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Faunal associations with drifting macrophytes and wrack accumulations in the nearshore of South ... PDF

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Faunal associations with drifting macrophytes and wrack accumulations in the nearshore of South Australian sandy beaches Ryan J. Baring BSc. Mar Biol (Hons), Flinders University A thesis submitted for the degree of Doctor of Philosophy (PhD) School of Biological Sciences, Faculty of Science and Engineering Flinders University September 2014 Table of Contents Summary .................................................................................................................................. iii Declaration.............................................................................................................................. vii Statement of contribution..................................................................................................... viii Acknowledgements .................................................................................................................. x Chapter 1: General Introduction ........................................................................................... 1 1. Drifting macrophytes in coastal ecosystems ………………………………………………. 1 2. Study background: Patchiness in time and space of drifting macrophytes …………………6 3. Macroinvertebrate assemblages associated with drifting macrophytes ……………….........7 4. Fish assemblages associated with drifting macrophytes ……………………………………9 5. Trophic interactions associated with drifting macrophytes ………………………………..12 6. Source to sink: The drift pathway ……………………………………………………........14 7. Aims of this project ………………………………………………………………………..18 Chapter 2: Steps for establishing precise estimates of floral and faunal distributions in patchy marine habitats: an example from nearshore marine environments ……………20 1. Introduction ………………………………………………………………………………..20 2. Methods ……………………………………………………………………………............27 3. Results ………………………………………………………………………………..........33 4. Discussion ……………………………………………………………………..…………..44 5. Conclusion …………………………………………………………………………….......48 Chapter 3: Storm versus calm: variation in fauna associated with drifting macrophytes in sandy beach surf zones …………………………………………………………………..50 1. Introduction ………………………………………………………………………………..50 2. Methods ……………………………………………………………………………..……..54 3. Results ……………………………………………………………………………………..61 4. Discussion ………………………………………………………………………….……...74 5. Conclusion …………………………………………………………………………………81 i Chapter 4: Trophic structure within surf-zone wrack accumulations at sandy beaches ………………………………………………………………………………………83 1. Introduction …………………………………………………………………………..........83 2. Methods ……………………………………………………………………………………88 3. Results ………………………………………………………………………………..........94 4. Discussion …………………………………………………………………………..........110 5. Conclusion …………………………………………………………………………..........118 Chapter 5: Drift dynamics of natural versus artificial seagrass at various distances from shore and initial colonisation by invertebrates and fish ………………………………...120 1. Introduction ………………………………………………………………………………121 2. Methods ……………………………………………………………………………..........124 3. Results ……………………………………………………………………………………134 4. Discussion …………………………………………………………………………..........155 5. Conclusion ………………………………………………………………………………..164 Chapter 6: General Discussion …………………………………………………………....166 1. Key findings ……………………………………………………………………………...167 2. Key areas for future research ……………………………………………………………..179 3. Management implications ……………………………………………………..…………180 4. Conclusion ………………………………………………………………………………..182 References ………………………………………………………………………………….185 Appendices …………………………………………………………………………………203 ii Summary The research in this thesis identifies and addresses some of the knowledge gaps surrounding the role that detached and drifting macrophytes play as habitat and potential food resources in the nearshore of sandy beaches. So far, very little research has been conducted in southern Australia to investigate if important detached macrophytes are to nearshore production. This is surprising considering the large volumes that regularly accumulate as wrack in the surf zone and on sandy beaches, and in some cases wrack is a semi-permanent feature. Very few beach management programs have considered the role of drifting macrophytes and wrack accumulations in coastal ecosystems, which is concerning when beach cleaning or wrack harvesting procedures are in place. Beach cleaning can occur up to multiple times a week along metropolitan Adelaide beaches, particularly during the warmer months. In South Australia, the amounts of wrack removed from beaches varies with approximately 50-110 tonnes of algae and 3000 tonnes of seagrass per year from the South East region of the state (PIRSA 2007). To establish efficient sampling designs for investigating drifting wrack and associated fauna in surf zones of sandy beaches, I undertook a carefully-planned pilot study to establish optimal-precision estimates for sampling macroinvertebrates and ichthyofauna. I investigated the precision of the mean for various numbers of sub-samples and replicates of seine nets to establish a larger sampling program. The pilot study showed that the processing time for individual seine-net samples could be reduced by 50 % using sub-sampling compared to double the amount of time taken to process the whole sample and the time taken to process replicate seine net samples could be reduced by 25 %, while maintaining acceptable precision. The pilot study and exploratory approaches used in Chapter 2 were essential for establishing precise estimates of flora and fauna associated with very patchy habitats, such as wrack in sandy-beach surf-zones. iii The effects of regular storms on drifting marine macrophytes, consequent accumulation of wrack (i.e. seagrass or macroalagae or both) and associated fauna in beach surf zones across three different regions was investigated. Results showed that the influence of storms may be more pronounced in sheltered coastal waters compared to more exposed coastlines, where biota could have adaptations to persist in larger swell conditions. Regional differences were more obvious than the impact of weather, with distinct macrophyte and macroinvertebrate species richness, abundances and assemblages occurring in each region. Fish were less aligned to the regional patterns identified for wrack or invertebrates. This study also showed that beach morphology, specifically grain size and beach slope, may be important to the ecology of surf zones. The new knowledge about the ecology of faunal associations in nearshore waters during storm and calm weather in multiple regions highlights the importance of surf-zone wrack as habitat for many fauna. A separate study was undertaken to investigate possible trophic pathways associated with wrack accumulations, which are little understood, particularly where wrack compositions vary or where wrack is a semi-permanent feature in sandy-beach surf-zones. In this study, I sampled the gut contents of fish and analysed