Macrophyte Phenology in a Temporarily Open/Closed Estuary Compared with a Permanently Open Estuary
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MACROPHYTE PHENOLOGY IN A TEMPORARILY OPEN/CLOSED ESTUARY COMPARED WITH A PERMANENTLY OPEN ESTUARY By Deborah Claire Vromans Submitted in fulfilment of the requirements for the degree Magister Scientiae in the Department of Botany Nelson Mandela Metropolitan University Port Elizabeth 2010 Supervisor: J.B. Adams Co-Supervisor: T. Riddin 1 DECLARATION In accordance with Rule G4.6.3, 4.6.3 A treatise/dissertation/thesis must be accompanied by a written declaration on the part of the candidate to the effect that it is his/her own work and that it has not previously been submitted for assessment to another University or for another qualification. However, material from publications by the candidate may be embodied in a treatise/dissertation/thesis. I, Deborah Vromans, student number 209090006, hereby declare that the dissertation for MSc (Botany) is my own work and that it has not previously been submitted for assessment to another University or for another qualification. Signature: ........................................ Date: ........................................25 January 2011 i ACKNOWLEDGEMENTS I am eternally greatful to Professor Janine Adams for putting her faith in me and for supporting my endeavour to undertake a Master of Science degree, without her this would not have been possible. I would like to sincerely thank Ms Taryn Riddin for her quiet kindness and unabiding patience with my seemingly endless throng of queries and her always timely responses. To Professor Eileen Campbell, I extend my appreciation for all the statistical support. To Professor Alan Whitfield, the National Research Foundation and Nelson Mandela Metropolitan University, I extend my thanks for financing this project. For my field and laboratory assistants, Mike Verala, Lucienne Human, Sabine Hoppe-Speer, Jacques Dennis and Amanda Khumalo, thank you for all your hard work and assistance at what seemed, at times, an endless task of wading in or filtering and weighing of muddy sediment. And last but not least to those at home that have supported me through this academic journey, Tom, your astute yet occasional troublesome truths, thank you. To my family, I would like to acknowledge your unwavering support, and to my new furry, four legged friends that always manage to put a smile on my face. In loving memory of my Layla and Roze. ii ABSTRACT Temporarily open/closed estuaries (TOCEs) are unpredictable environments that change in response to mouth condition, which is influenced by freshwater flooding or sea storm surges. The aim of the study was to determine whether macrophyte phenology in a TOCE was event driven rather than cyclically predictable and if it differed from permanently open estuaries (POEs). Macrophyte growth and flowering phenology in response to environmental conditions was investigated in the East Kleinemonde Estuary (TOCE) and the Kowie Estuary (POE) along the eastern coastline of South Africa. The lack of freshwater flooding due to low rainfall coupled with several overwash events resulted in a prolonged period of mouth closure from September 2008 to the end of this study period in 2010. This in turn caused the inundation of the supratidal and intertidal habitats, high water level (> 1.57 m amsl) and high salinity (30 - 42 ppt) in the TOCE. Principle Components Analysis showed that high water level and reduced sediments were the most significant environmental factors affecting macrophyte phenology. Macrophyte phenology in the POE was primarily driven by temperature, sediment redox potential and salinity. The saline high water level and reduced sediment significantly reduced macrophyte cover in all habitats in the TOCE. Macrophytes in the POE maintained high cover abundance due to seasonal re-growth compared to the TOCE where cover declined over the sampling period due to the high water level. Subsequent to water level dropping by as little as 11 - 20 cm in the TOCE, the intertidal species Sarcocornia tegetaria and Salicornia meyeriana completed their life-cycles and produced viable seeds within four and three months of germinating respectively. In contrast, the Sarcocornia hybrid and S. meyeriana in the POE took longer to complete their life-cycles, namely seven and nine months respectively, while S. tegataria did not germinate in situ but reproduced vegetatively despite producing seed. In the TOCE, the submerged species Ruppia cirrhosa and Chara vulgaris completed their life-cycles within five and three months and produced a maximum of 26 242 and 196 998 seeds m-2 respectively. Due to high water level and prolonged inundation, the reproductive periods were shorter for the intertidal and reed and sedge species in the TOCE compared to the POE. Seed output during the two reproduction periods