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Ghost Ponds” : Resurrecting Lost Ponds and Species to Assist Aquatic Biodiversity Conservation

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Ghost Ponds” : Resurrecting Lost Ponds and Species to Assist Aquatic Biodiversity Conservation “Ghost ponds” : Resurrecting lost ponds and species to assist aquatic biodiversity conservation Thesis submitted for the degree of Doctor of Philosophy UCL Department of Geography UCL 1 “ I took in February three table-spoonfuls of mud from three different points, beneath water, on the edge of a little pond; this mud when dry weighed only 6 and ¾ ounces; I kept it covered up in my study for six months, pulling up and counting each plant as it grew; the plants were of many kinds, and were altogether 537 in number; and yet the viscid mud was all contained in a breakfast cup!” Excerpt from: Charles Darwin. “The Origin of Species by Means of Natural Selection / Or, the Preservation of Favoured Races in the Struggle for Life, 6th Edition.” 2 Thesis declaration 3 Abstract Over the last century, around half of all ponds and small wetlands have been lost globally, predominantly from agricultural regions. These losses present a major conservation challenge for aquatic biodiversity and habitat connectivity. While the creation of new ponds is widely advocated as a means of restoring some of these lost habitats, this thesis presents an alternative approach – the resurrection of ‘ghost ponds’, historic ponds in-filled during agricultural intensification. Many aquatic organisms have evolved dormant propagules for surviving periods of habitat desiccation, and can remain viable for centuries buried in wetland sediments. While such long-term viability is well established in extant aquatic habitats, the fate of propagules buried within ghost ponds has remained unexplored. If their viability was proven, ghost ponds could present a crucial overlooked conservation resource. Here, I examine the potential conservation value of ghost pond resurrection within a typical agricultural UK landscape. First, historic declines in pond numbers and pond density are examined for this landscape, and the potential implications of these changes for aquatic biodiversity are explored using a space-for-time replacement approach. The conservation value of ghost ponds is then explored through a series of lab and field studies, that demonstrate the viability of propagules representing at least eight aquatic macrophyte species, after laying dormant for 50 – 150 years. Once resurrected, ghost ponds were quickly (<6 months) re-colonised by a diverse aquatic vegetation. With ghost ponds representing abundant, dormant time capsules for aquatic species in agricultural landscapes, these findings open up exciting new possibilities for aquatic habitat and biodiversity restoration on a global scale. 4 Acknowledgements Firstly, I would like to thank my supervisors Carl Sayer and Jan Axmacher for their continued support and patience, and their invaluable contribution to the ideas explored in this thesis. I would like to express my extreme gratitude to all of the people who have helped me in the field, in particular during the gruelling excavation and set-up of the study ponds and on-site mesocosms. Particular thanks go to Dom Arnold for his excellent digger-work, and to Ian Patmore, Phill Brown, Carl Sayer, Emily Smith, Phoebe Lewis, Bernard Dawson, and Henry Crawley, without whose tireless dedication to pond restoration and creation, this study would not have been possible. Thanks also go to Geoff Nobes for his assistance with the water beetle surveys, to Steve Lambert for his assistance with the microcosm experiment, and to Rachael Davies at the Millennium Seed Bank for her help with seed viability testing. Special thanks go to the landowners and managers who allowed me to dig up their fields, all in the name of pond science! To Nick Anema, the Jensen family, and Ross Haddow & the Stody Estate, a big thank you for taking part in this study. I also wish to thank the Natural Environment Research Council (NERC), Norfolk Biodiversity Partnership, and ENSIS Ltd. for funding this research. Finally, I would like to thank my friends and family, not just for their love and support, but because most of them also helped out in the field. Special thanks go to uncle-Phill for his exceptional botany skills, to my dad John for helping on numerous survey trips, and to Sarah Morgan and Dan Denis for their emotional support, both in the field and at home. 5 Contents Abstract…… .......................................................................................................................... 4 Acknowledgements ................................................................................................................ 5 List of tables ........................................................................................................................ 10 List of figures ....................................................................................................................... 12 Chapter 1 An introduction to ponds ................................................................................ 14 1.1 What is a pond? ................................................................................................... 14 1.2 Why are ponds important? - understanding an undervalued aquatic habitat ....... 17 1.2.1 Ponds & ecosystem services ........................................................................ 18 1.2.2 What makes ponds so biodiverse? ............................................................... 19 1.2.3 Biodiversity value of farmland ponds .......................................................... 23 1.3 Factors threatening the biodiversity value of ponds ............................................ 25 1.3.1 Threats relating to pond water quality ......................................................... 26 1.3.2 Threats relating to modification of a pond’s structure, shape, ………... or permanence .............................................................................................. 27 1.3.3 Invasive species threats to ponds ................................................................. 28 1.3.4 Pond loss through abandonment and terrestrialisation ................................ 30 1.3.5 Pond loss through deliberate in-filling ........................................................ 33 1.4 A brief introduction to habitat fragmentation – structural connectivity .............. 33 1.5 Functional connectivity, and an organism’s view of the landscape .................... 37 1.4 What is a ‘ghost pond’? ....................................................................................... 39 1.5 Thesis aims and objectives .................................................................................. 42 Chapter 2 Study sites ........................................................................................................ 47 2.1 The ponds of Norfolk ........................................................................................... 47 2.2 Study site selection .............................................................................................. 51 Chapter 3 Changes in an agricultural ‘pondscape’ – a history of pond loss and pondscape fragmentation. ......................................................................... 55 3.1 Introduction .......................................................................................................... 55 3.2 Chapter hypotheses .............................................................................................. 60 3.3 Methodology ........................................................................................................ 61 3.3.1 Ground truthing the Ordinance Survey maps .............................................. 61 3.3.2 Assessing changes in pond numbers ............................................................ 62 3.3.3 Assessing changes in pond density .............................................................. 64 3.3.4 Future potential – Adjusting analysis to incorporate spatial patterns and …………spatial autocorrelation in pondscape change. .............................................. 66 6 3.4 Results ................................................................................................................. 67 3.4.1 Ground truthing study .................................................................................. 67 3.4.2 Local scale changes in pond numbers ......................................................... 71 3.4.3 Landscape scale changes in pond numbers ................................................. 74 3.4.4 Local scale changes in pond density ........................................................... 75 3.4.5 Regional scale changes in pond numbers .................................................... 78 3.4.6 Regional scale changes in pond density ...................................................... 80 3.5.1 Ground truthing study .................................................................................. 85 3.5.2 Changes in pond numbers ........................................................................... 86 3.5.3 Ecological implications of the loss of different pond types ........................ 90 3.5.4 Ecological implications of changes in pond density and fragmentation …………of the pondscape .......................................................................................... 92 3.6 Conclusions ......................................................................................................... 97 Chapter 4 The ecological consequences of pond loss and pondscape fragmentation – …………….a space for time substitution approach. .......................................................... 99 4.1 Introduction
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