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Sommerfeltia sommerfeltia 2 R.H. 0kland & E. Bendiksen The vegetation of the forest-alpine transition in the Grunningsdalen area, Telemark, S. Norway. - 1985 sommerfeltia is owned and edited by the Botanical Garden and Museum, University of Oslo . SOMMER FELTIA is named in honour of the e minent Norwegian botanist and clergyman S~ren Christian Sommerfelt (1794- 1838). The generic name Sommerfeltia has been used in (1) the lichens by Florke 1827, now Solorina, (2) Leguminosae by Schumacher 1827, now Drepanocarpus, and (3) Compositae by :r.e ssing 1832, nom . cons. SOMMERFELTIA is a series of monographs in plant taxonomy, phytogeography, phytosociology, plant ecology, plant mo rphology, and evolutionary botany . Papers are by Norwegian authors. They are in English or, less often, in Norwegian with an Enqlish summary. An article must be 32 printed pages or more to be accepted. Authors not on the staff of the Botanical Ga rden and Museum in Oslo pay a page charge of NOK 20.00. SOMMERFELTIA appears at irregular intervals, one article per volume. Editor: Dr . Anders Danielsen. Editorial Board: Scientific staff of the Botanical Garden and Museum. Address: SOMMERFELTIA, Botanical Garden and Museum, University of Oslo, Trondheimsveien 23B, N-0562 Oslo 5, Norway. Order: On a standing order (payment on receipt of each volume) SOMMERFELTIA is supplied at 30% discount. Separate volumes are supplied at the following regular rates (1985 prices): Volumes of 32-42 pages NOK 0.80 per page Volumes of 43 - 60 pages N0K 0.65 per page Volumes of 61-120 pages NOK 0.45 per page Volumes o f more than 120 pages NOK 0 . 35 per page. sommerfeltia 2 R.H. 0kland & E. Bendiksen The vegetation of the forest-alpine transition in the Grunningsdalen area, Telemark, S. Norway. 1985 ISBN 82-9905-278-8 ISSN 0800-6865 Økland, R.H. & Bendiksen, E., 198�1_ The vegetation of the forest-alpine transition in the Grunningsdalen area, Telemark, S. Norway. - Sommerfeltia 2: 1-224. ISBN 82-9905-278--8. ISSN 0800-6865. This study is divided into two parts. The general part contains a review of theories of the nature of vegetation. It is concluded that present evidence points in the direction of species individuality and vegetational continuum as responses to continuous environmental gradients, on a regional, and mostly also on a local scale. Accordingly, a gradient approach to classification of the forest and alpine vegetation of the investigated area was designed as an alternative to traditional phytosociological classification. The importance of the concept of corresponding vegetation types in different regions is stressed. It is argued that four environmental gradients explain the major variation in Southern Norwegian forest and alpine vegetation. In the special part, the vegetation of the Grunningsdalen area is treated. Two gradients, the vertical gradient and the complex-gradient topographic moisture - snow cover, are considered sufficient to explain the major variation in forest and alpine vegetation in the area. The vegetation is classified into 16 site-types by division of the gradients into four zones (according to altitude) and four series (according to moisture - snow cover) by means of floristic criteria known to reflect particular ecological conditions. For each of the site-types a description of the vegetation and an extensive comparison with corresponding Fennoscandian vegetation is given. On the basis of results from the present study area regional variation in Southern Norwegian poor vegetation corresponding to the xeric, subxeric, and submesic series, and phytosociological classification in the forest­ alpine transition are discussed. Various numerical classification and ordination methods are used in an analysis of the floristic composition of the site-types and the autecology of the species. The main phytosociological gradient in the investigated vegetation runs from dry and high altitude to wet and low altitude, most closely approaching the moisture gradient. Diversity relations are discussed. It is strongly emphasized that a hierarchic system is unable to give a consistent classification of a vegetation that must be regarded as a multidimensional network of variation along environmental gradients. Viewed in the light of the results of this study, a gradient approach to classification seems most suitable for a variety of Fennoscandian ecosystems. Keywords: Alpine vegetation, Continuum, Diversity, Forest, Forest-alpine transition, Gradient reference frame, Norway, Numerical analysis, Regional variation, Site-types, Telemark. Rune Halvorsen Økland* and Egil Bendiksen, Botanical Garden and Museum, Univ. of Oslo, Trondheimsveien 23B, N-0562 Oslo 5, Norway {*present address: ØKOFORSK, Norwegian agricultural Univ., Box 64, N-1432 As-NLH, Norway]. SOMMERFELTIA 2 (1985) CONTENTS INTRODUCTION . 