Intertidal resource cultivation over millennia structures coastal biodiversity




Cox, Kieran D.

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Cultivation of marine ecosystems began in the early Holocene and has contributed vital resources to humans over millennia. Several more recent cultivation practices, however, erode biodiversity. Emerging lines of evidence indicate that certain resource management practices may promote favourable ecological conditions. Here, I use the co-occurrence of 24 First Nations clam gardens, shellfish aquaculture farms, and unmodified clam beaches to test several hypotheses concerning the ecological implications of managing intertidal bivalve populations. To so do, in 2015 and 2016, I surveyed epifaunal (surface) and bivalve communities and quantified each intertidal sites’ abiotic conditions, including sediment characteristics and substrate composition. In 2017, I generated three-dimensional models of each site using structure-from-motion photogrammetry and measured several aspects of habitat complexity. Statistical analyses use a combination of non-parametric multivariate statistics, multivariate regression trees, and random forests to quantify the extent to which the intertidal resource cultivation structures nearshore biodiversity Chapter 1 outlines a brief history of humanity's use of marine resources, the transition from extracting to cultivating aquatic taxa, and the emergences of the northeast Pacific’s most prevalent shellfish cultivation practices: clam gardens and shellfish farms. Chapter 2 evaluates the ability of epifaunal community assessment methods to capture species diversity by conducting a paired field experiment using four assessment methods: photo-quadrat, point-intercept, random subsampling, and full-quadrat assessments. Conducting each method concurrently within multiple intertidal sites allowed me to quantify the implications of varying sampling areas, subsampling, and photo surveys on detecting species diversity, abundance, and sample- and coverage-based biodiversity metrics. Species richness, density, and sample-based rarefaction varied between methods, despite assessments occurring at the same locations, with photo-quadrats detecting the lowest estimates and full-quadrat assessments the highest. Abundance estimates were consistent among methods, supporting the use of extrapolation. Coverage-based rarefaction and extrapolation curves confirmed that these dissimilarities were due to differences between the methods, not the sample completeness. The top-performing method, random subsampling, was used to conduct Chapter 4’s surveys. Chapter 3 examines the connection between shellfish biomass and the ecological conditions clam garden and shellfish farms foster. First, I established the methodological implications of varying sediment volume on the detection of bivalve diversity, abundance, shell length, and sample- and coverage-based biodiversity metrics. Similar to Chapter 2, this examination identified the most suitable method, which I used during the 2015 and 2016 bivalve surveys. The analyses quantified several interactions between each sites’ abiotic conditions and biological communities including, the influence of substrate composition, sediment characteristics, and physical complexity on bivalve communities, and if bivalve richness and habitat complexity facilitates increases in bivalve biomass. Chapter 4 quantifies the extent to which managing intertidal bivalves enhance habitat complexity, fostering increased diversity in the epifaunal communities. This chapter combines 2015, 2016, and 2017 surveys of the sites' epifaunal communities and habitat complexity metrics, including fractal dimension at four-resolutions and linear rugosity. Clam gardens enhance fine- and broad-scale complexity, while shellfish farms primarily increase fine-scale complexity, allowing for insights into parallel and divergent community responses. Chapter 5 presents an overview of shellfish as a marine subsidy to coastal terrestrial ecosystems along the Pacific coast of North America. I identified the vectors that transport shellfish-derived nutrients into coastal terrestrial environments, including birds, mammals, and over 13,000 years of marine resource use by local people. I also examined the abundance of shellfish-derived nutrients transported, the prolonged persistence of shellfish subsidies once deposited within terrestrial ecosystems, and the ecological implications for recipient ecosystems. Chapter 6 contextualizes the preceding chapters relative to the broader literature. The objective is to provide insight into how multiple shellfish cultivation systems influence biological communities, how ecological mechanisms facilitate biotic responses, and summarize the implications for conservation planning, Indigenous resource sovereignty, and biodiversity preservation. It also explores future work, specifically the need to support efforts that pair Indigenous knowledge, and ways of knowing with Western scientific insights to address conservation challenges.



Shellfish Mariculture, Species-Habitat Relationships, Resource Management, Spatial Subsidies, Structure-from-Motion Photogrammetry, Multivariate Random Forest