival was most strongly affected by the amount of time which chicks were left unattended by both parents and consequently exposed to predation by kelp gulls (Larus dominicanus) or to adverse weather conditions. Additionally, for females (but not males) there appeared to be a trade-off between foraging trip duration and chick survival. The Cape gannet appears to demonstrate a strategy whereby the costs of reproduction to the female are shifted towards male-dominated chick provisioning as the chick nears fledging. Drivers of time-activity budget variability such as chick age and parental sex therefore need to be considered if using data on foraging trip duration as a proxy of foraging effort and prey availability

Micro-structures, mineralogy, and chemistry of peritidal tufa stromatolites along the Eastern Cape coast

Peritidal tufa microbialites occurring along the coast near Port Elizabeth, South Africa have been investigated from multiple disciplines and are found to be similar to supratidal tufa deposits in South West Australia. Studies have been conducted on the biological factors, geomorphology, ecosystems, and associated water chemistry. However, to date no mineralogical, micro-fabric, or geochemical analyses have been reported on these tufa deposits. This work, carried out at a previously well-studied area, provides the first study of this kind on the tufa. Chapter 1 is a mineralogical and micro-fabric analysis of the tufa deposits near Port Elizabeth for the purpose of classification and contextualisation. Chapter 2 provides the first geochemical study of these peritidal microbialites. X-Ray diffraction (XRD) investigations reveal dominance of low-Mg calcite in the mineral make-up of the tufa. A micro-structure analysis via thin section exposes a number of fabrics, suggesting various micro-facies: phytoherm boundstone (layered), phytoherm framestone (non-layered), lithoclast, and minor metazoan tufa. Scanning electron microscopy (SEM) images show micro- to nano- scale variation in calcite grains and epitaxial forms of needlefibre calcite (NFC). The elemental composition of the tufa deposits were examined by X-ray fluorescence (XRF), Laser Ablation Inductively Coupled Plasma Mass Spectrometry (LA-ICPMS), and Scanning Electron Microscopy / Energy Dispersive X-Ray Spectroscopy (SEM/EDS). XRF and LA-ICP-MS were used to analyse the bulk rock chemistry while SEM/EDS was used to scrutinize specific areas within the tufa. These observations suggest the deposits are better classified as tufa microbialites (rather than exclusively “stromatolites”) and outline similarities and disparities to the micro-fabrics of supratidal tufa deposits in South West Australia, and Cape Morgan, South Africa. Here the Port Elizabeth tufa is shown to be similar, in terms of the dominant elements (O, Ca and to a lesser extent, Mg and Sr), to the Australian deposits despite subtle dissimilarities in water chemistry. Increasing trends toward more marine tufa for many elements are also shown here and can be explained by the interaction with increased amounts of sedimentary products and/or interactions with more saline water that contains a higher TDS (Total Dissolved Salts). This is also the first study to report needle-fibre calcite formation in stromatolites and the first to geochemically analyse modern peritidal microbialites

Fish utilization of surf-zones. Are they changing? A case study of the sheltered, warm-temperate King’s Beach

Surf-zone fish communities and their shifts over time are generally poorly understood. The aim of this study was to compare the current surf-zone fish assemblage at King’s Beach, South Africa, to a similar study conducted three decades ago, before the collapse of many exploited shore fishes in the region. Beach seine nets (mesh sizes of 10 and 50 mm) were used to target juvenile and adult fishes bimonthly from February to August 2011 over the high tide around sunset. A total of 14 species were recorded in both the 30mand 100mseine nets. The catch in these seine nets was dominated by Pomadasys olivaceus and Liza richardsonii, and this was significantly different to three decades ago, when P. olivaceus, Sarpa salpa and Diplodus capensis dominated the catch. Important linefish species belonging to the Sparidae and Sciaenidae families were significantly smaller and less abundant in this study. Two sparids, S. salpa and Lithognathus mormyrus, which made a large contribution to the surf-zone catch three decades ago were absent during this study. Reasons for the significant shifts in the surf-zone fish community, including overexploitation of the linefish and potential habitat modification, are discussed

Modern supratidal microbialites fed by groundwater: functional drivers, value and trajectories

Microbial mats were the dominant habitat type in shallow marine environments between the Palaeoarchean and Phanerozoic. Many of these (termed ‘microbialites’) calcified as they grew but such lithified mats are rare along modern coasts for reasons such as unsuitable water chemistry, destructive metazoan influences and competition with other reef-builders such as corals or macroalgae. Nonetheless, extant microbialites occur in unique coastal ecosystems such as the Exuma Cays, Bahamas or Lake Clifton and Hamelin Pool, Australia, where limitations such as calcium carbonate availability or destructive bioturbation are diminished. Along the coast of South Africa, extensive distributions of living microbialites (including layered stromatolites) have been discovered and described since the early 2000s. Unlike the Bahamian and Australian ecosystems, the South African microbialites form exclusively in the supratidal coastal zone at the convergence of emergent groundwater seepage. Similar systems were documented subsequently in southwestern Australia, Northern Ireland and the Scottish Hebrides, as recently as 2018, revealing that supratidal microbialites have a global distribution. This review uses the best-studied formations to contextualise formative drivers and processes of these supratidal ecosystems and highlight their geological, ecological and societal relevance

