ACTA PHYTOGEOGRAPHICA SUECICA 87 EDIDIT SVENSKA VAXTGEOGRAFISKA SALLSKAPET

Use and abuse of reindeer range

Olof Eriksson, Mikael Niva & Alexandro Caruso

lJPPSALA 2007

ACTA PHYTOGEOGRAPHICA SUECICA 87 EDIDIT SVENSKA V AXTGEOGRAFISKA SALLSKAPET

Use and abuse of reindeer range

Olof Eriksson, Mikael Niva and Alexandro Caruso

UPPSALA 2007 2

ISBN 978-91-7210-087-9 (paperback) ISBN 978-91-7210-487-7 (cloth) ISSN 0084-5914

Erik Sjogren Editor:

Cover illustration: Gullris - Solidago. Tuschteckning av Gunnar Ostman Inga, 2004.

© Eriksson, M. Niva & A. Caruso 2007 0.

Edidit: Svenska Vaxtgeografiska Sallskapet Villavagen 14 SE-752 36 Uppsala

Lay-out: Opulus Press AB Printed in Sweden by Fingraf, Sodertalje.

Acta Phytogeogr. Suec. 87 Abstract

Abstract. Eriksson, Niva, M. and Caruso, A. Use and abuse At all study sites, six adjacent plots were selected, half of which 0., of reindeer range. -Acta Phytogeogr. Suec. 87. Uppsala. were fenced to deny access to larger herbivores, and half were left open for grazing by herbivores. The composition of plant com­ all munities in the field, bottom and tree layer in plots was estimated In consequence of variations in geology and soils, in climate, in 1995-96, and re-estimated to four years later. three and in its wide extent in longitude, latitude and altitude, the Generally, marginal or no effects of enclosure were seen Scandinavian mountain chain exhibits major variations in natural on the vegetation communities, and there were no differences conditions. Nature is constantly influenced by processes that between vegetation types. include both natural forces and human activity. Up to the end of the 19th century, travellers in the montane In the early 1990s, there was an intense media debate about region, both Saami and outsiders, ocularly assessed the plant current damage to the montane vegetation, which many believed cover. As a rule, they reported a good supply of reindeer fodder they could observe. plants, especially lichen species. In 1992, the World Wide Foundation for Nature, WWF, invit­ From the end of the 19th century, there began to be obser­ ed representatives of responsible authorities, reindeer-husbandry vations of severely denuded lichen cover, especially in areas interests, voluntaryconservation bodies and interested researchers exposed to a veritable invasion of Saami and reindeer from the to a conference, which, somewhat erroneously, came to be called north-Norwegian and north-Finnish reindeer grazing areas. In­ the 'Reindeer grazing conference', but which included a spectrum cornersfrom those areas introduced an extensive form of reindeer of factors that can affectthe montane vegetation. husbandry, developed to suit conditions on the Finnmarksvidda One result of this conference was that, in 1993, WWF initi­ and in northernmost Finland, where large reindeer herds could ated a research project, extending over several years, intended readily find grazing on well-demarcated headlands and islands to provide information about temporal changes in montane during the snowfree season, without much supervision. vegetation. The conflictsof interest between the incomers, and indigenous Experimental areas distributed along the Swedish mountain Saami who wished to carry on an intensive fo rm of reindeer chain were selected: the southemmost are on Fulufjiillet in husbandry, with closely supervised herds, were great. From the Dalama,. and the nothemmost are ea. 15 S of Tavvavuoma beginning of the 20th century, state interventions, in the form of km in Swedish Lapland. (Some placenames are given in modem commissions of enquiry and fieldsurveys, were instituted. Their North-Saamish spelling in Appendix 2) The vegetation types aim was to resolve existing conflicts,to ensure a sustainable ac­ studied were Grass heath, Meadow with low herbs, Dry heath, cess to grazing, and satisfactory profitability. The results cannot Birch forest-heath type with lichens and Birch forest-heath with be said to have been satisfactory. mosses. These cover all major montane areas and are important grazing areas for reindeer.

Acta Phytogeogr. Suec. 87 4

Contents

Abstract 3

Preface 7

Acknowledgements 9

Introduction 10 Botanical research 13 Large-scale exploitation of mountain regions 18 Data for decision-making in Swedish-Norwegian reindeer grazing conventions - new knowledge of nature in mountain regions 19 National Parks 21 Present -day maps of the mountains 21

1. The reindeer 22

1.1 Origin 22

1.2 Reindeer feeding 25

Pastures in the mountain birch woodlands 26

1.3 Fertility 28

1.4 Diseases 28

1.5 Mortality 28

1.6 Slaughter 29

1. 7 Herd size over time 30

1.8 Reindeer domestication and handling 33

Saami and reindeer husbandry- some definitions 34 Husbandry of forest reindeer 34 Mountain reindeer husbandry 35 Good grazing - poor grazing 35 The Saamis and the authorities 35 The silver-mining era and the Saamis 37 The colonisation of the Saami territories 37 Regulations concerning the Saami territories 38 The boundary of the Saami territories 38

1. 9 Reindeer grazing and availability of grazing, fromthe time of Olaus Magnus 39

Reports from pioneers in Lapland 39 Linnaeus' travels in Dalama and Lappland 40 Zetterstedt's journey in Tome Lappmark 41 Zetterstedt's journey in Ume Lappmark 41 The Lapp bailiffs' yearbooks 42 The Lapp bailiffs in Norrbotten 42 Lapp bailiffsin Jamtland 42

Acta Phytogeogr. Suec. 87 5

The Saami view of the grazing situation 43 R. Hult's attempt to deduce the laws governing the composition of plant formations 45 Observations on reindeer grazing during the 20th century, by botanists working in the mountains 48 Tengvall's measurements of reindeer lichen growth 49

2. The WWF-project� Background 51 2.1 Project objectives and study sites 51

2.2 Geology 51

2.3 Physical geography 52

2.4 Climate and weather 52 Growing season 53 Humidity 54

2.5 Vegetation types 54 Grass heath 55 Meadow with low herbs 55 Dry heath 55 Mountain birch forests 56 Extent of vegetation types studied 57 Establishment of sample plots 59

3. The WWF-project:How it was carried out 59 Shrub and treelayer 60 Presence of herbivores 60 Data processing 60 Ordination 61 Cover 61 Frequency 62 Shrub and tree layer 63

4. The WWF-project: Results 64 4.1 Birch forest heath type with mosses and dry heath 64 Species present 64 Shannon's diversity index 65 Ordination 66 �� � 4.2 Birch forest-heath type with lichens 72 Species present 72 Shannon's diversity index 73 Cover 73 Frequency 74 Shrub- and tree-layer 74 Droppings 75

Acta Phytogeogr. Suec. 87 6

4.3 Ritsem meadow with low herbs and dry heath 75

Species present 75 Shannon's diversity index 76 Ordination 76 Frequency 78 Droppings 78

4.4 Tjuolmajaure and Puollanvare 79

Species present 79 Shannon's diversity index 79 Ordination 79 Cover 80

5. The WWF-project: Discussion 82 Species diversity 83 Ordinations 83 Cover of the species 84 Shrub and tree canopy cover 86

5.1. Concluding thoughts 87

References 89

AppendixI. Recorded species 95

Appendix11. Saami names for utility goods_, botanical names_, villages and study sites_, and list of persons who provided information 101

Acta Phytogeogr. Suec. 87 Preface

The Scandinavian mountain chain (the Scan des) or the Emergence of the National Park concept in the late Kjol ('Keel') (Ljungner 1948), part of which is situated 19th century led to some change of opinion, and a debate in Sweden, is a part of the Caledonian mountain chain, about the conservation of wilderness areas was started. which intermittently stretches from Ireland to northern In 1880, the discoverer of the Northwest Passage, the Greenland. It exhibits a wide range of variation with geologist and polar explorer A.E. NordenskiOld, brought regard to natural characteristics. This is partly because of up the question of Swedish National Parks. At that time differences in bedrock and soil type, climatic gradients in hydroelectric power and natural resources had begun to both north-south and east-west directions, as well as in be exploited by society, which had led to a more exten­ altitude. These differencesin abiotic conditions contribute sive communication system even in the mountain areas. to the occurrence of a large number of vegetation types. Improved communications then led to mountain tourism, Factors that most clearly form the vegetation are snow initially on a modest scale, but which, despite its taking protection and water. Over a short distance it is possible place only on foot, by ski or by rowing boat, was considered to note a shift from surfaces exposed to the wind, with by some as a potential environmental threat. sparsely occurringlichens, mosses and low-creeping dwarf shrubs, to slightly taller shrub-dwarf shrub vegetation or In 1909, the Swedish Parliament decreed that national even luxuriant vegetation. In the mountain birch forest parks similar to those in the United States (e.g. Yellowstone and on the open mountain, there are almost 600 vascular 1872) should be established in differentparts of the country, plant species, about 300 moss species and more than 600 to protect valuable environments for all time. Examples lichen species. in the mountain regions are Peljekaise, Sarek, Sanfjallet and Stora Sjofallet, all of which were established between As in other parts of Sweden, nature in the mountain 1909 and 1913. The last of these was severely reduced in chain has been influenced, and still is influenced, by both size when hydroelectric interests succeeded in persuading Parliament that its waterfalls could be exploited to produce natural forces and anthropogenic activities. Itis simultane­ ously exposed to exploitation by land-based industries, of electricity. Abisko National Park was established in 1919 which energy generation, mining, reindeer husbandry and and Vadvetjillo in 1920. mass tourism are the largest, the most space-requiring and those that possibly have the greatest effect on vegetation. Towards the end of the 20th century, a new debate on the Nature in these areas is also utilised by the local popula­ mountain regions emerged, this time based on observations tion and visitors from elsewhere, for recreation in different of accelerating wear-and-tear on mountain nature, reported forms on a smaller scale, e.g. fishingand hunting. Despite by devoted nature enthusiasts and by various research the exploitation pressure, the mountains are nonetheless projects. The root of this debate may possibly be found one of Sweden's most undisturbed environments and, in in the Lovhogen area, an isolated area of low mountains an international perspective, possess extremely important close to the border between the provinces of Dalama and natural values. Aronsson (1997), among others, even Harjedalento the west ofLillhardal, one of the home ranges speaks of 'Europe's last wilderness', but then probably in Sweden of the wild reindeer (Rangifer tarandus tarandus disregards northern Russia west of the Ural Mountains. L.). The last of these animals was reported to have been exterminated during the 1860s (Lonnberg 1909). Since that Anthropogenic influence on more or less extensive time, there have been no reindeer in the area, until the end areas in the mountains today can be traced far back into of the 1950s, when domestic or semi-wild reindeer (these history. Before the beginning of the 20th century,there was too were Rangifer tarandus tarandus L.), began to enter little discussion on whether anthropogenic activities could the area in increasingly large numbers. Their effect on the imply a risk of wear-and-tearon what was then considered part of the plant cover that was dominated by lichens was an almost infinite wilderness.At that time it was not even considerable (Hoglund 1970), and was later studied in valued to any extent, to judge from the name given to it greater detail by Hoglund & Eriksson (1973). by the authorities- state owned waste lands, 'kronoover­ loppsmark'- referring to areas retained by the Swedish Already in the 1980s, Nils G. Lundh, Funasdalen, was State when partitioning land in mountain regions. one of the foremost figures in the debate among nature­ conservation enthusiasts. With the help of a camera, he tried to record temporal changes in the mountain vegeta-

Acta Phytogeogr. Suec. 87 8 0. Erikssson et al.

tion cover, on both the Swedish and on the Norwegian on snow-free terrain in the open mountains, to minimize (reindeer-free) side of the international border. damage to the soil and vegetation. The right to travel by snow-scooter should be Lundh ( 1998) used older publications (Schmidt 1801, restricted to a considerably greater extent to established Backlin 1982) in an attempt to demonstrate that the vegeta­ tracks, in both the mountains and forest areas. tion changes were relatively recent and were related to the excessive grazing pressure and trampling caused by rein­ In 1992, the World-Wide Fund for Nature (WWF) ar­ deer. His views were supported by Kullman (1989), Boberg ranged a conference on 'The Reindeer Grazing Problem' (1992), Oksanen (1992) and Ihse & Allard (1995). at which representatives of the reindeer industry and the relevant authorities conducted a broad-ranging discussion Although the debate on wear-and-tear on mountain with nature conservationists and scientists. It became clear vegetation initially focussed on the excessive utilization that reindeer husbandry alone could not be accused of be­ of the soil cover by reindeer and reindeer management, ing responsible for the ongoing processes. A visible result other influential factors were also considered. Among of the conference was that the WWF's project committee other factors, Kullman (1979, 1989, 1998, etc.), who for decided to initiate a comprehensive research project, with many years had studied the relationship between climate Lennart Nyman as leader, which from the start became and vegetation at the tree-limit in the mountains, asserted known, rather unfortunately, as the 'reindeer grazing that an ongoing climatic deterioration was of great im­ project'. Certain influential factors other than reindeer portance in impoverishing the vegetation. Mechanical grazing land were also to be studied. wear caused by visitors and cross-country vehicles also came into focus, as revealed in studies by e.g. Borgegard The project was intended to demonstrate any changes et al. (1975), Emanuelsson (1984), Wallsten (1988) and in the vegetation within the study area along the moun­ Renman (1989). tain chain - both retrospectively and in the future. It was hoped that the facts obtained could be used, together The first study that reported acid precipitation in the with others, in planning land utilization by land-based mountains was published as early as 1971: Andersson et industries. The methods employed should also be useful al. ( 1971) reported that sensitive fish species had disap­ for the purposes of environmental monitoring. Thus, the peared from waters on Mt. Fulufjall by the mid-1960s. work should contribute to the long-term conservation of Similar reports later came from other southerly mountain today's mountain environment, and should also serve as areas. Degerman et al. (1992) presented a compilation of a platform for measures designed to return the mountain the knowledge available at that time on acidification of vegetation to its former richer state, principally with regard mountain environments. As regards terrestrial vegetation, to biomass. it was found that only a few reports had been published. However, various observations of presumed negative ef­ The project dealing with vegetation consists of two fects on the lichen cover were reported. sub-projects. A study of changes in plant communities, vegetation cover and erosion using remote analysis is be­ In 1995, at the request of the government, the Envi­ ing made through comparative interpretation of existing ronmental Advisory Committee (Miljovardsberedningen) infrared (IRF) aerial photographs from the early 1980s, submitted an analysis of the environmental situation, and and those more recently taken. The potential of satellite proposals for measures to be introduced for ensuring technology is also utilised, whereby mainly Landsat TM sustainable development in Sweden's mountain regions. pictures are comparedwith aerial photographs as part of an This work was based on existing enquiries and propos­ effortto develop the method (cf. Ihse&A llard 1995, Allard, als, various published scientific reports, reports from et al. 1998, Allard 2001). This sub-project has been carried national authorities and organizations, hearings with the on at the Department of Physical Geography, Stockholm local population and personal observations, etc. Particular University, and is reported separately (Allard 2003). attention was given to conditions in the southern part of the mountain chain. The ground-based sub-project that will be presented in what follows is based on repeated plant counts on From a botanical point-of-view, the following propos­ fenced and open sample areas situated in vegetation als, among others, are of interest: types commonly found in areas ranging from Fulufjallet, Dalama, in the south, to the region ofTavvavuoma (Dav­ Because it was a land-based industry, the reindeer vavuopmi), Lapland, in the north. This report covers the industry was required to shoulder especial responsibility initial phase and the first revision after 3-5 years. The for environmental considerations. cover and frequency of the plant species/groups present It was proposed to regulate the right to drive have been recorded. Among factors that may influence

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 9

species diversity and plant biomass, it has been possible Norrbotten, has been of great help, e.g. as coordinator and to record, to some extent, the effectof weather conditions coach when teams were to startfieldwork in differentplaces during the growing season and traces of the presence of at the same time. Dr. Ingvar Backeus and Willy Jungskar, large herbivores. A similar project in the Tavvavuoma Dept. of Plant Ecology, Uppsala University, helped us to area, started in 1967-1968, has been incorporated in this organise the initial phases of the computer processing. sub-project. The Department of Plant Ecology, Uppsala Professor Kurth Perttu, Dept. of Short Rotation Forestry, University, has been the base for this work. Swedish University of Agricultural Sciences (SLU), gave generous help in the handling of meteorological data. Professor Bernt Jones, SLU, helped us describe reindeer Acknowledgements deseases. Great help was given by Susanne Idivuoma and fila Svedell in checking the Saamish terminology. The main provider of funding is the World Wide Founda­ The translation from Swedish to English and correc­ tion for Nature -WWF.As a coordinator, Ola Jennersten at tion of our English was done by the late Nigel Rollison, WWF in Stockholm has had great patience with us. Con­ subsequently continued by Dr. Jeremy Flower-Ellis. The tributions, both economic and logistic, have been received Police Authorities in Gallivare and Kiruna generously from the County Administrations in Dalarna, Jamtland allowed us to use their excellent accommodations during and Norrbotten, and from the Swedish Environmental fieldwork at Ritsem and Jarama (Jarin).The editor of the Protection Agency. The Phytogeographical Society made Phytogeographic Society, docent Erik Sjogren, showed it possible for us to publish in its Acta Phytogeographica angelic patience and also offeredcontinual encouragement Suecica series. Professor emeritus Eddy van der Maarel during the writing process. and Professor Jon Agren at the Dept. of Plant Ecology, And last but not least, especial thanks go to our families, Uppsala University placed rooms at our disposal within for their love, understanding and indispensable practical the department;Jon assisted with statistical analysis during and emotional support during the all-too-lengthy writing the project. Senior-officerTuomoRaunistola (late), Section period. for Reindeer husbandry at the County Administration in We are most gratefulto all of you.

Uppsala 2007, Olof Eriksson, Mikael Niva and Alexandro Caruso.

Acta Phytogeogr. Suec. 87 Introduction

From runic engravings to a vegetation map based on Agrarian colonisation, principally in the formof animal remote analysis - the emergence of the Swedish moun­ husbandry combined with the hunting of wild reindeer, tain region from the shadows of history occurred in some Swedish mountain valleys during the Late Iron Age (Hansson 1997). Konigsson (1986) reports The earliest evidence we have of long-term human pollen evidence that grazing land was established during utilization of biological production in mountain regions is the Middle Ages near Fjallnas in Swedish Lapland. The found by archaeologists in remnants of habitations and on MittakUippen area in Harjedalenhas become an alarming stones, oftenwith game motifs. The artists were the hunters example of the currentover-utilization of reindeer grazing who followed the herds of wild beasts and harvested them land that already was in use for grazing during the Viking when the opportunity occurred. After southernScandinavia era (Wallin & Aronsson 1998). However, we must bear had become ice-free (ea. 13,500 B.C.), hunter-gatherer in mind that the elk (Alces alces) by far was the most im­ groups migrated into the area as early as 12,000 years portant large quarry during antiquity. This is testified by B.P., following the herds of wild reindeer. rich osteological material, e.g. at Vuollerim, Norrbotten More than 8,000 years ago, the oldest surviving work county, and numerous rock carvings, e.g. at Namforsen, of art in Scania, a reindeer hunter's tent and two reindeer, Angermanland county (Zackrisson 1992). was incised on a deer antler (Liljenberg 1991). Before Olof Rudbeck the Younger and Linnaeus, writ­ The oldest site excavated to date in northern Sweden ten descriptions dealing with the northern wildemesses, is situated north of lake Dumpokjauratj, east of Arjeplog, including the mountains, their inhabitants as well as their Lapland. Samples of charcoal were dated by Accelator animals and plants, were acquired through more or less Mass Spectrometry (AMS) radiocarbon dating, and found imaginative descriptions, frequently based on unreliable to be ea. 8,500 years old. A large number of Lithic Arti­ sources. The descriptions by Vergil and Tacitus, who wrote facts were found. Reindeer, elk and beaver were important during the early stages of our chronology about Ultima sources of food during different periods. Colonisation Thule, the unknown land situated in the farthest north, and is likely to have come from the north, the settlers being inhabited by remarkable creatures with peculiar ways of late descendants of the Komsa complex, who had a long living, are examples. Tacitus introduced the name 'fenni', northern subarctic history (Bergman, Olofsson, Hornberg, still used in North Norway, whereas the Finns there have Zachrisson & Hellberg 2004). retained their former name, 'kvaner'. The Fenni became The earliest finds in the mountains of south Norway renowned for their remarkable ability to use a pair of are about 9,000 years old (Skogland 1994 ). In the Swedish 'troll-wood' skis to pursue wild animals running over provinces of Harjedalen and Jfuntland, similar finds have snowfields. Thus, in the 6th century A.D., the Gothic been made, but are younger. author Procopius added the prefix 'skriti' to 'fenni'. The

The oldest known rock-carving representing a reindeer, from the Neolithic era, was found in 1658 at Glose in Alsen, Jamtland Fig. I. county (Hallstrom 1960).

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 11

. -

.,. - .... � � - - - -

Fig. The Kalvtrask skis were found in a peatland at Kalvtrask, Vasterbotten county. Their 14C-dated age is 5200 years. This find 11. ea. is the oldest known, and may be ea. 500 years older than the oldest Egyptian pyramid. The bindings consisted of straps bound around the foot and threaded through four vertical holes in the footrest. This type of ski is not otherwise known from Scandinavia, but was common in pmts of Siberia up to the present time, and may perhaps be regarded as one of many indications of early cultural contact with Eurasia (Astrom 1993). The 'shovel staff' (Saamish gaivo-soabbi)-an aid to progress as well as a means of testing the suitability of the snow-cover for grazing- is still occasionally used.

Saami were known as ski-finns - or skiers - far back in (Alfred's Orosius, ea. 890). The first part contained a history. Skis were, however, used long before the days geographical survey of Europe south of the Alps, a region of Procopius. As a proof of this, the 'Kalvtdiskskis' (fig. known to Orosius. Alfred supplemented the English version Astrom, 1993) can be seen, which together with a II, K. with a description of Europe north of the Alps. Here it is 'skovelstav' -an early form of ski-stick - were found interesting to find the description of North Scandinavian in 1924 during drainage work ea. 80 km east of Lycksele, conditions given by the Viking Chieftain Ottar, who lived Vasterbotten county. 'northernmost of all' in Halogaland, the northern part of In the early 5th century A.D., the Roman priest Oro­ P. Norway. Ottar visited Alfred's court around 890 A.D. He sius wrote a history book ('Historiae adversum Paganos') related that he had domestic animals, decoy reindeer, to which over the years became a classic in Europe. In the assist in the capture of wild reindeer, and 600 domesticated late 9th ce-ntury, Alfred the Great, King of Wessex (871- reindeer. He had travelled extensively and had a good 899), ordered the translation of Orosius into Old English know ledge of how the mountain S aamis lived, of taxes that had to be paid, and ofthose with whom barter was carried on. He had reached as far as the White Sea by ship, where the Bjarmar people had created a hunting-farming culture in the estuary of the River Dvina. Thus it is clear that the Vikings were acquainted with the northernmost regions of the Scandes and neighbouring areas of land. the early 13th century, the Danish historian Saxo In Grammaticus wrote on 'Lappia' and its inhabitants, the 'Lapps' in 'Gesta danorum'- The History ofthe Danes. Until the mid-16th century very little was published that extended our knowledge of nature within the region we now know as Kolen or the Scandes- the Scandinavian moun­ tain chain (cf. Lj ungner 1948). In 1539, an excellent map of the North - Carta Marina - was published in Ve nice. In 1555, this was followed by a large ethnographical work on the North- 'Historia de gentibus septentrionalibus'­ (The History of the Northern Peoples). The publications became of great importance as sources of knowledge for Ill. The 'Skridfinns'use curved, broad and smooth wooden Fig. many years. The author of both was Olaus Magnus, a planks bound to their feet when hunting the wild animals of the snowy mountains. The assistance of armed women is necessary Swedish Catholic priest who lived in exile in Rome after for success. - However, the men always decide how the catch the Reformation. His brother Johannes, who assisted with is allotted, and how it shall be used (0. Magnus, 1555). the work of publication, shared his fate.

Acta Phytogeogr. Suec. 87 12 0. Erikssson et al.

The description covers a wide spectrum, e.g. ethnog­ cultivated close relationships with national leaders, and raphy, economics, geography and natural history. The in 1671 was given the task of describing Lapland and author was clearly very well read and even had personal the Laps by Magnus Gabriel de la Gardie, at that time knowledge of parts of the country, such as that acquired Chancellor of the Realm. Priority was to be given to eth­ during a period as a bogus itinerant pedlar who travelled nological aspects, such as the value of the region to the as far north as Norrbotten. In 1519 he reached Pello in national economy. The result, 'Lapponia', was written in Tomedalen. He had a considerable amount of information Latin and first published in Frankfurt am Main in 1673. 1t about the Saamis, about their neighbours at that time, about was then rapidly published in several modem languages. ecology of reindeer and about their value to their owners However, itwould take almost three hundred years before and hunters. He knew that lichens were an important com­ it was published in Swedish, in the Acta Lapponica series ponent of the reindeer's winter diet, as well as that their issued by the Nordic Museum. hooves were particularly suitable for a life under snow­ Schefferus'knowledge ofLapland is not based on what covered conditions. The mountain chain that separated he himself saw. Apart from older sources such as Pliny, Sweden and Norway was known to him. Its name should Solinus Tacitus,Procopius, Saxo and the brothersJohannes be Mt. Doffra fjall - a name we today associate with a and Olaus Magnus, he used detailed reports on Lapland region in the centre of the Norwegian part of the Scandes and its living conditions prepared for him on the order of (Dovre). The 'History' consists of a remarkable mixture the Chancellor of the Realm, by clergymen working in of exact observation and loansfrom contemporary learned the northern parts of the country. Samuel Rheen, rector publications. Pliny the Elder (24-79 A.D.) was probably of Ranea, Johan Torneaus, a schoolmaster in Tornea, and the person from whom most loans were taken. Olaus Petri Niurenius, rector of Umea, submitted reports Up to the publication of Schefferus''La pponia' in 1673, that were most utilised. Their reports arecharacterised by knowledge of the mountain region was not increased to distinct objectivity.A detailed field notebook belonging to any particular extent. The botanical research and teaching the widely travelled Johannes Bureus (1568-1652), who that emerged were connected with the seats of learning. studied antiquities and languages (Bureus 1886), together Here, mention should be made of SigfridusAronus Forsius with oral records given by Saami youngsters studying in (ea. 1550-1624) who, at the turnof the 16th-17th century, Uppsala, supplemented the field material (G. Eriksson was active as a professor at Uppsala. His main interest 1969). was astronomy, but also botany (Forsius 1611). In fact, The term 'fjall' (mountains) was examined, together he was the first to publish botanical works in Swedish. with their extent and function as a protective border to In 1601-1602, he journeyed to Lapland, mainly with the Norway, along a line fromJamtland to the 'Ice Sea' (Arctic intention of studying border conditions between Sweden Ocean). Both open mountains and mountain regions cov­ and Norway, and to map the country up to Varangerfjord. ered with birch were described, as well as the coniferous This expedition probably made him more knowledgeable forests adjacent to themountains. The shrub layer, where about Lapland's nature than any of his contemporaries currant bushes were allowed to play a remarkably large (Eriksson, G. 1969). role, was superficially described. Berries found, as well In 1648, Johannes Schefferus(1621-1679), who was as some of the plants used by the Saami, e.g. Angelica born in Strassburg, was appointed Skyttean Professor archangelica L. and Rumex acetosa ssp. lapponicus Hiit., in oratory and politics at the academy in Uppsala. He were mentioned. The most important goals for reindeer

Lappish women transport their children to the priest differently in winter than in summer. In winter, the child is laid in a La­ Fig. IV. pland sleigh (Saamish gieres) drawn by a reindeer. In summer, she transports the child in a cradle (Saamish gietka) on the reindeer's back. (Schefferus 1956).

