Response to Changing Oceanography in the Dove Time Series: a Northumberland Plankton Community Study
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Response to Changing Oceanography in the Dove Time Series: a Northumberland Plankton Community Study Malcolm Charles Baptie Submitted for Doctor of Philosophy School of Marine Science and Technology November 2013 Abstract The Dove Time Series is a plankton monitoring station in the Northumberland coastal sea which has been sampled since 1969. Over the 20th century, major changes have occurred in the North Sea plankton which have in part correlated with oscillation in atmospheric mass over the Northern Hemisphere, the North Atlantic Oscillation (NAO). Westerly winds when the NAO is positive phase block northern low pressure systems and transport warm Atlantic water into the North Sea extending stratification, leading to greater phytoplankton biomass. Phytoplankton biomass in the central North Sea reached a sustained higher level after 1985. Phytoplankton, zooplankton and ichthyoplankton datasets were created or extended from the Dove Time Series to study the effect of oceanographic change at this location. There was a change to a high abundance community 10 years later, in 1995. The most important predictor of phytoplankton abundance was not the NAO index, but the Atlantic Meridional Oscillation (AMO), which exhibits 60-100 year and subordinate 11 and 14 year periodicity, describing a deviation from the long term sea surface temperature (SST) mean in the North Atlantic. Phytoplankton periodicity partly matched the 14 year period in the AMO, which correlates with a feedback mechanism of westerly versus northerly wind in the North Atlantic, regulating ocean-atmosphere heat flux. Zooplankton abundance was predicted by SST and ratio of maximum to minimum abundance by phytoplankton/AMO. Oceanographic conditions that were contemporary with the state of the AMO anomaly after 1995 promoted higher spring phytoplankton abundance and neritic copepod abundance peaks. Ichthyoplankton variability was not synchronous with these lower order changes, probably as a result of different effects on adult fish. The cyclical nature of the AMO means both low and high biomass communities observed in the Dove Time Series are part of one regime. i Acknowledgements I would like to thank my principal supervisor, Dr Jane Delany, for her support and her wisdom in guiding me during this study. Through her patient, assured and honest input, the reader has mercifully avoided the confused and disjointed collection of points they would otherwise have had to confront. I thank her for all of this, and for her friendship. Professor Matt Bentley deserves thanks for some kind words, spoken at the right time. Thanks are due to both Dr Richard Gowen and Dr Roy Sanderson for their criticism of the first submission. Their comments in person and on paper were valuable in reshaping and bringing focus to this thesis. Thanks are due to Dr Eileen Bresnan, of Marine Scotland Science, who was generous with her time and attention in assisting me in learning phytoplankton taxonomy, and continues to inform my views on plankton research. The crews of the R.V. Bernicia were of course crucial to the past and continuing success of the Dove Time Series, and for those I have worked with personally I thank them for going out in all weathers, and for their company and interest in my own part of the Dove Time Seres story. My gratitude goes to the Natural Environment Research Council for their funding during this PhD. Lastly my thanks go to my wife Suhaine, who has proved herself to have the patience of a saint while I have been working on this. She has my solemn promise that we'll get out the house now. ii Table of Contents Chapter 1. A Description of Oceanographic Change in the North Sea and the Responses of the Plankton...................................................................................................................1 1.1 The North Atlantic, North Sea, North Atlantic Oscillation, and the Atlantic Multidecadal Oscillation...............................................................................................1 1.2 Phytoplankton in the North Sea and North Atlantic.............................................11 1.3 Zooplankton in the North Sea and North Atlantic................................................20 1.4 Ichthyoplankton in the North Sea and North Atlantic..........................................28 1.5 Plankton science: field sampling, laboratory analysis and data interpretation of long term trends..........................................................................................................35 1.6 Thesis Aims...........................................................................................................41 Chapter 2: Long Term Hydrometeriological and Biological Variability and its Effect upon Phytoplankton at the Dove Time Series.................................................................42 2.1 Introduction...........................................................................................................42 2.1.1 Aims..............................................................................................................46 2.1.1.1 Null Hypotheses....................................................................................46 2.2 Methods.................................................................................................................47 2.2.1 Field sampling...............................................................................................47 2.2.2 Sample analysis and data manipulation........................................................47 2.2.3 Interpolation of missing values; consideration of approach..........................52 2.2.3.1 Interpolation stage 1: Stineman interpolation........................................53 2.2.3.2 Interpolation stage 2: ordinal logistic regression...................................54 2.2.3.3 Replacement of remaining short gaps...................................................55 2.2.4 Identification of species affinities.................................................................55 2.2.5 State space time series modelling.................................................................58 2.2.5.1 Extracting seasonal peaks from coarse resolution time series...............58 2.2.5.2 Describing interannual cycles and forecasting into the future..............60 2.2.5.3 Sensitivity analysis................................................................................62 2.2.5.4 Model validation....................................................................................62 2.3 Results...................................................................................................................63 2.3.1 Abundance and community composition......................................................63 2.3.1.1 Community composition.......................................................................66 2.3.1.2 Modelling and forecasting abundance...................................................74 2.3.2 Phenological change in abundance of each group........................................78 2.3.3 CCA linear constraints as predictors of abundance.......................................83 2.4 Discussion.............................................................................................................86 2.4.1 Changes in phytoplankton assemblage composition caused by changing hydro-meteorological conditions...........................................................................86 2.4.2 Forecast increases in spring phytoplankton and decreases in other groups..88 2.4.3 Wind and cloud regime promotes a longer and earlier growing season........88 2.4.4 DTS position likely shoreward of frontal zone for part of the year..............89 2.4.5 A persistent shift in phytoplankton abundance – for how long?...................90 2.4.6 Conclusion.....................................................................................................93 Chapter 3: Long Term Oceanographic, Meteorological and Biological Variability and iii Their Effects on Zooplankton at the Dove Time Series..................................................95 3.1 Introduction...........................................................................................................95 3.1.1 Aims..............................................................................................................98 3.2 Methods.................................................................................................................99 3.2.1 Field sampling...............................................................................................99 3.2.2 Laboratory analysis and use of FlowCAM...................................................99 3.2.3 Modelling and term selection......................................................................102 3.3 Results.................................................................................................................103 3.3.1 Time series structure and lack of breakpoint features in abundance...........103 3.3.2 Abundance and community composition....................................................105 3.3.3 Change in phenology of abundance peaks..................................................120 3.4 Discussion...........................................................................................................124 3.4.1 Conclusions.................................................................................................129 Chapter 4. Oceanographic, Meteorological and Biological Change and their Effects on a New Ichthyoplankton