Invertebrate Responses to Large-Scale Change: Impacts of Eutrophication and Cataclysmic Earthquake Events in a Southern New Zealand Estuary
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Invertebrate Responses to Large-Scale Change: Impacts of Eutrophication and Cataclysmic Earthquake Events in a Southern New Zealand Estuary A thesis submitted in partial fulfilment of the requirements for the degree of Doctorate of Philosophy in Ecology at the University of Canterbury, New Zealand Jennifer Erin Skilton 2013 Abstract Abstract Environmental stress and disturbance can affect the structure and functioning of marine ecosystems by altering their physical, chemical and biological features. In estuaries, benthic invertebrate communities play important roles in structuring sediments, influencing primary production and biogeochemical flux, and occupying key food web positions. Stress and disturbance can reduce species diversity, richness and abundance, with ecological theory predicting that biodiversity will be at its lowest soon after a disturbance with assemblages dominated by opportunistic species. The Avon-Heathcote Estuary in Christchurch New Zealand has provided a novel opportunity to examine the effects of stress, in the form of eutrophication, and disturbance, in the form of cataclysmic earthquake events, on the structure and functioning of an estuarine ecosystem. For more than 50 years, large quantities (up to 500,000m3/day) of treated wastewater were released into this estuary but in March 2010 this was diverted to an ocean outfall, thereby reducing the nutrient loading by around 90% to the estuary. This study was therefore initially focussed on the reversal of eutrophication and consequent effects on food web structure in the estuary as it responded to lower nutrients. In 2011, however, Christchurch was struck with a series of large earthquakes that greatly changed the estuary. Massive amounts of liquefied sediments, covering up to 65% of the estuary floor, were forced up from deep below the estuary, the estuary was tilted by up to a 50cm rise on one side and a corresponding drop on the other, and large quantities of raw sewage from broken wastewater infrastructure entered the estuary for up to nine months. This study was therefore a test of the potentially synergistic effects of nutrient reduction and earthquake disturbance on invertebrate communities, associated habitats and food web dynamics. Because there was considerable site-to-site heterogeneity in the estuary, the sites in this study were selected to represent a eutrophication gradient from relatively “clean” (where the influence of tidal flows was high) to highly impacted (near the historical discharge site). The study was structured around these sites, with components before the wastewater diversion, after the diversion but before the earthquakes, and after the earthquakes. The eutrophication gradient was reflected in the composition and isotopic chemistry of primary producer and invertebrate communities and the characteristics of sediments across the sample sites. Sites i Abstract closest to the former wastewater discharge pipe were the most eutrophic and had cohesive organic-rich, fine sediments and relatively depauperate communities dominated by the opportunistic taxa Capitellidae. The less-impacted sites had coarser, sandier sediments with fewer pollutants and far less organic matter than at the eutrophic sites, relatively high diversity and lower abundances of micro- and macro-algae. Sewage-derived nitrogen had became incorporated into the estuarine food web at the eutrophic sites, starting at the base of the food chain with benthic microalgae (BMA), which were found to use mostly sediment-derived nitrogen. Stable isotopic analysis showed that δ13C and δ15N values of most food sources and consumers varied spatially, temporally and in relation to the diversion of wastewater, whereas the earthquakes did not appear to affect the overall estuarine food web structure. This was seen particularly at the most eutrophic site, where isotopic signatures became more similar to the cleaner sites over two-and-a-half years after the diversion. New sediments (liquefaction) produced by the earthquakes were found to be coarser, have lower concentrations of heavy metals and less organic matter than old (existing) sediments. They also had fewer macroinvertebrate inhabitants initially after the earthquakes but most areas recovered to pre-earthquake abundance and diversity within two years. Field experiments showed that there were higher amounts of primary production and lower amounts of nutrient efflux from new sediments at the eutrophic sites after the earthquakes. Primary production was highest in new sediments due to the increased photosynthetic efficiency of BMA resulting from the increased permeability of new sediments allowing increased light penetration, enhanced vertical migration of BMA and the enhanced transport of oxygen and nutrients. The reduced efflux of NH4-N in new sediments indicated that the capping of a large portion of eutrophic old sediments with new sediments had reduced the release of legacy nutrients (originating from the historical discharge) from the sediments to the overlying water. Laboratory experiments using an array of species and old and new sediments showed that invertebrates altered levels of primary production and nutrient flux but effects varied among species. The mud snail Amphibola crenata and mud crab Austrohelice crassa were found to reduce primary production and BMA biomass through the consumption of BMA (both species) and its burial from bioturbation and the construction of burrows (Austrohelice). In contrast, the cockle Austrovenus stutchburyi did not significantly affect primary production and BMA biomass. These results show that changes in the structure of ii Abstract invertebrate communities resulting from disturbances can also have consequences for the functioning of the system. The major conclusions of this study were that the wastewater diversion had a major effect on food web dynamics and that the large quantities of clean and unpolluted new sediments introduced to the estuary during the earthquakes altered the recovery trajectory of the estuary, accelerating it at least throughout the duration of this study. This was largely through the ‘capping’ effect of the new liquefied, coarser-grained sediments as they dissipated across the estuary and covered much of the old organic-rich eutrophic sediments. For all aspects of this study, the largest changes occurred at the most eutrophic sites; however, the surrounding habitats were important as they provided the context for recovery of the estuary, particularly because of the very strong influence of sediments, their biogeochemistry, microalgal and macroalgal dynamics. There have been few studies documenting system level responses to eutrophication amelioration and to the best on my knowledge there are no other published studies examining the impacts of large earthquakes on benthic communities in an estuarine ecosystem. This research gives valuable insight and advancements in the scientific understanding of the effects that eutrophication recovery and large-scale disturbances can have on the ecology of a soft-sediment ecosystem. iii Table of Contents Table of Contents Abstract.......................................................................................................................i Table of Contents .....................................................................................................iv List of Figures..........................................................................................................viii List of Tables ...........................................................................................................xx Acknowledgements ..............................................................................................xxiv Preface....................................................................................................................xxvi Chapter One: General Introduction..........................................................................1 1.1. Large-scale change in the marine environment....................................................2 1.2. Effects of large-scale change on the structure and functioning of marine communities................................................................................................................4 1.3. Ecosystem resilience and recovery......................................................................6 1.4. Estuarine ecosystems..........................................................................................7 1.5. Objectives and aims.............................................................................................8 1.6. Study site............................................................................................................11 1.7. Key study species...............................................................................................18 1.7.1. Macroinvertebrates...............................................................................18 1.7.2. Marine plants........................................................................................20 Chapter Two: Estuarine Habitats and Communities: Spatial and Temporal Changes in Response to Eutrophication and Disturbance.................................23 2.1. Introduction……………………………………………………………………...…….24 2.2. Methods…………………………………………………………………….…........…28 2.2.1. Study sites............................................................................................28 2.2.2. Field monitoring and sampling protocols..............................................29