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 13

husbandry were also identified. Access to good natural link between Atlantis and the North became widespread. grazing in mountain valleys was emphasised, together with A few examples are given below: the presence of mires that could also be used for grazing. The question of their value as potential farmland was, Glysisvall- theoriginal form for Elysium, or the Elysian Fields, however, left unanswered (Schefferus 1673). the homes of the blissful, said to be found in the North dur­ Andreas Bureus (1571-1646), a cousin of J. Bureus, ing the summer. has been called the father of Swedish cartography. In 1611 lda fjall- according to Rudbeck, more or less the entire mountain chain, originating from the placename ldre. Homer's moun­ he published his first map (at a small scale) of the Kola tain Ida was said to be actually situated in the North. Peninsula and northern Scandinavia down to latitude ea. Helicons berg - Harjedalen 63°N. Enthusiastically supportedby King Gustav IIAdolf, Ararat - Areskutan he published a map covering all of Scandinavia and the Amazons - Hamai warriors and hunters of female gender, living neighbouring countries in 1626, 'Orb is arctoi nova et in a pronounced matriarchal society. They were said to have accurate delineatio' ('A new and accurate description of lived to the west of the Ripheic Mountains in northern Fin­ the arctic region'). Both maps are in conical projection, land. They were exterminated by Hercules, burntand placed and may be regarded as the first real maps showing the in three heaps at Amaisojiirvi, west of the Ripheic Mountains mountain regions. It was not until 1796 that they were (the name originating fromthe Swedish word ripa, meaning ptarmigan). That they should have been placed in three piles superseded, when the map made by Anton Swabo and parallels the three burial mounds at Gamla Uppsala. Clas Wallman of Vasterbotten (i.e. up to the watershed Hercules- according to Rudbeck, the first syllable refers to the between the rivers Tornealvand Kemialv) was published. Swedish word (host; military power), and -kules refers bar- This beautiful map was at the scale 1:440,000, and was to the masculine form of kulla, the word used in Dalarna included in S.G. Hermelin's countrywide atlas. when referring to a woman.

Olof Rudbeck's son, Olof the Younger (1660-1740) Botanical research inherited his father's interest in botany and was his equal Botanicalresearch in Sweden during the 17th century led to as an author. He began his academic studies in his home systematic floristicsand descriptive botany. Olof Rudbeck town ofUppsala at the age of thirteen.Botany and medicine the Elder ( 1630-1702) made a major contribution to this were his main subjects, and his father, who was of course development. He began his research career in medicine professor at the university and even Rector for a time, at Uppsala, and discovered the lymphatic system in the supervised his studies, which were supplemented with a mid-17th century. He published his results in 1653, shortly study visit, mainly concerningbotany, to England, Holland after the Dane, Thomas Bartholin, had independently and Germanyin 1687-1691. On his returnhe was appointed published very similar observations. Having studied in professor of medicine in succession to his father. the Netherlands during 1653-1654, where he studied not Economic interests, foremost those of the State, had only medicine but alsobotany, Rudbeck returnedto Upp­ takenearly travellers to the northernwilderness. Profitable sala, and soon became professor of medicine. He founded mining was the principal attraction. With Olof Rudbeck Sweden's first botanic garden, based partly on Dutch plant the Younger, purely scientificjourneys of discovery were material.His professorship was altered to cover both botany introduced in the north. During the summer of 1695, he and medicine. As a scientist, he represented the modern made a long journey of discovery in Lapland, including view of nature that emerged during the17th century and, as visits to the true mountain region in the Lulea and Tome regards botany, was formulated by, among others, Harvey parts of Lap land. and Malphigi. Rudbeck 's major contribution to botany was Rudbeck's research journey, which was so unusual at his Campus Elysii (1702), illustrations of all the world's that time, was intended to be described in a comprehen­ plants, organised so that prevailing errorsin nomenclature sive publication: 'Nora samolad sive Lapponia illustrata' and systematics could be corrected. Two volumes contain­ - 'Lapland illustrated'. Only one part, however, was ing several hundred illustrations had been printed, before published (in 1701), the part describing the journey from the extensive Uppsala fire of 1702 destroyed almost the Uppsala to the border with the province of Gastrikland. The entire collection of nearly seven thousand woodcuts,among firein U ppsala in 1702 destroyed a large part of the learned which were those to be used for illustrations of plants found production, such as notes, manuscripts, books and most in the mountain regions of Sweden. of the woodcuts. The library at Lovsta in NorthUppland Rudbeck is probably mainly known today for his contains, however, botanical material illustratedin several 'Atland', or 'Atlantica' (1672-1702), an historical work volumes. One of them covers some of Rudbeck's mountain in which he tried to demonstrate that Plato's poetic epic plants and other objects found in nature. Fragments of 'Atlantis' was in fact a description of Sweden, the origin Rudbeck's travel notes (Rudbeck 1695) can today be found of all culture from whence it spread to the rest of the in the Linnaean Society's library in London. These allow world. Through 'Atlantica', Rudbeck's belief about the us to glimpse some of his botanical observations.

Acta Phytogeogr. Suec. 87 14 Erikssson et al. 0.

During his travels in the Tornetrask area, Rudbeck found a large number of plants new to him. When consid­ ering that, on the advice of his guide, he never took more than twenty to thirty steps away from the path, his results are considerable. He also found reason to criticise Schef­ ferus, who had considered that Lapland was particularly deficientin species as regards botany. Whereas Schefferus, with the help of all his inform­ ants, had not managed to list more than 80 types of ' grass', Rudbeck had found more than three hundred during his firstjourney. In 1720 he published a list including the 95 most remarkable finds. Phenological observations were made concerning the flowering of certain herbs at Tornetrask, in Lulea and in Uppsala, as well as various observations of a more or less plant-geographical nature. In 'Acta literaria et scientiarium Sveciae' (1734), he described in words and illustrations the 'Cascavari' waterfall at Kvikkjokk. This is the first published illustration of nature in the Lule region of Lap land. However, interest in botany appears to have diminished gradually; perhaps the reason can be found in the losses caused by the Uppsala fire.Fo llowing the example of his fa­ ther, OlofRudbeck the Younger was also strongly attracted by the great Gothic vision. His father, for example, was influenced to such an extent that, in 'Atlantis ', he placed Paradise itself in vikkjokk. Lap land was, of course, the K former Elysian Fields where, according to ancient belief, Salix grew, and the relationship between Salix and the Swedish word 'salig' (blessed) was considered obvious. Rudbeck the Younger saw it as his duty to demonstrate that Gothic was the origin of all languages. He found, for example, clear relationships between Gothic, the Saam­ ish language and Hebrew. His interest in birds resulted in a richly illustrated bird book, which was not published until 1988. Linnaeus's life, particularly during his younger years, is well known to Swedes. However, a brief recapitulation may be useful in refreshing the memory. He was born in 1707, in a clergyman's home in rural Smaland. He received a relatively good education and spent a childhood close to nature under the inspiration of his father, who was inter­ ested both in horticulture and in the wild flora. Rothman, a teacher of him at the secondary school in Vaxjo stimu­ Fig. V. Cicerbita alpina (Swedish torta, Saamishjearja). Occurs lated his growing interest in botany, by telling about the in northern Sweden in meadow forests and tall-herb meadows, interesting botanical garden at Uppsala (which, however, from the lowland fore t areas to the mountains. It is readily eaten at that time was in rather poor shape). The next teacher by reindeer, elk and bears, and was an important vegetable for of importance was Kilian Stobaeus, who was Linnaeus's the Lapps. The young, peeled stalks were eaten both raw, and teacher in medicine and natural history during his short after being boiled together with reindeer milk (South saamisb time at the university of Lund (1727 -1728). gompa). The dried roots were used as a medicine rich in vitamin Linnaeus's move to Uppsala University in the autumn C, to treat scurvy (Rudbeck, the Younger 1720). 0. of 1728 gave him rather better opportunities to study medi­ cine and a better environment and personal circle of friends, that proved to be a good base for his early scientificwork in botany, its systematics, terminology and nomenclature.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 15

In the course of his journey along the Ume River to Fig. VI. Lycksele, Linnaeus illustrated Cladonia rangiferina var. alpestris (L.) Schaer, i.e. Cladina rangiferina (L.) Nyl., Cladina stellaris (Opiz.) Brodo or both, and Cladonia coccifera (L.) Willd. (Lin­ naeus 1732). The importance of lichens as reindeer fodder was already well-known, e.g. from Magnus (1555). 0.

Pedicularis flammea L. was illustrated by Linnaeus Fig. VII. on his journey between Virihaure and K vickjokk (Linnaeus Olof Celsius, Dean of the Cathedral, gave him economic 1732). support and introduced him to Olof Rudbeck the Younger, in whose home Linnaeus was given lodgings and access to his library. In 1732 the Royal Academy of Sciences (Du Rietz 1942). Similarly, his talent as an author was of placed its entire economic resources at his disposal for a the highest class, whereas his skills as an illustrator were scientific journey to Lapland, which he made during the less developed. Nonetheless, the sketches in his diary have same summer. The intention was 'to illustrate Lapland's a naive charm, and show evidence of a pronounced feeling natural history with regard to stones, soils, water, herbs, for important aspects of the objects illustrated. trees, grass, mosses, quadrupeds, birds, fish and insects, During the winter of 1733, Linnaeus visited Dalarna together with human diseases, health, nutrition, customs for the first time, to study mineralogy. During the follow­ and mode of living'. ing summer, at the request of the Provincial Governor An attempt to reach the mountain region of Vasterbot­ in Falun, Nils Reuterholm, he traversed the countryside ten during the early summer ended, quite understandably, together with friends and assistants in a search for 'rare in the waterlogged Lycksmyren swamp north-west of products of nature and economically unutilised resources Lycksele, after which the journey continued to Lulea and within the three realms of nature'. As regards botany, the furtherup the river Lilla Lule alv.The mountains northwest journeywas a disappointment. In Iter dalecarlicum ( 1953 ), of K vikkjokk gave hima rich reward for his efforts. Within he comments laconically that Sweden's southernmost the space of a few weeks above the coniferous tree limit, mountain region was much more deficientin species than he had collected material that formed the main part of his he had imagined. Flora Lapponica, printed inHolland in 1737. Contempo­ Several botanically interested physicians were among rary scholars considered this to be his masterpiece from Linnaeus's disciples. One of the more prominent was Lars the firstgreat period of publication. Nonetheless, there can Montin ( 1723-1785), a sharp-eyed fieldbotanist who bota­ be no doubt that Rudbeck's pioneering work was of great nised in the K vikkjokk mountains in 17 49. He was the first benefitto Linnaeus. Apart from a rich material of purely to find uncus biglumis. His journeyprovided material for a I botanical nature, Flora Lapponica contained a number of two dissertations, one on the bryophyte Sp lachnum luteum notes connected with the purpose of the journey.Linnaeus 's and the other, his graduate dissertation, on Saami diseases diary from the journey, Iter Lapponicum, was printed in and medicines. Another of these botanically interested London in 1811 and became a literary classic. doctors was Johan Otto Hagstrom ( 1716-1792). He re­ As an observer of all phenomena in nature, as well as ceived much praise from Linnaeus for his dissertation on of human living conditions and behaviour in a wide sense, wild flowers that bees prefer to visit (Pan apium, 1768). Linnaeus was remarkable. His division of topographic re­ Considerably earlier (1749), a visit to his home district gions into biological zones of differentextent, for example, of Jamtland had resulted in a dissertation on the region's was precise and even today endures in parts. He made an economic natural history. excellent differentiation between the mountain region's Linnaeus inspired his disciples to undertake adven­ alpine belts - the forested region, forests of mountain turous expeditions to distant countries, where they were birch, low mountains, mountain slopes and high mountains thought to function as their master's extended arms and

Acta Phytogeogr. Suec. 87 16 0. Erikssson et al.

Fig. VIII. Angelica archangelica. Common on moist sites in small valleys, and beside streams and springs. An important constituent of the spring and early-summer reindeer fodder. Formerly one of the Saarnis'most important vegetables (Saamish boska). Before flower­ ing, the peeled stalks were eaten raw. Like Rumex acetosa ssp. lapponicus (Saamish juopmu), it was used as a preservative in reindeer milk, which was stored for lengthy periods in springs. It also found an extensive pharmaceutical use as a herb for loosening phlegm and for treating cramp (Wahlenberg 1830). Drawing by L. L. Laestadius (1830).

senses, diligently collecting and describing not only true students, Johan Daniel Lundmark, a student of medicine, scientificaspects, but also those of benefitto society. Their and Olof Swartz, during an expedition to collect items interest in the mountain regions of Sweden appears to of botanical and zoological interest in Lapland in 1780. have been limited. One exception was, however, Daniel Unfortunately, it yielded little of scientificinterest. Olof Solander, born in Pi tea in 1733, who anived in Uppsala as Swartz later became one of Sweden's most prominent and a student in 1750. He was one ofLinnaeus 's most cherished widely travelled botanists. students, and went on research trips to Pitea Lappmark and Goran Wahlenberg (1780-1851) was appointed pro­ Norway in 1753, and later also visited the mountains. A fessor of medicine and botany in 1829. He extended our study trip to England led to his abandonment of Sweden know ledge of the Scandinavian world of plants. His Flora and Linnaeus, but with English support he achieved con­ lapponica (1812) and Flora Svecica (1824- 1826) proved siderable success in exploring the plant kingdom in the to be highly stimulating for research in floristics and sys­ southern hemisphere. Aboard the 'Endeavour', he took tematic botany. However, his foremost achievements were part in Captain Cook's first circumnavigation, when Tahiti, in plant geography, where he, together with Alexander von Tierra del Fuego, New Zealand and Eastern Australia were Humboldt, came to be regarded as a pioneer. Wahlenberg visited, yielding a rich collection of material. was a withdrawn and austere person, who largely kept to During the mid-1750s and onwards, the Royal Acad­ himself. He had no disciples who would continue along the emy of Sciences enthusiastically supported research paths he had staked out. However, Swedish plant geography expeditions to exotic countries, undertaken mainly by as he created it came to dominate the relatively modest Linnaeus 's disciples. Curiously enough, interest in discov­ research in this field during subsequent decades. ering more about their own country was weak. However, In the early 19th century, Wahlenberg made four bo­ an exception was the work carried out by two young tanical and geological expeditions to Lapland and northern

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 17

Norway. The Sulitelma Massif, at that time considered studied, among others, the climate and glaciers in the Sarek to be Sweden's highest mountain, especially attracted area, and carriedout advanced mapping, largely based on his interest In 1813, on the small mountain U nna Tuki photograrnmetry. In 1907 he was appointed professor of southeast of Virihaure, he made the first find of Arenaria geography at Uppsala University. Two years later, Parlia­ humifusa Wahlenb. in the Nordic countries, a species that ment decided that Sarek and Stora Sjofallet should be more than a century later came to be of great importance in national parks, something that mainly Hamberg should discussions on the early immigration of mountain flora. In be thanked for. the introduction to Flora lapponica ( 18 12) he gave an ac­ Thore C.E. Fries (1886-1930) spent several years re­ count of the main characteristicsin the plant-geographical searching Lapland's vegetation. In his doctoral dissertation structure of northern Sweden. He differentiated between 'Botanische Untersuchungen im Nordlichsten Schweden' the mountain vegetation zones, and correlated them with (1913), he distinguished between plant communities on climatic conditions. During the period 1812-18 14, he the basis of a new approach used in plant-geographical worked in the Swiss Alps and the Carpathian Mountains, studies in other Fennoscandian regions. The influence of in order to be able to compare conditions in the Swedish snowcover and reindeer grazing on the vegetation was mountains and those prevailing in corresponding regions also studied. elsewhere in Europe. Rutger Sernander( 1866-1944) was professor of Plant One of Wahlenberg's successors was Lars Levi Laes­ Ecology in U ppsala from 1908 to 1931. The dispersal tadius (1800-1861 ). He was born in Jackvik, parish of biology of plants and the development of climate and Arjeplog, his parents being of Saami descent. Laestadius the plant world in Scandinavia were his main fields of grew up in Kvikkjokk, read theology in Uppsala and was interest. His thorough knowledge of plant communities appointed vice-pastor in Arjeplog. In 1826 he moved to in northern countries allowed him to give an exception­ Karesuando as rector, then to Pajala in 1849. Laestadius ally detailed picture of the development of vegetation won great esteem as a botanist, particularly in floristics in Sweden. Semander's wide field of competence, and and plant geography. He conducted extensive journeys of enthusiastic personality, resulted in his being able to as­ botanical research, particularly in northernmost N orrland. semble a group of devoted disciples, the 'Uppsala School', In the early 19th century, he found Papaver laestadianum which enthusiastically fought to promote plant sociology Nordh. in the Paltsa district, which came to be an important during this period of 'Sturrn-und-Drang' in the field of item of evidence in connection with the Nunatak theory, i.e. plant geography. the discussion whether or not species were able to survive G.E. Du Rietz (1895-1967) was one of the more the most recent ice age on the top of mountain peaks that prominent disciples. He was Semander's successor as rose above the ice. In 1838, Laestadius participated in professor in Plant Ecology, 1934-1962. Du Rietz carried P. Gaimard's French expedition in Lapland, as a natural out comprehensive studies ranging from the mountains to history polyglot guide. He published a number of disserta­ the seacoast. Algae, lichens - particularly their reproduc­ tions dealing with Lapland's plants, as well as contributing tive conditions - and mosses (Sphagnum) were given drawings in Wahlenberg (1830), and providing important special attention. contributions to floras prepared by Wahlenberg, E. Fries T.A. Tengvall (1892-1946), who studied in the Sarek and Hartman.To gether with his family, he produced a large district, H. Smith (b. 1889), who worked in the mountains amount of high-quality herbarium material that was given of Jamtland and Harjedalen, and G. Samuelsson (1885- to major botanical collections in Europe. For example, 1944), who studied plants in the mountains ofDalarna, all Gaimard's expedition was given nearly 10,000 sheets. In followed in the footsteps of C. E. Fries. recognition of this, Laestadius received the French Legion Together with the above-mentioned researchers, of Honour (Larsson 1999). mention should be made of G. Bjorkman (b. 1898), Th. Theodor (Thore) Magnus Fries ( 1832-1913), professor Arvidsson (b. 1904) and especially S. Selander (b. 1891). of botany and practical economy at Uppsala University, They worked in the Pite and Lule Lappmark regions. Apart visited Ost-Finnmarken in Norway in 1857 and 1864 and, from plant ecology, Sten Selander involved himself deeply as one of the first, came to know the Arctic lichen flora in nature management, particularly in the preservation of thoroughly. Rutger Sernanderand the lichenologists A. H. undisturbed mountain nature. He was also a successful Magnusson and G .E. Du Rietz were three of his pupils. lyricist. Axel Hamberg (1863-1933) is probably the person Hugo Sjors (b. 1915) who succeeded G.E. Du Rietz, who has meant most for exploration of the Sarek area, was professor in Plant Ecology at Uppsala University partly through more than thirty years' work and partly from 1962 to 1980. His scientific production is extensive, through having inspired and supported a large number of and covers a wide field. He focussed on the ecology of researchers within many disciplines of science. He himself central Swedish mires, but also worked on several mire was a polyhistor, with roots in biology, geology, physical complexes in the mountains. Another of his research fields, geography, climatology and, in particular, glaciology. He of major importance for our knowledge of vegetation in

Acta Phytogeogr. Suec. 87 18 Erikssson et al. 0.

mountain regions, concerns studies associated with water 6 79 . systems chosen as potential objects for exploitation, e.g. the Angermanalv river. Olof Rune (b. 1919) is probably one of those who, in modern times, has been of major importance to research into the flora of mountain regions in southern Lapland, which previously attracted only limited research attention. The relatively few plants throughout the entire mountain chain that show a strong dependence on a substrate in­ fluenced by serpentines and other ultrabasic rocks, were studied in his doctoral thesis 'Plant life on serpentines and related rocks in North Sweden' (1953). Apart from the circle of researchers and explorers, the person who probably did most to spread knowledge of Sweden's northernmost regions was Selma Lagerlof, who published 'Nils Holgerssons underbara resa genom Sverige' (Nils Holgersson's wonderful journey through Sweden). Although this was a book intended for primary schools, it also appealed to older readers. Today, the book belongs to the Swedish national literary heritage. The book's climax was hinted at already at the start (in the southernmost province of Skane), when the wild geese with their exotic addresses were introduced to Nils: Mother Akka came from Kebnekaise, only relatively recently known as Sweden's highest mountain, Viisi came from the Ovik mountains, Kolme from Sarektjakko, etc. Selma Lagerlof's book has been translated into about thirty lan­ guages, and she was awarded the Nobel Prize in literature in 1909. Nils Holgersson may have given generations of readers, both Swedish and elsewhere, their only contact with the northern regions described in the book. Fig. IX. Astragalus alpinus L. (Saamish duottar-saphal)'There is no doubt that this species is one of the most palatable, and that Large-scale exploitation of mountain regions it is among the summer delicacies of the reindeer.' Drawing by L. L. Laestadius in Wahlenberg, C. 1830. The growth of the industrial world during the early 20th century made great demands on access to abundant energy and efficient transportation systems. In the context of botanical study was made at Stora Sjof allet before the dam Sweden's northernmountain regions, this implied, e.g. the was built, but that the reports were not published until many exploitation of hydroelectric resources in many of the large years later (Bjorkman 1939, 1965). Changes that occurred rivers, and the building of a railway to the Atlantic coast, subsequent to the completion of the four damming stages which was always ice-free. The 'Iron-ore Railway' from have been studied to a minor extent only. Kiruna to Riksgransen was opened in 1903, and electrified Stora Sjofallet National Park was considered by many in 1915. During the summer of 1911, the stretch of the 'In­ to be the finest in Sweden, perhaps even in Europe, and it land Railway' from Gallivare to Porjus was completed, and may be of interest to look a little closer at events leading in 1919 work started on the hydroelectric dam at Suorva on to its establishment and subsequent fate. Most of the great the river Stora Lule alv. The nature-protection organisation rivers draining the Swedish mountains are now regulated, at that time, which acquiesced in the destruction of large and in many cases the decision-making processes have parts of Stora Sjofallet National Park, subsequently was been fairly similar. subjected to massive criticism. In 1818, Abraham Roman gave the first relatively de­ Spin-off effects of hydroelectric exploitation were tailed description of Stora Sjofallet, in his book 'Beriittelser that some inventories of a biological nature were made. omNorrbotten och dess Lappmarker' (Tales ofNorrbotten A consistent feature was that fisheries biology inputs were and its Lappmarks (Saami districts). This was followed in given predominance, and that other biological studies were 1866 by the description given by Carl Anton Pettersson concentrated mainly to the littoral zones prior to regula­ in 'Lappland dess natur och folk' (Lapland, its nature and tion. It may be mentioned, for example, that a detailed people), and we may regard Pettersson as having, from a

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 19

Fig. X. Stora Sjofallet National Park. The Suorva reservoir at Suorva, seen from the 'Road to the West'. When the photograph was taken (30 September 1996), the water­ level was fairly low. Photo: Eriksson. 0.

tourist's point-of-view, 'discovered' Stora Sjofallet and the 50 m to about one kilometre. Normally, there is a belt of sev­ neighbouring mountains. The Swedish Tourist Association, eral hundred metres' width, almost devoid of higher plants founded in 1885, opened up the area to tourists in vari­ and animal life. Many species have disappeared, whereas ous ways, but sometimes in conflict with the spokesmen others have been drastically reduced. For example, within for nature protection. Stora Sjofallet National Park was the Sjofall area there were originally 532 different vascular established in 1909, but already in 1910 work started on plant species. Of these, 367 were in areas withdrawn from the Porjus power station some 85 km downstream. In a the park in the first stage of regulation. Seventeen species longer perspective, the regulation of the lake system above were totally exterminated, e.g. Rhododendron lapponicum Porjus was almost self-evident and, as might have been (Bjorkman 1939, 1963a,b). Lindberg (1970) estimated that expected, in 1920 Parliament decided that areas upstream 50 years' exploitation in the national parks had removed that would be affected by the building of the dam and other the living conditions for at least 100,000 pairs of ducks, regulation measures at Suorva should be withdrawn from waders, small birds, etc., together with mammals and birds the national park. of prey that were affected directly or indirectly. Abrahamsson ( 197 5) summarised some of the effects of This exploitation of an important mountain area natu­ regulation as follows: 'Three of the Stora Sjofall waterfalls rally could not take place without protest. Researchers such are completely dry (Laestadius's waterfall, Pettersson's as Axel Hamberg, G. Einar Du Rietz and Sten Selander waterfall and Diiben 's waterfall), and Widmark's waterfall reacted strongly. Abrahamsson (1975) expressed surprise and Hermelin's waterfall have only modest amounts of that the board of the Swedish Tourist Association (STF) what may be described as overspill water.' did not express an official standpoint concerningencroach­ Up to the early 1970s there were four stages during ments in the park. which the level between high water and low water, i.e. the regulation amplitude, was increased, to 30 m in the Data for decision-making in Swedish-Norwegian rein­ very large Suorva reservoir. Numerous smaller lakes and deer grazing conventions - new knowledge of nature waterways have disappeared. As a comparison, it may be in mountain regions mentioned that the natural amplitude in these water systems before regulation was 3.7 m. The water volume is about The 'Lapp codicil', i.e. the addendum to a treaty between two-thirds of that in Lake Vanem. The full reservoir is Denmark and Sweden that, since 1751, with supplements estimated to be ten times greater than the natural volume in 1883, had regulated the right of Swedish Saami to use at normal water level in the unregulated lakes upstream reindeer-grazing land in Norway, came to end in the early an of Suorva. 20th century ( 1907). It was replaced by a series of Swedish­ On account of its size and its orientation, largely NW­ Norwegian commissions that, under Finnish chairmanship, SE, the reservoir is often exposed to severe storms, which were charged with obtaining the knowledge necessary for would have been manageable for boat trafficon the smaller negotiations concerning a new convention. The work was lakes before regulation. largely conducted in the northernmost mountain region Depending on the character of the shoreline, a flooding within the Karesuando and Kiruna municipalities, and in zone has been created, which varies in width from perhaps Troms county in Norway.

Acta Phytogeogr. Suec. 87 20 0. Erikssson et al.

