Seagrass Meadows As Biodiversity and Productivity Hotspots
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Seagrass Meadows as Biodiversity and Productivity Hotspots New Zealand Aquatic Environment and Biodiversity Report No 137. M.A. Morrison, M.L. Lowe, C.M. Grant, P.J. Smith, G. Carbines, J. Reed, S.J. Bury, J. Brown ISSN 1179-6480 (online) ISBN 978-0-478-43764-5 (online) October 2014 Requests for further copies should be directed to: Publications Logistics Officer Ministry for Primary Industries PO Box 2526 WELLINGTON 6140 Email: [email protected] Telephone: 0800 00 83 33 Facsimile: 04-894 0300 This publication is also available on the Ministry for Primary Industries websites at: http://www.mpi.govt.nz/news-resources/publications.aspx http://fs.fish.govt.nz go to Document library/Research reports © Crown Copyright - Ministry for Primary Industries TABLE OF CONTENTS EXECUTIVE SUMMARY 1 1. OBJECTIVES 3 2. A GENERAL INTRODUCTION TO SEAGRASS SYSTEMS 3 2.1 Global understanding 3 2.2 New Zealand state of knowledge for Zostera muelleri 4 2.3 Seagrass connectivity and role as an ecosystem fuel 6 2.4 Seagrass replication and connectivity mechanisms 7 3. METHODOLOGY 8 3.1 Biogeographic Survey Work 8 3.2 Field operations 9 3.3 Laboratory Methods 13 3.4 Data analysis 16 3.5 Seagrass as an ecosystem fuel (isotopic analysis) 17 3.6 Seagrass replication mechanisms 19 4. RESULTS 21 4.1 Biodiversity of fish associated with seagrass across New Zealand 21 4.2 Spatial variations in composition and fish length of dominant species 25 4.3 Nursery role of seagrass for fish species 29 4.4 Seagrass characteristics 34 4.5 Multivariate community analysis of fish assemblages 37 4.6 Diet 42 4.7 Dietary changes with fish size and habitat 44 4.8 Biodiversity of invertebrates associated with seagrass across New Zealand 62 4.9 Seagrass associated infaunal invertebrate communities – multivariate 72 4.10 Seagrass as an ecosystem fuel component 82 Primary Producer Stable Isotope Signatures 82 4.11 Seagrass replication mechanisms 88 Primer evaluation 88 Genetic variation at the New Zealand wide scale 88 Genetic variation at the local scale 90 Sequencing selected fragments 93 5. DISCUSSION 93 5.1 Nursery role of seagrass habitats – fish 93 Spatial variation in fish abundance 93 Fish diet 95 Ontogenetic changes in diet 96 Habitat related changes in diet 96 Prey diversity 96 Variation of diet within seagrass meadows 97 Geographical setting: are all seagrass meadows equal? 97 Exclusivity of seagrass as nursery areas 98 5.2 Nursery role of seagrass habitats – shellfish 99 5.3 Faunal biodiversity and secondary production associated with seagrass habitats 99 5.4 Seagrass as a dominant ecosystem fuel 101 5.5 Seagrass replication and connectivity 101 6. RISK IDENTIFICATION AND APPRAISAL FRAMEWORK 102 6.1 Distribution 102 6.2 Historical distribution 105 6.3 Threats and stressors 106 6.4 Ecological appraisal and decision matrix for New Zealand seagrass meadows 107 7. CONCLUSIONS AND PRIORITIES FOR FUTURE RESEARCH 110 Implications for future research on fish nurseries 110 8. ACKNOWLEDGMENTS 112 9. REFERENCES 112 10. APPENDICES 123 10.1 Appendix 1 Stable Isotopes 123 10.2 Appendix 2: Operon 10-base oligonucleotide primers tested in Z. muelleri. 124 10.3 Appendix 3. Species list of infaunal invertebrates – New Zealand biogeographic survey of seagrass sites (mean density per core ± s.e.) 125 10.4 Appendix 4: Infaunal invertebrate community composition – PERMANOVA pairwise test133 10.5 Appendix 5: Species list of epifaunal invertebrates – New Zealand biogeographic seagrass survey (mean density per core ± s.e.). 141 10.6 Appendix 6: Epifaunal invertebrate community composition – PERMANOVA analysis 144 EXECUTIVE SUMMARY Morrison, M.A.; Lowe, M.L.; Grant, C.M.; Smith, P.J.; Carbines, G.; Reed, J.; Bury, S.J.; Brown, J. (2014). Seagrass meadows as biodiversity and productivity hotspots. New Zealand Aquatic Environment and Biodiversity Report No. 137. 147 p. A large scale survey of the communities associated with seagrass (Zostera muelleri) meadows and adjacent bare sediments for both fish and infaunal/epifaunal assemblages was undertaken across New Zealand to investigate any potential trends in biodiversity and secondary production with latitude or bioregion. Fish assemblages sampled from the nine locations revealed that subtidal seagrass meadows from northern New Zealand were important juvenile fish nurseries, particularly for species such as snapper and trevally. However, the relative fish nursery value of seagrass meadows varied strongly, dependent upon depth (tidal position), coast, landscape setting and latitude. South of Cook Strait, only spotties and piper were present, while seagrass meadows in Southland supported higher abundances of pipefish and juvenile leather jackets. From a strictly fisheries based