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The Importance of Edge Effects in Determining Fish Distributions in Patchy Seagrass Habitats

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The Importance of Edge Effects in Determining Fish Distributions in Patchy Seagrass Habitats The importance of edge effects in determining fish distributions in patchy seagrass habitats Timothy Malcolm Smith Thesis submitted in fulfilment of the requirements of the degree of Doctor of Philosophy at the University of Melbourne August 2009 Department of Zoology Faculty of Science The University of Melbourne Abstract Boundaries between adjacent habitats can create unique biotic and abiotic conditions, varying species compositions and abundances between the edge and interior of habitats. As habitats become fragmented, the relative amount of edge increases. Understanding the role that habitat edges have in determining species compositions and abundances is fundamental for conservation and management of habitats, particularly those under threat from fragmentation. Seagrass habitats are common nearshore habitats that harbour a rich and diverse faunal assemblage that are under threat worldwide from human disturbance. Human induced fragmentation, and the propensity of seagrass to form naturally patchy landscapes, makes it an ideal system to study the effects of edges on fauna. Evidence of fish displaying edge effects in seagrass habitats is equivocal. Assessment of fish edge effects was done by sampling seven positions within seagrass habitats at fine spatial scales. Strong, consistent patterns in fish distributions demonstrated clear edge effects both within and alongside seagrass at these sites. The total number of fish sampled was greater at the seaward seagrass edge than the seagrass middle, but there was little difference between the seagrass middle and the shoreward seagrass edge. Four individual fish species showed preferences for the seagrass edges. Further investigation revealed that patch size could influence the magnitude of edge effects in seagrass beds. Fish were sampled in ten variously sized seagrass patches in three positions within each patch. Two species showed variations in edge effects across patches which could be attributed to the area of the patch. Changes in patch size can influence the magnitude of edge effects that species display, suggesting that patch area effects (fish density varying with patch size) could be caused by edge effects. Food availability and predation are mechanisms commonly used to explain edge effect patterns. Gut analysis was done on Stigmatopora nigra sampled at the edge and middle of patches to determine if prey consumption varied between positions, and explain S. nigra distribution. There was little difference in prey consumed by S. nigra at the edge and middle of patches, suggesting that food was unlikely to be causing S. nigra edge 2 effects, or that the influence of prey distribution was being masked by other factors such as seagrass structure. Predator abundances and foraging efficiency may vary at the edge and middle of patches, and consequently influence the distribution of prey fish within patches. Underwater videos were placed at four positions within seagrass habitats to assess predator distributions. Predatory Australian salmon, Arripis spp., spend more time over adjacent sand than other positions, while small potential prey species (King George whiting, Sillaginodes punctata, recruits) appear to prefer the middle of seagrass patches, possibly to avoid encounters with salmon. To test if the predator-prey distributions reflected actual predation pressure, a tethering experiment was done to determine if predation was causing edge effects in small fishes. King George whiting recruits and pipefish (Stigmatopora spp.) were tethered at each of the four positions at different depths. Survival time of whiting recruits was greater in the middle of shallow seagrass patches than other positions. Few pipefish were preyed upon, and survival time was lower over sand adjacent to seagrass than at the seagrass edge or middle. Video footage revealed that salmon was the dominant predator of both whiting recruits and pipefish. The distribution of predators and associated predation can explain edge effects for some species (whiting) but other mechanisms, or a combination of mechanisms, are determining edge effects for other species (pipefish). Edge effects were common amongst fish species in seagrass habitats, and included permanent, temporary and predatory species. Patch size was found to influence the extent of the edge effect. There was little evidence to support prey consumption as an underlying mechanism causing higher fish abundances at the interior or edge of patches, however there was evidence that predation could be causing edge effects. Changes in fish distributions within seagrass patches due to patch size and predation when seagrass undergoes fragmentation need to be considered by not only ecologists, but also by managers in the development of plans for seagrass conservation. Future studies should investigate the relative contribution of different edge characteristics in determining the degree of seagrass edge effects. 3 Declaration This is to certify that: i. the thesis comprises only my original work towards the PhD, ii. due acknowledgment has been made in the text to all the material used, iii. the thesis is less than 100,000 words in length, exclusive of tables, maps, bibliographies and appendices. Signature________________ Date____________________ 4 Acknowledgments Firstly I would like to thank my supervisors Jeremy Hindell, Greg Jenkins, Mick Keough and Rod Connolly, who gave me the opportunity to undertake and complete this thesis. Their advice, patience, support and feedback over the years has been invaluable and without them I surely would never have been able to complete this journey. Thank you to everyone who has helped me in the field over the years, particularly sampling during cold, wet nights. Rod, Pete, Dave, Fabian, Hannah, Fi, Jess, Neils, Ted, Claire, Dad and Susi your help in the field will always be remembered. Thanks heaps Jules and Neil for reading over drafts of this thesis. To everyone at the Victorian Marine Science Consortium, Corey, Isla, Pete, Fi, Dave, Fabian, Lauren, Jay, Kathryn, Jo, Justin and particularly Rod and Liz who were always there to help and provided a great working atmosphere. To everyone that has helped me from the Marine and Freshwater Fisheries Research Institute, particularly Paul, Brent, Sean, Dave, Kade, Simon, Dave, Neil and Vicki your time and effort is greatly appreciated. Dave, Fabian, Camilla and Fi have been great housemates to talk over the trials and tribulations of marine science PhD’s and Paul and Ako gave me a bed when I couldn’t find anywhere else for which I am most thankful. I received financial support from Victorian branch of the Australian Marine Science Association for funding my trip to New Zealand to present at a conference, the Australian Research Council who funded the project and the University of Melbourne who provided me with a scholarship so that I could complete my work. To all my mates who have continually harassed and encouraged me, I have finally finished, thanks a lot boys!! Thanks Susi for all you support, encouragement and kindness throughout my time studying, without it things would have been a whole lot tougher. Tals, thanks for being there to help me finish this thesis. To Mum, Dad, Claire and the rest of my family, thanks for all the support, money, effort and encouragement, without it I might never have got to where I am today. 5 6 “One Fish, Two Fish Red Fish, Blue Fish” Dr Seuss 7 Contents List of Tables...........................................................................................................10 List of Figures..........................................................................................................12 1. General Introduction ............................................................................................15 Thesis Outline ..............................................................................................20 2. Edge effects on fish associated with seagrass and sand patches ............................22 Introduction..................................................................................................23 Methods .......................................................................................................25 Results .........................................................................................................29 Discussion....................................................................................................41 3. Seagrass patch size affects fish responses to edges ...............................................45 Introduction..................................................................................................46 Methods .......................................................................................................48 Results .........................................................................................................51 Discussion....................................................................................................56 4. Fine-scale spatial and temporal variations in diets of the pipefish Stigmatopora nigra within seagrass patches in Port Phillip Bay, Victoria, Australia................................61 Introduction..................................................................................................62 Methods .......................................................................................................64 Results .........................................................................................................65 Discussion....................................................................................................77
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