I ·I }

< !­ (.) a: �

Fig. XI. Part of the map by C.Th. Fries and E. Bergstrom, 'Map of the vegeta­ tion above the coniferous forest limit in the parishes of Karesuando and Norra Jukkasjarvi'. Eight dominant vegetation types, as well as boulder-strewn areas and snow-patches, are included. (App. C in E. Lonnberg 1909)

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 21

As an example of the planned work, it may be mentioned National Parks that the 1906 Reindeer Grazing Commission (Stockholm 1913) had created a picture of reindeer-grazing conditions The National Parks inthe mountains not only contribute to and migration patterns across national borders, by means providing -occasionally fragile- protection to valuable of interviews with Swedish and Norwegian Saami, local natural environments, but also provide freedom for the inhabitants and government officials. predatory fauna. An increased frequency of visitors, and The 1907 Commission (Stockholm 1909) focussed on publicity, have certainly increased the awareness of the the possibility of reducing Swedish reindeer herding on general public with regard to these important but limited Norwegian territoryby variousmeasures. Information was regions of the mountain chain. The parks also offer the collected during journeys along the migratory routes and opportunity to conduct research based on long-term series, in the spring and summer pastures in the mountains. e.g. concerning relationships between weather, climatic The Reindeer Commission of 1909 (Helsinki 1912) ini­ change, vegetation and fauna. Examples are the research tiated the creation of the widest biological knowledge base projects based on the Abisko Research Station (founded throughits own fieldwork.For two yearsthe vegetation was 1912) in the Abisko National Park and, to a lesser extent, studied, with emphasis on the reserves of reindeer grazing in the adjacent Vadvetjillo National Park. in differentseasonal grazing areas. A line transect model for enumeration, adapted to make it practically applicable, Present�day maps of the mountains was used. Great interest was shown in the phenology of forage plants, as well as in the location and nature of calv­ The first practically usable maps covering the entire moun­ ing sites, together with the timing of calving. tain chain are the Ordnance maps at a scale of 1:200,000. Details of the time at which a complete snow cover The first to be published - the Abisko map - appeared became established, and of its successive thawing, and of in 1886, and by 1923 the entire mountain chain had been the structural features of snow that negatively influence covered. Afterabout 50 years, the present-day topographi­ grazing, were also published. As an annex (originally con­ cal maps, based on interpretation of aerial photography, fidential) to the 1909 court proceedings concernedwith the have now superseded the Ordnance maps. The first of the reindeer-grazing question, a pamphlet entitled 'Om renarna new series covered the Kiruna area (1960-1961), and the och deras levnadsvanor' (On reindeer and their living final sheet of the mountain chain was published in 1978. conditions; Lonnberg 1909) was submitted. A vegetation The map scale is 1:50,000 or 1:100,000. These maps are map at the scale 1:200,000, showing the northernmostpart substantially more reliable than their predecessors. of the Swedish mountain chain, drawn by Th. Fries and E. A map of Sweden's solid geology was published in Bergstrom, was included. When considering the extent of 1958, the same year as the map of Sweden's Soil Types. the region and its inaccessibility, the compilation of this Both were published by the Geological Survey of Sweden. map may be regarded as a pioneering work of the highest The National Atlas of Sweden was published in 17 vol­ quality. The findings of the Commission were published umes between 1990 and 1996. Naturally, in addition to the in six volumes. abovementioned general maps, numerous detailed maps of Fries also participated in the Reindeer Grazing Com­ limited areas have been created over the years. During the 1970s, a progr e of aerial photography mission of 1913, where cooperation was established with amm a working group consisting of experienced Swedish, was started, covering all of Sweden on infrared film from Finnish and Norwegian researchers in the biological­ an altitude of 9,000 m. This material formed the basis of geographical sphere. This working group conducted the 'Vegetation map of the Swedish mountains', in colour fieldworkin 1914 and 1915. The results were presented and at a scale of 1 : 100,000, prepared by the Department of in 18 volumes (Stockholm, 1917). Probably none of the Physical Geography, Stockholm University, by Andersson earlier commissions that had worked to acquire data for et al. in 1978-1984. This work was commissioned by the the reindeer-grazing conventions could have presented so Swedish Environmental Protection Agency (EPA). The much know ledge on contemporary reindeer management National Land Survey, Lulea, is presently working with and on the biological conditions determined by natural the mapping of the remaining parts of the country, using conditions in the mountains, as did the 1909 and 1913 much the same approach. commissions. A new 'Convention between Sweden and When the National Land Survey's vegetation maps Norway regarding the right of migratory Lapps to reindeer finally cover areas along the borders of the mountains, grazing' became law in 1919 (Svensk forfattningssamling which were omitted in the EPA map, the vegetation of the 1919, No. 895). This has subsequently been prolonged in mountain chain may be considered to be largely known. stages. Anew Convention is presently being prepared, but when it will be approved is not known.

Acta Phytogeogr. Suec. 87 1. The reindeer

The reindeer was an important source of nutrients, 1.1 Origin both for the Neanderthals ea. 50,000 years ago, and for the The origin of the reindeer family is not fully known. Cro-Magnon people ea. 25-30,000 years ago, as evidenced However, it appears to have come from Alaska, Beringia by finds of bones and aJ.tefacts from ancient dwelling or from the mountain regions of northeastern Asia, more places. Cave paintings in the Dordogne, France, and at than 400,000 years ago (Banfield 1961). Altamira, Spain, are excellent examples of Palaeolithic Flagstad & R0ed (2003) used mitochondrial DNA art at its peak during the Magdalenian period 13,000 years sequences to establish that a large and coherent popula­ ago, and indicate the great importance of reindeer to the tion in Beringia, with dispersal far into Eurasia, appeared peoples of that time. to be the most important origin of the genus's gene pool. A period with relatively mild climate occurred about In addition, there appears to have been a small, isolated 16,000 years ago, and the ice began to recede towards the refugium in westernEurasia and a larger, well-defined one north. Southern Sweden became ice-free about 10,000 to the south of the North American ice-sheet during the years ago, and after a further 1500 years, it had been re­ Wichelian/Wisconsin glaciation. duced to a small remnant in Northern Sweden. Several subspecies have become differentiated from The reindeer, and other species we findtoday in the these dispersal areas and have spread over the Old and mountain fauna, e.g. certain rodents, the arctic fox and the New World, to both boreal and to alpine and arctic ptarmigan, followed the ice as it receded. Immigration to areas from lat. 45° N (northern China) to lat. 80° N (the Scandinavia took place during the later part of the 'Dryas Arctic islands). period' about 12,000 years ago. The Narke Sound, linking When the latest ice age began, about 1 90,000 years ago, the Yo ldia Sea with the Atlantic, cut off the Scandinavian the reindeer was already a pronounced feature of the Euro­ peninsula about 9,000 years ago and affected the expansion pean fauna. The ice had its greatest extent about 25-30,000 of the fauna towards the north. Reindeer were, however, years ago, and the reindeer had its corresponding greatest common on the Scandinavian tundra during the Aller0d dispersal 12-15,000 years ago. At that time, it dominated period (12-9,000 years ago). The tundra period came to an the ice-free tundra to the south of the ice, in what is today end with a warm period about 9-8,000 years ago, resulting Central and We stern Europe, including Ireland, England in the rapid establishment of coniferous and deciduous and large parts of the N orth Sea - at that time also tundra. forests and in the development of a fauna consisting of The reindeer co-existed with species such as the woolly elk, red deer, beavers and forest birds. Both humans and mammoth, hairy rhinoceros, musk ox, cave bear, cave reindeer moved successively towards more high-altitude hyena and arctic fox. The reindeer has managed to survive and nutrient-poor regions. the demise of most of its contemporary species. The melting of the land ice, especially in areas of north-

Fig. XII. The genus Rangifer occurs in a number of forms in the northernhemi sphere, e.g. Rangifer tarandus tarandus east of Seben­ a. Keul, Verkhoyansk Mountains, Republic of Sakha (formerly Yakutia). Photo: Eriksson, 0. September 1973; Rang?fe r tarandus tarandus at Tavva­ b. vuoma, Lainiovuoma Saami village, Swed­ ish Lapland. Photo: Eriksson, August 0. 10 1973; Rangife r tarandus groenlandicus at Ravs­ c. jon, S. Stromfjord, Greenland. Photo: 0. Eriksson, 17 August 1978; d. Rangifer tarandus platyrhynchus, fe male with calf, Svalbard. Photo: Anon.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 23

c

Acta Phytogeogr. Suec. 87 24 0. Erikssson et al.

A reindeer herd depicted ea. 10,000 years ago on a cave wall at Te yj at, France. The impression of a large number of animals Fig. XIII. in rapid motion is striking.

em Fennoscandia that melted early, probably permitted the Hardangervidda, Ottadalen, Snohetta and Rondane, there immigration of the tundra reindeer from the east, together still remain populations of wild reindeer, living under with the hunters who depended on them. Wild reindeer strictly regulated conditions. were now numerous also in the northernmostparts of the The Scandinavian domesticated reindeer, both the Scandinavian peninsula, and along the mountain chain mountain reindeer and the forest reindeer, can trace their down to Slima and ldre in Dalama.However, their numbers main origin to the Eurasian tundra reindeer, Rangifer decreased with time, depending on intensive hunting by tarandus tarandus L., which was described and named meat-demanding settlers as well as by Saami who wanted by Linnaeus in 1758. However, Hollsten (1774) reported to exterminate wild reindeer, which caused disorder by that, during the 18th century, there were wild forest rein­ mixing with the domesticated herds. The last wild reindeer deer of a different breed in the taiga around the Gulf of in Dalama were shot during the 1860s-1870s, whereas in Bothnia. These reindeer were reported to be much larger Norrbotten, they had probably already disappeared dur­ than the mountain reindeer, and had a darker coat. During ing the 1830s (L6nnberg 1909). However, in the southern the mating period they moved farther to the west from the parts of the Norwegian mountains, e.g. Setesdal Ve sthei, coastal regions into the area of the Lappmark (Saami ter­ ritory), where there were domesticated reindeer based on R. tarandus tarandus L. This intermingling might explain why some of the domesticated 'forest' reindeer, at least in the early 20th century, were somewhat larger than the domesticated 'mountain' reindeer. The wild forest reindeer (Rangifer tarandus fe nnicus Lonnberg) gradually was exterminated within most of its northern rangeup to the end of the 19th century. In east­ em Finland, Karelia and neighbouring regions of Russia, however, an increasing population still remains (Siivonen 1975, Nieminen 1980). Nonetheless, Lonnberg believed that descendents of the wild forest reindeer described by Holisten ( 1774) were to be found among the semi-wild 'domesticated' forest reindeer belonging to Sjocksjock's Saami village in the easternpart of J okkmokk parish. These were renowned for their size and attractive appearance, e.g. as mentioned in 'Proposals for regulations concerningthe Swedish Lapps and settlers in Sweden', a report published in Stockholm by a Parlia­ mentary Committee. (Forslag till Forordning angaende de Man and reindeer in an early rock-engraving at Fig. XIV. Zalavruoga, coast of the White Sea, Russia. A possible indica­ Svenska lappame och de bofasta i Sverige 1883) tion of an early migration from the east. After Skjenneberg & Sjockjock's Saami village was split in 1903 and amal­ Slagsvold ( 1968). gamated with the Jokkmokk and Gallivare forest Saami

Acta Phytogeogr. Suec. 87 se and abuse of reindeer range 25 V

villages. In 1945, Jokkrnokk was amalgamated with Serri serted by Lonnberg (1909). We can only speculate about and Udtja. The Rodingtdisk group in Udtja village became the conclusion he might have reached, had the material famous for their reindeer, which were much larger than included reliably identified Rodingtdisk skulls. Experi­ those normally found in other Saami villages. Lonnberg enced reindeer herders (pers. comm. Stefan Mikaelsson, ( 1909) observed them in the field, interviewed reindeer Rodingstrask andApmut-I var Kuoljok, Sirkas) assert that it owners and conducted taxonomic investigations, includ­ is still possible to distinguish reliably between descendents ing craniometric studies. Unfortunately, his material was of the ancient Rodingtrask line by sight. Regardless of limited. His conclusion was that the Rodingtdisk popula­ any conclusions reached, a study of family relationships tion of domesticated forest reindeer, although diluted, between domesticated mountain reindeer, 'normal' do­ probably originated from R. tarandus jennicus Lonnberg, mesticated forest reindeer, the Rodingtrask type and wild the Finnish-Karelian subspecies, whereas the domesticated Finnish forest reindeer (R. t. je nnicus), as inferred from mountain reindeer were taxonomically identical to R. mitochrondrial DNA sequences, would be illustrative. tarandus tarandus, as well as the domesticated 'forest' reindeer farther south in the reindeer-management area, e.g. Mausjaure or Mala. 1.2 Reindeer fe eding Espmark ( 1981) studied the taxonomic relationships between the domesticated mountain and forest reindeer As an herbivore, the reindeer is a fastidious generalist and in Sweden. Skull as well as antler measurements were opportunist, searching for the most valuable grazing avail­ made. The material studied consisted of more than 800 able while constantly on the move, generally against the skulls collected during the early 1950s from most of the wind, and consuming what it finds with great finesse. Its reindeer-management area, and stored in the Swedish menu consists of most of the alpine and forest flora; thus Museum of Natural History. Espmark's report does not examples are given only occasionally in what follows. reveal whether efforts were al so made to include material The types of vegetation used in mountain reindeer from the Rodingtrask population. His conclusion was that husbandry, principally during spring, summer and au­ the two groups of domesticated reindeer are not completely tumn, are all within the year-round territories where the identical, but that the differences are not as distinct as as- Reindeer Husbandry Act of 1977 is applied throughout

XV. Even on sites exposed to heavy grazing pressure, patches with abundant lichen growth can be observed. Topography, snow­ Fig. depth, etc., limit access to such sites by the reindeer. Cladina or Cladonia spp. On steeply inclined slabs in the Birch forest heath type with mosses east of Seukokjaure, Sorkaitum Saami village. Photo: Eriksson. 15 July 1998. 0.

Acta Phytogeogr. Suec. 87 26 Erikssson et al. 0.

Fig. XVI. Among other species, Ep ilobium angustifolium and Solidago virgaurea (Saamish horbma and beatnatnjuovccarassi) are in bloom at the height of summer, providing high-quality forage near the tree line. Birch fo rest heath type with mosses, east of Seukokjaure, Si:irkaitumSa ami village. Photo: Eriksson, 1 August 1997. 0. the year (Geijer 2003), and in the low-alpine belt and the lows (Salix spp.), especially those that have more or less upper parts of the subalpine mountain birch woodlands. green, non-hirsute, leaves, such as Salix phylicifolia and The types of vegetation included in the present study can S. hastata, have great value for grazing in the early stages, all be found within these zones. and are considered to be among the most attractive (L. Baer, The tree limit (Saami: orda) is one of the most im­ Idivuoma, pers. comm., Eriksson unpubl, Warenberg P. 0. portant natural lines of demarcation that influence the et al. 1997). Willows of similar value, but developing later presence of reindeer Blind, Idivuoma, pers. comm., and smaller in size, are S. reticulata and S. herbacea, the (0. P. Ruong 1982). The low-mountain area at higher altitudes occurrence of which can, to some extent, be associated provides early grazing on thawed summits and southern with thawed snowdrifts. slopes, as well as on the sheltered, yet easily monitored and fairly regularly used calving grounds. However, the Pastures in the mountain birch woodlands grazing there is often sparse. Lichens such as Cetraria spp., Cladonia spp., Cladina spp., Sterocaulon spp., and shrubs The mountain birch woodlands (close to the tree limit) such as Arctostaphylos spp., Va ccinium spp., Empetrum provide protection in bad weather and both lichen and green hermaphroditum and last-year's grasses, dominate initially grazing during the snow-free period. Here can be found (Lonnberg 1909, Palo 1981). Betula nana has somewhat more or less the same grazing plants as within the low of a special status as a grazing plant. The top shoots are alpine zone. Mountain birch leaves provide a considerable cleaned of bark, buds and young leaves. The twigs are nutrient supplement and, in addition, a number of nutritious broken off more or less where wood and bark make up herbs flower during July-August, e.g. Ep ilobium angusti­ roughly similar amounts ofbiomass. Traces of grazing are jo lium, Melampyrum sylvaticum, Solidago virgaurea, and seen in the white-scraped remaining twigs. Cicerbita alpina. These plants are of great importance, and As the spring and early summer progress, fresh grazing are grazed when the myriads of insects permit. of graminaceous plants becomes available, e.g. Festuca Fungi such as Leccinum versipelle, L. rotundifolia, ovina, Deschampsia cespitosa, D.flexuosa, Anthoxanthum Suillus bovinus, S. variegatus and Lactarius trivialis are odoratum, A. odoratum subsp. alpinum, Phleum alpinum, searched for and grazed in ten sively during the late summer and early autumn (Lonnberg 1909, Skjenneberg Slags­ ]uncus tri.fidus and Carex bigelowii. Thawed stretches of & wetland and springs provide species such as Eriophorum void 1968, 0. Eriksson, unpubl.). Dry fragments can even vaginatum, Angelica archangelica, Menyanthes trifoliata be identified in rumen contents from the winter-grazing and sedges. Roots and rhizomes are also grazed. Wil- period. However, the moisture content of fungi is high

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 27

Fig. XVII. If the ground-lichens are inaccessible to the reindeer, tree lichens become important as a food source. Intensive grazing on birch can, however, lead to digestive problems, because fragments of birch bark are ingested and accumulate in the digestive canal (personal communication, Lars Baer). Birch forest heath type with mosses and scattered Uingfjallet, Idre Saami village. Photo: Pinus sylvestris. 0. Eriksson, 15 July 1998

(ea. 90%) and the content of essential nutrients is low. The In years when weather conditions (i.e. snow) make it proteins that are present are principally enzymes, of high impossible to move to, or to remain in, the winter-grazing quality owing to their content of all essential amino acids. areas, the spring and autumn areas may also be used as In addition to large amounts of phosphorus and potassium, winter grazing. The lichens and shrubs associated with the many fungi have valuable concentrations of calcium, zinc, ground layer are now supplemented by epilithic lichens, iron, manganese and selenium. The partiality of reindeer such as Cetraria hepatizon, Hyp ogymnia alpicola, H. for a diet of fungi is probably conditioned by the mineral intestiformis, Parmelia centrifuga and saxatilis. The P. 1968, content (Skjenneberg & Slagsvold Holmberg & grazing of epilithic lichens can be detected through the Marklund 1996). wear caused by mineral particles, which appears to have Apart from the migrations from winter to summer graz­ a disastrous effect on the front and eye-teeth of the lower ing areas via the spring and autumn grazing pastures, an jaw, thereby shortening the life of the animal (Skuncke 1958, 1968). altitudinal migration can be seen in the mountain area of Skjenneberg & Slagsvold some Saamivillages. This extends from the spring pastures The tree layer contains epiphytic lichens. On the branches of the low-alpine zone, down to the early-thawed pastures of the mountain birch, the most common lichens are possibly along valley bottoms below the forest limit, and then moves Cetraria sepincola and Bryoriasim plicior, whereas on the upwards following the thaw and budbreak, driven by the trunks the lichen that is most clearly visible is Parmelia oli­ hatching of mosquitoes (0. Blind, pers. comm.). vacea, owing to its deep olive-green to brown-black colour. Within the summer-grazing area, there is a daily al­ It is abundant above the level that it generally covered by titudinal migration, between good pastures in the valley snow during the winter. Other frequent species are Parmelia bottoms, and resting places on snow-patches or on wind­ sulcata and Physcia stellaris. swept summits, where the reindeer find cool conditions and The intensive utilisation almost throughout the year a minimum of irritating insects (Kuhmunen 1974). of plants sensitive to wear-and-tear is likely to have a de­ The reindeer normally make use of the plant commu­ structive effect on the vegetation, which consists of many nities close to the tree limit during the snow-free period, species with long regeneration periods. partly during a rather short period, connected with the traditional reindeer migrations between winter and sum­ mer areas during the calving period in May, and partly during the autumn, when the herd is often left to graze more or less freely.

Acta Phytogeogr. Suec. 87 28 0. Erikssson et al.

Fertility affect other organs and parts of the body, causing severe 1.3 necrotic lesions that often contribute to the death of the Skuncke (1973) reported that 80% of the female reindeer animal. In the Giillivare district between two and three in the reindeer research herd normally gave birth to a calf thousand reindeer died during the summer and autumn of annually, from their second year of life until their tenth or 1866 (Skjenneberg & Slagsvold 1968). thirteenth year. Skjenneberg & Skogsvold (1968) reported Anthrax is a serious bacterial disease that afflictsmany 70-80% calving by Norwegian domestic reindeer in the species of herbivorous animal. It is caused by Bacillus same age interval. anthracis. Mortality among untreated animals is 5-20%. The disease has not been verifiedwith certaintyon reindeer in Scandinavia. In Russia, a number of severe reindeer Diseases epizootia have been confirmed as anthrax. For example, 1.4 there was a massive outbreak on the Jamal peninsula in The reindeer's mode of life, the relatively sparse groups or 1911, when more than 100,000 reindeer died (Dratjinski herds present in large areas, and the length of time during 1914, Skjenneberg & Slagsvold 1968). the year for which the ground is covered by snow or ice, Brucellosis is a disease that affects many different result in the most common diseases' being of minor im­ species of animal, including man. There are various Bru­ portance (BerglOf 1923, Skjenneberg & Slagsvold 1968). cella species; the one that causes disease and abortions in Infectious diseases can, however, become a major problem reindeer is Brucella suis biotype 4, earlier named Brucella if the reindeer are kept under more intensive management rangifer. Brucellosis occurs in most areas where reindeer or confined in pens. The Reindeer Grazing Convention of are found, but not in the N ordic countries. The disease is a 1919 (Svensk fo rfattningssamling 1919 N:r 895 §158-167), problem for reindeer husbandry, on account of its stealthy provides information on how epizootia such as reindeer development and continuous losses, but also because it is plague, reindeer disease, anthrax and infectious hoof dis­ a zoonotic disease (Rehbinder & Nikander 1999). ease should be dealt with when Swedish reindeer graze in Among parasites that cause disease in reindeer, the Norway. Reindeer plague (Pestis tarandi) is a disease that meningeal worm, Elaphostrongylus rangiferi, is the one occurred frequently during the 18th and 19th centuries. In capable of causing the largest losses. Eggs and larvae are the mid-1760s, there was a particularly severe outbreak transported through the veins to the spinal cord and beneath in the reindeer herds. Many Saarni lost almost all their or within the cerebral membranes. Locomotion disorders reindeer, and began instead to fishin the Norwegian fjords. occur, leading to the death of the animal. In some years As a result, the population in Jukkasjiirvi was reduced there are large losses among calves, but also among adults over a 25-year period, from 1300 persons to fewer than (Rehbinder & Nikander 1999). 900 (Ruong 1982). In 1896, thousands of reindeer died in Air-bornepests, such as mosquitoes, gnats, horseflies, northern Lapland, probably because of an infection with skin warble(H ypoderma tarandi), and throat warble ( Ce­ Clostridium septicum (Skjenneberg & Slagsvold 1968, phenemyia trompe) areassumed to be well known and are Rehbinder & Nikander 1999). In 1911, reindeer plague not discussed further here. broke out in the Arjeplog mountains, killing the entire calf population (Ruong 1982). Reindeer pasteurellosis (Septicaemia haemorrhagia) 1.5 Mortality is caused by bacteria belonging to the Pasteurella group. In 1912 and 1913, there were severe outbreaks, extending All efforts to calculate the number of reindeer that die frornArjeplog northwards. Between one and two thousand each year suffer from large errors in the source material, reindeer died. In 1924, there were outbreaks in Jiimtland. and thus provide rather unreliable results. The original ex­ The summer of 1959 was exceptionally hot and dry in perimental method was to search for carcasses in the field, southern Scandinavia. Several hundred reindeer, most of which, for practical reasons, is never 100% reliable. During them calves, died on account of pasteurellosis in about the summer, carcasses disappear rapidly, e.g. because of the same areas as were affected by the 19 12- 13 outbreak. drowning, predators or scavengers. During winter, the snow In one Saami village, the losses amounted to 70% of the cover may hide a carcass (Haglund 1966, 1968, Bjiirvall one-year-old calves (Skjenneberg & Slagsvold 1968). et al. 1990). Radiotelemetry, i.e., searching by means of Contagious hoof disease, caused by infection with radio signals from reindeer fitted with transmitters, has Fusobacterium necrophorum, was a common disease been used in research in the Saami villages of Umbyn and of reindeer during the days of intensive management Jillak:aska,but for economic reasons, only a few animals (Rehbinder & Nikander 1999). Linnaeus wrote in Iter could be used (Bjiirvall et al. 1990). lapponicum (1732) about this disease in the mountains of As regards adult reindeer, studies in Umbyn and Jruddi­ Lapland. The Saami name 'slubbo' refers to the swollen kaska Saamivillages gave the result that, among females, an­ foot and means, quite simply, 'club'. The infection can also nual mortality is probably ea. 2.8% (Bjiirvallet al. 1990).

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 29

Table lA. Reindeer killed by predators in the study areas and in Sweden in 1986 - 1994 (Nilsson, 2004)

Saami Village 1986/87 1987/88 1988/89 1989/90 1990/91 199 1192 1992/93 1993/94

Lainiovuoma 20 10 59 22 51 107 52 123 Saarivuoma 104 90 85 65 483 155 6 209 Mellanbyn 26 61 125 64 114 33 70 113 Sorkaitum 38 243 214 179 311 85 328 132 Mittadalen 461 558 381 760 318 340 470 176 Idre 32 67 149 179 28 10 10 26

Total in Sweden 5012 6859 6910 8252 9030 8152 12495 5491

Table Reindeer killed by predators in 1993/1 994 divided among study areas, reindeer gender and/or species of predator (Nilsson lB. 2004).

Saami Village Male Female Adult, Unknown unknown age and Unknown gender Calf gender Wolverine Lynx Bear Eagle predator Total

Lainiovuoma 87 12 110 4 2 5 2 123 24 Saarivuoma 138 21 11 37 2 199 6 3 209 Mellanbyn 51 23 39 82 21 10 113 Sorkaitum 78 12 6 82 37 11 2 132 40 Mittadalen 42 12 112 4 9 141 4 21 1 176 ldre 9 6 11 24 2 26

Calf survival is strongly influenced by the condition ficultyto findall carcasses.The cause of death can probably of the reindeer cows, which is governed, e.g. by access to, be determined with much greater reliability (Rehbinder & and quality of, grazing both before and during pregnancy Nikander 1999, Bjarvall et al. 1990). Compensation from (Rehbinder & Nikander 1999). Factors such as late arrival the State for reindeer killed by predators was paid up to at the calving areas, low temperatures, precipitation, long­ 1993/94 on the basis of carcasses found. From 1994/95, lasting snow cover, difficulty in fording swollen rivers compensation has been based upon the number of bears, when the thaw is late, as well as predation, strongly affect wolverines, lynxes and wolves within each Saami village the survival of new-borncalves (Bergerud 1980, Reimers (Svensk rennliring, 1999). 1989, Bjarvall et al. 1990). When the firstproblems in life have been successfully overcome, the calf must then find access to the right sort of forage in sufficient amounts 1.6 Slaughter during the following autumn and winter. The results of the study of reindeer losses in Umbyn The slaughter percentage in Swedish reindeer husbandry and JillakaskaSaami villages (Bjarvall et al. 1990) indi­ varies with time and is decided by, e.g. the preferences cate that annual calf mortality during 1982-1986 was, on of the owners, the requirements of the market and the average, 13.1% in Jillakaska and 13.5% in Umbyn. In Jillakaska, predators were responsible for 66% of these losses, and in Umbyn the corresponding figure was 75 %. Table lC. The total grazing area in the Saarni villages and the maximum number of reindeer allowed (Svensk rennaring 1999*, Among calves that die during the summer, 60% are the re­ Swedish-Norwegian Reindeer Grazing Commission of 1964t). sult of predation and 15-20% are due to illness, drowning, starvation, etc. (Rehbinder & Nikander 1999). The Board Saami village Area Max. no. of of Agriculture estimates that annual losses of adult animals (km2) reindeer 1988 in Swedish reindeer husbandry amount to 10%. Half ofthe Lainiovuoma 3 497 10 500 animals lost die during the summer (Lantbruksstyrelsen t Saarivuoma 3 513 7000 1985). However, in 1988 predators killed 58,000 reindeer t Mellanbyn 3 355 7 000 * (Svensk rennaring 1999). Sorkaitum 4 879 8 000 * The extent of predation over time is very difficultto Mittadalen 3 902 5 000 * estimate with acceptable precision, on account of the dif- Idre 2 883 2 700 *

Acta Phytogeogr. Suec. 87 30 0. Erikssson et al.