the δ13C and δ15N stable isotope signatures of fish, macroinvertebrates and macrophytes. Results showed that fish may be using wrack as habitat but are likely to be foraging widely over multiple habitats away from wrack accumulations. More evidence exists that some invertebrates such as grazers may be using wrack accumulations for both habitat and food. Although preliminary, this study identified baselines for trophic pathways associated with wrack accumulations in these sandy-beach surf-zones, which can be applied to more directed studies in future. More directed studies of specific trophic pathways, other food sources and fauna in nearby habitats (e.g. benthic fauna) would provide better understanding of the trophic pathways within wrack accumulations and also linkages between wrack accumulations and nearby habitats. iv Previously, no studies had investigated the surface drift dynamics of detached seagrass. Existing studies have mainly focused on larger macroalgae such as kelps or fucoids in open-ocean systems, rather than along sheltered coastlines. Here, the trajectories of detached natural and artificial seagrass were investigated to establish the drift pathways and short-term colonisation of fauna to drifting macrophytes close to shore. Various release distances from shore in gulf waters in South Australia were compared. Tagged seagrass generally travelled in the same direction as tides but patterns were variable across sampling days and different distances from shore. Many tagged seagrass units released close to shore were stranded on beaches after six hours but the eventual stranding of drifting macrophytes further offshore was more difficult to quantify. The colonisation of macroinvertebrates and fish to tagged seagrass was compared to fauna found from in situ drifting (natural) macrophytes. The results of the drifting macrophyte study also showed that colonisation by fish to drifting macrophytes are more likely to be for habitat, rather than food. Also, more macroinvertebrates may colonise drifting macrophytes at distances closer to shore, possibly due to shallower depths and thus increased proximity of source habitats. This study provides further knowledge about macrophyte drift dynamics, the short-term colonisation of fauna and habitat function along nearshore coastlines. This is essential for further understanding of the ecological significance of allochthonous material arriving on shorelines. The research in this thesis was strategically planned to investigate and better understand the dynamic movement of nearshore drifting macrophytes, their arrival into surf zones and the fauna that utilise the drifting material as habitat and a food resource. This thesis had an equal amount of research effort placed on the study of macrophytes, macroinvertebrates and fish across multiple regions that were variously dominated by macroalgae and/or seagrasses. The research encompasses some of the few studies anywhere v where much wrack in the surf zone is from seagrasses. Information gained from this research will be useful for future investigation of the ecological function and management of nearshore ecosystems, particularly where processes such as beach cleaning occur. Future management of allochthonous inputs such as wrack in the nearshore zone is particularly important in southern Australia where it has received no attention in previous beach management programs. vi Declaration I certify that this thesis does not incorporate without acknowledgement any material previously submitted for a degree or diploma at any university; and to the best of my knowledge and belief it does not contain any material previously published or written by another person except where due reference is made in the text. Ryan Baring vii Statement of Contribution This thesis was prepared as a series of manuscripts (Chapters 2-5) which are either in the process of being, or will in the future be, submitted to various scientific journals. Prof. Peter Fairweather and Dr Rebecca Lester, as my primary supervisors, provided conceptual and methodological input throughout the writing of this thesis. Dr Charlie Huveneers as my secondary supervisor assisted with input into drafting of some chapters at the later stages of writing. Chapter 2 will be submitted in the near future to a methods based journal, which is yet to be decided. This manuscript is co-authored by my supervisors, Prof. Peter Fairweather, who provided assistance to me with conceptual design, data analyses and editing the manuscript, and Dr Rebecca Lester, who also provided assistance with the writing of R script and subsequent data analyses. I reviewed the literature and undertook all field data collection, laboratory processing and species identification, data analyses and manuscript preparation. My proportional contribution to this manuscript was 85 %, Dr Rebecca Lester contributed 10 % and Prof. Peter Fairweather contributed 5 %. . Chapter 3 has been published in Journal of Experimental Marine Biology and Ecology (see Baring et al. 2014) . This manuscript is co-authored by my supervisors, Prof. Peter Fairweather and Dr Rebecca Lester, who provided assistance with conceptual design, data analyses and editing the manuscript. I reviewed the literature and undertook all field data collection, laboratory processing and species identification, data analyses and manuscript preparation. My contribution to this manuscript was 90 %, and Prof. Peter Fairweather and Dr Rebecca Lester contributed 5 % each. viii Chapter 4 will be submitted in the near future to Estuaries and Coasts. This manuscript is co- authored by my supervisors, Prof. Peter Fairweather and Dr Rebecca Lester, who provided assistance with conceptual design, data analyses and editing the manuscript. Dr Charlie Huveneers provided advice with editing of the manuscript. I undertook all field data collection, laboratory processing, data analyses and manuscript preparation. My contribution to this manuscript was 89 %, with Prof. Peter Fairweather and Dr Rebecca Lester contributed 5 % each and Dr Charlie Huveneers 1 %. Chapter 5 will be split into two manuscripts in the near future, with one manuscript focusing upon the drift movement of detached macrophytes and the second manuscript focusing on colonisation and biological interactions with associated fauna offshore. The two manuscripts will be submitted to Estuarine, Coastal and Shelf Science and Marine Ecology Progress Series, respectively. These manuscripts will be co-authored by my supervisors, Prof. Peter Fairweather and Dr Rebecca Lester, who provided assistance with conceptual design, data analyses and editing the manuscript. I undertook all field data collection, laboratory processing, data analyses and manuscript preparation. My contribution to this manuscript was 90 %, and Prof. Peter Fairweather and Dr Rebecca Lester contributed 5 % each. ix

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Thank you to Flinders technical staff Sandra Marshall, Phil Michelsen, derived material to food webs between the swash zone and into beach
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