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PDF (Volume 1) Durham E-Theses Phytosociology and community boundaries of the British heath foundation Bridgewater, P. How to cite: Bridgewater, P. (1970) Phytosociology and community boundaries of the British heath foundation, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/9690/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk Phytosociology and Community Boundaries of the British Heath Formation. A thesis submitted by P. Bridgewater, B,Sc. (Dunelm), to the University of Durham, for the Degree of Doctor of Philosophy. Department of Botany, University Science Laboratories, South Road, DUKiAL'l. June. 1970. This thesis, which is entirely the result of my own work, has not been accepted for any degree, and is not being submitted concurrently in candidature for any other degree. CONTENTS Abstract Acknowle dgeme nt s Species nomenclature SECTION I. Introduction The project outlined, some notes of the Zurich-Montpellier system of phytosociology, Ideas to date on Heathland Vegetation in Britain and Europe. SECTION II. Comparative Methodology A review of several techniques for vegetation analysis, using a standard data set. SECTION III. Vegetation Boundaries An analysis of the nature and function of some boundaries between plant communities. SECTION IV. Phytogeography of the British Heath Formation. Production of pflytogeographical 'spectra' for heathland vegetation, and erection of Heathland provinces. SECTION V. Community descriptions The ecology of the vegetation units described. SECTION VI. Historical Phytosociology Use of standard phytosociological techniques and information to illustrate the recent vegetation in areas now urbanised, etc. Bibliography Appendices I - VI are to be found in Vol. II. r ABSTRACT A review of phytosociological techniques, including the Zurich-iiontpellier (Z-M) system, Association analysis and Simple ordination is made, using a standard data set. The complementary nature of the results obtained is noted. As classification erects divisions and boundaries in a data set, and the prime method used in the analysis of Heath vegetation is classificatory, evidence for the prescence of ' real1 vegetation boundaries is presented, together with a discussion of their nature and function. Heathland vegetation in Britain has been classified using the Z-*l phytosociological system, and four main types have been distinguished, which are equated with alliances at present in use in Europe. These are; Erica cine re a Heath: Ulicion nanae (Duvijnd, I944)em. San den Bergen. Calluna vulgaris Heath: Calluno-Genisition pilosae (Duvignd, 1944) Vaccinium myrtillus Heath: Llyrtillion bore ale (BScher, 1943). Erica tetralix Heath: Ericion tetralicis (Schwick, 1933). The first three are typical of Dry heaths, whereas the fourth indicates Uet heath. A phytogeographical analysis of the Heath vegetation gives further validity to these four types, indic• ating the dependance of the three dry 'Heaths' on species which are geographically restricted. Use of the Z-M system and some other techniques to indicate the nature of recent vegetation in areas now urbanised or agri- culturalised is made, using South Gloucestershire as such an area. (ii) SPECIES NOMENCLATURE. Species are named from the following sources; Lichens James, P.W. - A new check-list of British Lichens. The Lichenologist, (3), 1965* Hepatics Jones, E.W. - An annotated list of British Hepatics. Trans. B.B.S. , 3, 1958. Mosses Richards, P.W. and Wallace, E.C. An annotated list of British Mosses. Trans. B.B.S., 1, 1950. Vascular plants Dandy, J.E. - List of British Vascular Plants, London, 1958, together with changes noted in Dandy, J.E. - Nomenclatural changes in 'List of British Vascular Plants'. Watsonia, (7) , 1969. Species named from other sources are; Zygogonium ericetorum. Kfltz. Helianthemum nummularium, (L) Miller. Pilosella offioinarum. O.H. and F.W. Schultz. P. peleterana (Merat). C.H. and F.W. Schultz. In the text and tables the following abbreviated names are used; Agrostis montana for A. canina ssp. montana. Sarothamnus maritimus for S. scoparius ssp. maritimus. Hypnum ericetorum for H. cupressiforme var. ericetorum. Also, the two species Hieracium vulgatum and Rubus fruticosus are named to the infra specific section, where possible (i.e. R. discolores) (i) A0KNQT?7LEDGEI,IENTS. I wish to thank Professor D. Boulter for the provision of research facilities in the Botany Department, University