Time-activity budgets of seabirds in relation to sex, chick age and time

<p>This data-set expands upon http://dx.doi.org/10.6084/m9.figshare.1044267 and provides a detailed summary of foraging trip durations, nest attendance bouts and provisioning rates for Cape gannets at Bird Island, Algoa Bay, South Africa over two breeding seasons (2011/2012 and 2012/2013). Time-activity budgets (TABs) are presented in the CSV files according to chick age, parent sex and time of bout initiation.</p

Automated and manual time-activity budget data of colonial seabirds

<p>These data were collected at Bird Island, Algoa Bay, South Africa during the 2011/2012 and 2012/2013 Cape gannet breeding seasons. The CSV files contain data of signal strength tests for VHF transmitters (in three configurations) at increasing distances from a VHF receiver, chick growth measurements, as well as time-activity budget (TAB) data obtained automatically from VHF receiver data, from direct observations (of both VHF-equipped Cape gannets and control birds) and from Cape gannets equipped with temperature-depth recorders (TDRs).</p

Coastal urban reliance on groundwater during drought cycles: Opportunities, threats and state of knowledge

Urbanisation and population growth are concentrated on the coast with approximately 40% of the human population living within 100 km of the ocean. The freshwater systems on which coastal urban areas rely are vulnerable to bidirectional pressures including coastal processes such as sea-level rise and coastal erosion coupled with land use changes and pollution occurring in inland catchment areas. These threats are likely to be amplified in the future under climate change conditions and more frequent and severe drought periods are expected to jeopardise already constrained water supply systems. Groundwater is used as a freshwater resource globally and is especially important as a conjunctive supply during drought periods due to perceived buffer capabilities. However, several threats impact coastal aquifers due to over-abstraction, such as salinisation, land subsidence and groundwater flooding and often these subterranean resources are “out of sight and out of mind” when it comes to management strategies. Here, we present an assessment of current issues and management options relevant to coastal aquifers using recent literature. These insights provide knowledge on global issues relevant to groundwater resources, especially regarding water use during droughts. This is exemplified using a South African case study of two metropolitan municipalities that have experienced or are experiencing severe multi-year droughts. Both municipalities have grappled with the depletion of surface water resources, which constitutes the bulk of the local water supply systems. Consequently, groundwater resources have been explored as an augmentation strategy. Although groundwater resources may be useful in alleviating drought effects, it is crucial that a local understanding of the aquifers is developed through baseline hydrological studies and long-term monitoring. Furthermore, unregistered groundwater use needs to be quantified. Finally, a holistic groundwater management view, and the communication thereof, is required to ensure the sustainable management of coastal aquifers

Harmful algal blooms of Heterosigma akashiwo and environmental features regulate Mesodinium cf. rubrum abundance in eutrophic conditions

Functional drivers of phytoplankton that can potentially form harmful algal blooms (HABs) are important to understand given the increased prevalence of anthropogenic modification and pressure on coastal habitats. However, teasing these drivers apart from other influences is problematic in natural systems, while laboratory assessments often fail to replicate relevant natural conditions. One such potential bloom-forming species complex highlighted globally is Mesodinium cf. rubrum, a planktonic ciliate. This species occurs persistently in the Sundays Estuary in South Africa yet has never been observed to “bloom” (&gt; 1,000 cell.ml −1). Modified by artificial nutrient-rich baseflow conditions, the Sundays Estuary provides a unique Southern Hemisphere case study to identify the autecological drivers of this ciliate due to artificial seasonally “controlled” abiotic environmental conditions. This study utilised a three-year monitoring dataset (899 samples) to assess the drivers of M. cf. rubrum using a generalised modelling approach. Key abiotic variables that influenced population abundance were season and salinity, with M. cf. rubrum populations peaking in summer and spring and preferring polyhaline salinity regions (&gt;18) with pronounced water column salinity stratification, especially in warmer months. This was reflected in the diel vertical migration (DVM) behaviour of this species, demonstrating its ability to utilise the optimal daylight photosynthetic surface conditions and high-nutrient bottom waters at night. The only phytoplankton groups clearly associated with M. cf. rubrum were Raphidophyceae and Cryptophyceae. Although M. cf. rubrum reflects a niche overlap with the dominant HAB-forming phytoplankton in the estuary (the raphidophyte, Heterosigma akashiwo), its reduced competitive abilities restrict its abundance. In contrast, the mixotrophic foraging behaviour of M. cf. rubrum exerts a top-down control on cryptophyte prey abundance, yet, the limited availability of these prey resources (mean &lt; 300 cells ml −1) seemingly inhibits the formation of red-water accumulations. Hydrodynamic variability is necessary to ensure that no single phytoplankton HAB-forming taxa outcompetes the rest. These results confirm aspects of the autecology of M. cf. rubrum related to salinity associations and DVM behaviour and contribute to a global understanding of managing HABs in estuaries. </p