Effects of Marine Reserve Protection on Adjacent Non-Protected Populations in New Zealand
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Effects of Marine Reserve Protection on Adjacent Non-protected Populations in New Zealand by Daniela Díaz-Guisado A thesis submitted to Victoria University of Wellington in fulfilment of the requirements for the degree of Doctor of Philosophy in Marine Biology. 2014 This thesis was conducted under the supervision of: Associate Professor James J. Bell (Primary supervisor) Victoria University of Wellington, Wellington, New Zealand and Professor Jonathan P.A. Gardner (Secondary supervisor) Victoria University of Wellington, Wellington, New Zealand Abstract Marine reserves (MRs) have been established in many parts of the globe for a variety of reasons and there is an increasing body of evidence that indicates they provide a wide range of benefits that can extend beyond their boundaries. In the present study, the biological effects of protection provided by MRs in New Zealand were evaluated, particularly focusing on the potential impacts of reserves on non-protected areas in terms of export of biomass. First, the biological response of two exploited species to MR protection in New Zealand was quantified by comparing meta-analysis results based on response ratio (RR) analysis and Hedges’ g statistics. Then, effect of MR size and age on those biological responses was determined. Most MRs supported a greater density of larger individuals than unprotected areas. Results indicated that the benefits provided by MRs scale with reserve size. Also, MR age explained a significant amount of the variation in the density and length of both species. Comparison of the performance of RRs with Hedges’ g revealed that RR analysis is an appropriate alternative to Hedges’ g statistic for meta-analyses of MR effectiveness because of its ease of use and interpretation. Then, a 14-year time series of fish density data was analyzed to determine early changes in a multi-species fish assemblage inside the Taputeranga Marine Reserve (TMR) compared to adjacent fishing grounds using a Before and After Control-Impact i Paired Series (BACIPS) design. This analysis was performed in order to detect changes in fish density due to protection. Commercial, recreational and traditional fisheries are important in this region and the biomasses of several exploited species have been substantially depleted as a result of fishing. The exclusion of fishing from the area should enable at least some species to recover inside the reserve, as has happened in other reserves in New Zealand. The faster growing, more productive species, and those that have been heavily exploited are expected to recover within a few years. Early changes in density were evident in the area protected by the TMR for most of the species surveyed in terms of the effect size analysis. However, most of the changes were too small to be detected with the statistical analyses that were performed. To determine the most appropriate methodology to be used in a later survey in the study area, two Baited Underwater Video (BUV) methodologies (Horizontal versus Vertical set-up) were compared in terms of their ability to record the density and size of reef fish. Results indicated that both the horizontal and vertical BUV techniques are able to detect both conspicuous and cryptic species and both techniques were effective in the detection of carnivorous species, especially large predatory species such as blue cod, but also effective in the detection of fish species that have been overestimated in terms of abundance by other methodologies. The horizontal BUV technique seems to be a better technique for evaluating reef fish size, especially when measuring large fish that exhibit highly aggressive behaviour. The horizontal BUV technique was later used in conjunction with the Underwater Visual Census (UVC) technique to assess the effects of the protection provided by the TMR. A multispecies analysis was carried out to detect any differences in density and length of fish between reserve and fished areas and to detect gradients of fish density across reserve boundaries that could be related to the occurrence of spillover from the reserve to adjacent fished areas. Density gradients provide indirect evidence of spillover, defined as the movement of adult individuals from reserve to adjacent non-protected areas. Little evidence consistent with a positive effect of reserve protection in the TMR was found. Also, little evidence of spillover was found, with theexception of two target species (blue cod and blue moki). In contrast with the findings of previous studies, density gradients were found for both sedentary and vagile species. These results are consistent with the occurrence of density independent ii spillover that is expected to occur as soon as the density inside reserve areas is higher compared to fished areas. To further understand the patterns of fish movement relative to the effect of protection provided by