Potentials and Limitations of Testate Amoebae from Tidal Marshes As Bio-Indicators of Environmental Change in the Scheldt Estuary

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Potentials and Limitations of Testate Amoebae from Tidal Marshes As Bio-Indicators of Environmental Change in the Scheldt Estuary Faculteit Wetenschappen Departement Biologie Onderzoeksgroep Ecosysteembeheer Potentials and limitations of testate amoebae from tidal marshes as bio-indicators of environmental change in the Scheldt estuary Potentieel en grenzen van het gebruik van thecamoeben uit schorren als bio-indicatoren voor omgevingsveranderingen in het Schelde estuarium Proefschrift voorgelegd tot het behalen van de graad doctor in de Wetenschappen aan de Universiteit Antwerpen te verdedigen door Marijke OOMS Promotor: Prof. Dr. Stijn Temmerman Co-promotor: Prof. em. Dr. Louis Beyens Antwerpen, 2013 Contents Samenvatting 5 Summary 9 Chapter 1 13 Introduction Chapter 2 37 Testate amoebae as estuarine water-level indicators Chapter 3 65 Testaceae of a brackish tidal marsh Chapter 4 95 Comparison of testate amoebae assemblages along a salinity gradient Chapter 5 115 Sea level reconstruction and selective preservation of testaceae Chapter 6 149 Role of testate amoebae in Si cycle Chapter 7 173 General Discussion References 187 Appendices 203 Abbreviations 233 Dankwoord 237 Samenvatting De huidige stijging van het zeeniveau wordt in estuaria versterkt door een combinatie van de getijwerking en antropogene modificaties aan het estuariene stroombekken. Al deze aanpassingen hebben een diepere landinwaartse doordringing en verdere opstuwing van de getijgolf tot gevolg. Dit resulteert in een verhoogd overstromingsrisico van haven- en kuststeden. Om het overstromingsrisico te doen dalen wordt in vele estuaria extra waterbergingscapaciteit (door ontpoldering) voorzien voor tijden van verhoogd overstromingsrisico (vb. tijdens stormen). Reconstructie van vroegere waterpeilveranderingen kan helpen om de toekomstige waterpeilveranderingen binnen het estuarium te voorspellen. Waterpeilveranderingen kunnen gereconstrueerd worden aan de hand van protisten (o.a. foraminiferen, diatomeeën en thecamoeben) waarvan de omhulsels (schaaltjes), die de soortspecifieke determinatiekenmerken dragen, kunnen fossiliseren. Er zijn reeds meerdere studies verschenen die aantonen dat deze protisten goede proxies zijn voor het reconstrueren van zeespiegelveranderingen. Deze reconstructies zijn gebaseerd op het gebruik van een transferfunctie, welke de relatie tussen moderne protistengemeenschappen en waterpeilveranderingen gebruikt om vroegere waterpeilveranderingen af te leiden uit fossiele protistengemeenschappen. In deze studie wordt gefocust op het gebruik van thecamoeben als proxy voor de reconstructie van waterpeilveranderingen binnen het Schelde-estuarium. Thecamoeben behoren tot de protozoa en zijn geschaalde amoeben (gemiddelde grootte tussen 20 µm - 200 µm) die voorkomen in de schorbodem. Ze hebben een korte generatietijd ( ± 60 generaties jaar-1), wat maakt dat ze snel reageren op omgevingsveranderingen en hun schaaltjes, met soortspecifieke eigenschappen, kunnen fossiliseren. Samenvatting Moderne thecamoebensoortensamenstellingen van een zoetwaterschor (Notelaarschor), brakwaterschor (Groot buitenschoor) en zoutwaterschor (Paulina) werden onderzocht. Op het zoutwaterschor werden onvoldoende thecamoeben teruggevonden om te spreken van thecamoebengemeenschappen (450 schaaltjes g-1). De thecamoebensoortensamenstellingen van het zoet- en brakwaterschor varieerde in de eerste plaats met schorhoogte t.o.v. gemiddelde hoogwaterpeil. Deze relatie werd gebruikt om een “zoete” en “brakke” transferfunctie voor waterpeilveranderingen te maken. De resulterende transferfuncties hebben een nauwkeurigheid die vergelijkbaar is met andere gepubliceerde transferfuncties voor zeespiegelreconstructies. Sub-fossiele thecamoebengemeenschappen werden onderzocht in boorkernen van het zoetwaterschor (Notelaar). De zoetwater transferfunctie, gebaseerd op de moderne data van de Notelaar, werd gebruikt om vroegere waterpeilveranderingen te reconstrueren. Uit vergelijking van de gereconstrueerde waterpeilveranderingen met gemeten waterpeilveranderingen, is duidelijk geworden dat de reconstructiemethode op basis van fossiele schaaltjes werkte tot een diepte van 50 cm (~ 1965). De limitatie van de transferfunctie werd veroorzaakt door een snelle daling in de concentratie van fossiele thecamoeben in de schorbodem, waardoor nog maar weinig schaaltjes overblijven dieper dan 50 cm. Onze studie van de afname van fossiele thecamoebenconcentraties met de diepte hebben aan het licht gebracht dat er mogelijk selectieve oplossing is, waarbij voornamelijk thecamoeben met biogene silica schaaltjes (idiosome thecamoeben) snel verdwijnen. Aangezien de biogene idiosome thecamoebenschaaltjes snel oplossen, is de relatie tussen de silicium cyclus en thecamoeben verder onderzocht. 