of Durham, for the duration of this project. Also, I am extremely grateful to the numerous people who helped with accommodation and guidance in the field work of the project, particularly Dr. D. Shimwell, University of Hull, Dr. S. Woodell, University of Oxford and Mr. M. Horwood, Nature Conservancy, Dorset. My special gratitude is due to Dr. J. de Smidt, Institute of Plant Systematics, State University of Utrecht, for his hos• pitality, and helpful ideas on the classification scheme, and the European context of British Heath. Some field work from the Shropshire area is the result of members attending a field course in phytosociology at Preston Montford Field Centre, conducted by Professor R. Tttxen and Fr. J. J. Moore - whose further help and advice is gratefully acknowledged. Thanks are also due to my Research colleagues - particularly Mr. F. Pollett and Mr. C. Marshall for hours of thought provoking discussion.1 In particular, I should like to thank my Supervisor, Dr. D. J. Bellamy, for his helpful guidance and enthusiasm through• out the project. Finally, I should like to thank the N.E.R.C. for the provision of financial assistance throughout the project. I I. INTRODUCTION. 1.1. Aims of the project. The major part of this project has been a study, over as wide a range as possible, of vegetation belonging to the British heath formation and the construction of a classification of this veg• etation, in relation to existing information on the continent of Europe. In addition, studies have been made on the nature of 'boundaries' within the Heath formation, and between vegetation of the Heath formation and other formations. At the outset it was realised that soi^c restriction! would have to be imposed on the Geographical areas studied. To this end Britain (mainland Scotland, England and "ales, with associated Islands and the Isle of Han) was taken as the unit area, and Ireland excluded. The extreme aceanityof this latter country, with its attendant wide range of Heathland vegetation made its inclusion untenable at this stage. An additional part of the project was to test the effect of several differing 'numerical* methods against a standard data set. This introduction deals with some theoretical aspects of the Zfirich-IIontpellier (Z.LI.) School of Phytosociology, existing "synsys- tematics" of the Heath formation, and the results of previous work attempted in Britain on the heath formation. 1.2. Terminology. 'Phytosociology' is usually regarded, in English speaking countries, as the study (quantitative or qualitative) of plant communities, and that is the sense in which it is used from now on. The term Formation has been used and defined many times in the early ecological literature. First used by G-risebach (1838) with reference to the controlling influence of marcoclimate on vegetation, V/eaver and Clements (1929) reinforced this idea, terming formation as "a fully developed, or climax, community of a natural area in which the essential climatic relations are similar, or identical - It is a product of the climate and controlled by it." Raunkiaer (1910) distinguished between Plant climatology, "the science dealing with the distribution of the major types of world vegetation, controlled by climate" and the Theory_of Formations "dealing with the grouping of Individual species of the Flora on a given substratum". Moss (1907, 1911) used the idea that all associations (his sense) that are developed on the same habitat constitute a 'formation^ (i.e. all communities developed over chalk, including grassland, scrub chalk heath, beechwood etc., would constitute a 'formation'). TCith respect to this useage of the term, Tansley (1920) observed; "So far as England is concerned this is a perfectly good, workable, objective classification, affording an excellent conspectus of the actual veget• ation." Though this may be true, the system is obviously untenable in an international context, and, it would be difficult to reconcile it with existing International Systems. Tansley (1939) synthesised some of these differing viewpoint and defined formations as "relatively stable communities determined.by edaphic or biotic factors, each marked by the dominance of a charac• teristic lifa-form, but divisible into Geographical regions." Dansereau (1957), further simplified this into the following definition "one or more plant communities exhibiting a definite structure and physiognomy;
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