Ecology Écologie des of the subarctic régions regions subarctiques Proceedings of the Helsinki symposium , Actes du colloque d’Helsinki % unesco Ecology and conservation / Écologie et conservation / Titles in this series Dans cette collection I. Ecology of the subarctic regions. Proceedings of the Helsinki symposium 1 &cologie des régions subarctiques. Actes du colloque d’Helsinki II. Methods of study in soil ecology. Proceedings of the Paris symposium 1 Méthodes d’étude de l’écologie des sols. Acies du colloque de Paris III. Resources of the Biosphere. Proceeding of the Paris Conference (in preparation) PubIished in 1970 by the United Nations Educational, Scientific and Cultural Organization Place de Fontenoy, 75 Pari~-7~ Printed by Imprimeries Réunies de Chambéry Publié en 1970 par l'organisation des Nations Unies pour l'éducation, la science et la culture place de Fontenoy, 75 Paris-7e Imprimeries Réunies de Chambéry 0 Unesco 1970 Printed in France SC.SS/XXIV1. /AF Preface Préface The promotion of research in plant and animal ecology La promotion de la recherche concernant l’écologie has been the objective of many symposia organized végétale et animale a fait l’objet d’un grand nombre or sponsored by Unesco and is one of the main fields de colloques organisés OU patronnés par l’Unesco, et of research towards which the Organization directs c’est un des principaux domaines de la recherche dont its attention in relation to its activities in natural s’occupe l’organisation dans le cadre de ses activités resources research. While previous work has been relatives aux ressources naturelles. Les travaux anté- concentrated on problems in the arid zone and in the rieurs ont porté essentiellement sur les problèmes de humid tropics, the present volume is devoted to a la zone aride et de la zone tropicale humide, mais le completely different environment: the subarctic présent volume est consacré à un milieu entièrement environment. différent: celui des régions subarctiques. The area of transition between the temperate zone La région intermédiaire qui est située entre la zone and the Arctic tundra, comprising tracts of forest, fen tempérée et la toundra arctique et qui comprend des and heathland in north-western Canada, Alaska, forêts, des marécages et des landes (nord-ouest du northern Scandinavia, northern Russia and Siberia, Canada, Alaska, Scandinavie septentrionale, nord de was the subject of a symposium organized by the la Russie et Sibérie) a fait l’objet d’un colloque orga- Government of Finland and Unesco in co-operation nisé par le gouvernement finlandais et l’Unesco, en with the International Geographical Union. It was coopération avec l’Union géographique internationale. held in Otaniemi (near Helsinki) from 25 July to Ce colloque s’est tenu à Otaniemi (près d’Helsinki) 3 August 1966, and was attended by 70 scientists from du 25 juillet au 3 août 1966 et a réuni soixante-dix 13 countries. The opening speeches were given by savants de treize pays. Les discours d’ouverture ont the Finnish Minister of Education,M r. R. H. Oittinen été prononcés par le ministre finlandais de l’éducation, and by Professor Paavo Kallio, of the University of M. R. H. Oittinen, et par le professeur Paavo Kallio, - Turku. de l’université de Turku. The papers presented were divided into nine sec- Les mémoires qui ont été présentés étaient répartis tions : subarctic definition; meteorology of subarctic en neuf sections: définition de la zone subarctique; regions; snow cover as an ecological factor;w eathering météorologie des régions subarctiques ; le tapis neigeux and geomorphological processes ; permafrost as an en tant que facteur écologique; phénomènes météoro- ecological factor; main features of soil-forming pro- logiques et géomorphologiques; le pergélisol en tant cesses; ecology of subarctic vegetation; ecology of que facteur écologique ; principaux aspects des phé- important species of the subarctic fauna; conservation nomènes de formation du sol; écologie de la végétation of nature and rational use of renewable natural subarctique; écologie des espèces importantes de la resources of subarctic regions. faune subarctique; conservation de la nature et uti- It is hoped that the publication of these papers will lisation rationnelle des ressources naturelles renou- lead to a better understanding of the physical condi- velables des régions subarctiques. tions of the regions under study and will open up new On espère que la publication de ces mémoires contri- perspectives for further research and practical appli- buera à faire mieux comprendre les conditions