Saarivuoma sameby Lainiovuoma sameby 16000 20000

14000 - ______-- "\ '+-! 15000 0 .... ______...... i I \:\\ . .... 4) \ . . -- � �:8000 "-- -=-�\ .. / " 10000 .. --- '-:c-- .8i ...... / ·-. 6000 · -· · a ".., �---1--- - � · ..� - - · · - 4000 --- -- ______·-·- -- -·- --·-- .... 5000 � - z - - ..i 2000 ------

0 ' ' 19 19 19 19 19 19 19 19 19 19 19 19 19 19 19 20 0 19 19 19 19 19 19 19 19 19 19 19 19 19. 19 19 20 25 30 35 40 45 so 55 60 65 70 75 80 85 90 95 00 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 00

Year Year

Sorkaitum sameby Mellanbyn sameby

12000 1 00 ------. 00 10000 '+-! · -- · 0 .... - �·\-· -- --- .... G) 8000 - ;/_---'""'_-_-_-_-_-,_-_-�_-_-__ ------";l -- _ · '\ - ---- :. .8]s ..... 6000 ------/ - - ...... ---- i' - � - 4000 ...... i e ·· ·· 2000 \; 0 0 - 19�· 19 19/ 19 19 19 19� 19 19 19 19 19 19 19 19 20 19�---�--�------� 19 19 19 19 19 19 19 19 19 19 19 19 19 19 20 25 30 35 40 45 50 55 65 70 75 80 85 90 95 00 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 00 60 Year Ye ar

Mittadalens sameby nya sameby Idre

12000 .------·----, 5000 -�------� --- 10000 . t .... 8000 .... G) G)-86000 - ,.Q .=.... a 4ooo - i e - -- /' 2000 - - - - � - ��-�::::==-�"""''i::·::c.L"•.. / . 0 - 0 19�-���--�-:�--��--� 19 19 19 19 19 19 19 19 19 19 19 20 19 19 19 19 19 19 19 19 19 19 19 19 20 40 45 50 55 60 65 70 75 80 85 90 95 00 40 45 50 55 60 65 70 75 80 85 90 95 00

Year Year la. Number of reindeer in the winter population during most of the 20th century within the Saami villages Saarivuoma, Lainio­ Fig. vuoma, Sorkaitum, Mellanbyn, Mittadalen and Idre new sameby (County Administration in Norrbotten and Jamtland). Short, dotted line indicates the maximum permitted herd size in Mittadalen and ldre (Svensk rennliring 1999). - (Sameby: Saami village)

availability of grazing. Statistics are unreliable, owing to Herd size over time deficiencies in reports submitted to the Boardof Agricul­ 1.7 ture. In 1993/94, the reports stated that 98,300 reindeer The genus Rangifer, whether wild, tame or allowed to run were slaughtered, of which 39% were calves and 61% were wild, is present within large parts of the northern hemi­ adults. A comparison with the report from 1998/99 shows sphere's boreal and arctic regions, where it is exposed to that slaughtering had been halved during the intervening different influences imposed either by nature or by man. period; 49,092 animals were slaughtered, of which 60% Diseases and lack of forage, together with predation, have were calves and 40% were adults. The slaughter of calves long been regarded as important factors that limit the size had thus increased by 21% (Svensk rennaring, 1999). of animal populations (Fowler 1987, Andersson 1982). However, in the arctic world the islands often lack predators and the reindeer herds are regulated by access to grazing, weather fluctuations and, to some extent, hunting (Vibe 1967, 1982, White et al. 1981). In mainland areas, the populations are additionally exposed to slaughter

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 31

Fig. XVIII. In Sakha (Yakutia), the reindeer is still used for riding and draught purposes more or less throughout the year. Photo: 0. Eriksson september 1973 or hunting of varying intensity (Syroechkovskii 1984, Tj aerga, wind crust, hard wind-packed snow. Bergerud 1980, 1983). Reduced grazing land, forestry Ts uongo, thaw crust, a thick layer of thawed and later frozen and forest fires are other influencing factors (Eriksson packed snow. & Snow depths of more than 80 cm. Raunistola, 1990, 1993, Eriksson et al. 1987, Eriksson 0. (The 1909 Reindeer Grazing Commission 1912, Ruong 1964, 1979, 1980, 1984a, Klein 1982, Miller 1976, Uggla 1958). N. Skum, pers. comm., Eriksson 1976) In this way, the reindeer populations that are exposed to 0. predation commonly loose 50% or more of their calves. As can be seen from Figure lA, the number of reindeer Abandonment of calves and physiological or medical has fluctuated widely within 'our' research Saami villages disorders have a lesser influence on numbers (Eloranta during the 20th century. The explanations are probably the & Nierninen 1986, Nordkvist 1980, Bergerud 1971). For same as those that governed the variations in the national the entire Swedish reindeer herd, the variation in numbers population. The neighbouring villages Lainiovuoma and during the past century or so has been about 25%, around Saarivuoma had similar reindeer population peaks around a mean level of about 225,000 reindeer in the winter 1935 and 1955, after which the number of reindeer in population. The winter population refers to the number Saarivuoma decreased to about 6,000 during the 1970s, of animals following the annual cull, but before calving before increasing again to about 12,000 during the 1980s. (Svensk rennaring 1999). Lainiovuoma had a very uniform number of reindeer dur­ The variation is cyclic, with peaks about every 25 ing the period 1960-1990, about 10,000 head. During the years. The dips in the 1920s, 1940s, 1960s and 1970s 1990s, both villages suffered severe reindeer losses on were mainly caused by long winter periods, with snow account of difficult winter-grazing conditions and poor conditions that hindered migrations without breaking up calving results. the herds, and which completely or partly prevented the Sorkaitum and Mellanbyn had the best figuresduring reindeer from reaching ground grazing. Examples of such the 20th century, with 8-10,000 reindeer around 1930, snow conditions are: 1955 and 1990, but with large decreases in-between and towards the end of the 20th century. Sorkaitum, in particu­ Tj uohki, ice cover after rain on cold open ground (Swedish lar, suffered severe losses, for the same reasons as those 'ften'). that affectedLainovuoma and Saarivuoma before the turn Skarta, thawed, packed snow cover directly on the ground, encapsulating the ground vegetation. of the century (Svensk renniiring 1999). Ts aevvi, an inner layer of snow inside the main snow cover at The Mittadalen Saami village had a uniform number temperatures around 0 °C, that prevents the reindeer from of reindeer up to the middle of the 1970s, around 3,000 smelling the presence of ground vegetation. head, after which the herd increased rapidly, and during

Acta Phytogeogr. Suec. 87 32 0. Erikssson et al.

Fig. XIX. Female reindeer (vajor) were milked to a limited extent until the 1980s. At an earlier stage, reindeer milk was important in the diet and in the domestic economy. Reindeer cheese is nutritious and can be stored, and is therefore an attractive commercial product. According to Ruong (1982), the protein content is 10.9%, the fat content 17.1% and the milk-sugar content 3.8%. The photograph was taken at Vivungi, in Vittangi forest Saami community, by Eriksson in Julyl968. 0.

Fig. XX. When the reindeer are gathered in summer in enclosures for marking and occasional milking, the irritation caused by biting insects is reduced with the help of smudge fires (Saamish suovva). Viikusjarvi, Vittangi forest Saami community. Photo: Eriksson, 0. July 1 969.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 33

the 1980s and mid- 1990s numbered around 9,000 animals. Tungus type: (East of Lake Baikal) Pack -carrying;Dra ught; Rid­ Subsequently, a decision by the village council led to a ing without stirrups; Milking; Herding with the help of dogs. considerable reduction. Idre new Saami village was reorganized in 1984, after a Koryak type: (North-Easterntype) Draught; Hunting with the help of decoy reindeer; Herding without dogs. long depression during which only sporadic reindeer counts were made. The new village's reindeer numbers decreased Sayan type: (West ofLake Baikal) Pack-carrying; Riding with sharply during the 1980s, and during the 1990s thenumber saddle and stirrups;Milking; No hunting with the help of decoys; was relatively even (around 3,000), which corresponds to No herding with dogs. the maximum number permitted. Riding in the strict sense did not occur among the Saami, but children and persons enfeebled by age could occasionally be carried on reindeer-back during a migration 1.8 Reindeer domestication and (Ruong 1982). Milking has been practised sporadically by handling most reindeer-herding peoples (op. cit.). The systematic use of milk occurred only within the Saami-, Tungus- and The utilisation of the genus Rangifer in a more or less Sayan-type of reindeer husbandry. For example in the Vit­ domesticated form is originally associated with the Old tangi Saami village, parish ofKiruna, reindeer were milked World. This includes attempts during the 19th and 20th until the 1980s, to maintain the tradition (L. Berggren, pers. centuries to introduce reindeer husbandry as an alterna­ comm.). The employment of tame reindeer as decoys in tive occupation for native peoples in North America and the hunting of wild reindeer is a method used by the three on Greenland (Skjenneberg & Slagsvold 1968). Marco northerly types, Saami, Samoyed and the north-eastern Polo (1967) mentions a tribe dwelling in the Lake Baikal (Koryak) type (Vainshtein 1980). region during the latter part of the 13th century, which The origin and development of methods of reindeer owned reindeer and utilised them as riding animals. Laufer husbandry and equipment follow several paths of develop­ ( 1917) discovered, in still older Chinese annals, notices ment or combinations of such paths. Examples of these referring to reindeer husbandry on the northern boundary are innovation and development within a given population, of the Chinese empire. From a knowledge of linguistic knowledge transfer between reindeer-herding peoples and and literary sources, he dates the beginning of reindeer the copying of types of equipment, terminology, etc., from husbandryto the beginning of the present era, in the Baikal neighboring peoples who have domesticated other spe­ region. This evidence probably refers to the Soyots, hunter­ cies of animals. Examples of this are riding, draught- and gatherers on a tributary of the River Ye nisey in the Sayan transport- equipment, such as saddle and stirrups, bridle Mountains, who also carried on some reindeer husbandry. and pack-saddle from horse-based cultures, and sledges Their herds were small, and rarely comprised more than adapted for pulling by reindeer, adopted frompeoples who 50 animals, which, however, were large and readily tamed. use dogs as draught animals (Wiklund 1919). They were used as beasts of burden, for riding, as decoys The reasons for the adoption of reindeer-herding and for milking, but rarely kept for slaughter (Wiklund within peoples originally dependent on hunting and fish­ 1919). Vainshtein (1980) is of the opinion that the reindeer ing have been the subject of lively discussion during the has been domesticated in Siberia since at least the last past century. During the first half of the century, a range of millennium B. C. hypotheses was presented by ethnologists and philologists. In the course of time, a number of peoples within the Lundmark (1982) commented on the shortage of reliable distribution area of the reindeer in Eurasia have developed source material for these hypotheses. Since the mid- 1950s, various forms of domestication, which have led to various research has been placed on a considerably firmer founda­ degrees of tameness.If meat production is the sole aim, then tion fo llowing Tegengren's (1952) and Hultblad's (1968) naturally a lower degree of tameness is required (but docil­ thorough analysis of the historical source material from ity at the level of the herd), compared with what is required Finnish and Swedish Lapland, respectively. Zackrisson of reindeer used as riding or draught animals. Vainshtein ( 1976) was responsible for the introduction of archaeologi­ ( 1980) illustrated, on the basis of significant cultural traits, cal source material into the discussion. In recent research areas of utilisation among reindeer-herding peoples (here into the introduction of reindeer husbandry, the ecological reproduced in a somewhat simplified form): context has been studied by means of palaeoecological methods (Aronsson 1991 ). According to Fj ellheim (2004), the shift in the natural economy of the Saami may have Saami type: Pack-carrying; Draught; Milking; Herding with the been brought about by environmental factors, originating help of dogs; Hunting with the help of decoy reindeer. in climatic changes in the 16th and 17th centuries, which Samoyed type:(W. Siberia) Draught; Hunting with the help of made it advantageous to change from a hunting-based decoy reindeer; Herding with the help of dogs; No milking. economy to nomadic reindeer husbandry. Ingold (1980),

Acta Phytogeogr. Suec. 87 34 0. Erikssson et al.

Lundmark (1998) and others consider, however, the tran­ between the summer and winter grazing areas are often of sition to be a product of changes in human society. Thus considerable length (op. cit.). the introduction, and subsequent expansion, of reindeer NorthernSaamis -Southem Saamis: These names are principally husbandry might be explained by a process of ecologi­ based on linguistic differences. A long history of contact has also given rise to some cultural variation, which should not, cal adaptation and economic specialisation. In the inland however, be regarded as a sharply defined cultural bound­ areas and the mountains, reindeer husbandry, combined (op.cit.). The valley of the Lule river may be seen as ary with hunting and fishing, was the natural development, an approximate boundary, since Saamis living north of it while agriculture advanced in the coastal areas of northern principally were in contact with a settled population who Sweden (Aronsson 1995). spoke a Finnish dialect (Mienkielin), while those who lived The earliest mention of the reindeer as a domestic ani­ south of it had their externalcontacts with Swedish speakers mal in Scandinavia can be found in the narrationof Ottar, (Marainen 1984). from the latter part of the 9th century (Alfred's Orosius). Semi-nomadism: A special and original form of small-scale Ottar told of the ownership of decoy reindeer and of a herd reindeer husbandry, combined with hunting, fishingand some wage labour, which has been common throughout northern of 600 tame reindeer. Some scholars, e.g. Manker (1947) Eurasia. Migrations are usually of limited extent (Ruong and Ruong (1982), consider that some form of reindeer 1982, Aronsson 1991, 1995). husbandry already existed during the late Iron Age, whereas Reindeer nomadism: Herds of tame reindeer are the basisof the others, e.g. Steckzen (1964) and Selinge (1982), would date economy. Migrations between summer and winter grazing its origin considerably later. Lundmark ( 1982) accepts the areas are often of considerable length (Lundmark 1982, possibility that small herds of reindeer may have been used Rheen 1983). for hunting, as decoys and as beasts of burden, even before Intensive reindeer husbandry: Indicates a multi-facetted utilisa­ the introduction of direct domestication during the 17th tion of the reindeer and reindeer products: reindeer are used century. This view appears to be supported by Zackrisson as draught- and pack-animals, for milking, as a source of meat and hides, and for selective slaughter (Paine 1972). (1984). Elucidation of the distinction between reindeer Extensive reindeer husbandry: This implies a tendency towards nomadism and a Saami hunting culture is important to the sale of animals for slaughter (op. cit.). the understanding of reindeer husbandry in earlier times. By the term 'reindeer nomadism' it is to be understood that the herd of tame reindeer is the basis of the economy. In general, it may be said that intensive reindeer hus­ This leads to a way of life in which the Saamis follow bandry is aimed at a natural economy, whereas extensive their herds throughout the year, and live mainly on them. reindeer husbandry leads to a monetary economy. The In the hunter society which preceded reindeer nomadism, meaning of the terms 'intensive' and 'extensive' fluctu­ there were also reindeer, but these had a subsidiary func­ ates somewhat, however, depending on whether they tion within an economy based on hunting, fishing and the are associated with the herding of reindeer in contrast to gathering of food plants (Lundmark 1982). Hunter-fisher reindeer husbandry as a business. When associated with peoples have lived in northern Scandinavia for thousands herding, they imply a varying degree of tameness, and of years-the earliest traces of dwelling sites in the inland when associated with business, they tend to indicate dif­ areas date from7000-6000 B.C. (Aronsson 1995, Kuoljok ferent degrees of utilisation. Among both forest Saamis 1998). The Saami identity developed among them from and mountain Saamis there exist both reindeer nomads and the 6th century A.D.,. when 'external' competition from semi-nomads, with a varying length of migration, as well other peoples increased (op. cit.). as hunters, fishers and other occupational groups. They can also be northern Saamis or southern Saamis. Within the Saami village, reindeer husbandry can be intensive Saami and reindeer husbandry - some definitions in some respects and in others, extensive, e.g. as regards The Saami culture exhibits a wide range of variation in time migration, herding and provision of fodder. and space. A number of terms has been used to describe these variations. Not all such terms are unequivocal, hence Husbandry of forest reindeer some lack of clarity, and misunderstanding, have arisen (cf.Lundmark 1985, Aronsson 1991 and Skold 1992). To The entire annual cycle is passed in the taiga. During the characterise their adaptation to the environment, the fol­ snow-free period, areas which provide fresh forage adjacent lowing terms are used here: to wetlands are grazed. When there is a snow cover, areas of coniferous forest which provide ground- and arboreal lichens as well low brush and a few graminoids, are grazed. Forest Saamis: (earlier called Spruce Lapps, Forest Lapps). These Originally, the reindeer herds were small, but were kept make their living in the taiga throughout the year (Ruong together. Herds were moved over fairly short distances 1982, Aronsson 1995). Mountain Saamis: (Mountain Lapps). These live above the co­ between milking areas with good grazing, and with some niferous forest limit for much of the year. Their migrations means of reducing disturbances caused by biting insects,

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 35

e.g. smudge fires, and-for preference-good fishing. In The Saamis and the authorities autumn and winter, herd movements were determined by One of the contacts with the outside world, which in time the availability of opportunities for hunting, and access to came to be burdensome to the Saamis, and strongly influ­ grazing. Within a family, the division of labour was such enced their way of life, was their tax liability, earlier men­ that the women were responsible from an early stage for tioned by Ottar and by the authors of the Icelandic sagas dealing with the animals, while the men were occupied with (Ruong 1982). The boundaries between nation-states were hunting and trapping. As the number of reindeer increased however, the men became increasingly involved with th not as yet delimited, hence the Saamis, at worst, could face � demands from tax collectors-doubling as traders -from reindeer (Ruong 1944, Marklund 2004). The transition to the chieftainly domains in northern Norway, from Finn­ the reindeer nomadism of the forest Saamis (with hunting ish and Russian areas, and from Sweden. Novgorod was and trappingas more or less important adjuncts) continued long the most prominent power in upper Norrland, and throughout the 18th century. At the turn of the 18/ 19th cen­ based its tax-collection and its trading system on a series turies, a clear shift was evident (Marklund 2004). Ruong of intermediaries. Local elites acted as an extension of the ( 1944) considers, however, that the forest Saamis at that authority ofNovgorod in the area around the White Sea and stage did not strategically choose major change, but that the Gulfs of Bothnia and Finland during the 13th century they attempted to maintain balance withinthe system. As an (Lundmark 1998). Furs, prized throughout Europe, hides example, it may be mentioned that a 'big reindeer owner' in the Arvidsjaur Saami village owned 60-80 animals at and hide-based products were both the most important trade items and the basis for taxation from an early stage. the end of the 18th century (Marklund2004 ). The source material concerning the earliest Swedish presence in northernmost Fennoscandia is meagre, and is Mountain reindeer husbandry based on hunting and trade expeditions, which long trans­ gressed present-day national boundaries. essence, the The migration patternof the reindeer is followed. Between In land was considered to be 'waste' before the 14th century, spring and autumn, the mountain grazings are utilised, from but was sporadically crossed by Saamis, who had been the tree limit (Saamish: 'orda') upwards to the mountain exposed to military incursions by peoples 'from the east'. heaths. The winter is normally passed in the lichen-bearing After peace with Novgorod was declared at Noteborg in coniferous forests. 1323, the Swedish Crown aimed at securing its suzerainty As early as the 16th century, the Saamis were divided over what had now become Swedish-Finnish territory- and into 'spruce Lapps', i.e. forest Saamis, and 'mountain naturally to benefit from whatever taxes that could be de­ Lapps ', who mainly lived in the mountain areas. The lat­ manded. The Novgorod system of tax collection having ter lived on reindeer products to a greater extent than the been seen to be inefficient, it was decided to establish a former. Hunting and fishing were of lesser importance to new system. This was admittedly based on a body of tax them (Rheen 1983). Developments during the 17th century, collectors- 'birkarls' -who earlier had served Novgorod, towards full-blown nomadism, resulted in their migration but who now derived their authority from, and were di­ areas having a narrowly elongated shape, approximately rectly responsible to, the Swedish Crown. The origin and orientated in an east-west direction, and adapted to the designation of these 'birkarls' has long been the subject of migration pattern of the reindeer and to the course of the lively debate by many scholars. One theory was that they rivers which formed the boundaries of the areas (Beach originated in Pirkala parish near Tampere in Finland. 1981, Ruong 1982). Their herds also came to be consider­ An attempt was even made to equate the word 'birkarl' with the ably larger than those of the forest Saamis. Rheen ( 1983), name of the settlement Birka, and with animals-beaver, then priest at J okkmokk, wrote regarding tame reindeer in 'bjur'-which provided a highly regarded The 'birkarls' the 17th century that 'Tame reindeer are they that increase fur. are first mentioned in a document dated 1328, in which it is the Lapps' reindeer, of which many Lapps own 100, indeed stipulated that their travel and trading activities should not 1 ,000, and some more ... the Lapps own many more female be disturbed, and neither should the hunting activities of than male animals, so that the Lapp who owns 100 females the Saamis. The 'birkarls' are described in the document as has no more than 20 or 30 males, since it is through the a group of wealthy farmers residing in the coastal areas of females that their herds increase ...' . the parishes of Pite,Lule, Tome and Kalix, who possessed the privilege oftrading with the Saamis and of taxing them, Good grazing-poor grazing subject to the payment of fees to the Crown. This system This classification can include many usable forage plants appears to have operated with little change up to the middle of the 16thcentury. Its main beneficiarieswere undoubtedly within a given seasonal grazing area. It can also cover e 'birkarls', since the annual fee to the Crown was only accessibility, which usually is determined by snow con­ � e1ght marten pelts and 1280 squirrel pelts. Before 1528, the ditions- snow depth, density, hardness, etc. 'birkarls' paid only half that fee for their privileges.

Acta Phytogeogr. Suec. 87 36 0. Erikssson et al.

King Gustavus Vasa laboured untiringly to increase the Saami household was to contribute two reindeer and eight revenues of the Crown. Among such efforts were succes­ 'lispund' of dried fish by way of tax (1 'lispund' equals 6.8 sive increases in tax liability. Even the Saamis were liable kg; 8 'lispund' corresponded to ea. 150 kg live weight). In to pay tax by families. In 1553, individual tax assessments addition, the Crown was to have a tithe of reindeer calves were introduced in the province of Vasterbotten, i.e. over and a tithe of dried fish. Such a tax demand, framed in terms more or less the entire extent of the current provinces of of foodstuffs, was a serious blow to the Saami economy. N orrbotten and Vasterbotten. In Pite-, Lule- and Tome The reindeer demanded by the Crown as tax were initially Lappmarks, 301 'tax Lapps', i.e. Saami families, were retained in the Saami areas, with the help of hired Saamis registered for taxation, at the same time as they were taxed as herders. In 1610, the Crown's reindeer, which were at values which in total far exceeded the fees previously carefully recorded by the tax-collectors, amounted to ea. paid to the Crown by the 'birkarls'. 1,000 in Lule Lappmark alone. They were successively The new realisation of the tax-paying capacity of the slaughtered and the meat was sent to Stockholm. King Saami areas led to major changes. The 'birkarls' lost their Charles IX showed a close interest in reindeer husbandry. right of taxation, but on payment of a fee, were confirmed Large herds of Crown reindeer in the Saami areas were in their old right to trade with the Saamis. After some time, one of his visions for increasing the exploitation of the one 'birkarl' in each Saami area became a tax-collector resources of the Saami territory. Attempts to farm reindeer for the Crown, and was given detailed instructions to ex­ in the province of Medelpad were a signal failure. tract, with the help of the taxation rolls, the tax due from The new tax led to a crisis in the economy, since the 'his' tax-liable Saamis, and to deliver an account and tax Saami population had already outgrown the limits of the revenues to the Treasury in Stockholm. trapper society. The wretched condition of the Saamis is The tax levied on each family was, as a rule, one or documented in reports from the tax-collectors and in peti­ two bundles of furs (squirrel pelts; one bundle equals 80 tions to the King from, among others, priests active in the pelts), or one or two marten pelts, and possibly, a quan­ Saami territories. The tax revenues decreased successively, tity of dried fish. In other words, not an especially heavy and in 1620 it was necessary to halve the tax, for lack of a burden. The steady increase in the number of Saamis li­ sufficient tax base. After a few years it was further reduced, able for tax indicates a degree of prosperity towards the and lay at the level of two 'lispund' of dried fish or one beginning of the 17th century. Trade with the Crown and thaler or two pairs of shoes of tanned hide. with the 'birkarls' provided an influx of goods, money The supply crisis led quite rapidly to innovations among and precious metals (mainly silver). The transactions took the forest Saamis. When the old economy, based on hunting place in connection with winter fairs, which were held at and trapping, no longer sufficed,there wasa gradual transi­ the winter dwelling-places at a time of year when, owing to tion to a strategy based on a greater year-round utilisation the weather, hunting and trapping were less profitable. The of the reindeer. This was undoubtedly reinforced by the addition of purchased flourand butter introduced agricul­ other products that nature could provide, such as fish, game tural produce to the diet, and tided over a period of dietary and vegetables. It is here possible to speak of a gradual insufficiency, which could have fatal consequences. transition from a trapper society to a semi-nomadic way Trapping, hunting and the initially small reindeer herds of life. Forms of contact between Saami society and the had earlier sufficed for the needs of the sparse population. outside world, other than the purely fiscal,may also have Increased population size, increased trade and the intro­ contributedto steering development away from a hunter­ duction of goods produced outside the Saami areas, led fisher culture, via various intermediate forms, to reindeer to difficulties when the supply of game and fish began to nomadism and present-day reindeer husbandry. decrease as a result of increased pressure on the natural From 1500 onwards there is evidence for the division resource base, and possibly also in consequence of climatic of the Saami territories into very large community hold­ deterioration. ings, sijdas, which were more or less circular in shape, The fiscal demands of the Crown could be met during and adapted to the trapper society's utilisation of its re­ the 16th century, so long as they primarily consisted of sources. The holding comprised the fishing-waters, hunt­ furs, which in any case were little used in Saami society, ing-grounds, etc., required for supplying the community's where reindeer hides were more important. At the begin­ needs throughout the year. This division had probably ning of the 17th century, the Crown abruptly changed its arisen far earlier than the 16th century. Within the holding, taxation policy. The reason for this was that Sweden was which could comprise about one hundred members, smaller at war with Poland, and that the campaign of 1601 had areas were allotted to individual families. Each family pos­ been a failure as a result of the lack of food supplies. In sessed, as a rule, several tracts of land, between which it March 1602, King Charles IX decreed a new tax on the migrated as necessitated by the seasonal food supply. Saamis, which was to be paid in reindeer and in increased Within the sijda there was originally a single, common quotas of dried fish. After some transitional difficulties, dwelling site, dalvvesijd, to which the families returned a new tax ordinance came into force from 1607. Every in autumn when communal hunts for wild reindeer and

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 37

other labour-intensive activities took place, and which was of Saami extraction (and incidentally the only Saami were the prerequisite for winter survival. Mid-winter was ever ennobled), strove hard with the help of his contacts a quieter period, which provided opportunities for social in the government to improve the situation of the rein­ activity. Matters of common interest were takenup by the deer drivers. At the start, his efforts were fruitless. The sijda council, which was made up of the heads of families Saamis fledeven from Lule Lappmark, especially after a and functioned as a legal body. Judgements were given, new ore body was discoved at Alkavare, near Kedkevare, disputes were ventilated, land was re-allocated in accord­ and an increased need for transport was therefore to be ance with changes in the needs of families or because new expected. The number of Saamis registered as resident families had been formed. The council also dealt with in Lule Lappmark declined during the mid-17th century, external contacts, with other sijdas, with traders and tax from almost 200 to 65. officials, who before the 16th century could originate from At the beginning of the 18th century, the mines at Ked­ several sources. The natural boundaries of the sijdas were kevare and Alkavare were almost worked out. The Saamis well known to all concerned. had returned, and the population of Lule Lappmark was The changed tax policies of the Crown also affected once more at its 1650 level. To ease pressure on the Saamis the sijda system. In 1602, King Charles IX decided that from mines and smelting-works with a large demand for the Saamis should attend fairs at fixed sites every winter. transportation, Johan Grahn suggested that the Saami ter­ There they would pay tax, and when churches had been ritories should be opened to colonisation by pioneers from built, would also attend services. The administration of an agricultural background, and who were familiar with justice was also associated with the fairs. Traders were the use of the horse as a draught animal. This was done prohibited from doing business at the old dalvvesijdas. In out of consideration for the Saamis-he did not foresee its consequences. 1611, King Charles IX died, and with him died the interest of the Swedish Crown in the intensive exploitation of the Saamis for tax purposes. The colonisation of the Saami territories

As was mentioned in the Introduction, the utilisation of the The silver-mining era and the Saamis Saami territoriesfor agriculture is, in an historical perspec­ In 1634, argentiferous ore was discovered on Nasafjall, tive, a very recent event. From the 16th century onwards, which lies near the Norwegian border in the present-day the Crown's interest in establishing a settled, agricultural Arjeplog municipality. In Stockholm, where knowledge population in the 'waste areas' increased; this was a pre­ of the riches seized by the Spaniards in the New World, requisite for the assertion of territorial rights over the land, including silver, was widespread, it was supposed that which increased in importance with the discovery of silver Nasafjall was Sweden's own new Eldorado, which must be deposits in Pite and Lule Lappmark. The inclination among exploited as quickly as possible. As has already been noted, the population in general to colonise the Northlands was, Sweden's many wars had strained the Crown's finances. however, limited. It increased when, in 1673, the Crown The collector of Saami tax in the Pite district was made promulgated the 'Lappmarksplakatet', which provided for responsible for ensuring, with the help of hired Saamis and benefits coupled to settlement in the Saami territories; tax their pack- and draught-reindeer, that ore was transported exemption for 15 years and relief from military service, from the mine to the smelting-works at Silbojokk at lake those which probably were of greatest importance. Sadvajaure, and thence to the coast at Pitea, a roadless Colonisation increased successively, and with it the fre­ distance of ea. 400 km. In compensation for this, a tax quency of disputes between the Saami and the colonisers, reduction and other appropriate benefits were to be offered. principally regarding the right to fisheries. These disputes However, it proved difficult to recruit a sufficientnumber were often ventilated in court, where at the start the Saami of reindeer drivers, which led to an unsuccessful attempt to side of the argument was successful, since the assessors solve the transport problem by coercion. As a last resort, were usually Saami, who were aware of the problems of the Saamis assigned to transport duties reacted by fleeing obtaining a livelihood. Cases brought by Saami plaintiffs from Pite Lappmark over the Norwegian border, or down often involved unlawful entry into fisheries. The colonis­ to the coast of the Gulf of Bothnia. The mine ran at a loss ers made claims concerning damage to hay by passing for a number of years, until Danish troops invaded from reindeer herds, the hay having been left unprotected on the Norwegian side of the border and burnt down the mine wooden hay-drying racks, to be brought in later when a buildings, which were never afterwards rebuilt. snow-cover facilitated transport. The damage consisted The Crown had discovered a new source of silver in in the overturning of hay-drying racks, and trampling and the mountain Kedkevare, north-west of Kvikkjokk. Min­ fouling of the hay when the reindeer attempted to reach for­ ing began during the 17th century, and in this case also, age plants, such as Equisetum spp., Menyanthes trifoliata the Saamis were to be responsible for transporting the ore. and Potentilla palustris, which were mixed with the hay. The governor ofVasterbotten, Johan Grahn, who himself

Acta Phytogeogr. Suec. 87 38 0. Erikssson et al.