6 Samenvatting Schorren spelen een belangrijke rol in de Si cyclus, omdat zij de hoeveelheid opgelost silicium in het estuarium aanvullen in tijden van tekorten om zo toxische algenbloeien tegen te gaan. Tot nu toe is de afkomst van dit opgelost silicium toegeschreven aan oplossing van diatomeeën en fytolieten (biogene siliciumstructuren van planten). Ons onderzoek van proportionele afname van idiosome thecamoebenschaaltjes, diatomeeën en organisch materiaal gehalte (als proxy voor fytolieten) in de bodem, heeft aangetoond dat thecamoeben een belangrijke rol spelen in de Si cyclering. Hun relatief snelle oplossing zorgt ervoor dat ze gemakkelijk bijdragen bij het export van opgeloste silicium naar het estuarium in tijden van opgeloste silicium limitatie. We kunnen besluiten dat er een goede relatie is tussen moderne thecamoebensoortensamenstellingen en de hoogteligging van de schorbodem ten opzichte van het gemiddelde hoogwaterpeil. De thecamoebenschaaltjes fossilizeren echter slecht en er komt selectieve bewaring van thecamoeben in de schorbodem voor. Hierdoor zijn thecamoeben toch geen geschikte bio-indicatoren voor het reconstrueren van waterpeilveranderingen in estuaria. Ons verkennend onderzoek over de bijdrage van thecamoeben in schorsedimenten aan de Si cyclus zijn veelbelovend. Verdere studie naar de rol van thecamoeben in de Si cyclus is nodig om een beter beeld te krijgen van hun algehele invloed op de Si cyclus. 7 Summary The present sea-level rise is enhanced in estuaries due to a combination of tidal waves and estuarine anthropogenic modifications, like embankments, channeling and dredging. All these modifications result in further inland penetration and amplification of tidal waves. The increase in tidal amplitude leads to a higher flooding risk of harbor and coastal cities. In order to reduce flood risks, extra water bearing capacity (by restoring floodplain) is foreseen for times of enhanced flooding risk (e.g. storm surges). Reconstruction of past water level changes can help to predict future water level changes within the estuary. Water level changes can be reconstructed using protists (e.g. foraminifera, diatoms, testate amoebae) that have shells which preserve in the soil and carry the species specific traits. Multiple studies have been published that acknowledge protists as good proxies for the reconstruction of sea level changes. The reconstructions are based on the use of a transfer function. A transfer function uses the relationship between modern protist assemblages and water level changes (environmental variable) to infer past water level changes from fossil protist assemblages. This PhD study focuses on the use of testate amoebae as proxy for the reconstruction of water level changes of the Scheldt estuary. Testate amoebae, belonging to the protozoa, are shelled amoebae (average size between 20 µm - 200 µm) that habituate the marsh sediments. Their short generation time ( ± 60 generations year-1) makes that they react quick to environmental changes and their shells, with species specific traits, can fossilize. Modern testate amoebae assemblages of a freshwater tidal marsh (Notelaar), brackish marsh (Groot buitenschoor) and salt marsh (Paulina) were investigated. The found testate amoebae concentrations Summary of the salt marsh Paulina were too low to speak of testate amoebae assemblages ( ± 450 shells g-1). Modern testate amoebae assemblages of the freshwater tidal marsh and brackish marsh varied mainly with marsh elevation relative to mean high water level. This relationship is exploited to make a “freshwater” and “brackish” transfer function. The resulting transfer functions have a comparable accuracy for water level reconstructions to sea level reconstruction transfer functions. Sub-fossil testate amoebae assemblages were investigated in sediment cores of the freshwater tidal marsh (Notelaar). The freshwater transfer function, based on the modern data of the Notelaar, was used to infer past water level changes of this location at the estuary. Comparison of the reconstructed water level changes with measured water level changes showed that the transfer function gives reliable water level reconstruction, based on fossil testate amoebae shells, until a depth of 50 cm (~ 1965). The limitation of the use of the transfer function was related to the quick disappearance of fossil testate amoebae in the soil. Investigation of the decrease in fossil testate amoebae with depth have demonstrated that there might be selective dissolution. Mainly testate amoebae that have biogenic shells (idiosomic testate amoebae) disappear fast out of the fossil record. The finding of quick dissolution of idiosomic testate amoebae shells, made us investigate the relationship between silica cycle and testate amoebae. It is known that marshes play an important role in the Si cycle by providing dissolved Si to the estuary in times of depletion,
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