phy- cations. Many of the papers which were the result of siques des régions étudiées et ouvrira de nouvelles highly specialized research were presented in a way perspectives en ce qui concerne la recherche et les such as to arouse interest among colleagues working applications pratiques. Un grand nombre de mémoires, in entirely different fields. qui étaient l’aboutissement de recherches hautement Among the major points arising from the discussions spécialisées, ont été présentés de manière à éveiller were the need for the establishment of protected l’intérêt de collègues travaillant dans des domaines control areas ; for international treaties protecting entièrement différents. migratory birds, whales, seals, etc., and for a better Les discussions ont fait ressortir notamment la understanding of cyclic fluctuations of subarctic nécessité de créer des zones de contrôle protégées, de mammals and birds, the prime problem of boreal conclure des traités internationaux visant à protéger terrestrial ecology. The meeting urged that research les oiseaux migrateurs, les baleines, les phoques, etc., should be carried out on productivity and sustained et de mieux comprendre les fluctuations cycliques yield of subarctic fauna and flora and that meteoro- des mammifères et des oiseaux subarctiques, problème logical observations be extended and improved for capital de l’écologie terrestre boréale. Les participants microclimatological studies in the subarctic zones. ont estimé qu’il faudrait, d’une part, effectuer des Following the presentation of papers and the dis- recherches sur la productivité et le rendement per- cussions a visit was arranged to the University of manent de la faune et de lafloresubarctiques et’d’autre Turku, and a four-day field excursion to Lapland to part, multiplier et améliorer les observations météo- visit the Kevo Research Station. An alternative field rologiques en vue de l’étude microclimatologique excursion was arranged for zoologists to the Oulanka des zones subarctiques. Biological Station of the University of Oulu. Après la présentation des mémoires et la conclusion In presenting this volume, Unesco wishes to offer des débats, ont eu lieu une visite à l’université de its thanks to the participants in the symposium whose Turku et une excursion de quatre jours en Laponie papers have made it possible, and to Professor Kallio à destination de la station de recherche de Kevo. Les and the members of the Finnish organizing committee zoologistes, pour leur part, ont eu la possibilité de for their generous and efficient organization of the participer à une autre excursion jusqu’à la station symposium. biologique d’oulanka, qui dépend de l’université The responsibility for the selection and presentation d’Oulu. of facts and for opinions expressed rests with the L’Unesco tient à exprimer sa reconnaissance aux authors. membres du colloque qui, grâce à leurs mémoires, lui ont permis de publier le présent volume, et à remercier le professeur Kallio et le comité d’organisation fin- landais pour la générosité et l’efficacité avec lesquelles ils ont organisé le colloque. Les auteurs sont seuls responsahles du choix et de la présentation des faits, ainsi que des opinions exprimées. Contents Table des matières J. Bliithgen Problems of definition and geographical differentiation of the Subarctic with special regard to northern Europe . 11 Problèmes de délfinition et de différenciation géographique du Su barctique, spécialement en Europe septentrionale [Résumé] . 30 B. A. Tikhomirov Forest limits as the most important biogeographical boundary in the North . 35 Importance des limites forestières en tant que frontière biogéographique dans le Nord [Résumé] . 38 I. M. Dolgin Subarctic meteorology . 41 Météorologie subarctique [Résumé] . .4-5 9 S. Huovila Some features of the microclimate within hilly regions in Finland . 63 Quelques caractéristiques du microclimat dans les régions accidentées de Finlande [Résumé] . 65 H. Odin and K. Perttu Radiation measurements near the forest limit in northern Sweden. . 67 Mesure du rayonnement aux limites de la forêt dans la Suède septentrionale [Résumé] . 77 R. Sarvas Temperature sum as a restricting factor in the development of forest in the Subarctic . 79 La température globale en tant que facteur restrictif dans le développement des forêts subarctiques [Résumé] . 81 W. O. Pruitt Jr. Some ecological aspects of snow . 83 Quelques aspects écologiques de la neige [Résumé] 't- 97 A. Corte Bioecological aspects of the snow plant communities of Cape Spring, Argentine Antarctica . . 