The boundary decision affecting Denmark­ Regulations concerning the Saami territories 1751 Norway and Sweden-Finland By the mid-18th century, the conflict of interests had reached such proportions that the Crown, in 1749, promul­ In 1738, a boundary commission began work on defin­ gated a set of regulations applicable to the Saami territories, ing a national boundary between Denmark-Norway and which aimed at defining their utilisation. The colonisers Sweden-Finland. The boundary was mainly to fo llow were primarily required to carry on cultivation and animal the watershed along the Scandes. The local populations, husbandry. They were given limited hunting rights within including the Saamis, were given an opportunity of present­ an approximate 5-km radius from the smallholding. Fish­ ing their views on the proposed alignment of the boundary. ing was permitted within the same area, including waters In Finnmark, in the farthest north, Sweden withdrew its over which the Saamis had rights. The Saamis themselves claims to the parishes of Kautokeino and Karasjokk, but were required to confinetheir hunting, fishingand reindeer received in compensation the parishes of ldre and Sarna husbandry to the areas subject to taxation. in the province of Dalama, which formerly had belonged Up to the mid-18th century, the large-scale reindeer to Denmark (Ruong 1982). nomadism that had begun during the 17th century, had The border divided the Saami lands into a Danish and developed into a profitable occupation, which gave the a Swedish sphere of interest. In the Saami context, there Saamis trading advatages over the still limited settled popu­ was initially no real border, and it was therefore of little lation. From the latter part of the 18th century onwards, importance. The mountain range (the Scandes) was already decisive changes took place; colonists arrived in increasing divided into different sijdas for late spring grazing and numbers, and the proportion of Saamis in the population early autumn grazing. The sijda areas extended across the of the inland and mountain areas declined. The importance Scandes, defined by the alignment of the Swedish valley of reindeer husbandry to the economy also declined in step systems (Marainen 1984 ). with the increased importance of other forms of enterprise. Two codicils were appended to the boundary treaty, The situation of the Saamis now deteriorated markedly, which was signed in Stromstad in 1751 by representatives mainly in consequence of the measures taken by the au­ of both kingdoms. In one codicil (Lapp-codicil of 1751 ), thorities. For example, the Saamis' chances of succeeding the rights and obligations of the migratory Saamis in both in legal disputes with colonisers were sharply curtailed. kingdoms were defined. In spring and autumn, the Saamis, Within Saami society itself, several negative changes oc­ in accordance with ancient custom, were permitted to move curred. To begin with, the reindeer herds suffered severe with their reindeer from one kingdom to the other. Without losses. From the mid-18th to the beginning of the 19th regard to the border, they were permitted to utilise both century, years of dearth in reindeer husbandry repeatedly land and shore for the sojourn of both people and animals. occurred. In Jokkmokk parish, for example, two-thirds of They were to be received amicably, and protected even if the reindeer herds perished in 1756. Even the years 1786, a state of war existed. They were to pay tax only in the 1796, 1813-1816, 1830-1837 and 1843 appear to have kingdom in which they were registered as resident. been afflicted. As a result, the Saami population of Lule The codicils, with their humane spirit, are remarkable Lappmark decreased by half in the latter part of the 18th for their time and have been called the Saami Magna century and during the firsthalf of the 19th century. Charta. They constitute a kind of charter of rights for the The crisis in reindeer husbandry was probably caused Saamis in Sweden and Norway, which, however, has partly by a combination of factors, e.g. by a shortage of grazing been invalidated by subsequent historical events. due to weather conditions, by increased losses of animals to predators, and by disease among the reindeer. There The boundary of the Saami territories may also have been an imbalance between the number of reindeer and current methods of husbandry, whereby Afterthe mid-1 8thcentury, the Swedish Crown came to regard the reindeer herds were concentrated and the hinds were all land that was not subject to Saami taxation, as belonging milked on overgrazed pastures, with a consequent risk of to the state. The intention was to promote the establishment disease transmission. That the milking areas and grazings of colonists, even where the natural circumstances were not were heavily exploited, is illustrated by the fact that, fr om directly favourable. One means was to prevent the coastal the end of the 18th century, disputes between Saamis con­ farmers, who from mediaeval times had hunted andfished in cerning the areas liable for tax were increasingly decided the inland areas without much regard for thesijda, from com­ by the courts. peting withthe Saamis and colonists. In17 51, a boundarywas established, the 'Boundary of Saami Territories' which cor­ responded, by andlarge, with the boundary which at present separates the provinces of Norrbotten and Vasterbotten from Lapland. The land west of this boundary (i.e. Lapland) was to be utilised only by colonists and Saamis.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 39

Cultivation limit Information concerning reindeer forage plants was given by, among others, Samuel Rheen (1671), who During the 19th century, the pace of colonisation greatly for many years was a priest in the parish of Jokkmokk: increased, which was to the disadvantage of the Saamis, 'towards autumn, when the ground is snow-covered , the who lost rel.ndeer pasture in the form of wetland- and reindeer mainly seek the white moss (i.e. Cladina spp.), meadow-plants, which were mowed by the colonists as which is found both on the mountains and in the forest winter fodder. Access to fisheries was essential to the areas. The reindeer dig for this with their forefeet, by colonists, especially during the establishment phase, and throwing aside the snow and consuming the little they had always been essential to the Saamis. Intrusions into can find beneath it. ' fishing waters were consequently a severe blow. N. Lundius from Pite Lappmark, a student in Uppsala In 1873, the 'limit of cultivation' was set out. This was (son of the firstpriest of Saami birth, Andreas Petri Lundius, intended to separatecultivable land from the montane areas from 1640 minister in Arvidsjaur), related that 'During in the provinces of Norrbotten and Vasterbotten. West the period of snow, the reindeer subsist on a particular of this boundary, no colonisation was to take place, and kind of white moss (i.e. Cladina spp.), which grows in the Saamis were to have sole rights throughout the year. large quantities both on the mountains and in the forests East of the boundary, they were permitted to keep their throughout Lap land.' In the work 'Descrip tio Lapponiae' reindeer between October and April. The boundary never (1675), he relates that 'in Umea Lappmark, the Mountain gave the intended protection to the Saamis. Numerous Lapps travel to the fair on TwelfthNight (Epiphany). When colonists already existed within the limits of the area, and they pass through the ( tax)lands of the Spruce Lapps, they permission continued to be granted for new settlements. ask leave to pursue wild reindeer. Many a Spruce Lapp When the value of the natural resources in the mountain accomodates up to ten Mountain Lapps, who may stay areas (minerals, water-power and forest) became apparent, with their reindeer until Lady Day (Annunciation Day), large-scaleexploitation began, which was in manyrespects as there is more winter grazing on the lands of the Spruce to the detriment of Saami interests. Lapps than on their own. The reindeer do not graze on the moss in summer, but on green plants that we call 'misar grass', which grow in water; the root is thick, and above it Reindeer grazing and availability of grow three leaves, which branch off one from another (this 1.9 probably refers to Menyanthes trifoliata). In summer, both grazing, from the time of Olaus Magnus reindeer and Lapps willingly eat Angelica archangelica, which grows beside rapids.' (op. cit.) Also P.M. Martiniere, a French physician, who trav­ Reports from pioneers in Lapland elled in Lapland in the mid-17th century, gives an account Earlytravellers or residents (usually clergy) in the reindeer­ (Schefferus 1673) of the grazing by reindeer of lichens, ' grazing areas have left posterity few accounts of grazing ... which everywhere are found in vast quantities'. sites and of the availability of grazing. Olof Rudbeck the Younger was the first to undertake The earliest, scanty, information about reindeer graz­ a true research expedition to the northernmostparts of the ing during the period of snow cover may be that given kingdom. In the summer of 1695, he travelled as far as by Olaus Magnus ( 1555). 'The food of this animal (i.e. lake Tornetdisk. In his journal (Rudbeck 1695), he gives the reindeer), consists of the white lichens on the hills, some information about the choice of forage plants by especially in winter. ' reindeer at different seasons, and about their behaviour: Olaus Graan served for about 30 years as a priest in the 'where food is concerned, they eat not only 'moss', i.e. Pi tea town and rural parishes, and was well informed about lichens, but also the leaves of trees and shrubs that grow conditions in the Lapp ish areas.In his 'Relation' ( 1672), he in the mountains, and diverse herbs and grass, as well as gave an account of the reindeer's habit of straying widely various kinds of fungus, especially Lactarius species. It is during the autumn in search of the most attractive fungi. remarkable that, notwithstanding the abundance of green The difficultyof keeping the herds together at that time, and forage, they are not as fat in summer as they can be in of protecting them against predators, was mentioned. winter, when they graze on the white moss as long as the In 1673, Johannes Schefferuspublished his work 'Lap­ snow-depth and absence of an ice-crust permit. But when ponia', compiled on the orders of the Chancellor, Magnus they are unable to reach the ground-forage, they turntheir Gabriel de la Gardie. This was intended to describe both attention to the grey and black moss that hangs like a beard natural conditions in Lapland, and the inhabitants' way of from tree-branches, and which is called lichen. If reindeer life, and was to focus on the value of the region to society. come upon human urine, they lick it up with a good ap­ On the orders of the Chancellor, information was provided petite. When reindeer calves wish to suckle, or become by clergy resident in Lapland, and by university students separated from their mothers, they call to them with a sound of Saami origin. that resembles that made by tame swine. Their hooves are

Acta Phytogeogr. Suec. 87 40 0. Erikssson et al.

of great importance to the ability of reindeer to move in a. This lichen is the most important of the plants, that are the terrain without losing their fo othold. If a reindeer is found in Lapland. startled, or otherwise moves of its own volition, it moves b. Within the plant kingdom, there is no other species by preference against the wind.' (op.cit.) that appears in such masses, especially in the forest areas, where the sterile, sandy and gravelly heaths are covered with sparse pine forest. In such areas one can pass through Linnaeus' travels in Dalarna and Lappland areas 20-30 km long, where the ground is white as snow In the summer of 1734, Linnaeus undertook his expedition and covered only by this lichen. to Dalarna. His travel report was not After the ravages of forest fire,the ground may lie quite (Iter Dalecarlicum) c. published in Swedish until 1889, and an annotated edition dry and bare, but where all other vegetation refuses to appeared in 1953. return, the reindeer lichens thrive abundantly, and after In western Dalarna he encountered lichen-covered six or more years attain their normal size. areas, both in the coniferous forest and on the mountains. d. . . . the reindeer exist during the summer mainly on Many of the observations concerning the utilisation of grass. Then they abandon for preference the warm, forested lichens are familiar from Flora Lapponica (1737). valleys and climb to the windy mountains, covered by As regards the region, the reindeer forage plants and perpetual snow, where they pass the warm midday hours their use, he recounts: 'In the area around Sama, the only on the snowy mountains . ... cereal sown is Hordeum vulgare (barley), which often is To be able to watch their animals, their owners too, the damaged by frost. The mainstay of life here is principally Lapps, pass the summer on the mountains. At the begin­ hunting, fishing and cattle-herding. No state taxes are paid ning of autumn, both Lapps and reindeer, the former by the here. The Lapps, who carry on reindeer husbandry, would severe cold, the latter by the lack of forage, are compelled be more profitableto the state in this parish'. to leave this abode of cold ... The farmers must each year collect large quantities e. Even the settlers (whose way of life much resembles of reindeer lichen as winter fodder for the cattle. Every that of our farmers) comprehend the benefits of using this husbandman collects 100-200 loads of reindeer lichen lichen. During rainy weather they scrape off with a many­ with the help of an iron rake, usually in rainy weather. He toothed rake this plant, which clings together and is readily takes home loads of lichen in winter, and feeds them to loosened from the ground. They gather it into heaps and the cattle instead of hay. This keeps them alive, but does use it during the winter, after it has been moistened with a not fatten them. The regeneration of the lichens takes little water, as an excellent fodder for their cattle. 20 years. Where, however, the reindeer has fed on good f. The reindeer lichen is exceedingly common in almost lichen areas, their regeneration takes only one year. The all Lap land's forested areas, but less abundant in the moun­ mountains are the Lapps' summer dwelling, and the forest tains. The mountains in Dalarna, by contrast, are almost their winter dwelling. Both, in other words, are utilised. completely covered by this lichen, Lichen rangiferinus By producing reindeer products, the Lapp would greatly ( =Cladina rangiferina ), Lichen niveus (= Cetraria nivalis), benefitthe province. The settlers in Lap land and the Lapps Empetrum and Betula nana, the round-leaved birch . could very well co-exist. In summer, the reindeer do not . . . On the heaths in Alvdalen there are reindeer lichens, harm the meadows, and in winter they need no hay. Any which in moist weather are collected as winter fodder for competition need only affect hunting and fishing ... It was the cattle. remarkable to see here that the top of the large, white rein­ ... The great quantities of Calluna vulgaris (ling) found deer lichen was so dry, that one needed only touch it with here, appear to be capable of collection and of feeding to the fingers, forit to fall into dust, but internally it was not the stock as fodder, as also is horse-dung, which in winter only soft and juicy, but also tough and moist ... One had is used as fodder. expected to see more new and unusual species here than ... South of Siirna, we entered a dense forest, which in the Lappish mountains, since the area lies so far to the on the ground was snow-white with lichen, but the southward, but one was sadly deceived. Barely one-tenth branches of the trees were quite black and softby reason of the species found in Lapland were found here, even of a black, beardlike lichen, which hung down from their though the mountains were white with lichens, such as lower parts . never were seen in Lapland. The mountains here were as . . . From Hostnadden and Langfjiillet,the distant Viister­ dry as those in Lapland were moist. Shed antlers of wild dalsfjiillenwere visible in the east. Farther to the west lay reindeer were visible on the fields.' In Flora Lapponica a large group of mountains, above which the twin peaks of (1737), Linnaeus gives information about the reindeer Sulufjiillet on the Norwegian side appeared. From (Stor­ lichens and their importance both to the Saarni, and to vattes-) Hagna many mountains were visible to the north the settlers. and northwest, of which Swucku was the foremost. The This is a selection of his information concerning the whole of this area was largely white, covered by reindeer reindeer's forage plants: lichens,. but the higher is the mountain, the less (sparser,

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 41

lower) also is the white lichen. concerningreindeer forage were somewhat sparing: 'The Judging by the accounts of journeys and scientific mountains on either side of the south end of lake Tornetdi.sk publications, interest in the quantitative aspects of reindeer were covered with reindeer lichen (Cladina rangiferina) forage plants was limited during most of the 19th century. and Stereocaulon paschale. The upper reaches of the Alta With some exceptions, the observations made were fairly river were so shallow, that travel by boat was impossible. fragmentary. Exceptions were those made by e.g. Wahlen­ The journey therefore continued on foot towards the berg ( 1804) and Hult ( 1881 ). It may be claimed, with some Kautokeino river, initially through forests of pine and justification, that Hult was the first to attempt to quantify birch. Later, after they had gained height, they traversed the plants that provide fo od for reindeer. a stretch of ea. 40 along a fairly level mountain ridge. km At the urging of the mining minister Baron S.G. Her­ Open, dry areas were covered by Cetraria nivalis. 'Those melin, during the summer of 1802 Goran Wahlenberg areas and hills covered by this plant, show up white even undertook a mainly botanical expedition in Kemi Lapp­ at a long distance. In these mountains, it is incomparably mark, within the parishes ofUtsjoki, Enare and Sodankyla. taller and thicker, and the separate lichen stands richer, than At that time, these parishes were part of the province of in more southerly areas. This lichen too is a sought -after Vasterbotten, and were occupied by the sijdas which food for the reindeer, and could doubtless be employed as subsequently invaded Karesuando and Jukkasjarvi. His a healthy food for humans.' ( op.cit.) report (Wahlenberg 1804) also contained information of From Kautokeino, it was possible to travel upstream a geographical and economic nature. by boat ea. 15 km along the Kautokeino river, which later He found Utsjoki parish to consist of a very broken became too shallow. The journeyto Karesuando continued mountainous area. 'The hilltops extend just above the birch on foot, to begin with over fairly level terrain covered with forest limit, and are covered with reindeer lichen to the sum­ Cladina rangiferina, Cetraria islandica, Stereocaulon mit. Especially at Mandujaure in the Utsjoki valley system, paschale and Empetrum hermaphroditum. the hillsides are entirely covered with reindeer lichens, Subsequently, the land became more boggy. From so that they appear quite white at a distance through the Karesuando, the return journey began by boat along the sparse birch forest. Utsjoki parish is one of the best lichen border river. areas in the north; these extend parallel with the eastern ide of the Scandes, and constitute the s regia subalpina Zetterstedt's journey in Ume Lappmark Lapponiae of botanical nomenclature. Utsjoki has for this reason always been one of the Lapp congregations richest In 1832, Zetterstedt made a round trip in Ume Lappmark, in reindeer. Almost all true Reindeer Lapps are Swedish up the Ume river across to the source of the Angerman citizens. During winter, which is the greater part of the river and downstream to Asele, thence via Lycksele to his year, they reside within Swedish territory, where they find home in Lund. The primary aim was to collect insects, but an abundance of reindeer moss. In this may be found the also birds. In the course of the journey, a compendious explanation of the relative liveliness of the winter season diary was kept, containing both Zetterstedt's own observa­ in the Lappish areas of Sweden. In Enare and Utsjoki, tions and those of a multitude of informants, concerning Wahlenberg found no noteworthy resources of reindeer soils, economy, living conditions, etc., within parts of the forage for year-round use, nor was there any noteworthy parishes of Lycksele Lappmark. In 1833, the travel report reindeer husbandry. ' (op. cit.) was published (Zetterstedt 1833), and as regards reindeer forage, was still less detailed than Zetterstedt (1822). A few observations were made on lichen resources: At Zetterstedt's journey in Torne Lappmark Gaskeluokta on the south side of lake Storuman, ea. 40 In the summer of 1881, the professor of zoology at the km northwest of Stensele, Cetraria islandica occurred University ofLund, Wilhelm Zetterstedt, travelled inTome sparsely, and epidendric lichens were common, especially Lappmark and adjacent areas of Norway and Finland. His on dead trees. Cladina spp., Cladonia spp. and Stereocau­ route mainly followed the Tome river and lake Tometrask, lon paschale were abundant, and epidendric lichens were the Norwegian coast in Trams and Finnmark counties and common, especially on dead trees. southwards via theAlta and Kautokeino rivers to Palojarvi. The vegetation at lake Voj msjo resembled that at Storu­ Thence he travelled along the Palojoki river to the Muonio man. Between lake Torvsjo and Asele, it was noted that and Tome rivers. extensive areas afforest had been burnt, although this was In the course of the journey,excursions were undertaken illegal-spruce forest to create good grazing, pine forest to to collect both scientific information and 'specimens'. In­ bum offthe lichens to the detriment of reindeer husbandry, formation concerningthe economy of the districts through which for reasons of competition was unwelcome in the which he passed was noted. His travel report (Zetterstedt area. (op. cit.) 1822) was published the following year. From what follows, it is evident that observations

Acta Phytogeogr. Suec. 87 42 0. Erikssson et al.

The Lapp bailiffs'yearbooks from year to year to use the lichen grazings in the lowlands ever earlier in autumn (Hultin, J. 1908). The new taxation law of 1695 stipulated that every 'Lapp J.O. Holm reported in 1917 that, at the beginning of village' should have its District Clerk, who, together with the 20th century, the Lapps in the villages north of lake the Lapp Bailiff, should allocate and receive the common Tometrask-Tome river had begun to exhibit an increas­ tax impost on the members of the community, as decided ingly marked unwillingness to migrate any great distance by the state (Beach 1981). During the 18th century, there southwards for the winter within their winter grazing was a large increase in the number of tax-Lapps and in areas. They remained by preference close to the line Juk­ taxland. The number of disputes concerning questions of kasjarvi-Karesuando. land ownership and disposition, which reached the courts, The consequence was that the areas west of the winter presided over by the Lapp Bailiffs,culminated during the grazings' uppermost parts were too heavily exploited, and 1750s. They provided the early administration by bailiffs became impoverished. In the winter grazings that had with valuable information about changes in the circum­ remained unused, perhaps for 20 years, the lichen cover stances of reindeer husbandry, which called for adjust­ had, however, reached a considerable thickness. The main ment of existing rules (op. cit.). From the end of the 19th reasons for the unwillingness to migrate fartherwere said to century until the mid-20th century, the archives contain a be business connecions with sedentaryvillagers, proximity number of the annual reports submitted by the Lapp bail­ to the town of Kiruna where provisions were purchased, iffs in Norrbotten and Jamtland to their respective county the schooling of the children in the villages of Jukkasjarvi administrative boards. The reports from Vasterbotten are, and Lannavaara, and the risk of controversies with farmers however, absent from the archives. The reports contain in 'the lower country'. The source of the disagreements scattered notices about grazing areas and their utilisation. was the damage that could be caused by the reindeer along Some examples: their migration route, to unprotected hayracks which often were left by farmers on the hay-mires for later collection The Lapp bailiffs in Norrbotten and transport over the snow. Farmers might also wish to protect lichen areas close to the villages, since they needed In 1892, Forsstrom reported that it was possible that there it as fo dder both for their own draught reindeer and for F. had been more reindeer in Arjeplog in former times than at domestic livestock during severe winters. To maintain the time of writing, but that a reduced availability of lichen controlover the reindeer and to avoid damage, was almost grazing areas had reduced the number of reindeer. Itseemed impossible, especially during difficultgrazing conditions indisputable that the reindeer lichens in Arjeplog had been when the reindeer became scattered over wide areas owing most evenly distributed and grown most abundantly within to a lack of forage. the mountain area, in a tract ea. 40 km wide eastwards from the Swedish-Norwegian border. Spring, summer and Lapp bailiffs in Jamtland autumn grazing, and in former times certainly even winter grazing, must have resulted in the near-exhaustion of the In the report for 1899, the Lapp bailiff Axel Franden reindeer lichens by grazing and trampling. reported to the county administration in Jiimtland that In the 1907-1908 report, he told that the Lapps in Juk­ ten to twelve unfavourable years for reindeer husbandry, kasjarvi and Enontekis could, as a rule, over-winter west of withflen, bodni-vihki or both (Eriksson 1976), which had Jukkasjarvichurch village and 50 km west of Karesuando hindered access to reindeer lichens from under the snow, during the 1830s, because the number of reindeer was so had caused large numbers of reindeer to die of hunger or to small that the lichen grazings in the mountains sufficed. In become dispersed in all directions while seeking accessible 1892, the situation was such that the majority of reindeer forage. Many of these unsupervised, stray reindeer had were compelled to move to the winter grazings far down in fallen victim to predators. Among the Saami,it was claimed Pajala parish. longA period of spring and autumn grazing that a few wolves could kill as many as 30 reindeer in a in a 40-km broad zone east of the border had, in addition, night. The number of reindeer, which at the end of 1890, caused the grazing areas to be so heavily grazed down and possibly with some exaggeration, had been estimated at trampled, that scarcely a trace of reindeer lichen ( Cladina 60-70,000, and at the beginning of 1896 at 27,000, was spp.) could be seen (op. cit.). estimated to be 12,000 in 1899. In the report for 1907-1908, J. Hultin reported that in Tourists, who appeared in increasing numbers each recent years, the Lapps in Karesuando, Jukkasjarvi and a year, were criticised for having contributed in their own large proportion of Gallivare parish had begun to move way to the decline of reindeer husbandry, by luring the the reindeer herds far too early from the summer grazings Lapps away from their nomadic life to an existence as down to the autumn lichen grazings, in consequence of porters and guides. which the lichens were both grazed and trampled down. An important cause of the decline in the reindeer stocks As a result of this abuse, the Lapps would be compelled was to be found in the decrease or even disappearance of

