ACTA PHYTOGEOGRAPHICA SUECICA 53 EDIDIT SVENSKA VAXTGEOGRAFISKA SALLSKAPET Plant Cover and Environment of Steep Hillsides in Pite Lappmark By Jim Lundqvist UPPSALA 1968 ALMQVIST & WIKSELLS BOKTRYCKERI AB ACTA PHYTOGEOGRAPHICA SUECICA 53 Plant Cover and Environment of Steep Hillsides in Pite Lappmark avec un resume en francais By Jim Lundqvist By due permission of the Faculty of Science of the University of Uppsala to be publicly discussed at the Institute of Ecological Botany (Vaxtbiologiska I nstitutionen), on May 24, 1968, at 10 a.m., for the degree of Doctor of Philosophy UPPSALA 1968 Illustrations printed with contribution from Langmanska Kulturfonden Printed in Sweden by Almqvist & Wiksells Boktryckeri AB Uppsala 1968 To the Memory of my Mother CONTENTS PREFACE 7 THERMOPHILOUS PLANT COMMUN ITIES OUTSIDE THE HILLSIDE ECOSYSTEM ...10 3 INTRODUCT£0 9 The flora and vegetation of the Merk Gorge . 103 The flora and vegetation in an old river valley 105 ENVIRONMENT 13 Concluding remarks on the study of the ve- getation . 106 Introductory remarks . 13 Altitude . .. 14 HILLSI DES LOCATED IN THE CALEDONIAN 15 Bedrock and soil . AREA . 107 The development of talus slopes 15 The chemical properties of the bedrock and the 1. Mt. Akkapakte . 107 soil ......... 18 2. Mt. E Ramanpakte 108 Local climate and sun exposure . 29 3. Mt. W Ramanpakte 108 Introduction . 29 4. Paktesuolo . 109 Measuring methods and gauges. Sources of 5. Mt. Ardnapakte 109 error ........... 32 6. Mt. Kebnevare . 110 The radiation climate of mountain slopes 33 7. Mt. Kaldopakte 111 The thermometer screens and their placing . 36 8. Mt. Jokkokvarats . 111 Monthly mean temperatures of the stations and 9. Mt. Laisvare . 112 the dependence on the weather . 37 10. Mt. Fabmevare . 112 Minimum temperatures . 42 11. Mt. Korpberget 113 Maximum temperatures 44 12. Mt. Markepakte 113 Soil temperatures 48 13. Mt. Storberget . 114 The snow conditions during thaw. 50 14. Mt. Alep Nadok 116 Phenology ........ 52 15. Mt. Staburknosen 116 Supply of water . 56 16. Mt. Aistjakk . 117 Precipitation and interception . 56 HILLSIDES LOCATED IN THE AREA OF PRI- Surface and sub-surface waters 59 MARY ROCKS . 118 THE FLORA OF THE HILLSIDES . 64 17. Mt. Pellavardo . 118 18. Mt. Lulep Istjakk . 118 Introduction . 64 19. Mt. Vuornats 120 The floraof vascular plants 64 20. Mt. Jutavare . 120 The bryophyte flora 74 21. Mt. Dalktjapakte 121 Conclusions 82 22. Mt. Ailesvare 121 23. Mt. MAfevare 122 THE VEGETATION 83 24. Mt. Valbmapuoda 122 25. Mt. Kuottavare 122 Introduction . 83 26. Mt. E Starpatsvare 123 Terminology and methodology 83 27. Mt. Fisktraskl:erget 124 The vegetation types of the hillsides 86 28. Mt. Harrejaurliden 124 Poeto-Lactucetum . 87 29. Mt. Brunberget . 125 Tortulo-Poetum . 88 30. Mt. Barberget . 126 Stands with Vaccinium vitis-idaea or other 31. Mt. Gaddaberget 126 dwarf shrubs 92 32. Mt. Halberget 127 Rhytidietum 93 94 Dicrano-Polytrichetum RESUME 128 A comparison with the vegetation on talus slopes in Sikilsdalen . 96 REFERENCES 147 Zonation of vegetation on tal us slopes 96 The succession of vegetation 101 TABLES 1-2, 5, 13-14, 17-3 1 (behind the text) Acta Phytogeogr Suec 53 PREFA CE The principal problems regarding the complex of possesse lists of vascular plants from them. They are biocoeno es and factors comprising the hillside eco­ hardly of any definite value in the present discussion system were summarized by Du Rietz (1954, pp. 174 and have only to some extent been commented on in et seq.). He gave a review of the earlier plant eco­ this book. The material will be published later by logical and botanical literature and the principal theo­ G. Wistrand who will also make a study of special ries and known facts with regard to the environmen­ groups of plants on the basi of the distribution type tal factors. Du Rietz (op. cit., p. 183) concluded that in N Scandinavia only. "what would be desirable at the present time would The hillsides in other parts of Fennoscandia vis­ be a differentiated analysis of the plant communities ited by the present author have been important for a of the hillside vegetation in a way performed by wider view of the hillside ecosystem, but, with the Nordhagen (1943, pp. 542-567) . and a funda­ probable exception of the hillsides of Sikilsdalen, S mental special study of the ecology of every species Norway, they have only very briefly been commented of the hillside" (translated from the Swedish). on. Especially important have been visit to the The aims of the present investigations were prin­ Kuusamo region in Finland, the Porsanger and Alta cipally to investigate the environmental factors and regions in Finnmark, N Norway, and the valleys of the phytocoenoses or special pioneer communities in Junkerdalen and Dunderlandsdalen close to the the way proposed and suggested by Du Rietz. The boundary between Pite Lappmark and