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The Biodiversity of the Wealden Ghyll Woodlands: Species Richness, Abundance and Distribution Patterns in a Rare and Fragmented Habitat

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The Biodiversity of the Wealden Ghyll Woodlands: Species Richness, Abundance and Distribution Patterns in a Rare and Fragmented Habitat The biodiversity of the Wealden ghyll woodlands: species richness, abundance and distribution patterns in a rare and fragmented habitat Andrew R. Flint A thesis submitted in partial fulfilment of the requirements of the University of Brighton for the degree of Doctor of Philosophy March 2014 The University of Brighton - For my mum - Dungeon-Ghyll Force William Wordsworth Along the river's stony marge The sand-lark chants a joyous song; The thrush is busy in the wood, And carols loud and strong. A thousand lambs are on the rocks, All newly born! both earth and sky Keep jubilee, and more than all, Those boys with their green coronal; They never hear the cry, That plaintive cry! which up the hill Comes from the depth of Dungeon-Ghyll. Said Walter, leaping from the ground, "Down to the stump of yon old yew We'll for our whistles run a race." --Away the shepherds flew; They leapt--they ran--and when they came Right opposite to Dungeon-Ghyll, Seeing that he should lose the prize, "Stop!" to his comrade Walter cries-- James stopped with no good will: Said Walter then, exulting, "Here You'll find a task for half a year. "Cross, if you dare, where I shall cross-- Come on, and tread where I shall tread." The other took him at his word, And followed as he led. It was a spot which you may see If ever you to Langdale go; Into a chasm a mighty block Hath fallen, and made a bridge of rock: The gulf is deep below; And, in a basin black and small, Receives a lofty waterfall. Wordsworth’s notes: Title: "Ghyll," in the dialect of Cumberland and Westmoreland, is a short and, for the most part, a steep narrow valley, with a stream running through it. "Force" is the word universally employed in these dialects for waterfall. Abstract The Wealden ghyll woodlands are associated with unique plant assemblages that include nationally rare bryophyte species with oceanic affiliations. The identification and monitoring of this type of ‘priority’ habitat, recognised as important in terms of regional and national biodiversity, is a central facet of the UK Biodiversity Action Plan (UKBAP). Despite the acknowledged importance of ghyll woodlands for non-vascular plant species, previous studies attempting to examine and characterise the ghyll woodlands have neglected to include these bryophyte communities. This research identifies and characterises the Wealden ghyll woodlands through an examination of the spatial and temporal distributions of bryophyte and flowering plant species. The research also seeks to provide baseline data against which biodiversity levels can monitored. In order to identify and contextualise the importance of ghyll woodland in terms of regional biodiversity, survey data was collected from other types of ancient woodland throughout the region for comparative analysis. The study involved the collection of species and environmental data from a total of 1440 random quadrats from 60 survey sites situated throughout the Weald, as well as the use of archive survey data collected during two 20 year periods (1951-1970 and 1976-1995). A number of statistical approaches including general linear modelling, ANOSIM, Mann- Whitney U and Spearman rank correlation analysis were used to identify the environmental correlates of spatial and temporal changes in species distributions. Spatial analysis indicated that ghyll woodland is restricted to the stream valleys themselves which were significantly richer in bryophyte and flowering plant species than the surrounding woodlands. NVC classifications assigned to the ghylls indicated the presence of ‘oceanic’ plant communities that are associated with damp, humid microclimatic conditions. A number of authors have explained the presence of oceanic bryophytes within the ghylls as being the result of a damp, humid microclimate present within the stream valleys. However, the study found no significant differences between climatic conditions within the ghyll valleys and those in the surrounding ancient woodlands. ANOSIM analysis indicated that community composition was influenced by site substrate, with clay and sandstone ghyll woodlands containing significantly different plant communities. Chi-squared analysis identified a temporal increase in the ratio of oceanic bryophytes and ancient woodland indicator flowering plant species during the study period. Analysis of Ellenberg indicator values indicated a move towards more shade- tolerant plant communities within the ghyll woodlands. The patchily distributed ghyll woodlands were examined for signs of habitat fragmentation through genetic analysis of the bryophyte Conocephalum conicum (Great Scented Liverwort) using the random amplification polymorphic DNA technique (RAPD). Wright’s fixation index (FST) and Nei’s coefficient of gene variation (GST) both indicated a loss of genetic diversity characteristic of genetic isolation. A Mantel test based on Nei’s genetic distance values indicated that the genetic isolation observed was not correlated with the geographical distance between populations. The study indicated that temporal changes are occurring in the composition of ghyll woodland plant communities and that bryophyte populations are displaying symptoms of genetic isolation. The study illustrates the importance of some form of monitoring program if the biodiversity value of these sites is to be maintained. iv Table of Contents LIST OF FIGURES ..................................................................................................................... VIII LIST OF TABLES .......................................................................................................................... IX APPENDICES ................................................................................................................................ XI ACKNOWLEDGEMENTS ........................................................................................................... XII CHAPTER 1 INTRODUCTION .................................................................................................... 1 1.1 The Wealden ghyll woodlands ........................................................................................................... 1 1.2 The formation of the Wealden ghyll woodlands ................................................................................ 3 1.2.1 The significance of the Wealden geology ....................................................................................... 3 1.2.2 The history of the Wealden woodlands .......................................................................................... 5 1.3 Ancient woodland within the Weald .................................................................................................. 9 1.4 The biodiversity of ghyll woodlands ................................................................................................. 11 1.4.1 Ghyll woodland bryophytes .......................................................................................................... 11 1.4.2 Ghyll woodland vascular plants .................................................................................................... 13 1.4.3 Ghyll woodland fauna ................................................................................................................... 14 1.5 The major threats to Wealden ghyll woodland biodiversity ............................................................. 14 1.5.1 Changes in woodland management .............................................................................................. 14 1.5.2 Climate change .............................................................................................................................. 15 1.5.3 The effects of habitat loss and woodland fragmentation ............................................................. 16 1.5.4 The abundance of ghyll woodland sites in the Weald .................................................................. 17 1.6 Study objectives ............................................................................................................................... 18 1.7 Thesis structure ............................................................................................................................... 18 2 A SPATIAL INVESTIGATION OF THE RICHNESS, ABUNDANCE AND COMPOSITION OF GHYLL WOODLAND VEGETATION .................................................. 21 2.1 Introduction ..................................................................................................................................... 21 2.1.1 Identifying ghyll woodland plant communities ............................................................................. 21 2.1.2 The distribution of plant species and the ecological niche ........................................................... 23 2.1.3 Environmental factors influencing the distribution of plant species ............................................ 24 2.2 Aims ................................................................................................................................................. 30 2.3 Methods .......................................................................................................................................... 30 v 2.3.1 Study area ..................................................................................................................................... 30 2.3.2 Site selection ................................................................................................................................. 30
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