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 43

reindeer lichen in the Mountain Lapp villages, with the the entire family comes to dwell throughout the year, and exception of Tannas and Idre, where there still was good where the children in particular learn habits that make access to lichens. The blame for the decline in the lichen them idle, and incapable of the nomadic life, and of be­ carpets was ascribed to the difficult snow conditions during coming true Reindeer Lapps; (3) the transformation of a series of years, which were said to have affected lichen reindeer husbandry to comprise solely meat production, growth negatively; to trampling during the snow-free pe­ whereby the Lapps' contact with the reindeer is reduced riodby free-roaming reindeer flocks;and to an abundance to a minimum.' of small rodents, which were claimed to destroy 'the root The report for 1933-34 was written by Waldemar system'. Complaints were also made regarding grazing Gardham (1935). As to the number of reindeer, he writes, competition by Saami, who recently had become settlers, among other remarks: 'On the basis of the stocklists and who themselves kept domestic livestock as well as (24,370 animals) and the 1934 reindeer inventory (29,465 providing grazing during the summer for about 200 of animals), the reindeer stock can be estimated at between the farmers' horses. The horses were permitted to stray 25,000 and 30,000 animals. On the one hand, this number freely on the mountain heaths, with trampling damage as is greater than the Lapps' lichen grazings in the mountains a consequence. In thereport for 1900, Franden reiterated can stand in the long term, and on the other hand, it is the fear that reindeer husbandry would collapse if, as in more than is compatible with ordered, friction-free and the previous decade, the reindeer herds were permitted to economic reindeer husbandry. ' Most of the Lapp bailiffs roam freely on the mountains and in the submontane areas, undertook voluntarily to reduce the reindeer numbers to trampling andcrushing the lichens which, on warm summer the level set by the Crown. days, were thin and fragile. For advice, Franden wrote to his colleagues, the bailiffs in Norrbotten and Vasterbot­ The Saami view of the grazing situation ten. The bailiff in Norrbotten, J. Hultin, considered that the number of reindeer had periodically been too great, so Johan Olofsson Turi was a Saami, who was born near that the annual growth of the lichens had been less than the Kautokeino in the mid-19th century and who migrated need for forage. The lichen areas had been grazed down to Sweden. At an early stage in his life he was a reindeer annually, so that they finally resembled those devastated herder, though later his predominant interest was hunting by forest fire. and fishing,as well as being a source of income in which The Lapp bailiff Fredrik Burman in Vasterbotten the major predators were a central fe ature. His know ledge wrote: 'Even in Vasterbotten, the Lapps complain that the of nature and reindeer husbandry in Tome Lappmarkwas reindeer lichens are disappearing from the mountains. It great. In 1908, he obtained the support necessary for him seems improbable that the formation of an ice crust on the to be able to carry out his dream project-a book about ground should be the cause; such things have occurred at all the Mountain Saamis and their life as he had experienced times. The mountain lemming is, however, commoner than it. The principal supporter of the undertaking was Emilie formerly, and may be an appreciable factor. Perhaps, too, Demant Hatt, a Danish woman with a great interest in the Lapps, in their work with the reindeer nowadays, are Saami life, and the manager of the mining company less afraid of trampling on dry summer days than formerly Kiirunavaara-Luossavaara AB , Dr. Hj almar Lundbohm. was the case. This appears to be one of the most likely Lundbohm exercised a great influence on society and causes of the disappearance of the reindeer lichens.' cultural life in northern Sweden. In the report for 1911 , the bailiffAbraham S taaf writes: Turi had a command of written and spoken north-Saam­ 'The Lapps' care of the reindeer is lax. The increasing ish and Finnish, but less so of Swedish and Norwegian. tendency to let the reindeer fend for themselves is a danger The book 'Muittalus samid birra' (A book about the life to the future existence of the reindeer-herding population. of the Saami) was published in Copenhagen in 1910, and It is not enough that they, by their indifference, cause comprised both the Saamish text and a translation into intolerable conditions in the contacts between the settled Danish by Emilie Demant Hatt. A Swedish translation population and the Lapps. It also contributes to the fact that 'En bok om samernas liv', appeared in 1917. The extracts the reindeer lichen, especially in the dry summer months, is below deal with reindeer grazing: more than needfully broken and destroyed by the trampling 'When the reindeer lichen is finished, no other food of the reindeer; and since the lichen is so greatly reduced grows for the reindeer. And the reindeer lichen is finished within the county, it is necessary for the Lapps to strive for no other reason than that the reindeer have consumed in every way to preserve their mountains.' it. the 1913 report (Staaf 1913), he writes: 'As the prin­ -- Nowadays there is little more than the berrybushes In cipal causes of the currentdecline of reindeer husbandry, up here, where there are so many reindeer, and therefore I would put forward: one must take great care to keep the herd together or to (1) the Lapps' upbringing, both in the home and at herd the reindeer. school; (2) their inclination to construct log houses where -- The Lapps have now begun to think about their fu-

Acta Phytogeogr. Suec. 87 44 0. Erikssson et al.

ture, and they can now see that the reindeer no longer can are said to have been equally scraped bare of reindeer live on these areas in Jukkasjarvi and Karesuando, since lichens, as were many other areas, a circumstance which the reindeer lichen is finished; now the young sons of the justified the transition within the Saami villages to joint Lapps are beginning to look for other occupations. reindeer herding during the spring and summer (S.E.T. --Forty years ago, there was still beautiful white lichen No. 4, 1920). here in the Kattavuoma area (east of lake Tometrask), so In S.E.T. No. 4, 1921, the editor writes regarding the that the whole ground was white. And at that time there Oviksfjall mountains, west of lake Storsjon in Jamtland, were not many nomad villages inTal ma, nor in the whole that: of Jukkasjiirvi parish; but when more Lapps began to move in from Karesuando and Kautokeino, the reindeer 'By their nature the Oviksfjall mountains give the im­ lichen decreased year by year. And now, when there are pression of a fertilelandscape, both the high mountains and so many mountain lemmings, they too have eaten much the forest areas around them. The reindeer lichen (South lichen, and in this way the reindeer lichen has decreased, Saamish 'viste') is indeed grazed down, but here it occurs and is still decreasing. Of this there is proof: if one counts incomparably more abundantly than on the Vasterbotten back 25 years, it is easy to recall that there was still plenty mountains and to the northward of them.' of lichen in Talma and in the entire parish of Jukkasjarvi, and now all is, as it were, burnt. In S.E.T. No. 4, 1921, the reindeer herder Jonas Torkels­ --A Saami knows well enough what his own livelihood son relates that: 'I came to the Oviksfjall mountains from is, and he would not wish for any other; but when the Hotagen in 1873. At that time, it seemed as if no Saamis Crown has taken the land from the Saami and given it to had dwelt on the Oviksfjall mountains. The reindeer lichen the farmers, the Crown no longer has power over that land. was as good as untouched.' In S.E.T. No. 3, 1922, Torkel And thus the Saamis now see that the land on which they Thomasson wrote: 'The fact is that the reindeer lichens on may live and graze reindeer, on that land the reindeer can our grazing grounds, principally in the mountains-with no longer live, for the reindeer lichen is entirely gone.' few exceptions-is nowadays grazed down, to the extent The newspaper Samefolkets Egen Tidning (S.E.T.) that the support given to reindeer husbandry by these began to appear regularly in 1919. Its main aim was 'to grazing areas is no more than marginal. Do the natural give its attention to the Saamis, and to see to their eco­ preconditions for pursuing reindeer husbandry exist in nomic and cultural interests.' The editor-in-chief from its our country?' inception and for many years was Torkel Thomasson, a The reindeer-grazing convention between Sweden and Saami, born near Fatmomakke in Vasterbotten county. Norway, which came into force on 1 January 1923, had Originally a reindeer herder, as an adult he acquired some some consequences for reindeer husbandry in northern legal training. Dalarna (Idre), and to some extent also, in the adjacent Over the years, S.E.T. often published field reports areas of Harjedalen. The reason was that reindeer strayed concerning reindeer forage and reindeer husbandry in the into neighbouring areas of Norway from time to time broad sense. In addition to the editor, the authors were during the snow-free period, which was not regarded active reindeer-herders and others with an insight into favourably by the local people. It was suggested to the questions relating to reindeer husbandry. county administration in Jamtland that Idre should be The Lapp Commission of 1919, appointed by the gov­ closed to future reindeer husbandry. (S.E.T. No. 3, 1925). ernmentto provide a basis for new legislation concerning (cf. Linnaeus 1953) reindeer husbandry, the poor-law organisation and the Naturally, the Saamis in Idre objected to this, and argued nomad schools administration, made an extensive journey for the construction of a fence along the Norwegian border. throughout the Saami districts in the summer of 1920, to This would also prevent the movement of reindeer from the canvass the views of the Saami population. The editor of direction ofTannas, which also occurred.There was some S.E.T. took part in this, and in his travel diary, published support for the suggestion from the county administration in S.E.T. Nos. 3 and 4, 1920, gave some information about in Falun. The desire of the Saamis in Idre to remain there current grazing conditions. Some of this informationwa s, may be explained by the fact that the area has Sweden's however, at second hand. richest lichen grazings, as well as on the mountains, with a carrying capacity for many more reindeer than there The heathland areas in the coniferous forest between were in the area at the time. There was, in addition, the Gallivare and Soppero appeared to him to be heavily advantage that the area was rarely affected by the snow grazed; there were only traces of reindeer lichens. The conditions that hinder grazing, in the form of a surface ice­ same observation was made along the road between Ovre crust (jlen, tj uohki), which are feared in the northernparts Soppero and Karesuando, where short-stemmed, crooked of the reindeer-grazing region (S.E.T. No. 1, 1926). mountain birch had increasingly usurped the place of co­ The opposition of the Saamis led to court proceedings. niferous trees in the forests. The mountains in the north The lower court found in their favour, but the superior court

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 45

sen was to estimate the abundance of the formations and species according to a five-point scale:

Rr and r rare 1 and 2 occasional 3 and 4 scattered 5-7 frequent 8-10 abundant

Fig. IB. (In Hult 1881) Investigation sites: 1. - KittiHi.,2, - Turtola, The basic formations were. 3. - Kittila-Isovaara. Pine forests with a complete lichen cover. I. Coniferous trees II. Broadleaved trees Ill. Shrubs IV. Dwarf shrubs V. Grass found in favour of the county administration in Jamtland. In VI. Herbs VII. Creepers (absent) a judgement dated 28 March 1930, the affected Saamis wer VIII. White mosses found guilty of illegal reindeer grazing in ldre, and were IX. Liverworts required to evacuate the area no later than 15 December, X. Lichens and to remove to Tossasen Saami village, which, from the point-of-view of reindeer husbandry, was a far poorer The results from each sample area were presented option (S.E.T. No. 4, 1930). partly as grid diagrams divided into horizontal and vertical 'By all accounts, reindeer husbandry in this countryhas cells. The horizontal direction showed the stratificationof now passed its zenith. There are signs that it is declining. the vegetation and its abundance. The vertical direction The overcrowding in Arjeplog has led to grazed-down showed the basic formation. Among the most northerly lichen fields, and it is no longer attractive for young Saami sites investigated were three pine stands situated at KittiHi, to devote themselves to reindeer husbandry, with some Turtola and KittiHi lsovaara. exceptions. The perpetual conflict over the convention As is shown in Fig. IB (In Hult 1881), the cover of (between Sweden and Norway, concerning questions of ground lichens was rich. At KittiHi, Stereocaulon paschale reindeer grazing) andmuch else, causes one to be doubtful.' occurred abundantly. Cladonia spp., Cladina spp., Cetraria (S.E.T. No. 3, 1932; LarsAndersson-Spellok, among the spp. were occasional. At Turtola, Cladina arbuscula was Saami villagers appointed assistant bailiff in Semisjaur­ abundant, C. rangiferina frequent, Nephroma arcticum Nj arg Saami village). was occasional and Stereocaulon sp., Cladonia spp. and 'There is overcrowding and the lichen fieldsare totally others were sparse. grazed down, which indeed is the situation not only within At KittiHi-lsovaara, Cladina arbuscula was frequent, Arjeplog but also in the areas north and south of it. ' (S.E.T. and Cladonia spp., Cladina rangiferina, C. stellaris and No. 3, 1932). Stereocaulon spp. and others were scattered.

R. Hult's attempt to deduce the laws governing the The results of an inventory of four pine heaths in the vicinity of Kolari, between Turtola and KittiHi, and fr om composition of plant formations exposed bedrock areas above the tree limit on YUistunturi R. Hult, together with Hj . Hjelt, took part in 1877 in a at ea. 900 m above sea level also showed a complete lichen botanical expedition to northernmost Osterbotten and the cover with a similar species assemblage. This gives reason western part of Kemi Lappmark, Finland (Hult 1881 ). to infer that the grazing pressure in this area, close to the At the time (until 1889), the studied area was still open Swedish reindeer-husbandry region, was low at the end to 'Swedish' reindeer husbandry, carried on by Finnish of the 19th century. Saamis, who had recently obtained Swedish citizenship, and who consequently could migrate to Enare as well as to Norway. In addition to the 'Swedicised' Saamis, there were reindeer-herding Saamis who had retained Finnish nationality. Hult's goal was to contribute to the investigation of 'the laws that determine the composition of plant forma­ tions (vegetation types).' The aim thus was not to assess the availability of reindeer forage, but the result is to some extent illuminating even in that respect. The method cho-

Acta Phytogeogr. Suec. 87 46 0. Erikssson et al.

The 1909 Commission on Reindeer Grazing Lands as follows: ( 1) small remains (or sparse traces) of lichen The 1909 Commission on Reindeer Grazing Lands carried (2) remains (or traces) of lichen out field studies in 1910 and 1911 regarding the supply of, (3) lichen of abundance 1 and access to, reindeer forage within the Saami villages (4) lichen of abundance 2, etc., up to abundance 10 in Tome Lappmark which make use of grazing lands in Norway during the summer. The studies were part of the By lichen of abundance class 1 (y= 1) is understood preparatory work for a new convention between Sweden a lichen grazing that may indeed be grazed down, but and Norway, covering the trans-border reindeer husbandry. nonetheless no more severely than that it can be considered The inventory methods and some results, taken from 'Pro­ satisfactory when snow and ground conditions are not a ceedings of the 1909 Commission on Reindeer Grazing hindrance to grazing. -Lichen of abundance class 2 (y=2) Lands', are summarised below: provides good grazing when grazing conditions are not a hindrance. - Lichen of abundance class 3 (y=3) can be I. In consideration of differences in topography, soil regarded as very good grazing, which provides necessary and vegetation, and with regard to their varyingimportance nutriment, even when ground conditions are a hindrance. to the reindeer husbandry, the study areas are divided into -Lichens of abundance classes 4-8 have been encountered zones exceptionally, and then only on small patches, which prob­ ably are protected from reindeer grazing. Small remains (1) The coniferous forest zone (S-zone) or sparse traces of lichen and remains or traces of lichen (2) The mountain birch forest zone (FS-zone) may on average be regarded as equivalent to 1/3 and 2/3 (3) The montane heath zone (V-zone) of class y= 1. The sub-classes most commonly represented (4) The high-mountain zone (H-zone) in the study area are 'small remains or sparse traces' and 'remains or traces'. They occur both on sites heavily grazed II. The fieldworkwas based on line transects. For prac­ by reindeer, and in moss-dwarf shrub vegetation in the tical reasons, e.g. accessibility, the transects could not be form of small lichen patches with taller fruiting bodies. laid out parallel. Emphasis was placed on 'all forest types, Lichens of abundance classl, and exceptionally of class terrain and extent of vegetation in which they might be y=2-3 mainly occur in forested areas. represented within each individual area'. The inventory of the lichenresources within the moun­ tain birch forest and montane heath zones in the Saami Ill. 'The percentage occupied by the forage types villages Talma, Saarivuoma, Lainiovuoma and Konkama, lichen and grass-herb forage, and their quality, as also of the 'northern villages', is summarised in the tables below. moss, dwarf shrubs and rock exposures along the transect, Information concerninglichen grazings in these villages, was recorded directly. ' Occasionally, when the transects which has been obtained under other circumstances and on traversed dry sites to too large an extent, some adjustments other occasions, will be presented in a later section. were made, following discussion. 'These adjustments were The zone of mountain birch forests, the FS-zone, ap­ made in each individual case according to the unanimous pears to correspond most closely to the mountain birch decision of the commission.' forests described by Andersson et al. (1978-1984) and Pahlsson ( 1998). The zone of montane heaths (the V­ IV. The inventory distinguished four main classes: zone) is comparable with the heaths described by the same authors. This zone is absent from the surveyed part (1) Sites unsuitable for grazing of Talma. (2) Moss and dwarf shrubs As Tables 2A-D show, the lichen resources in 'the (3) Lichen grazings mountain birch forest' and 'montane heaths' were very (4) Grass-herb grazings limited. The commission's assessment was that the re­ sources in none of the villages corresponded even remotely Restriction to these main classes was determined by to the requirements of the existing numbers of reindeer. difficulties in correctly distinguishing between all of the 'The nomad Lapps' reindeer within the studied areas graze transitional forms that exist. It may be of particular inter­ very hard. It is sometimes difficult to understand how est to touch upon the commission's definition of lichen lichens over such great areas came to be so thoroughly grazing grounds: grazed down by reindeer. ' ( 1909 Commission on Reindeer ' ... in the term lichen grazing grounds the commission Grazing Lands). includes all sites that are covered to varying degrees with The representatives of the Saami villages interviewed ground lichens, that are suitable for reindeer grazing'. by the commission claimed, quite unanimously, that the ruined grazings on the Swedish side of the border were the The relative value of the lichen ground was classified result of the hordes of reindeer that had poured in from the

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 47

Table 2A. Talma Saarni village. Lichen resources in the mountain birch forest zone (FS-zone).

FS-zone

Lichen grazings km2 % of area of lichen grazings % of area of region

Small remains (or sparse traces) 544.0 84.8 58.8 Remains (or traces) 104.5 14.9 6.6 Abundance =1 53.6 7.6 3.4

Table 2B. Saarivuoma Saarni village. Lichen resources in the mountain birch forest zone (FS-zone) and montane heath zone (V-zone), respectively.

FS-zone Lichen grazings Ian2 % of area of lichen grazi.ngs % of area of region

Small remains (or sparse traces) 583.9 77.5 34.3 Remains (or traces) 94. 1 13.7 9.5 Abundance =1 10.4 1.1 1.5

V-zone Small remains (or sparse traces) 390.1 97.7 51.3 Remains (or traces) 8.9 2.2 1.2 Abundance =1 0.2 0.1 0.0

Table 2C. Lainiovuoma Saarni village. Lichen resources in the mountain birch forest zone (FS-zone) and montane heath zone (V­ zone).

FS-zone Lichen grazings % of area of lichen grazings % of area of region

Small remains (or sparse traces) 305.5 73 .2 51.9 Remains (or traces) 79.6 19.1 13.6 Abundance = 1 32.1 7.7 5.5 Abundance =2 0.2

V-zone Small remains (or sparse traces) 347.4 96.2 47.0 Remains (or traces) 27.8 7.4 3.8

Table 2D. Konkama Saarni village. Lichen resources in the mountain birch forest zone (FS-zone) and montane heath zone (V-zone).

FS-zone Lichen grazings Ian2 % of area of lichen grazings % of area of region

Small remains (or sparse traces) 311.5 64.4 48.1 Remains (or traces) 107.4 22.2 16.6 Abundance = 1 59.3 12.2 9.2 Abundance =2 4.7 1.0 0.7 Abundance =3 0.8 0.2 0.1

V-zone Small remains (or sparse traces) 547.4 91.7 45.3 Remains (or traces) 43.9 7.4 3.6 Abundance =1 5.6 0.9 0.5

north. 'Aged Lapps relate that in their youth (before the lichen grazings after the closure of the Finnish border mid-19th century) the land was still white with lichens, but (1889). Then, the large herds of the Rommavuoma and that two or three decades after the incursion of the Kauto­ Suondavaara Saamis, which in former times had regularly keino Lapps in the 1850s and 1860s, this began to decrease spent the winter in Finland, could no longer be taken there, markedly. ' Still more striking was the deterioration of the for fear of the brusque methods of the Finnish officials.

Acta Phytogeogr. Suec. 87 48 0. Erikssson et al.

These reindeer, too, must now graze in Sweden. Table 3A. Plant cover on sample plots in three lichen-rich vegeta­ 'Formerly, every area was worked over by the herds tion types (Fries 1913): I. Empetrum-rich lichen birch forest at Kelottijarvi, 30 km of Karesuando; Heath-like lichen but once during the winter. It was also possible to leave WNW 11. birch fo rest. Tavvaskaite, S slope; Lichen-rich Empetrum some areas completely untouched. At the present time, Ill. heath, ofKiepenjilla bythe 'Barrier lake'. Ve getation layer: an area is worked over both during the migration to the W a=upper tree layer (> 6 m); b=lower tree layer (2-6 m); c=shrub coniferous forests, and in the late-winter migration to the layer (0.8-2 m); d=upper field layer (0.3-0.8m) ; e=intermediate mountains. Indeed, it happens that some areas are worked field layer (0.1-0.3 m); f=lower field layer (0.03-0.1 m); over three or four times during the winter, by herds from g=ground layer (0-0.03 m). several villages.' (op. cit.) In the opinion of the Saamis, it was necessary to reduce Ve getation the numbers of reindeer, so that the grazing grounds would layer Species I. 11. Ill. not be worked over more than once each winter, thus avoid­ b Betula odorata 2 1 ing further deterioration, which threatens to make reindeer c Betula odorata 2 2 husbandry impossible. Furthermore, it would be desirable d Calamagrostis lapponica that some areas should be left untouched in some years, e Festuca ovina to recover. The number of reindeer must nevertheless be ]uncus tri.fidus reduced. Some consider a reduction by half, others by at Luzula arcuata least one-third. In their opinion, it might be possible to to Calamagrostis lapponica attain the same result if the area of winter grazings were f Empetrum nigrum 2 2 3 doubled and, above all else, if they were protected from Arctostaphylos alpina Lycopodium complanatum grazing by forest reindeer throughout the year (ibid.) Betula nana 1 ,5 Va ccinium vitis idaea 1 Observations on reindeer grazingduring the 20th cen­ Myrtillus uliginosa 1 ,5 estuca ovina 1 tury, by botanists working in the mountains F Pedicularis lapponica g Polytrichum juniperinum During the summers of 1905-1912, Thore C.E. Fries, partly + + + +Polytrichum cfr. piliferum 3 accompanied by and in cooperation with E. Bergstrom and + + Dicranum sp. 1 3 3,5 S. Martensson, carried out extensive studies of vegetation + Cladonia rangiferina alpestris 3 conditions in Tome Lappmark. In the summer of 1908, + + Cladonia rangiferina vulgaris + the reindeer's choice of forage plants was investigated. In + + Cladonia rangiferina silvatica 3 4 + the winter of 1909, long journeys were made in the study Stereocaulon paschale 2 area, in company with reindeer-herding Saamis. The aim Lecanora tartarea 1 ,5 3 was to study both reindeer husbandry in winter, and the Cetraria cucullata 1 + Cetraria islandica effects of the snow-cover on the vegetation. + Among the results of this work can be mentioned a Cetraria nivalis 3 3 vegetation map tailored to the needs of reindeer husbandry, Cladonia bellidijlora Cladonia cenotea at a scale of 1:200,000 (Fries & Bergstrom 1909, in Lon­ Cladonia coccife ra nberg 1909, Fig. sections on the reindeer's choice of XI), Cladonia decorticata forage plants and on conditions for reindeer husbandry in Cladonia deformis the 'northern villages' in Lonnberg (1909), and a doctoral Cladoniafurcata crispata + thesis (Fries 1913). Cladonia gracilis cornuta In Lonnberg 's publication ( 1909), Fries concluded that Cladonia pyxidata 1 1 the winter grazings in the coniferous forest region sufficed Cladonia uncia/is 2 3 for the then existing stock of reindeer in the 'northern Lecanora hypnorum villages'. However, grazing pressure was intense, as was Lecanora tartarea Pertusaria dactylina 1 shown by the shortness of the lichens. As regards the Phyllocladien sterile crustaceous lichens 3 + autumn and spring grazings above the coniferous forest Alectoria ochroleuca rigida 1,5 limit, the situation was worse; forage was very scarce. It Alectoria divergens consisted of sparse and short lichens, clumps of Empetrum Alectoria nigricans hermaphroditum and scattered tussocks of grass- where Solorina crocea there was any vegetation at all. Through the trampling of Peltigera rufescens the reindeer, and the effects of the wind, a large part of Thamnolia vermicularis the ground was devoid of all plant cover. Cover: !=occasional; 2=scattered; 3=sparse; 4=rich; 5=abundant. In his doctoral thesis (Fries 1913), he gave an account of the vegetation above the coniferous forest limit in the

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 49

Table 3B. Bergfors 1: heath-like lichen birch forest, 487 m a.s.l.; Bergfors 2: My rtillus nigra-rich lichen birch forest, 485 m a.s.l.;

Koppen\sen: on lichen-rich Empetrum nigr -heath, 430 m a.s.l.; Riksgransen 1: on lichen-rich My rtillus ig -heath , 470 m a.s.l.; a n ra Riksgransen 2: on lichen-rich My rtillus nigr -heath, 480 a.s.l. (for explanations see Table 3C). a m

Bergfors Bergfors Kopperasen: Riksgransen Riksgransen 1: 2: I 2 h c h c h c h c c h

Cladina stellaris E T 31 E E C. rangiferina 15 E E 29 T T 39 T 41 C. arbuscula/C. mitis s 30 T s 26 s 41 s 39 Cladonia uncialis T T E T T Stereocaulon paschale T- S s 24 24 E 30 s 30 s 40

northernpart ofTorneLappmark. Hult's scale (Hult 1881) ponica heath. The occurrence in the bottom layer of a was the basis for the description of cover and for the divi­ large number of lichen species of varying value as forage sion into vegetation strata. plants, was noted. On half of the plots,. important forage The descriptions of the vegetation were related to species occurred scattered or even richly, but were of low 'representative' sample plots of varying size in existing stature. plant communities. As examples here have been chosen the Fries considered that the role of reindeer as a major description of the lichen cover in the lichen-rich mountain contributor to the degradative effects of wind erosion on birch forest-valuable for grazing -and the lichen-rich the lichen heaths of Tome Lappmarkwas beyond doubt. On montane heaths, respectively. the extensive, sandy plateaux in the alpine region, intensive Nine types of lichen-rich mountain birch forest were reindeer grazing and wind erosion together had given rise to distinguished, on the basis of the characteristics of the large, bare patches. To this process, Fries' contemporaries tree layer, the presence of a shrub layer, the dominant could bear witness, for they had during their lifetime seen species of the field layer, and lichen dominance in the the vegetation disappear from areas where the reindeer bottom layer. lichens previously had formed a continuous carpet. On one sample plot, Cladina spp. and Cetraria nivalis were scattered in the lowest fieldlayer. In the bottom layer TengvaU 's measurements of reindeer lichen growth of all plots, there were single/scattered individuals of a considerable number of lichen species. In the years 1916-1919, commissioned by the Interior Fifteen types of lichen heath were distinguished. The Ministry, T.A. Tengvall investigated lichen growth in the classification was based on the dominance of lichens and northernmost Saami areas (Tengvall 1928). The method vascular plants, respectively, in the field and bottom layer. used was serial measurement of the height increment of The absence of vascularplants gave a sixteenth type-the species protected fromreindeer grazing. Enclosed sample pure lichen heath. On two only of the plots classified as plots were laid out at seven sites along the railway between lichen heath did lichens appear in the field layer, viz. in Gilllivareand Riksgransen, three of which were below the lichen-rich Cassiope tetragona heath and Diapensia lap- coniferous forest limit. In the present context, attention

Table 3C. The sample plots along the south side of lake Tornetrask, at the start of the project in 1914. Lichen cover and height. Cover (c), E=occasional, T=scattered, S=sparse, R=rich, Y=abundant and Height (h) in mm of lichen thalli. Bergfors 1: heath-like lichen birch forest, 487 m a.s.l. Bergfors 2: My rtillus nigra-rich lichen birch forest, 485 m a.s.l. Kopperasen: on lichen-rich Empetrum ni ra-heath, 430 m a.s.l. g Riksgransen 1: on lichen-rich My rtillus ni r -heath , 470 m a.s.l. Riksgransen 2: on lichen-rich My rtillus nigra-heath, 480 m a.s.l. g a

Bergfors Bergfors Kopperasen Riksgransen Riksgransen 1 2 1 2 h h c c h c c h c h

Cladina stellaris E T 31 E E C. rangiferina E E 29 T 15 T 39 T 41 C. arbuscula/C. mitis s 30 T s 26 s 41 s 39 Cladonia uncialis T T E T T Stereocaulon paschale -S s 24 T 24 E 30 s 30 s 40

Acta Phytogeogr. Suec. 87 50 0. Erikssson et al.