101 Aspects bio-écologiques des communautés de plantes des neiges au cap Spring, Argentine antarctique [Résumé] . . 104 A. Rapp Some geomorphological processes in cold climates . . 105 Etude de certains processus géomorphologiques dans les régions à climat froid [Résumé] . . 113 J. Biidel Denudation and river erosion in the “zone of pronounced valley forma- tion” on South-east Spitsbergen . . 115 Dénudation et érosion fluviale dans la zone de formation accusée de vallées au Spitzberg du Sud-Est [Résumé] . . 117 A. Jahn Soil movements under the influence of freezing . . 119 Les mouvements du sol sous l’inyuence du gel [Résumé] . . 122 J. Malaurie Complexité des notions de faciès morphologique arctique et subarctique (nord-ouest et centre ouest du Groenland). Géographie boréale et anthro- pologie: fondements physiques des notions de lieu de territoire . . 125 Complexity of the terms LLArctic” and “Subarctic” as notions of morpho- logical aspect (North-westa nd Middle-west Greenland) [Summary] . 128 R. J. E. Brown Permafrost as an ecological factor in the Subarctic . . 129 Le pergélisol, facteur écologique de la région subarctique [Résumé] . . 138 T. L. Péwé Permafrost and vegetation on flood-plainso f subarctic rivers (Alaska) : [a summary]. . 141 Le pergélisol et la végétation dans les plaines inondables des cours d’eau subarctiques (Alaska) [Résumé] . . 141 M. Salmi Investigations on palsas in Finnish Lapland . . 143 Recherches sur les hydrolaccolithes (palses) de la Laponie Jinlandaise [Résumé] . . 151 E. Schenk Permafrost and frost structures in the subarctic area . 155 Formation et structures du pergélisol [Résumé] . . 158 H. Svensson Frozen-ground morphology of northeasternmost Norway . . 161 Morphologie des sols gelés dans l’extrême nord-est de la Norvège [RBsumé]. P7 P. L. Johnson Remote sensing as an ecological tool . . 169 La détection à distance en écologie [Résumé] . . 185 J. C. F. Tedrow Soils of the subarctic regions . 189 Les sols des régions subarctiques [Résumé] . . 199 F. di Castri Soil ecosystems in subantarctic regions . . 207 R. Covarrubias Écosystèmes du sol dans les régions subantarctiques [Résumé] . . 221 E. Hajek L. A. Viereck Soil temperatures in river bottom stands in interior Alaska . . 223 La température du sol dans les peuplements forestiers du fond des vallées à l’intérieur de l’Alaska [Résumé] . . 232 I. Hustich On the study of the ecology of subarctic vegetation . 235 Sur l’étude de l’écologie de la végétation subarctique [Résumé] . . 238 D. J. Bellamy A critical limit of primary production for the survival of arctic alpine W.M . Tickle plants in the northern Pennines of England . . 241 Un seuil critique de la production primaire pour la survie de plantes alpines arctiques dans les Pennines septentrionales (Angleterre) [Résumé]. 246 E. Einarsson Plant ecology and succession in some nunataks in the Vatnajökull glacier in South-east Iceland . . 247 Phyto-écologie et évolution du tapis végétal dans certains nunataks du gla- cier de Vatnajökull, dans le sud-est de l’Islande [Résumé] . . 254 A. N. Formozov Ecologie des plus importantes espèces de la faune subarctique . 257 Ecology of the major species of subarctic fauna [Summary] . 273 A. G. Loughrey The ecology and population dynamics of the barren-ground caribou in J. P. Kelsall Canada 275 Le caribou des toundras du Canada [Résumé] . 279 V. A. Peiponen Animal activity patterns under subarctic summer conditions . 281 Organisation de l’activité des animaux en fonction des caractéristiques de l’été subarctique [Résumé] . 286 E. Aubert de la Riie Les îles Saint-Pierre et Miquelon, une enclave subarctique méridionale 289 The St. Pierre and Miquelon Islands, an enclave of the southern Subarctic [Summary] . 292 P. Mikola Forests and forestry in subarctic regions . 295 Les forets et la sylviculture dans les régions subarctiques [Résumé] . 300 R. Kalliola National parks and nature reserves in subarctic regions . 303 Parcs nationaux et réserves naturelles dans les régions subarctiques [Résumé] 307 I. G. Simmons Problems of the conservation of relict arctic and subarctic species in Britain 309 Conservation des survivances botaniques arctiques et subarctiques en Grande-Bretagne [Résumé] . 317 R. Ruuhijärvi I Subarctic peatlands and their utilization 319 Les tourbières subarctiques et leur utilisation [Résumé] . -. .* -. 325 E. Ehlers The rational use of forests and bogs, in view of comparative observations in north-western Canada and northern Finland . 327 Utilisation rationnelle des forets et des marécages d’après des observations comparées effectuéesd ans le nord-ouest du Canada et le nord de la Finlande [Résumé] . 