varied between the two estuaries. Sarcocornia decumbens and S. tegetaria produced a substantially higher number of seeds in the TOCE compared to the POE, namely 0 - 102 847 versus 20 661 - 48 576 seeds m-²; and 7 001 - 45 542 versus 1 587 – 16 958 seeds m-² respectively. Seed output in the TOCE was significantly higher in S. tegetaria during the second reproduction period despite the significantly lower plant cover, which may be a function of the stressful environment in the TOCE. Seed production of S. meyeriana was significantly higher in the POE compared to the TOCE, with 264 224 - 640 292 compared with 24 050 - 27 643 seeds m-², due to higher plant cover in the POE. The research suggests that macrophyte phenology in the TOCE was significantly influenced by mouth condition. Further, macrophytes were able to demonstrate considerable phenotypic plasticity in response to changing and unfavourable environmental conditions. These data can be used in mouth management plans and freshwater requirement studies in TOCEs to ensure that macrophytes can complete their life-cycles and produce viable seeds for the safeguarding of habitat persistence and ecological processes. In impacted estuaries where artificial mouth opening is practised and the macrophytes have been severely degraded or extirpated, management should ensure that the intertidal and supratidal habitats are not inundated during peak flowering and seed production periods i.e. late spring to early autumn (November to March). iii TABLE OF CONTENTS DECLARATION……………………………………………………………………………………………………….i ACKNOWLEDGEMENTS……………………………………………………………………………………………ii ABSTRACT ........................................................................................................................................... .iii LIST OF FIGURES……………………………………………………………………………………………………vi LIST OF TABLES……………………………………………………………………………………………………..x 1. CHAPTER 1: INTRODUCTION ....................................................................................................... 1 2. CHAPTER 2: LITERATURE REVIEW ............................................................................................. 6 2.1 Estuaries……………………………………………………………………………………………………..6 2.1.1 Temporarily open/closed estuaries compared with permanently open estuaries…………...6 2.2 Macrophyte habitats………………………………………………………………………………………..10 2.2.1 Salt marsh .................................................................................................................... ..10 2.2.2 Reeds and sedges .......................................................................................................... 13 2.2.3 Submerged macrophytes ................................................................................................ 15 2.3 Effect of environmental conditions on the phenology of macrophytes…………………………… ….19 2.3.1 Climate: Seasonal changes in temperature, rainfall, irradiance and photoperiod .............. 19 2.3.2 Salinity ............................................................................................................................ 22 2.3.3 Water regime .................................................................................................................. 25 2.3.4 Tidal exchange ............................................................................................................... 30 2.3.5 Redox potential ............................................................................................................... 33 2.3.6 Sediment organic matter ................................................................................................. 37 2.3.7 Sediment moisture content.............................................................................................. 40 2.3.8 pH .................................................................................................................................. 44 2.3.9 Light, turbidity and temperature ....................................................................................... 47 2.4 Seed viability…………………………………………………………………………………………….…51 2.5 Synopsis…………………………………………………………………………………………………….57 3. CHAPTER 3: MATERIALS AND METHODS .......................................................................... ……59 3.1 Location of study sites……………………………………………………………………………………..59 3.2 Climate of study sites………………………………………………………………………………….......59 3.3 The East Kleinemonde Estuary…………………………………………………………………….........60 3.3.1 Physical characteristics ............................................................................................... …60 3.3.2 Macrophytes ................................................................................................................... 61 3.4 The Kowie Estuary…………………………………………………………………………………………62 3.4.1 Physical characteristics ..................................................................................................