will be concentrated to the plots above this limit, i.e. those grazing, and then usually during the snow-free period. situated along an east-west line along the south side of The number of reindeer that grazed the area during the lake Tometdisk, viz. summer is uncertain. In 1945, when reindeer inventories Cladina stellaris was absent from the plot at Kop­ were carried out, over 13,000 reindeer appear to have penisen. On the other plots, the cover of the lichens most grazed there, and of these 3--4,000 had habitually strayed valuable as reindeer forage was estimated. In summary, it in from Vaisaluokta in the north. may be stated that the stock of lichens within the research Selander considered that reindeer grazing within his area was small. The degree of cover was in the interval study area did not affect the vegetation to the extent that E to T. Cladina arbuscula/C. mitis and Stereocaulon pas­ might be suggested by the large number of reindeer. The chale, however, occurred somewhat more richly and had reason for this may have been both the tendency of the rein­ slightly taller thalli. deer to roam widely, their light grazing pressure compared Harald Smith (1920) carried out fieldwork of a bo­ with that of domestic livestock, and their sophisticated tanical nature in the years 1908-1919, with emphasis on choice of forage plants and plant parts ( cf Lonnberg 1909, plant-sociological studies in the high-mountain areas of p. 144; Du Rietz 1924, p. 95). southwestern Jamtland and northwest Harjedalen. Fries The observations concerningthe relationship between (1913) had grouped the heath series plant communities reindeer trampling and wind erosion and exposure of the into two parallel sub-series-the lichen-rich and the moss­ soil, which Fries (1913, p. 252) had made, were not shared rich. Smith found that this division was not clear within by Selander, who did not note this phenomenon within his his study area, with the exception of the most continental study area, which to him appeared to be poor in lichens. part-Vattafjall-Rogstoten. Otherwise, he largely failed to G. Degelius (1943) studied during a few weeks in findtypical lichen-rich heath, naturally with the exception the summer of 1941 the lichen flora and vegetation in of heaths of wind-barren type. the mountains east and south of Virihaure, an area that At the start of the study, intensive grazing by exces­ was included in Selander's (op. cit.) study area. Degelius sively large reindeer herds had entirely ruined existing considered that the Virihaure area is a part of the mountain heaths. Only on limited areas of block-rich terrain, inac­ chain that is heavily utilised for reindeer grazing, which cessible to reindeer, was it possible to see well developed contributes to the absence of major, continuous lichen specimens of the larger lichens. Between 1913-1915, the heaths. reindeer herds were heavily reduced, at the same time as The factors that principally shape the lichen flora and the management routines were tightened. As a result, Smith vegetation around Virihaure are, according to Degelius: considered that he could see a notable increase in lichen biomass on the heaths he had classified as lichen-rich, 1. Easily weathered and more or less lime-rich bedrock. despite the low growth rate. In comparisonwith the lichen 2. A limited occurrence of mountain birch forest. 3. The moist climate, which, especially to the westward, gives heaths of the Rogen-Vattafjall area, they must, however, rise to areas in which the snow thaws late or not at all. be considered as markedly poor in lichens. 4. Reindeer grazing. The first, and froma botanical point of view, possibly 5. The temporary and quite insignificant Saami settlements, the best studied of our montane areas, together with the which nevertheless lead to some removal of wood and of Abisko mountains, is that part of Lule Lappmark around material for building fences and enclosures, which influences K vikkjokk and Virihaure. the sparsely occurring birch forests. Olov Rudbeck the Yo unger and Linnaeus were the first outside visitors, and after them, the number of visi­ The late thaw, which affects some partsof the area, as tors to this part of the mountains is almost incalculable. well as the intensive spring reindeer grazing in other areas, The first investigation of the flora in Lule Lapp@mark prevents the developments typical of the more continental of a modem plant-geographical nature, was made by G. montane areas, where reindeer grazing does not occur ­ Wahlenberg (1812). e.g. on Fulufjall and Dovre (ibid.). During the 1940s, S. Selander (1950) carried out plant­ geographical fieldwork in the area between the Sarek and Stora Sjofallet national parks, which had been the subject of intensive study by Tengwall (1920, 1924, 1925) and Bjorkman (1924, 1939). The limit in the south was the areas in which Arwidsson (1926, 1943) and Wistrand (1962) had worked. Selander's study area covered parts of three Saami vil­ lages-Tuorpon, Jillak.askaand Sirkas, all three of which had a varying use as migration routes, as spring, summer and autumn residences, and - naturally-for reindeer

Acta Phytogeogr. Suec. 87 2. The WWF-project, Background

Project objectives and study sites Map 1), were incorporated into the project, to illustrate 2.1 vegetation development over a longer period. The objective of the main project was to document long­ term changes in mountain vegetation. Results obtained Geology should be useful, for example, in the development of 2.2 methods for environmental monitoring, and in studies of Table 4 lists the position, height above sea level, solid the utilisation of grazing in reindeer husbandry. Conse­ geology and soil type of all the experimental sites. At all quently, it was appropriate for the work to be concentrated on types of vegetation thatare susceptible to wear-and-tear, those that are of great importance for grazing, those that cover large areas, and that can be reliably identified on the available photographs from different periods. To IR provide comparisons, sample plots were laid out in types of vegetation considered less susceptible to wear-and-tear, e.g. moss-rich forests of mountain birch, grass heaths and sub-alpine meadows (Emanuelsson 1984). To fulfil the aims of the project, a reference group of five experimental areas, spread along the mountain chain, was selected. Reindeer husbandry and nature-conservation interests were included, together with access to vegetation maps and available older IR aerial photographs. These last t;'-Ostereund were also intended to serve as a basis for remote-sensing studies, to be carried out at the Department of Physical

Geography, Stockholm University (Ihse & Allard 1995). N The final report on those studies was published by Al­ lard (2003). The five experimental areas (see map) were given the working names Tavvavuoma, Ritsem, Sanfjallet, �l Langfjhllet and Fulufjallet, and were reconnoitred during Map 1. Location of experimental areas in the WWF-projectalong the s er of 1993. The southernmost areas are on Fu­ the Scandinavian mountain chain. The experimental areas are umm located in vegetation types which are assessed to be susceptible lufjallet in Dalarna, and the northernmost are 15 ea. km to deterioration, of importance for grazing, covering large areas, south of Tavvavuoma in Lap land. and are identifiable on available IR-photographs. The areas are In 1967-1968, several paired sample plots (a fenced (working name/Saami village) : 1. Tjuolmajaure/Lainiovuoma, 2. and an open) were laid out in Tavvavuoma (0. Eriksson et Poullanvare!Lainiovuoma, 3. Tavvavuoma!Lainiovuoma, 4. Rit­ al. 1998), and in connection with fieldwork in 1996, the sem/Mellanbyn, 5. Ritsem/Sorkaitum, 6. Sanfjilllet/Mittadalen, two in best condition, Puollanvare and Tjuolmajaure (see 7. Langfjallet/Idre Nya Sameby, and 8. Fulufjallet.

Table 4. Position, height above sea level, solid geology and soil type (Berggrundskarta over Norrbottens Ian 2000 (BD), Berggrund­ skarta over Norrbottenfjallens norra del l965, Karta over berggrunden i Jamtlands lan 1969 (Z), Jordartskarta over Kopparbergs lan 1948 (W). National Atlas of Sweden 1994).

Locality Latitude, longitude ma.s.l. Bedrock Soil type

Tavvavuoma (T), Bd-prov. 68° 20 ', 21° 15' 500-575 Metagranodiorite, Till and weathered deposits above (incl. Poullanvare) Metatonalite timber line Ritsem (R), Bd-prov. 67° 45 '' 17° 40' 725-800 Micaschists - , - Sonfjallet (S), Z-prov. 62° 20', 13° 55' 910-940 Vemdalen Quartzite Gravelly wave-washed till, formation high boulder frequency , Umgfjallet W-prov. 62° 05 , 12° 25' 800-1010 Dala granite Gravelly till (L), Fulufjallet (F), W-prov. 61° 30', 12° 25' 860-930 Sandstone Gravelly till Tjuolmajaure (Tj), Bd-prov. 68° 30'' 20° 24' 865 Gneiss Till and weathered deposits above timber line

Acta Phytogeogr. Suec. 87 52 0. Erikssson et al.

sites except Ritsem, the bedrock is severely weathered and mountain chain functions as a divider between two large chemically acidic. The sites thus lack the potential to form a climate zones, the Atlantic and the Continental. The warm suitable substrate for rich vegetation. The soils have podsol westerly winds from the Atlantic are forced upwards,and profilesthroughout, with till and, in some cases, weathered slowed down, by the mountain chain, which has the re­ soil at the base. The raw humus layer on the lichen-rich sult that their effecton areas to the east is less than if the vegetation types is thin (1-4 cm), whereas it is slightly mountains were not there. Correspondingly,the mountains thicker in the moss-rich stands of mountain birch. prevent the easterly winds from inner Eurasia from affect­ The Ritsem site lies within an area of bedrock made up ing the climate too strongly to the west of the mountain of mica schist that is soft,relatively easily weathered and chain. As a result, relatively low summer temperatures locally calciferous, which influences the vegetation. characterize the mountain region, which has to do with the normal decrease of temperature with height. The winter temperature, however, is relatively high, since it often is 2.3 Physical geography higher at higher altitudes than in the valley bottoms in these regions, and because of the maritime influence of The sites studied were based on aggregates of both abiotic the sea to the west (Angstrom 1974). Comparisons on and biotic parameters. Both Tj uolmajaure and Ritsem an annual basis between calculated temperatures within are situated within the Northern High Mountain area of areas at certain latitudes and certain heights above sea Nordland Trams and Lapland. They consist of rounded level, with those actually measured, reveal anomalies low mountains, interspersed with mountain plateaux and that, when transferred to maps, show zones with similar valleys. The solid geology and soil types are described in temperature climate. Table 4. Mountain birch woodlands become increasingly The study sites at Tjuolmajaure and Tavvavuoma are more frequent to the south and east. situated within an extensive continental-subcontinental Tavvavuoma is part of the continental forests of area in the northernmostpart ofFennoscandia (Sjars 1990, Finnmark and Lapland. The area is characterized largely Oksanen & Virtanen 1995). Low winter temperatures and by subalpine-subarctic mountain birch woodlands, with relatively little precipitation, which falls as snow during open montane plant communities at higher altitudes. The half the year, characterize the region. Permafrost, in the area consist largely of expanses of mountain and hilly form of palsa hummocks, is abundant. Inmany respects, the terrain, up to ea. 700 m a.s.l. experimental sites at Samjallet, Langfjallet and Fulufjallet Sanfjallet, Langfjallet and Fulufjallet all lie within the are situated in a similar continental-subcontinental region southernmountain area, which is characterized as rounded with its centre in Norway, but which extends over northern mountains on poor bedrock, with intermediatelevel areas Dalarna and southwest Harjedalen (Oksanen 1998, Sjors (Anon. 1977). 1999). The site at Ritsem is located at a high altitude, and the western mountains do not provide an effective barrier to the westerly winds; consequently the climate in this area 2.4 Climate and weather is to some extent oceanic (Oksanen 1998). Continentality can be calculated as the sum of the Northwestern Europe, particularly Scandinavia, benefits temperature difference between July and January, and during the colder parts of the year by an inflow of mild between day and night during June (Raab & Vedin 1995). air from the Atlantic (Sjors 1999). In many respects, the The Ritsem areawill then have an index in the range 25-35,

500

-400 E e -• Tawa ; 300 --•--- Ritsem 0 ____.,__Son� 200 ---tr- Lang� ui e -a--- Fulu1j 11. 100

1990 1992 1994 1996 1998 2000 Year Figure 2. Precipitation during the growing season (daily temperatures > +3°C) for the years 1990-2000. The precipitation data were obtained from meteorological stations in Sweden and Norway, not far from the study sites (Table 2). Precipitation (mm) is presented as mean values.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 53

whereas the other study sites are in the range 35-40. By 150 . r------·-·······-·-·····-: comparison, the most continental area in Sweden, situated i n1961-1990 i 120 . i I in the inland parts of Norrbotten and Vasterbotten, has an i l-��-��-�� ! index in the range 40-45, whereas the most maritime area, ! '8 90 along the southwest coast, has 20-25. ·c & To calculate the length of the growing season for each en 60 c study site, the air temperature was weighted with the dis­ �·� 30 tance to the nearest meteorological station, according to the following formula: 0 T /d2 Tawa Ritsem Son1j Langlj Fulufj = L i i T 2 Study sites I, lldi 3. The length of the growing season (days >3°C) during where T is the weighted temperature at the site, is the Fig. Tj 1961-1990 (normal data), and for 1990-2000. temperature at station i, and di is the distance (km) between station i and the site, as shown in Table 5.

exist, because different plants have differing abilities to Growing season withstand extreme temperatures. Good examples are the The growing season, i.e. the period when vegetation is most ground-living Bryorialichen species, with adark -coloured active with regard to reproduction, growth and dispersal, thallus that can be photosynthetically active in small melt is usually defined as that part of the year when the daily cavities under a thin snow cover, even if the temperature temperature exceeds, on average, a certain limit. Angstrom is low. A condition for this, however, is that there is some ( 197 4) gives this as the period(days) during which the daily contact with layers of airclose to the soil, and sufficiently mean temperature continuously exceeds +3 oc it ends intensive sunshine (Larson & Kershaw 1975). ; when the temperature again falls below the threshold value. The length of the growing seasons during 1961-1990 Raab & Vedin ( 1995) use the threshold value 5 °C. Both of were calculated according to Moren & Perttu (1994), and these values are, in fact, based on temperatures considered is given in Fig. 3 as the mean of the period's length for relevant for agriculture and forestry. A generally applicable vegetation types at the differentstudy sites. threshold value for the mountain region probably does not

Table 5. Distance (km) between study sites and the meteorological stations used in determining the length of the growing season for each locality.

Meteorological station Sites

Tavvavuoma Ritsem Sonfjlillet Langfjallet Fulufjallet

Katterjiikk 131 Naimakka 38 Karesuando 52

Ritsem 15 Nikkaluokta 57 Satis 49 Stora sj ofallet 49

Hede 14 KlovsjohOjden 46 Lofsdalen 33 Dravagen 33 ldre 89 33

Roros 72 Drevso 25

Sarna 26 Grundforsen 28 Alvdalen 70

Acta Phytogeogr. Suec. 87 54 0. Erikssson et al.

Humidity 2.5 Vegetation types Precipitation and evaporation, being dependent on tem­ perature, and runoff in large or small waterways, together The underlying bedrock and its weathering capacity influ­ largely decide the level of moisture in the soil, and thus also ences the composition of the vegetation. On easily weath­ the character of the climatic zone. Humidity is of greatest ered basic bedrocks, e.g. limestone, dolomite and mica importance during the growing season. De Martonne ( 1926 schist, all of which are nutrient-rich, the vegetation is richer in species than on the more acidic and less readily weath­ a,b,c) created an aridity index (more correctly a humidity index) based on the formula: ered bedrocks, e.g. granite, gneiss, quartziteand sandstone. Even if lichens do not actively take up nutrients from the soil, they are nevertheless influencedby the bedrock. This p -­ is because the minerals influence the composition of the H = T+10 surface water, and thus the nutrients available, in addition to the fact that competition frommosses and vascular plants where P is the annual precipitation (mm) and Tis the often is considerable on nutrient-rich sites. mean annual temperature CC). The constant 10 is used to In selecting the vegetation types within the study sites, reduce the influenceof temperature on the index slightly, use was made of Ve getationskarta over de svenska fjiil­ and to make it more useful in colder regions. len (Andersson et al., 1978-1984) and Ve getationskarta According to Angstrom (1953a), de Martonne's for­ over norra Hiirjedalens skogsland (Rafstedt 1987). The mula can also be used in a modified form to calculate vegetation classifications used in these maps are based on humidity during periods other than an entire year, e.g. for physiognomic and plant-sociological-ecological factors, the growing season. as well as on what it is possible to map from available infrared aerial photographs. The vegetation types studied in this project cover all large mountain areas, and are also important grazing areas for reindeer. The linkage of

is the humidity during the period, n = the number of Hv v the localities to the aerial photographs, the desirability of days in the period, = the precipitation and the mean Nv Tv gaining access on the ground, and the need for large areas, temperature during the period. De Martonne's formula have sometimes caused the requirementfor homogeneous possibly gives slightly too high humidity values for some vegetation types to be disregarded. mountain areas where the excess of precipitation over The study sites situated above the tree limit, i.e. on evaporation rapidly runs off on account of the ground's grass heaths, dry heaths, dry heaths with lichens and mead­ irregularity (Angstrom 1953a). Nonetheless, on the whole ows with herbs, lie within the low alpine zone, which is the maps of the areas agree fairly well with those that show delimited at its lower boundary by the woodland formed temperature during comparable periods (ibid. 1953a). by Betula pubescens var. czerepanovii. It has a vertical B. Eriksson (1986) calculated the share of the precipi­ extent of about 300 m, up to the level at which Va ccinium tation that did not evaporate, but which could benefit the myrtillus ceases to dominate, even in situations where there vegetation. This is based on the difference (mm) between is a reliable, late, thaw (Sjors 1999). precipitation and evapotranspiration. Table 6 shows the The present localities lie in the altitude interval 570 m humidity climate of the study sites during the growing sea­ (Puollanvare dry heath) to 1010 m (Lfmgfjallet grass heath), son, according to de Martonne and Eriksson, respectively. a range of 440 m which is approximately equivalent to a According to Eriksson, the three southernmost study sites difference of six degrees of latitude. The plant cover on are in much moister climatic zones than they are according the heaths may be more or less fragmented: this depends, to de Martonne. e.g. on the degree of exposure, possible trampling and

Table 6. Humidity at the study sites during the growing season.

Study site Humidity index Character of area2J Humidity Character of climate region 1l (mm)3l 4l

Tj uolmajaure 28-32 Continental 0-50 We akly humid Tavvavuoma 28-32 Continental 0-50 We akly humid Ritsem 48-52 Humid 150-200 Very stronglyhumid Sonfjallet 36-40 Moderately dry 100- 150 Strongly humid U.ngfjallet 44-48 Subhumid 150-200 Ve ry strongly humid Fulufjallet 44-48 Subhumid 150-200 Very strongly humid

Martonne in Angstrom (1953a); 2l Hesselman in B. Eriksson (1986); 3l B. Eriksson (1986); Lundin (2003). 1l 4l

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 55

Fig. XXI. A peacefully grazing reindeer outside an enclosed sample plot on Langfjallet, being observed by the motor-borne, wandering and botanising human being, yet another important factor that affects the mountain vegetation. The photograph was taken in connection with the revision of the grass heath plot on U\ngfjallet, in the summer of 1998. grazing by reindeer (Sjors 1956, 1971, Andersson et al. within the low alpine region. It changes to species-rich 1978-1 984, Oksanen 1992). plant communities if influenced by lime (Andersson et al. 1978-1984): Poa alpina-Ranunculus acris meadows with e.g. Bistorta vipara, Parnassia palustris and Bartsia Grass heath alpina (Sjors 1999). The abundant (mobile) seepage water Thin vegetation, dominated by Carex bigelowii, Des­ gives the meadow vegetation at Mellanbyn the occasional champsiajlexuosa, Festuca ovina andNardus stricta. There appearance of a fen. is a gradual transition to dry heath. It occurs mainly in the higher and northern parts of the mountains (Andersson et Dry heath al. 1978-1984) , and diffe rs from the meadow by being well irrigated only during and immediately after snowmelt. Dominance oflow dwarf shrubs, e.g. Va ccinium myrtillus, The study site consists of a snow-protected heath of Empetrum hermaphroditum and low Betula nana with Nardus stricta-C. bigelowii type, Mat-grass type (Pablsson some narrow-leaved grasses. The closure of the plant 1998), that corresponds closest to D. jlexuosa-N. stricta cover is generally low, but varies with snow cover. The heath (Sjors 1956). The grass heath has a fairly small dry heath type covers large areas in the low alpine region, extent, but has been included in the study for long-term and changes to species-rich plant communities under the observations of whether it can change, after the cessation influence of lime. of reindeer grazing, to a wind heath of type Cladina alpes­ In the southern mountains, low-growing Calluna tris (Pahlsson 1998), or to a wind heath of type Mountain vulgaris dominates, with some Empetrum nigrum, Va c­ cinium vitis-idaea, myrtillus, Arctostaphylos alpina Crowberry (Empetrum hermaphroditum) which, accord­ V. ing to H. Mellkvist, County Administration, Falun (pers. and a number of lanceolate grasses. The bottom layer i comm.), formerly dominated the area. dominated by Cladonia and Cladina spp. The closure of the plant cover is generally low. A variant in wind-exposed positions is the Dry heath with lichens, where Cladina Meadow with low herbs spp. and Cetraria islandica dominate the plant cover, with Grass- and herb-dominated vegetation types restricted some Empetrum hermaphroditum, Calluna vulgaris, Phyl­ to soils with relatively long-term protection by snow, lodoce caerulea and Arctostaphylos alpina (Andersson et and often with mobile soil water. The extent is largest al. 1978-1984).

Acta Phytogeogr. Suec. 87 56 0. Erikssson et al.

Fig. XXII. The revision of the control plot on Poullanvare in the summer of 1996. The vegetation type is Dry heath. The heath vegetation is dominated by crowberry (Empetrum), and the dominant lichen species is Stereocaulon paschale. In the mountain birch forest below the mountain, large areas damaged by the larvae of the autumnal moths (Oporinia/Epirrita and Operophthera) are visible.

The study areas on Dry heath at Tj uolmajaure, Tavva­ development, but can be influencede.g . by shifts in climate vuoma (including Puollanvare ), Sanfjallet and Langfjallet and by the mass occurrence of insects, such as Oporina are of Mountain Crowberry type (Pahlsson 1998), which autumnata and Operophthera brumata (Tenow 1972). The corresponds to Flechtenreiche Empetrum ass. (Fries 1913) . defoliated branches during the winter provide inadequate Va ccinium myrtillus and Cladina spp. benefitfrom protec­ protection against snow, and in consequence, windbroken tion against wind and snow. On Fulufjallet, only sporadi­ trees as well as deep snowdrifts can be seen close together cally visited by reindeer herds, as well as Langfjallet and (Sjors 1971). Sanfjallet, of which the latter two are grazed by reindeer, The present study sites below the tree limit are situ­ there is Dry heath with lichens (Andersson 1982) in some ated in high-lying parts ofthe mountain birch region. The areas outside our sites. Th is corresponds to the Cladina altitude varies between 580 m (Tavvavuoma) and 940 m alpestris type (Pahl son 1998) and Flechtenreiche Betula a.s.l. (Sanfjallet). At Tavvavuoma and on Sanfjallet, the nana-Gebtisch-ass. (Fries 1913). tree layer is completely dominated by Betula pubescens var. czerepanovii, whereas minor occurrences of conifers can be noted on Langfjallet and Sanfjallet. Mountain birch forests Pahlsson ( 1998) describes four types and seven variants The zone of mountain birch forests in Sweden, covering of mountain birch forest. The present study concerns two, about 22,000 km2, is largely a northern oceanic phenom­ viz. Mountain birch forest-dwarf shrub type, which most enon that forms a distinct altitudinal zone between the closely corresponds to birch forest heath type with lichens coniferous forest limit and the open mountain. Betula (Andersson et al. 1978-1984), and Mountain birch forest pubescens var. czerepanovii completely dominates the of dwarf shrub-grass type, which corresponds to birch tree layer. forest heath type with mosses (op.cit.). Mountain birch forests are found not only in the The study areas containing the birch forest heath type Scandes, on the Kola peninsula and in the northern Urals, with lichens are situated at Tavvavuoma, 520 m a.s.l., but also on a minor scale on southwestern Greenland, and on Fulufjallet at 860 m a.s.l. (cf Table 4). They cover southern Iceland and possibly in Scotland (Sjors 1956). fairly large areas, mainly in the north, east and inner parts Several researchers, e.g. Hahmet-Ahti (1963) and Sjors of the mountain chain. On poor and dry land trees of this (1999) today include the mountain birch forests in the type often have multiple stems, are thin, crooked and fairly heterogeneous, northern boreal subzone-however, 4-6 m tall (Carlsson et al. 1999). Juniperus communis as a part that almost entirely lacks conifers. The mountain may form a sparse shrub layer. Dwarf shrubs, mainly birch forest is a relatively stable final stage in vegetation Calluna vulgaris and Empetrum spp., are found only in

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 57

patches. The plant cover of the bottom layer is domi­ as proposed by Pablsson ( 1998), it is a result of long-term, nated by lichens, mainly Cladina spp. (Andersson et al. intensive grazing by reindeer. 1978-84). Fries (1913) coined the name Empetrum-reicher Dry heath is found, as is grass heath, along the entire Flechtenbirkenwald. Nordhagen (1943) used the name mountain range, and covers largeareas almost everywhere. Betuletum empetro-cladinosum, alternatively Betuletum To a lesser extent it is also found on open areas slightly empetro-hylocomiosum. Sjors ( 1971) called it a lichen-rich below the tree limit, where it can form undergrowth in the mountain birch forest. mountain forests (Sjors 1956). The birch forest heath type with mosses (Andersson Meadows with low herbs form a rather sparsely occur­ et al. 1978-1984 ), used in some of the present studies, is ring vegetation type. Only on the map-sheets Sitasjaure found at Tavvavuoma and on Slmfjallet, Uingfjallet and and Fatmomakke does the type cover more than 10% of Fulufjallet. The altitude varies from460 m (Tavvavuoma) the ground surface. This vegetation type is restricted to to 880 m a.s.l. (Fulufjallet). As regards size, it is the larg­ snow-protected ground with mobile ground water (Sjors est of all mountain birch forest types (cf. Table 4), and 1956). is found throughout the entire mountain chain (Pablsson Birch forest-heath types are found along the entire 1998). This type of forest is low, and has an undergrowth mountain range, and in northern Norrbotten, also to the of dwarf shrub species such as Va ccinium myrtillus and east of it. They cover about 22,000 km2. In dry locali­ Empetrum spp., as well as grasses such as Deschampsia ties with little warmth, they are low (4-6 m) and mainly flexuosa and herbs, e.g. Solidago virgaurea and Cornus multiple-stemmed. Single stems, up to 12 m tall, are found suecica. Mosses are more common than lichens in the in moist, nutrient-rich sites with good snow protection bottom layer (Pablsson 1998). Fries (1913) calls it Moos­ (Carlsson et al. 1999). birkenwald. Nordhagen ( 1943) calls it Betuletum myrtillo­ Birch forest-heath types with lichens have their greatest hylocomiosum, Hahmet-Ahti ( 1963) subalpine Empetrum extent in northern Norrbotten, whereas the birch forest­ myrtillus type or subalpine Empetrum type, and Sjors calls heath type with mosses is found along the entire mountain it a blueberry-rich mountain birch forest, or a mountain range, although to a lesser extent in the far south. crowberry birch forest. The dry heath and the birch forest-heath type with lichens at Tavvavuoma and the 'old' dwarf-shrub heath at Puollanvare are all within Saarivuoma Saami village. Extent of vegetation types studied Tj uolmjaure's 'old dwarf-shrubheath' and Tavvavuoma's An estimate of the proportion of the selected types of moss-rich mountain birch forest arewithin Lainiovuoma vegetation in the mountain plant cover is given in Table Saami village (here 'old' refers to experiments initiated in 7. It has been compiled using data from Andersson et 1967-1968). The dwarf-shrub heath at Ritsem is within al. (1978-1984), Rafstedt (1987) and Albertsson et al. Sorkaitum Saami village, but the meadow is within Mel­ (1986-87). lanbyn. Sanfjallet lies within Mittadalen Saami village, The 'mountain area' was delimited by Andersson et and Langfjallet within Idre new Saami village. Fulufjallet al. (1978-1984 ). It should be noted that some of the low is completely outside the reindeer-husbandry area, but mountains along the eastern edgeof the mountain range within the former range of the wild reindeer (Rangifer fall outside the map that covers montane vegetation. Raf­ tarandus L.), now extinct (Lonnberg 1909). This area is stedt (1987) gives an example of a vegetation map of the now included in the National Park established in 2002. area to the east of those that are covered by the map of The seven study sites are situated within six Saami villages montane vegetation. It illustrates the vegetation on the low with varying sizes of reindeer herd. In purely legal terms, mountains Sanfjallet, Ve mdalsfjallen and KlOvsjofjallen, all study sites in the Tavvavuoma area are situated within and that within the surrounding area of northern boreal Lainiovouma Saami village. However, a fence which subzone 'forest' as defined by Sjors (1999). separates the Saami villages and which does not entirely A picture of the northern boreal subzone's mountain follow the administrative boundary, leads to the result that birch forests outside the mountain range is given by Al­ Saarivuoma's reindeer graze on the dwarf-shrub heath and bertsson et al. ( 1986-1987). These forests cover ea. 1,900 in the moss-rich mountain birch forest in Tavvavuoma, as km2 in northern Norrbotten. The low mountain area Pes­ well as on Puollanvare's dwarf-shrub heath. sinki, with extensive reserves of primeval montane forest, is in this area. Within the northernboreal zone, these forests extend as far as the Norwegian Sea and to the coast of the Barents Sea (Tuhkanen 1980). Grass heath is found to some extent along the entire mountain range, but has its greatest extent farthest north and in central and southern Norrbotten. It is not known whether this extent is conditioned by climate or whether,

Acta Phytogeogr. Suec. 87 58 0. Erikssson et al.