333 E. Schenk On the string formation in the aapa moors and raised bogs of Finland . 335 L’origine des tourbières réticulées (aapamoors) [Résumé] . . 340 G. Sirén Remarks on current silvicultural research in the Subarctic of Finland . 343 Remarques sur les recherches effectuéese n sylviculture dans la région sub- arctique de Finlande [Résumé] . 348 T. Pritchard International scientific co-operation in conservation with special refer- ence to peatlands . 349 General discussion and conclusions / Discussion générale et conclusions 351 List of participants / Liste des participants . 363 Problems of definition and geographical differentiation of the Subarctic with special regard to northern Europe Joachim Blüthgen I should like to express my appreciation for being the terminological structure for “the Subarctic”, to invited to read the introductory paper which will set combine them and to integrate them into this struc- the scene and provide the frame for the work of this ture according to their due weight. symposium on the Subarctic. I a m aware that I can The geographical term “subarctic” is one of those deal with only some of the problems attached to this geographical designations of areas which are often very complex and variably used term. The more I used for the characterization of transitional zones worked on the subject, the more I doubted whether between core areas. Delimiting them, however, is it is possible at present to give a satisfactory definition only possible in a complex fashion, i.e., by a combina- of this primarily geographical term which is also used tion of different distinctive features. They are defined in numerous neighbouring sciences though each gives in different ways, if ever. With subarctic we can it a different meaning. Although my task is to try group subpolar, subtropical, suboceanic, subconti- to define and subdivide the Subarctic I must confess nental, subalpine. These “sub” areas form every one that this-due to contributions which are still contra- of them a transitional zone at the periphery of those dictory-is not quite objectively possible as yet. It core areas which are formed by differentiation of the reflects inevitably a great deal of personal opinion. I ask Earth according to latitude, surface, and height, and therefore that my comments be regarded as a still of whose names they are derived (arctic, tropics, etc.). incomplete contribution. It is bound to be of a prelim- But it is not possible to deduce clearly from their inary character since only much more painstaking names whether they form parts of these areas or investigations into the countless contributions to whether they are areas of their own outside them. this subject could lead, in the near future, to a solu- These terms, which derive their names from geogra- tion of this terminological problem. Besides, more phical situations, are now liable to be connected with extensive contacts with competent specialists over the contents of various systems : plant-geographical, the world are a primary necessity. climatogeographical, pedological, ecological, etc., or The entanglement of terms is far more complex they may be used as terms in landscape or in the than I originally assumed. Therefore I should espe- geography of population, human settlement, or econ- cially like to acknowledge gratefully the proposal omy. Their delimitation is not only different in the made by Unesco to open the symposium with this various branches of learning, but even within one and topic. It really corresponds to an urgently felt need the same field there exist serious discrepancies between to avoid as much as possible all misunderstandings of different authors, according to which characteristic what is meant by the term “Subarctic”. Thereby is stressed. When we look into geographical litera- help is offered to all those specialists doing research ture we see at once the numerous different uses of on special problems which are in any way connected these terms. The same holds true in those many with the Subarctic, so that they might eventually be neighbouring sciences on which geography is bound able to refer the results they have obtained to the to rely. right or at least a conventionally agreed upon frame. In the following pages I shall try to throw some It will be the specific task of the geographer to form light on the term subarctic and its use until now in building-blocks from all the numerous individual various fields of learning and according to the geo- results and interpretations, building-blocks to erect graphical differentiation of its area, especially in 11