view-point, this means that northern subtidal seagrass meadows are of much higher economic value. A small number of species such as yellow-eyed mullet and mottled triplefin had more universal distributions. These results show that the value of a given habitat type is contextual, being affected by factors such as biogeography and local setting, as well as habitat quality (e.g., seagrass blade height and density, water depth, and patchiness). Ontogenetic dietary shifts were evident for the majority of fish species, with meiofaunal crustaceans (0.5–1 mm) predominating in the diet for fish less than 25 mm in size. Newly recruited fish exhibited an obligatory planktivorous stage, with a gradual transition to a diet of larger crustaceans such as gammaridean amphipods (particularly for subtidal meadows), mysids and caridean shrimps and crabs. Habitat-related differences in diet were also evident, reflecting benthic prey availability and diversity. The role of seagrass habitat on the composition of faunal invertebrate communities (both infaunal and epifaunal) is complex and highly variable spatially. The presence of seagrass does not always equate to higher abundance, species richness or secondary production when compared to local bare or sand habitats. In terms of secondary production derived from seagrass compared to bare habitats, no consistent latitudinal trends were apparent for associated invertebrate communities. Subtidal seagrass sites were not identified as having higher secondary production values when compared to their intertidal seagrass counterparts, in contrast to it’s fish nursery values (north of Cook Strait). Random amplified polymorphic DNA (RAPD) markers were evaluated for detecting genetic variation in Z. muelleri, the seagrass species found in New Zealand, and subsequently used to estimate genetic variation within and among regions. Significant genetic differentiation was found among seven regional populations. There was no evidence for a simple isolation by distance model at the national level, or within the east coast, or North Island sites. The high level of regional genetic differentiation is indicative of limited gene flow, with little long distance (over 100 km) dispersal of seeds or vegetative parts of plants between widely separated geographic regions. At the local scale (less than 1 km) the majority of genetic variation was found within sites (over 86%) and there were shared composite genotypes among sites, indicative of clonal reproductive strategy. Investigation of the stable isotope signatures of seagrasses along with other sources of primary production from a relatively pristine harbour (Rangaunu Harbour) and a harbour which is known to be influenced by anthropogenic pollution sources (Kaipara Harbour), provided preliminary results that indicate that seagrass was not the main ecosystem fuel for either of these harbours, as higher order Ministry for Primary Industries Seagrass Meadows as Biodiversity and Productivity Hotspots 1 consumers had stable isotope values for carbon which fell in between those for seagrass and phytoplankton sources. An ecological appraisal framework is provided to aid managers in objectively ranking seagrass meadows of varying ecological value based on the findings of this current report. 2 Seagrass Meadows as Biodiversity and Productivity Hotspots Ministry for Primary Industries 1. OBJECTIVES 1 Complete a national bio-geographic assessment of seagrass associated biodiversity. 2 Quantify seagrass connectivity with surrounding marine landscapes through nursery functions and detritus export. 3 Quantify seagrass replication/connectivity mechanisms: reproductive or clonal? 4 Develop an appraisal model for seagrass systems in New Zealand. 2. A GENERAL INTRODUCTION TO SEAGRASS SYSTEMS 2.1 Global understanding Seagrasses are a unique group of flowering plants that exist fully submerged in the sea. Seagrasses are distributed globally, but unlike terrestrial angiosperms exhibit low taxonomic diversity (approximately 60 species worldwide), with 12 genera. All species share similar architecture and physiology, and perform similar ecosystem functions. Seagrasses are a characteristic component of many coastal areas ranging from subarctic to temperate and equatorial regions, reaching their most southerly global distribution at Stewart Island, New Zealand (Hemminga & Duarte 2000, Turner & Schwarz 2006). Seagrasses commonly occur in sheltered areas, away from strong currents and wave action, on a variety of substrata ranging from mud through to sand and bedrock (Hemminga & Duarte 2000, Green & Short 2003). However, the most extensive meadows are found on soft substrata, often forming continuous expanses over several square kilometres. Alternatively, they can form mosaics of discrete patches (often in areas with more wind-generated