MRs, spatial differences in density, length and survival of blue cod inside the TMR and adjacent fishing grounds and the movement patterns of the species across the boundaries of the reserve through a capture-mark-recapture (CMR) analysis were examined. CMR studies can provide direct evidence of spillover. Evidence of a positive effect of reserve protection in the TMR for blue cod in terms of increased density, length and survival in reserve areas was found. Also, evidence of high site fidelity of blue cod in both reserve and fished areas, with the majority of individual moving only short distances was found. However, the potential for this species to also travel long distances (>100 km) was confirmed, suggesting the possibility for spillover of the species from reserved to fished areas. Overall, the results of my thesis indicate that New Zealand MRs, consistent with a large body of earlier evidence, are having positive effects on the abundance and size of the species that afford protection to. These results also highlight that both MR age and area are important factors determining the response to protection both in terms of the effects within reserves and on adjacent non-protected areas. Finally, my results highlight the fact that the greater benefits in terms of increased abundance and size, and also movement across reserve boundaries, are obtained for highly exploited species that can potentially move between areas. iii Dedicated to my son; Daniel iv Acknowledgments The completion of this dissertation was made possible through the support of many people that have contributed to my continued development as person and as a scientist. First of all, I am deeply grateful of both my supervisors, Dr James J. Bell and Professor Jonathan P.A. Gardner, for their guidance, encouragement and patience. Much of the fieldwork necessary for the completion of the present study was only possible with help of many people. I would particularly like to acknowledge the support of the technicians at the Victoria University Coastal Ecology Laboratory (VUCEL), John Van der Sman and Daniel McNaughtan. Also, many thanks to Don Nelson, for his skills as a skipper, for passing on local knowledge of the Wellington area and spending long hours onboard the VUW vessel. Thanks to the many field assistants that lent me a hand in the field, often at short notice and thanks to all my dive buddies specially Cesar Cardenas and Tyler Eddy. I am also grateful to my boat skippers, especially to Danelle Lekan, Jenny Oliver, Alejandro Perez-Matus and Ian Geeson. Also thanks to all those friends that helped me during the many field trips particularly to Sergio Carrasco, Mauricio Cifuentes, Paul Mensink, David Burr, Ryan Hughes, and Hayden Smith. I would also like to thank the Department of Conservation, especially Helen Ketles, Debbie Freeman, John Henry, John Adams, Danica Stent and Hawea Tomoana for their support during the monitoring survey of the TMR. Tyler Eddy providing me with some v of the data I used in the analysis of the effects of protection in the TMR. Thanks also to Russell Cole, Robert Davidson, Debbie Freeman, Shane Kely, Alison Macdiarmid, Anjali Pande, Rob Stewart and Carl Struthers for providing me with much of the data I used in the meta-analysis in chapter one and for their comments as co-authors of a published manuscript. Also a special thanks goes to Malcolm Francis (National Institute of Water and Atmospheric Research, NIWA) for teaching me the tagging techniques I used in the capture-mark-recapture (CMR) survey. I am also grateful to Tim Jones, Shane Geange, and Mauricio Cifuentes for their statistical advice, and also to Dalice Sim for helping me to getting started with the statistical software R and providing valuable statistical advice. Thanks to Shirley Pledger for introducing me to the software program MARK and guiding me through the analysis of the CMR data in chapter six. I am grateful to my colleagues at VUCEL for their support and for making my time in New Zealand so enjoyable. A great group of friends have also supported me throughout this process, especially Claudia Campos, Rodrigo Gomez, Gelant Vilches, Leasa Mead, David Irvine and Julie and Steve Gianoutsos who always lent me a hand when I needed it. This dissertation was possible with financial support of CONICYT (Chile) and Victoria University of Wellington (VUW - CONICYT scholarship). Finally, I would like to thank my family, my sister Alejandra and her family for their support, my dad Nelson for his encouragement and especially my mum for teaching me the passion for learning and for being a great example to me. Also, I wish to thank my son Daniel for accompanying me in this journey and for being part of my life. vi vii Contributions to this Thesis Chapter two Dr. Russell G. Cole, Dr. Debbie J. Freeman, Dr Shane Kelly, Dr. Anjali Pande, Dr. Alison MacDiarmid A, Rob Stewart, Carl Struthers M.S., and Rob J. Davidson provided some of the data used in the meta-analysis of New Zealand MRs performed in chapter two.