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The Holocene History of Pinus Sylvestris Woodland in the Mar Lodge

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The Holocene History of Pinus Sylvestris Woodland in the Mar Lodge The Holocene history of Pinus sylvestris woodland in the Mar Lodge Estate, Cairngorms, Eastern Scotland Danny Paterson Institute of Natural Sciences, University of Stirling, Stirling, UK Submitted for the degree of Doctor of Philosophy November 2011 i Statement of Originality I hereby confirm that this research was carried out by the undersigned and that all research material has been duly referenced and cited. Danny Paterson, December 2010 ii Acknowledgements This thesis has taken rather a long time to complete, there’s a long list of people who’ve helped me, several of whom I’ve actually forgotten as they moved on or completed and left and I didn’t. Thanks and apologies to all people with whom I’ve shared an office or lab space and have now vanished from my mind. Thanks for the company to those whose names I can still place, Helen Shaw, Sue Bowen, Althea Davies. Throughout the three changes of departmental name: Department of Environmental Science, School of Biological and Environmental Sciences and now the Institute of Natural Sciences, help from laboratory, IT and cartographic staff, Helen Ewen, Scott Jackson, Bill Jamieson and John McArthur has been invaluable. Bob McCulloch helped with coring and together with Eileen Tisdall also provided guidance regarding pollen preparation and colorimetric analysis. Richard Tipping has provided excellent and cheerful supervision, and David Paterson very helpfully proof read the dissertation. The NTS staff in Mar Lodge have also changed during the time I’ve been working on this project, but special thanks go to Peter Holden and Alister Clunas on the Estate and also to Shannon Fraser. Colin Edwards of Forestry Research has also been very helpful with aspects of pine ecology. Lastly Shirley has waited (fairly) patiently as the project spent a long time apparently going nowhere, provided support when it seemed to be going into reverse and also proof read the whole thing. Thank you for that. iii Abstract This thesis investigates the past extent, structure and dynamics of Mar Lodge Caledonian pine (Pinus sylvestris L.) woodland, near Braemar in the south-eastern Cairngorms. The geographical extent and Holocene history of the Scottish pinewoods are generally understood, but the Mar pinewoods are relatively obscure. This thesis is concerned with the Holocene history of the Mar pinewoods; the timing and reasons for first appearance, the increase in abundance of Pinus to become a dominant species, the spatial extent of the woodland and its structure and form. The investigation includes changes to the woodland assemblage during its fragmentation and disappearance in the late Holocene and possible influences on the woodland from people living in the area. At the heart of this thesis is an understanding of the factors underpinning the ecology of Pinus and the response of the species to competition with other taxa. This is related to the spatial and temporal changes in climate that contribute to the location and development of Pinus in Scotland and Mar Lodge. Areas comparable to Mar Lodge are defined as ‘core areas’ of pine woodland rather than ‘native areas’. This avoids the necessity of considering every short period of colonisation by Pinus in areas distal to large populations. The location, extent, form and behaviour of woodland according to macro sub-fossils and micro sub-fossils is used to define core woodland as those with a long presence of Pinus, often continuing to the present day. Areas with a long history but no extant population are regarded as peripheral areas. This thesis consists of extensive palaeoecological investigations of three peat sequences: from within extant pine woodland (Doire Bhraghad), from just beyond its edge (White Bridge) and from peat with sub-fossil pine stumps located 10km west of the modern range of Pinus (Geldie Lodge). A range of techniques, including loss of mass on ignition and colorimetric light transmission analysis are applied to the peat, but palynological techniques form the basis of the investigation. Stomatal counts are used in iv conjunction with pollen counts to explore the process of Pinus colonisation, and its increase in abundance to form woodland. Pinus percentage and influx, together with the ratio of arboreal to non-arboreal pollen and the percentage of Empetrum are used to define the density of the woodland canopy. The stability of the Doire Bhraghad assemblage confirms the area as core Pinus woodland. Pinus is present from c. 9600 cal BP and dominates woodland from c. 9150 cal BP. Woodland here is a closed, solely Pinus canopy from c. 8600 until 4000 cal BP. Arrival of Pinus at Geldie Lodge is undated but occurs before c. 7550 cal BP. Woodland is always more open; Pinus is co-dominant with Betula, showing affinity with other peripheral areas. Pinus woodland fragments at all Mar Lodge sites from c. 3900 cal BP, disappearing from Geldie Lodge by c. 2800 cal BP and White Bridge by c. 1900 cal BP. Calluna replaces Pinus as the dominant species at all three sites. The disappearance of Pinus is thought to relate to regional climatic change toward wetter conditions. At Geldie Lodge a prior Coleopteran study suggests Pinus growing on the mire surface to be small and short lived. These may not have been the only trees growing in the area but they perhaps contributed to the major fluctuations in arboreal and non- arboreal pollen. Early canopy fluctuations (c. 7550 to 6000 cal BP) at Geldie Lodge may be related to Mesolithic human activity; there is stronger evidence of human presence from c. 4000 cal BP, possibly including cereal cultivation. Evidence from Doire Bhraghad and White Bridge is indicative only of low intensity grazing activity. It is unlikely that human activity instigated the fragmentation and disappearance of woodland, but may have contributed to the process. v Contents List of Figures ........................................................................................................... - 3 - List of Tables ............................................................................................................ - 5 - 1 Thesis rationale and organisation........................................................................... - 7 - 2 Defining the past presence and abundance of Pinus in Scotland: a review of the literature ....................................................................................................................... 10 2.1 Introduction ........................................................................................................ 10 2.2 Preferred conditions for growth of Pinus sylvestris ........................................... 12 2.3 Native woodland, core areas and peripheries .................................................... 17 2.4. Reconstructing past presence, extent and behaviour of Scottish Pinus woodland .................................................................................................................................. 24 2.5 Macroscopic plant remains ................................................................................ 25 2.5.1 Research history and background ............................................................... 25 2.5.2 Woodland and environmental reconstruction ............................................. 26 2.6. Microscopic plant remains: history, interpretive problems and reconstructions .................................................................................................................................. 34 2.6.1 History of research ...................................................................................... 34 2.6.2 Defining source areas of pollen .................................................................. 37 2.6.3 Pinus pollen and pollen diagrams ............................................................... 38 2.6.4 First appearance of Pinus in Scotland from microscopic remains .............. 44 2.6.5 Factors influencing first appearance of Pinus and woodland expansion .... 47 2.6.6 Separating local establishment and expansion ............................................ 48 2.6.7 Associated taxa ........................................................................................... 50 2.6.8 Visual appearance of the trees and woodland ............................................. 54 2.6.9 Demise ........................................................................................................ 57 2.7 Other proxy data for past Pinus woodland ........................................................ 61 2.8 Summary of Chapter 2, hypotheses and aims for the project ............................ 62 2.8.1 Woodland history ........................................................................................ 62 2.8.2 Core areas of woodland .............................................................................. 62 2.8.3 First Appearance and woodland expansion ................................................ 63 2.8.4 Woodland density ....................................................................................... 64 2.8.5 Woodland diversity and competitive interactions....................................... 64 2.8.6 Use of recently developed palynological methods ..................................... 65 2.8.7 Timescales................................................................................................... 65 2.8.8 Other proxies for woodland ........................................................................ 66 2.8.9 Anthropogenic activity...............................................................................
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