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Disentangling the Effects of Disturbance and Habitat Size on Stream Community Structure

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Disentangling the Effects of Disturbance and Habitat Size on Stream Community Structure Disentangling the effects of disturbance and habitat size on stream community structure _____________________________________________________________________ A thesis submitted for the Degree of Doctor of Philosophy in Ecology at the University of Canterbury by Phillip Jellyman ____________________________________________________________________ University of Canterbury Christchurch, New Zealand 2011 In loving memory of my friend Matthew Langley 1983 – 2009 Table of contents ___________________________________________________________ Abstract ....................................................................................................................... 1 Chapter 1. General Introduction .............................................................................. 5 Chapter 2. Is disturbance a main driver of variability in stream fish communities? Abstract ............................................................................................................... 15 Introduction ......................................................................................................... 16 Methods ............................................................................................................... 19 Results ................................................................................................................. 30 Discussion ........................................................................................................... 40 Appendices .......................................................................................................... 46 Chapter 3. Disturbance-mediated prey assemblages determine fish community structure Abstract ............................................................................................................... 53 Introduction ......................................................................................................... 54 Methods ............................................................................................................... 57 Results ................................................................................................................. 70 Discussion ........................................................................................................... 82 Appendices .......................................................................................................... 90 Chapter 4. Interactive influences of habitat size and disturbance on top predator body size in stream fishes Abstract ............................................................................................................... 93 Introduction ......................................................................................................... 94 Methods ............................................................................................................... 97 Results ................................................................................................................. 108 Discussion ........................................................................................................... 118 Appendices .......................................................................................................... 127 Chapter 5. Habitat size and external prey subsidies drive changes in stream food-web structure Abstract ............................................................................................................... 135 Introduction ......................................................................................................... 136 Methods ............................................................................................................... 139 Results ................................................................................................................. 146 Discussion ........................................................................................................... 153 Chapter 6. General discussion: disentangling the drivers of stream food-web structure... ............................................................................................................................. 163 Acknowledgements ..................................................................................................... 183 References ................................................................................................................... 187 Abstract ________________________________________________________________ Our ability to predict community responses to environmental stress remains limited. To address this issue, I investigated how species abundance, community composition and food- web structure varied across abiotic gradients (principally disturbance and habitat size) in New Zealand streams. In surveys, community composition, biomass and richness were all strongly influenced by flood-related habitat disturbance, although disturbance influenced each trophic level via different mechanisms. Experiments indicated that macroinvertebrate prey communities were primarily structured by physical disturbance effects, whereas predatory fish communities were structured by physical disturbance effects and disturbance-mediated changes to prey communities. Prey community biomass and composition affected fish species identity and abundance and an in situ stream channel experiment suggested that prey communities were structured by trade-offs between resisting biotic interactions in physically stable environments and successfully exploiting highly disturbed habitats. The prey community traits associated with different disturbance regimes then directly influenced the composition and predatory impact of the resultant fish communities. In addition to disturbance-mediated biotic interactions, abiotic gradients also provided strong selection pressures on predatory fish communities. In particular, disturbance and habitat size strongly influenced predator community responses (e.g., biomass and maximum body size) in surveys and experiments. However, a habitat‟s capacity to support predator community biomass was largely determined by its size. Food-web structure changed with habitat size; small streams supported more prey than predator biomass, whereas large streams had inverted biomass pyramids (i.e., more predator than prey biomass). Similar relationships between food-web structure and habitat size were found in grassland and forested streams, but terrestrial invertebrate subsidies meant that forested streams supported more predator biomass per unit area than grassland stream food webs. My results indicate that human actions resulting in habitat loss (e.g., water abstraction or river impoundment) and increases in flood-related disturbance events (e.g., climate change) are likely to have significant impacts on stream food webs, ultimately leading to habitats that support smaller fish communities (i.e., less biomass, smaller body size). This means that ecologists and managers will need to consider the separate, interactive and indirect effects of disturbance and habitat size on ecological communities if we are to accurately predict and manage food-web responses to global environmental change. 1 2 Frontispiece. A view of the alpine headwaters of the upper Waimakariri River (Photo credit: Angus McIntosh). 3 4 Chapter One General Introduction Globally, ecosystems are experiencing unprecedented levels of pressure due to pervasive impacts from land-use intensification, habitat fragmentation, species invasions, over exploitation and climate warming (Vitousek et al. 1997, IPCC 2001, Foley et al. 2005, Banavar & Maritan 2009). Consequently, one of the main challenges confronting ecologists is to understand and predict how the composition of biological communities may vary given rapid global change (Agrawal et al. 2007). Global change drivers are already having significant impacts on ecosystems (Didham et al. 2007, Tylianakis et al. 2008), and some of the strongest effects are being observed in freshwater environments (e.g., freshwater biodiversity is declining at greater rates than even the most affected terrestrial ecosystems, Ricciardi & Rasmussen 1999). In particular, riverine ecosystems have been subjected to extensive anthropogenic alteration for water, energy, transportation and recreational needs (Nilsson et al. 2005). Whilst it is widely recognised that these actions have significant impacts on biodiversity, understanding of the ecological consequences is limited (Banavar & Maritan 2009). Our inability to predict such consequences arises from significant “gaps” in our current knowledge of community ecology. In reviewing ecological knowledge, Agrawal et al. (2007) identified understanding how abiotic and biotic contexts shape species distribution, abundance and interaction strength over space or time as the major limitation to advances in general ecological theory. To address this knowledge gap, I investigated how species abundance, community composition and food-web structure varied across abiotic gradients (principally disturbance and habitat size) and under different biotic contexts in New Zealand streams. 5 Disturbance and its influence on stream food webs Disturbance is one of the major abiotic factors determining community composition because it alters the relative importance of biotic and abiotic processes (e.g., Peckarsky 1983, Menge & Sutherland 1987). Biotic interactions (e.g., predation, competition) are predicted to strongly influence communities in low disturbance (i.e., benign) conditions,
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