Nordland, N environmental factors were studied with the aim of Norway. giving the background of the later mainly syneco­ On the suggestion of my teacher, the late Profes­ logical discussion of the book and not in the sense of sor G. E. Du Rietz, who had recently published the producing causal facts about adaptation or require­ inspiring paper just discussed, I decided (in 1956) ment. In Du Rietz' defini'tion of the term synecology to make an investigation of the plant ecology of the was included the study of habitat factors, their eco­ hillsides of Pite Lappmark. As soon as my academic logical amplitudes and the interrelationships between degree was obtained (in 1958) I started out with my the habitat factors and the plant communities. Stress field work. My studies were always generously en­ was laid on the study of both the climatic and couraged by my former teacher, who acted a a edaphic aspect . Autecological aspect remained less con tant ource of knowledge and information. I studied. deeply regret that he did not ee the work completed. As regard the phytocoenoses and the fragmentary Professor H. Sjors has always taken great interest occurrence of only some single individuals grouped in the problems of the ecological discussion and ha together because of similar demands on abiotic habi­ kindly revised the manuscript. tat, the principal aim was to define and classify I am much obliged to my teachers in meteorology, pecial units. This was done without regard to habi­ Professor T. Bergeron and Professor G. H. Lilje­ tat but on the basis of exclusive, preferential, dif­ quist, for much advice and interest in my climato­ ferential and other indicator species. logical studies. The colleages at the Meteorological In a few cases a comparative study was performed Institute, Uppsala, I thank for all kindness and help of some atypical warm sites outside the hillside given me during past years. ecosystem but containing a few southern species. Dr. G. Wistrand has been an inspiring comrade This was done to supplement the study of the inter­ and friend always with valuable views on the plant relations existing between a "southern" flora and its geography of Pite Lappmark. extreme northern habitats. Several specialists have given me valuable help in Some hillsides in Pite Lappmark, especially of ex­ checking the material of vascular plants. This is posures other than near S, were not investigated by above all the case with Professor J. A. Nannfeldt. I the present author, although a regional survey was also thank him for discussions and for information regarded as important. Some hillsides, mainly of S on the distribution of Dryopteris assimilis. I also re­ aspect, have been visited only, by G. Wistrand who ceived help from Professor N. Hylander and Dr. H. Acta Phytogeogr Suec 53 8 Plant cover and environment in Pile Lappmark Smith; Professor E. Almquist and Mr. S. Norden­ tuffs) and Rikets Allmanna Kartverk for giving me stam, Crown Forester, determined the material of some height values of mountain summits. H ieracium which will be published later. For all service received at the Institute of Eco­ In the field of bryology, many scientists have given logical Botany, Uppsala, I am especially grateful. me valuable help in the checking of difficult pecies. Without the inspiring environment, the kindness and In earlier years, Dr. 0. Martensson gave me much the assistance received through the years this work help. Later Mrs. E. Nyholm, Dr. S. Arnell, Dr. E. would not have been completed. I include in my Sjogren, Dr. E. von Krusenstjerna and Dr. T. Ko­ thanks all visitor to the Institute with which I have ponen have given me special assistance. Dr. P. 0. had many inspiring discussions. Nyman informed me on the bryology of the NW During my field work, including my visits to Fin­ alpine and subalpine area of Pite Lappmark. Dr. H. land and Norway, I have come in contact with many Persson gave me information on his routes of travel­ people who have received my wife and me very ling in Pite Lappmark in 1928 and 1930. The Kee­ kindly. My father assisted me in the field especially pers of the Botanical Museums at Lund, at Gothen­ in early years. burg and at Helsinki informed me on bryophyte ma­ In the summer of 1967 I had the opportunity of terial from Pite Lappmark. taking part in the International Phytogeographical Dr. G. Sandberg has given me inspiring advice Excursion through the Jura, organized by Professor especially in connection with his lecture series. Dr. Dr. R. TUxen. I am indebted to Professor Dr. A. A. Rapp I thank for discussions on the development Quantin, Professor Dr. J. Gehu, Besan9on, and Dr. of tal us slopes, and Dr. P. Holmgren for discussions J. L. Richard, Neuchatel, for rich information on the on the rate of photosynthesis under different condi­ dependence of vegetation on slope exposure in the tions of environment.