7 AIMS 8 REALIZATION 8 PREVIOUS BOTANICAL INVESTIGATIONS IN THE AREA 9 NOMENCLATURE 9 GENERAL PART - COMMENTS ON THE LITERATURE MAJOR PHYTDSDCIDLOGICAL APPROACHES APPLIED TD FENNDSCANDIAN FOREST AND ALPINE VEGETATION BASIC CONCEPTS AND TERMINOLOGY ........................... 10 THE UPPSALA AND BRAUN-BLANQUET SCHOOLS 10 The Uppsala school - history and concepts 10 The Braun-Blanquet approach - history and concepts 11 Merging of the Uppsala and Braun-Blanquet schools 12 The Braun-Blanquet approach to regional variation in vegetation 12 Regional approaches to forest and alpine vegetation in Scandinavia 13 THE FINNISH SCHOOL AND FOREST SITE-TYPES 14 History and concepts 14 Regional approaches by the Finnish school 14 THEORIES ON THE NATURE OF VEGETATION HITH PARTICULAR EMPHASIS ON IMPLICATIONS FOR REGIONAL APPROACHES TO VEGETATIONAL VARIATION ..................................... 16 THEORIES ON THE NATURE OF VEGETATION 16 EVIDENCE ON THEORIES, WITH SPECIAL REFERENCE TO SCANDINAVIA AND REGIONAL VARIATION 17 ASSESSMENT 19 IMPLICATIONS FOR LOCAL AND REGIONAL APPROACHES TO VARIATION IN THE COMPOSITION OF VEGETATION 19 A GRADIENT REFERENCE FRAHE FOR ANALYSIS OF REGIONAL VARIATION IN SOUTHERN NDRHEGI AN FOREST AND ALPINE VEGETATION ................................................. 21 TERMINOLOGY OF GRADIENTS 21 SOME COMMENTS ON CURRENT APPROACHES TO LOCAL AND REGIONAL VARIATION IN VEGETATION 21 A PROPOSAL FOR A REGIONAL APPROACH TO VARIATION IN SOUTHERN NORWEGIAN VEGETATION 22 REGIONAL GRADIENTS 23 The zonal gradient 23 The sectional gradient 31 LOCAL COMPLEX-GRADIENTS 31 The complex-gradient nutritient status 32 The complex-gradient topographic moisture-snow cover 33 SPECIAL PART - THE FOREST-ALPINE TRANSITION IN THE GRUNNINGSDALEN AREA THE INVESTIGATION AREA ..................................... 41 DELIMITATION 41 SOMMERFELTIA 2 (1985) GEOLOGY 4?, TOPOGRAPHY 43 CLIMATE 43 Precipitation 43 Temperature 44 Humidity 44 Oceanicity 45 Wind 45 Snow cover 46 SOIL 46 FLORISTIC PHYTOGEOGRAPHY 46 The western element 47 Warmth-demanding elements 47 The eastern element 48 The alpine element 48 The importance of the elements 50 HUMAN INFLUENCE 50 PRINCIPLES OF CLASSIFICATION IN THIS HORK .................. 52 THE DIRECT GRADIENT APPROACH 52 GRADIENTS IN FOREST AND ALPINE VEGETATION OF THE INVESTIGATED AREA 53 A DIRECT GRADIENT APPROACH TO FOREST AND ALPINE VEGETATION IN THE GRUNNINGSDALEN AREA 53 DIVISION OF THE VERTICAL GRADIENT 55 DIVISION OF THE COMPLEX-GRADIENT TOPOGRAPHIC MOISTURE-SNOW COVER 56 MATERIAL AND SAMPLING PROCEDURE ............................ 58 DESCRIPTION OF INVESTIGATED TRANSECTS ...................... 59 DESCRIPTION OF THE VEGETATION OF THE SITE-TYPES AND COMPARISON HITH CORRESPONDING VEGETATION DESCRIBED BY OTHER AUTHORS .............................................. 62 XERIC SERIES 62 Middle boreal zone 62 Upper boreal zone 67 Low alpine zone 74 Middle alpine zone 77 Survey of the xeric series 79 SUBXERIC SERIES 82 Middle boreal zone 82 Upper boreal zone 85 Low alpine zone 89 Middle alpine zone 91 Survey of the subxeric series 97 SUBMESIC SERIES 100 Middle boreal zone 100 Upper boreal zone 103 Low alpine zone 107 Middle alpine zone 109 Survey of the submesic series 113 MESIC SERIES 115 Middle boreal zone 115 Upper boreal zone 117 Low alpine zone 120 Middle alpine zone 124 Survey of the mesic series 125 SOMMERFELTIA 2 (1985) REGIONAL VARIATION IN SOUTHERN NORWEGIAN POOR FOREST AND ALPINE VEGETATION ALONG THE OCEANICITY-CONTINENTALITY GRADIENT .................................................. 132 MATERIAL AND METHODS 132 RESULTS AND DISCUSSION 132 Regional variation of the xeric series 132 Regional variation of the subxeric series 137 Regional variation of the submesic series 143 Regional variation of the mesic series 147 Concluding remarks on the vegetation of Grunningsdalen in relation to regional variation 148 MATERIAL AND METHODS FOR NUMERICAL ANALYSIS ............... 149 NUMERICAL ANALYSIS OF VEGETATION 149 Material and data preparation 149 Similarity between site-types 149 Cluster analysis of site-types 150 Ordination of site-types 150 NUMERICAL ANALYSIS OF AUTECOLOGY 151 Material and data preparation 151 Similarity between species distributions 151 Cluster analysis of species 151 Ordination of species 151 HOMOTONEITY AND DIVERSITY 152 Homotoneity 152 Within site-type diversity 153 Within gradient diversity 153 Between site-type diversity along gradient 154 RESULTS OF NUMERICAL ANALYSIS ............................. 155 VEGETATION 155 Similarity between site-types 155 Cluster analysis of site-types 155 Ordination of site-types 159 AUTECOLOGY 163 Cluster analysis of species 163 Ordination of species 164 Interpretation of GA-groups 166 HOMOTONEITY AND DIVERSITY 171 Homotoneity
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