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Durham E-Theses Durham E-Theses Experimental taxonomy of some members of the Teesdale ora Elkington, Trevor Thomas How to cite: Elkington, Trevor Thomas (1962) Experimental taxonomy of some members of the Teesdale ora, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/9314/ 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 EXPERIMENTAL TAXONOMY OF SOME MEMBERS OF THE TEESDALE FLORA. BY TREVOR THOMAS ELKINGTON B.Sc. (DUNELM) Being a thesis presented in candidature for the Degree of Doctor of Philosophy in the University of Durham 1962. ACKNOWLEDGEMENTS. I wish to thank Professor D.H. Valentine for suggesting that the Teesdale flora would be a rewarding field for study and for the interest he has taken in supervising my work. I should also like to thank D. Briggs and M.J, Harvey for their stimulating thoughts, and also for their help and that of many other friends and colleagues in collecting material and for giving other help to me. I would especially like to acknowledge the help of A. Eddy in determining many of the bryophytes, and the. technical staff of the Botany Department, Durham Colleges for their help and cheerfulness on many occasions. I would also like to express my gratitude to Mr. D.W. Gilchrist Shirlaw for both providing facilities for carrying out the soil analyses at the Durham School of Agriculture, Houghall, and for many valuable discussions on this, subject. I wish to thank Professor T.W. Bocher for his helpful comments and the provision of facilities for finishing this thesis at the Institut for Planteanatomi og Cytologi, K0benhavn, and also to M.C. Lewis for much practical help at this time. The work was carried out during the tenure of a Research Studentship awarded by the Nature Conservancy, whom I should also like to thank for the generous way in which they have supported the fieldwork which these investigations have entailed. INDEX Chapter Page I Introduction. 1. II Geographical Distribution. 15. Ill Cytology. 30. Introduction. 30. Methods. 31. Results. 32. Discussion. 51. IV Studies on individual species. 53. Introduction. 53. Gentiana verna, 54. Taxonomy. 54. Description. 54. Relationships. 55. General Biology. 56. Variation. 60. Potentilla fruticosa. 78. Taxonomy. 78. Description. 79. General Biology. 80. Variation. 83. Myosotis alpestris. 91. Description. 91. Relationships. 92. General Biology. 33. Variation. 95. Dryas octopetala. 100. Description. 100. Taxonomy and Relationships 102. General Biology. 104. Variation. 108. Introduction. 108. Leaf size and shape. 109. Leaf hair and gland characters. 118. V Ecology. 130. Introduction, 130. Vegetation and floristics. 131. Collection of data. 131. Analysis of data. 132. Gentiana verna. 133. Potentilla fruticosa. 137. Myosotis alpestris. 140. Dryas octopetala. 143. Bartsia alpina. 150. Soils. 153. Collection and storage. 153. Analysis methods. 154. Results. 159. Introduction, 159. Gentiana verna. 160. Potentilla fruticosa. 162. Myosotis alpestria. 164. Dryas octopetala. 164. Bartsia alpina. 167. Discussion. 168. Climate. 168. VI General Discussion. 174. Bibliography, 1 CHAPTER I. Introduction The flora of Teesdale first became known to British botanists in the first half of the nineteenth century, although some of the first records were made as early as the end of the eighteenth century by John Binks, a miner living in the area, and were communicated by the Rev. Harriman, Even earlier than this Ray (1689) knew of Potentilla fruticosa from the Tees valley. There are a number of early accounts of the area of which probably the best known are those of the Backhouses published in 1843 and 1844. Prom this time the valley of the upper Tees from Middleton in Teesdale to its source on Cross Pell including the surrounding hills has become famous as one of the richest botanical localities in Britain, especially renowned for the abundance of montane, and rare plants, and perhaps only rivalled in Britain by Ben Lawers and Glen Glova in Scotland. By the end of the first half of the nineteenth century the majority of the flowering plants of the area had been discovered, although this process has continued until the present day, Alopecuinis alpinus only having been confirmed in the area in 1959 by Ratcliffe and Eddy. In the second half 2. of the century interest was aroused in the geographical distribution of the British flora and this was greatly stimulated by the publication of Watson's Cybele Britannica (1847) and Topographical Botany (1873). Interest in arctic plants was also strong at -this time and the distributional types and the species present were analysed in detail by Hooker (1861). As a result of these studies it was realised that there were, in Teesdale, representatives of a wide variety of types of geographical distributions ranging from species with a southern and continental distribution such as Helianthemum canum to arctic-alpine species such as Dryas octopetala. A detailed description of some of the distribu• tional patterns exhibited by species in Teesdale is given here in chapter II. It was also in the middle of the last century that theories were first formed as to the origin of the British flora and here one of the best known pioneer studies was that by Forbes (1846), who put forward the idea that the British flora was a result of immigration since the Ice Age. The same idea was also developed by Darwin who expounded it in detail in the 'Origin of Species' (1859). It was gradually realised that the Teesdale flora presented somewhat special problems in relation to this theory of migration since in the area 3. were a large number of species with northern distribution patterns, some of which were not found in the Scottish mountains and a few nowhere else in Britain, As a result another theory was brought forward to explain this and other localities rich in such disjunct species. This is the perglacial survival theory which seeks to explain such con• centrations of these species as a result of their survival, through the time of glaciation, on unglaciated areas or nunataks in situ. This is to be contrasted with the other alternative, that the total British flora is a result of immigration after the Glacial Period. The arguments are well set out in the discussion of the Royal Society in 1935 on the origin and relationship of the British flora. The views of the proponents of the perglacial survival theory are well summed up by Wilmott (1935) who writes "It seems incredible that these species could have migrated over intervening areas from their nearest stations, have settled on unglaciated Teesdale and have died out over the routes travelled. They are there because they survived glaciation there." The evidence on which a judgement may be made between these two arrival times is mainly indirect. It has been shown by Raistrick (1931) that it is probable that the summits of Cross Pell and Mickle Pell and some of the fells at the head 4. of Weardale were unglaciated. It is conjectural whether plants could have survived on them through the glacial maximum. Such nunataks would probably consist of shoulders of bare rock which would be extremely exposed at all times. It has been pointed out in support of the per-glacial survival theory that nunataka in Greenland at the present day sometimes bear quite a rich flora. It is therefore of interest to see how such nunataks compare with the above picture. BScher (1956) has separated four types of refugia in East Greenland. These are (a) nunatak refugia, mountains projecting through the ice and surrounded by ice in all directions; (b) semi- nunatak refugia, mountains facing the sea on one side, but otherwise surrounded by ice; (c) cross-valley refugia, those containing cross valleys cut out during earlier glaciations, but later unglaciated due to local changes in the drainage of the ice cap; (d) parallel valley refugia, valleys with the same direction as the main valleys (or fjords) and situated between two parallel valleys which drain off the neighbouring ice cap and receive most of the local glaciers. Since in Teesdale all the surrounding areas bear evidence of glaciation it would seem that only the first type, the nunatak refugia, are valid as a comparison. The flora on these nunataks has been shown by Dahl (1946) to be very dependent 5, on the firn line. If this is below the foot of the nunatak, only areas where the snow is blown clear will be clear of ice and such areas bear a very poor flora. Dahl has called these antarctic nunataks. Manley (1959) has considered that in the p;O3t-Aller0d recession in the Lake District, that with an annual rainfall of 70 - 80" the firn line would be in the region of 2300', and also that the annual precipitation would be similar to the present day. It is very difficult to visualise what conditions in full glacial times were like, but it seems possible that the firn line would be as low or lower than this.
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