Table 7. The presence and cover of vegetation types. From: Ve getationskartan over de svenska fj iillen (Andersson al. 1978-1984), et Ve getationskarta over norra Hiirjedalens skogsland (Rafstedt 1987) and Ve getationskarta bladen Karesuando, Soppero and Lannavaara (Albertsson al. 1986-1987). et

Birch forest - Birch forest - Meadow Heath type Heath type with with with County Map Area Grass heath Dry heath low herbs lichens mosses Notes I

km2 % % % km2 % %

Bd 1 Treriksroset 3550 444 12.5 1402 39.5 53 1.5 284 3.5 694 8 Tj uolmajaure3l Tavvavuoma3l Bd 2Abisko 3060 337 11 459 15 275 9 0 0 214 7 Bd 3 Rensjon 2366 24 1 308 13 <12 <0.5 450 19 544 23 Bd 31L Karesuando 417.4 0 0 9.4 2 0 0 14.9 3.6 99.9 24 one sheet Bd 30K Soppero 2500 21.7 236.1 9.4 6 <1 128 5.1 858 34 four sheets Bd 30L Lannavaara 2594.4 0 0 68.8 0.3 0.1 <1 63.3 2.5 746. 1 28.8 four sheets Bd 4 Sitasjaure 3110 559 12 591 19 389 12.5 0 0 187 6 Ritsem3l Bd 5 Kebnekajse 2500 375 15 338 13.5 175 7 25 200 8 Bd 6 Virijaure 2980 536 18 656 22 253 8.5 0 0 <15 <0.5 Bd 7 Sarek 3500 438 12.5 648 18.5 53 1.5 35 403 11.5 6. Bd 8 Sulitelma 1900 266 14 266 14 95 5 0 0 114 6 Bd 9 Kvikkjokk 2500 375 15 275 11 75 3 25 1 225 9 Bd 10 Pieljekaise 3170 380 12 443 14 159 5 0 0 666 21 Ac 11 Umfors 2487 137 5.5 560 22.5 187 7.5 0 0 460 18.5 Ac 12 Ammamas 2500 125 5 500 20 125 5 0 0 600 24 Ac 13 Tarna 2950 103 3.5 339 11.5 148 5 0 0 738 25 Ac 14 Fatmomakke 3560 178 5 409 5 196 11.5 0 0 712 20 Z 15 Frostviken 3320 17 0.5 338 10 68 2 0 0 406 12 Z 16 Hak.afot 1670 8.5 0.5 142 8.5 25 1.5 0 0 159 9.5 17 Kolasen 3350 100 3 636 19 67 2 0 0 67 2 z Z 18 Storlien 2600 156 6 624 24 130 5 0 0 130 5 Z 19 Are 3100 155 5 465 15 31 <1 31 <1 310 10 Z 20 Funasdalen 3030 61 2 758 25 91 3 91 3 333 11 Z 18 D.E.N Harjedalens 19 D. E skogsland 5625 34 0.6 360 6.4 0 0 73 1.3 180 3.2 one sheet Si'mfjallet 3l W 21 ldre 2920 29 321 11 <15 <0.5 29 58 2 Langfjallet 3l W 22 Saten 3190 48 1.5 239 7.52) 32 1 16 0.5 64 2 Fulufjatlet 3l

1) Bd - Norrbottens Uin. AC - Vasterbottens liin. Z - Jiimtlands liin. W - Dalarnas liin; 2) Including 3 % dry lichen dominated heath; 3) Study site.

Acta Phytogeogr. Suec. 87 3. The WWF-project: How it was carried out

Establishment of sample plots external influences, and it was decided that a system of open sample plots could be used. During 1995, however, At all study sites except Puollanvare and Tj uolmaj aure, a reindeer were seen on the mountain and droppings were subjective choice was made in 1994 of six adjacent plots, observed. As a result, the sample plots were modified in 25x25 m, for each vegetation type, in which-the vegetation 1996 to consist of three open and three fenced plots per was considered to be as similar as possible. Ridges and vegetation type on all study sites. crests were avoided because they make up only a minor At Puollanvare and Tj uolmajaure, a pair of plots part of the total area, and because they are largely affected 12.5x 12.5 m (one fenced and the other open) were laid out by both weather and wear-and-tear.The plots were laid out on each site in 1967-68 by Eriksson. The identification with the diagonals in a north -south and east-west direction 0. markings in the terrain were similar to those at other sites and were delimited by corner crosses and cornermarkers (i.e. in the project). Table 8 lists vegetation types, of steel h ered into the soil. To avoid edge effects, all WWF amm height above sea level, plot size, number of plots, year of records were later made on 22x22 m net areas, the corners establishment, and year of second inventory of each type of which were also marked with steel pegs. of vegetation, for all study sites. A preliminary ordination approach, bas�d on collected Within each fenced plot, a maximum of eight subjec­ field data, provided a basis for dividing the six sampling tively chosen 0.5x0.5 m quadrats were selected, and per­ plots into pairs with similar vegetation, where one plot was manently marked for photography at each revision. These 170 fenced and the other left open. Fences consisted of quadrats were intended to illustrate the type of vegetation cm high reindeer fencing material. Each relevant Nature that dominated in the original situation, and subsequently Management unit decided the type of fence post used. to illustrate any changes in plant composition, degree of Small herbivores, rodents and insects, still had free e.g. cover and frequency of various species in the field layer, access to the plots. that might have occurred. When the project began in 1994, the vegetation on Fulufjlilletwas considered not to be exposed to measurable

Table s. Vegetation type, height above sea level, plot size, number of plots (o=open f=fenced), year(s) of establishment,. and yearof + second inventory of each type of vegetation, for all study sites.

Site Vegetation type Alt. area No. of plots Established 2nd inventory (m) (m) (o+f)

Tavvavuoma Dry heath 580 25 25 3+3 1995 1999 X Birch forest heath type with lichens 520 25 25 3+3 1995 1999 X Birch forest heath type with mosses 460 25 25 3+3 1995 1999 X

Ritsem Dry heath 840 25 25 3+3 1995 1997 X Meadow with low herbs 820 25 25 3+3 1995 1999 X

Sanfjallet Dry heath 910 25 25 3+3 1995+1996 1998 X Birch forest heath type with mosses 940 25 25 3+3 1995+1996 1998 X

Dry heath 840 25 25 3+3 1995 1998 Langfjallet X Grass heath 1010 25 25 3+3 1995 1998 X Birch forest heath type with mosses 800 25 25 3+3 1995 1998 X

Fulufjallet Dry heath 930 25 25 3+3 1995+1996 1999 X Birch forest heath type with lichens 860 25 25 3+3 1995+1996 1999 X Birch forest heath type with mosses 880 25 25 3+3 1995+1996 1999 X

Poullanvare Dry heath 570 12.5 12.5 1 + 1 1967 1997,1999 X

Tj uolmajaure Dry heath 865 12.5 12.5 1 +I 1968 1997,1999 X

Acta Phytogeogr. Suec. 87 60 0. Erikssson et al.

Composition of the ground layer Data processing

To assist studies of the field and bottom layers, each net Descriptions of the composition of a plant community area was mapped and divided into 484 subplots of lxl m that include only the number of taxonomic groups (e.g. (O.SxO.S m at Puollanvare and Tj uolmajaure). Of these, species) cause vital information on community structure to 20 subplots were randomly chosen and measured, starting be ignored. No knowledge is obtained on whether certain from the cornermar kings of the net plots. The percentage species are common or rare. To obtain a measure of species cover of each individual species, of litter, mineral soil, etc., richness in a community, it is customary to use various on the subplots was then estimated. diversity indices, which not only consider the number of Where species determination and quantification were taxa, but also individual degrees of cover, biomass per unit impossible without destructive sampling� some species area, or distribution of cover between taxa. were grouped together. One example is the group of In the present study, use was made of Shannon's di­ juvenile Cladonia/Cladina lichens, which are difficultto versity index (Shannon 1948), whereby it was possible to determine in the field. Consequently, in what follows, the calculate species diversity at the different study sites, as term 'species' may also include genus, species group or well as the types of vegetation according to the formula: other class. T. = In H -"""-� _LT ____L.. i=I T, T,ot Shrub and treelayer where = Shannon's diversity index, = cover of a H r; species or group of species, = total cover of all species To be able to describe the shrub and tree layer in the net Tt ot and groups of species, and s = the number of identified plots for the birch forest-heath type with mosses and birch species/groups of species. The number of taxa found, and forest-heath type with lichens, the detailed plan was used to how evenly they are distributed in the ground layer, thus select sixteen areas S.SxS.S m, of which six were randomly determines the value of The greater the number of chosen, and were measured from the corners of the net H. taxonomic groups, and the more evenly distributed they plot. Estimates were then made of species composition, are, the higher is the index value. In actual communities, degree of cover, height, diameter (base, breast, crown) His often between 1 and 6 (Stiling 1996), and maximum and individual density. Grazing pressure on each subplot diversity occurs when all species have degrees of cover was also determined. that are of the same magnitude, i.e., = ln Hmax s. Before the various statistical tests were carried out, Presence of herbivores the recorded species were divided into groups according to Table 9. The subdivision was based on the occurrence The presence over time of hares, reindeer and elk within of the species. Those that occur in most vegetation types the studied plant communities was estimated by count­ or which are characteristic of a certain vegetation type, or ing droppings (Lavsund 1975) in six areas 5.5x5.5 m per both, were not grouped, but were treated individually. One 25x25 m plot. Theage of thedr oppings was not considered. vegetation type, Meadow with low herbs in Mellanbyn,was Traces of browsing in the shrub and tree layer were also given its own grouping, because its species composition noted. These records were not made at the Puollanvare differed more compared with that of the other types. and Tjuolmajaure sites. In the statistical processing of the collected data, the vegetation types birch forest heath type with mosses and

Table Shannon's diversity index for fenced and open plots in the moss-rich birch forest and the dry dwarf-shrub heath at Tavvavuoma, 9. Sonfj ill.let, Langfja.Iletand Fulufjallet. In each vegetation type and at each site three fenced and three open plots were studied, each with an area of 22 m x 22 m.

Ve getation type Plots Tavvavuoma Sonfjallet Uingfjallet Fulufjallet

Birch forest with mosses Fenced 1995 1.867 2.399 2.715 2.661 1) 1) Fenced 1999 2.259 2.409 2) 2.471 2) 2.779 Open 1995 1.883 2.405 2.639 2.880 Open 1999 1.948 2.508 2.532 2l 2.724 2l Dry heath Fenced 1995 2.291 1.757 2.807 2.423 1) 1) Fenced 1999 2.894 2.381 2l 2.858 2l 2.483 Open 1995 2.273 1.718 2.625 2.464 Open 1999 2.924 2.311 2) 2.866 2) 2.575

= Inventory, 1996; 1) 2) = Inventory, 1998.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 61

Table 10. Conversion table for thepercentage cover of species. limit, but which nevertheless are very dissimilar. We can then make another line, on which the plots can be located Cover in percent Value in PCA in a coordinate system with two axes, still with vegetation

0.0001 -2.0 2 similarities as the common base. We would then probably 2. 1 - 3.0 3 have few meadow plots at one end, and heath plots at the 3.1 - 5.0 4 other end of this axis. 5.1 - 12.5 5 Thus thefinal result will be a two-dimensional ordina­ 12.6 - 25.0 6 25.1 - 50.0 7 tion diagram, with points that identify the differentplots. 50.1 -75.0 8 The closer two plots are to each other, the greater is their 75.1 - 100 9 similarity in vegetation. Species can also be included in the diagram in the same way. Species that usually occur together will then lie in the same part of the diagram. An arrow that starts from the centre represents each species dry heath were analysed simultaneously. Thus a a balanced and ambient variable. The arrow for a certain species layout, with four replicates for each vegetation type, was points towards those plots on which it is most abundant. obtained. The dry heath in Ritsem was not included in this The length of the arrow is proportional to the degree of layout, and was treated separately, because this site lacked cover of the species. If many short arrows point towards the vegetation type birch forest heath type with mosses. the same group of plots, one can conclude that those plots The vegetation types birch forest heath type with lichens, have a high diversity. The direction in which an arrowfor a grass heath and meadow with low herbs, were also treated certain ambient variable points is the direction in which this individually; the same applies to the sites at Puollanvare variable increases. If the arrows for two or more ambient and Tj uolmajaure. variables point in the same direction, we may conclude that those variables are positively correlated. The diagram may thus be regarded as a graphical Ordination summary of basic data. As was mentioned above, plots To obtain a picture of changes in the composition of the with similar vegetation are situated close together in the vegetation over the years, we carriedout a PCA (principal diagram. This means that we can obtain an idea of the component analysis) using the program CANOCO version magnitude of changes in vegetation with time, by seeing 4.0 (ter Braak & Smilauer 1998) for each vegetation type, how much the plots have moved in the diagram from year with all study sites included. An exception was the dry to year. One can also study the effect of different environ­ heath at Ritsem, which was treated individually, separately mental variables on the vegetation composition. from the other dry heath localities. Mathematical descriptions of ordination methods can S The percentage cover of the species/species groups was be found ine.g. ter Braak & milauer ( 1998), Orl6ci ( 1978) revised according to a 9-degree scale (Table 10). and 0kland (1990). All species were present in the analysis. Cover by shrubs, shrub height, cover by trees, tree height and drop­ Cover pings were used as passive environmental variables. PCA, which is an ordination method for indirect gra­ To make a closer study of changes in vegetation with time, dient analysis, implies that the two least similar sample we compared the cover values of the species/species groups plots are chosen and placed one at each end of a line; other from the two inventory occasions. The statistical method plots are then arranged along the same axis, depending on used was a mixed-model ANOVA from the program their floristic similarityto the plots at each end. It should package SAS (SAS Institute, 2001). The intention of an be noted that the plots are not arranged along a gradient ANOVA is to search for significant differences between in the terrain, although it is reasonable to assume that they mean values, by comparing variances. will nonetheless be dispersed along the most important In a study in which several variables or variable groups environmental gradient, since the gradient should correlate are to be compared, an analysis of variance is preferable, with the vegetation's composition. e.g. to the t-test. A reason for this is that the number of t­ Naturally, the gradient concerneddepends on the plots tests increases as a function of the number of groups, and at the ends of the line. We may assume, for example, that the more tests are performed, the higher is the probability the gradient is from the birch forest to the open mountain. of making at least one type 1 error in the analysis, i.e. of This gradient may take on differentappearance s, depending rejecting the null hypothesis. upon topography, soil type, etc. Probably, some of the plots The term 'mixed' means that the model tested consists in our imaginary study are in heath vegetation and others of both fixed variables and random variables. A random in meadow vegetation. Somewhere in the middle of the variable is characterised by normally consisting of a line we may then find plots which all are close to the tree random selection, e.g. from a population, if a population

Acta Phytogeogr. Suec. 87 62 0. Erikssson et al.

Table 11. Models for mixed-model AN VA conducted on birch forest heath type with mosses ( =BFM) and dry heath ( =D H) together, as 0 well as individually on birch forest heath type with lichens (=BFL), Ritsem dry heath (=DH), meadow with low herb (=meadow), grass heath ( =GH), Puollanvare ( =Poul) and Tjuolmajaure ( =Tjuol), where the degree of cover of a species/species group is tested against the influencing factors and the interactions between them.

Model l Model 2 Model 3 Model 4 BFM +DH BFL Ritsem, DH, meadow, GH Poul., Tj uol. year year year year site site plot(tre atment) treatment veg plot(site*treatment) treatment plot(site*veg*treatment) treatment year* treatment treatment Year*plot(treatment) year*plot(site*treatment) Year*treatment year*veg year* site year* site year*tre atment year* treatment site*treatment veg*treatment site*treatment year* site*treatment year*veg*treatment year*site*treatment site* veg*treatment year*veg*site year*veg*site*treatment

study is concerned. In the present case, the 20 subplots that had a total cover of more than 100%. The transformed were randomly chosen within a plot (25x25 m), which degrees of cover then constituted the response variable, means that a 'sample plot' is a random variable. A fixed which we analysed against different influencing factors variable, on the other hand, has the same value or level and against interactions between the factors. The birch throughout. The time of the inventory (year 0 and year forest heath type with mosses and dry heath were tested 1, respectively), the treatment (fenced or open area, re­ together, whereas the dry heath at Ritsem, birch forest spectively), the site and type of vegetation, are the fixed heath type with lichens, meadow with low herb, grass variables in our study. heath and the sites at Puollanvare and Tjoulmavare, were For the ANOV A, we converted the cover values for the tested individually. The complete models for all analyses species and species groups according to the formula: are given in Table 11. With the species/species groups model l, thatshowed Transformed degree of cover = arcsin %cover/100 in -J significant differences in any of the interactions year x Since it is not possible to calculate the arc sin of numbers treatment, year x type of vegetation x treatment, year x larger than 1 , the value 1 00 was used for species groups site x treatment and year x type of vegetation x site x treat­ ment, additional models were constructed and analysed. These analyses were made for each type of vegetation, but for one site at a time. In a similar manner we treated the Table 12. Model for the mixed-model ANOVA used on the species/species groups in model 2, that showed significant shrub and tree layer in birch forest heath type with mosses and differences in any of the interactions year x treatment and lichens. yearxsitextreatment. The model we used for these tests was the same as model 3 in Table 11. Here, the interaction Factors yearxtreatment is the most interesting, because it can give year an indication of any changes in the degree of cover with site time as a response to the treatment. The same approach plot(site*treatment) was also used for model 4. treatment

Ye ar* site Frequency Ye ar*treatment Ye ar*area(site*treatment) We also investigated whether there had been any changes Site*treatment in the occurrence (frequency) of a species/species group. Ye ar*site*treatment The analysis was made by means of the logistic regres-

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 63

sion module from the SAS package (SAS Institute, 2001), ANOVA, we were mainly interested in the interaction year treatment. and we used the same species and species group as for the x analysis of variance (Table 9). Logistic regression is a variant of common linear Shrub and tree layer regression, and can be used when the response variable is binary, i.e. it can only assume two values (generally 1 and To trace changes in the shrub and tree layer between years 0). The regression examines the relationship between one in the moss-rich and lichen-rich birch forests, we used a or more independent variables, and the probability that a mixed-model ANOVA. We compared the crown cover of response variable will assume a certain value. Examples of shrubs and trees individually, as also the basal area (m2 a binary variable may be mortality(living = 1, dead = 0), or ha-1) of the tree layer. All individuals taller than 200 cm the frequency of a species (occurrence = 1, non-occurrence were treated as trees. The tests were made on each veg­ = 0). Use can also be made of proportions, when they are etation type individually; the model ANOVA is shown in based on binary data. Table 12. We used the occurrence of a species/species group The variables that showed significantresults in any of on a large plot as the response variable. For all sites, the the interactions year x treatment and year x site x treat­ same logistic regressions were then calculated, whereby ment were used in a further ANOVA. This time we tested each type of vegetation was treated separately. We tested each site individually, using the model: year, treatment, the response variable against year, area, and treatment, plot( treatment) and year x treatment. and against the interaction year x treatment. As with the

Acta Phytogeogr. Suec. 87 64 0. Erikssson et al.

4. The WWF-project: Results

Birch forest heath type with mosses the two inventories in Sanfjallet's moss-rich birch forest. 4. 1 Arctostaphylos alpinus was not found at the inventories in and dry heath 1995/96, but was found on the open plots in 1998. Phyl­ lodoce caerulea and Cetraria ericetorum are two new Birch forest heath type with mosses are found at the four species on the fenced plots. localities Tavvavuoma, Sanfjallet, Langfjallet and Fulufjru­ In Langfjailet's birch forest-heath type with mosses, let, whereas dry dwarf-shrub heath is the only type found Cetraria nivalis was found on the fenced plots only in 1995 , on all sites studied by the project. As mentioned above, but had disappeared at re-inventory in 1998. Nardus stricta the dry heath at Ritsem was treated separately. disappeared from the open plots between the inventories, whereas it appeared on the fenced plots. Festuca ovina, Species present Linnea borealis, Arctostaphylos alpina, Nephroma arcti­ cum and Peltigera aphthosa were not found on the open Table 13 shows the number of taxa (species, species plots in 1995, but were found at re-inventory in 1998; the groups, genus, etc.) found on the two inventory occasions same applied to Festuca ovina on the fenced plots. at the different study sites and vegetation types. All taxa At Fulufjall, Eriophorum vaginatum was found only are listed in the Appendix I. The fact that some species on the open plots and Hieracium spp. only on the fenced that are difficultto identify (e.g. juvenile lichens belong­ plots in 1995. Neither was found in 1999. Arctostaphylos ing to the genus Cladonia) were grouped together when alpina was not present in 1995, but had arrived on the it was not possible to determine the species, makes for open plots in 1999. difficulty in deciding from the species list whether such a The dry heaths at the different sites generally had a species actually is present, or whether it has disappeared larger turnoverof species than the birch forest-heath type over the years. with mosses. It may be noted that at Tavvavuoma, Phyl­ In the Tavvavuoma birch forest-heath type with mosses, lodoce caerulea, Diphasiastrum complanatum and Hier­ we found Salix herbacea, Rubus chamaemorus, Cetraria acium spp. were found only on the fenced plots in 1995, ericetorum and Bartsia alpina only in the fenced plots in and that they had disappeared by the time of re-inventory 1995, whereas Cladina stellaris had disappeared from the in 1999. Pedicularis lapponica, on the other hand, was open plots during the years. Huperzia selago and Lycopo­ found only on the open plots in 1995, but had disappeared dium clavatum were not found in 1995, but when the plot by 1999, and Deschampsiaflexuosadisappeared only from was re-inventoried they were found on the open plots, and the fenced plots between the two inventories. Examples the latter species on both the open and fenced plots. Other of newly arrived species on the fenced plots are Solidago species that had arrived by the time of the 1999 inventory virgaurea, Call una vulgaris, Va ccinium myrtillus and Fes­ were Festuca ovina and Equisetum sylvaticum in the fenced tuca ovina, whereas Agrostis mertensii was a new species plots and Stereocaulon paschale on the open plots. on the open plots. Solidagovirgaurea, Stereocaulon paschale and Cetraria In the dry heath at Sfmfjallet, Pleurozium schreberi ericetorum had disappeared from the open plots between was found on both fenced and open plots in 1995/6, but

Table 13. Number of taxa for fenced and open areas in the moss-rich birch forest and in the dry dwarf-shrub heath at Tavvavuoma, Sonfjiillet, Langfjallet and Fulufjallet. In each vegetation type and in each study site three fenced and three open areas were studied, each with an area of 22 x 22 m m.

Vegetation type Plots Tavvavuoma Sonfjallet La.ngfjiillet Fulufjiillet

Birch forest with mosses Fenced 1995 43 24 1) 39 31 I) Fenced 1999 27 2) 43 2l 39 30 Open 1995 37 24 36 33 Open 1999 40 29 2 4Pl 35 ) Dry heath Fenced 1995 30 19 30 30 I) 1) Fenced 1999 41 31 2) 39 2) 30 Open 1995 33 18 32 27 Open 1999 43 29 2l 39 2) 29

1) = Inventory, 1996;

2) = Inventory, 1998.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 65

05 Fulufjlillet ..0.. c Ulngfjallet <> 0:1 Shrubheight Sonfjilllet. 0 O:lcI 0� � Ta vva vuomaQ !:5 0�1• / � 0:1 I:J l• •u C1" 0;1 ....

Shrub cover 06

... ��. 0 � • Droppings ·a ·· ''" ... C1>�

• "''•• 1:3 1:1 ;: o:l :t oa •"" ,uO e t�.e. )!..a I \ 1:6 001:2 . ,,, ;&15. ,0:1 (}.J A .....:o:4 Tn•> <:

0 ·�

-1 .0 +1 .0 -1 .0 +1 .0

4. Ordination diagram (PCA) for study plots in birch forest-heath type with mosses (a) and on the dry heath (b) at Tavvavuoma, Fig. Sanfjallet, Langfjallet and Fulufjallet. The first figure in the identification codes of the plots refers to the year of inventory (O=first inventory, !=re-inventory), whereas the second figure refers to the plot number. Black symbols represent fenced plots. only on the fenced plots in 1998. New species/groups of was found on the fenced plots in 1996, but not in 1999, species on the dry heath are Hieracium spp. on open plots whereas that species had arrived on the open plots at re­ and Juncus trijidus and Loiseleuria procumbens on the inventory. Arctostaphylos alpina is another species found fenced plots. Cladina rangiferinaand Phyllodoce caerulea on the open plots in 1999, having been absent in 1996. are new species on both fenced and open plots. Nardus stricta was found neither on the fenced nor on the Arctostaphylos alpina and Juncus trifidus were found on open plots in 1995/96, but was found on the fenced plots the open plots in theLangf jaJ.let dry heath in 1995, but not at re-inventory. in 1998. Festuca ovina, Andromedapo lifolia and Tr ientalis europaea were absent from the open plots in 1995, but had Shannon's diversity index arrived by thetime of the re-inventoryin 1998. ]uncus trifidus was a new species on the fenced plots in 1998. Table 14 shows species diversity for the fenced and open On the Fulufjall dry heath, Stereocaulon paschale was plots, calculated according to formula 1. In Tavvavuoma's found only on the open plots in 1995. Tr ientalis europaea birch forest-heath type with mosses, diversity increased

Table 14. Shannon's diversity index for fenced and open plots in the moss-rich birch forest and the dry dwarf-shrub heath at Tavva­ vuoma, Sonfjallet, Uingfjallet and Fulufjallet. In each vegetation type and at each site three fenced and three open plots were studied, each with an area of 22 m x 22 m.

Vegetation type Plots Tavvavuoma Sonfjallet Umgfjallet Fulufjallet

Birch for. mosses Fenced 1995 1.867 2.399 2.715 2.661 1) w I) 2.259 2.409 2.471 2.779 Fenced 1999 2l 2l Open 1995 1.883 2.405 2.639 2.880 Open 1999 1.948 2.508 2.532 2.724 2) 2l Dry heath Fenced 1995 2.291 1.757 2.807 2.423 I) I) Fenced 1999 2.894 2.381 2l 2.858 2l 2.483 Open 1995 2.273 1.718 2.625 2.464

Open 1999 2.924 2.311 2l 2.866 2l 2.575 =Inventory, 1996; 1) =Inventory, 1998. 2)

Acta Phytogeogr. Suec. 87 66 0. Erikssson et al.

during the years, but the increase was significantly greater and V. vitis-idaea are the most common forms of dwarf on the fenced plots (p<0.05, Mixed-model ANOVA). On shrub, and Deschampsiajfexuosa is almost the only grass FulufjiHlet, diversity increased on the fenced plots, but de­ present. The result of the analyses of variance shows that creased on the open plots (p<0.05, Mixed-modelANOVA), the degree of cover of Deschampsia flexuosa, Va ccinium and on Langfjallet, species diversity generally decreased, myrtillus and V. vitis-idaea changed significantlyover the without any differences between fenced and open plots years, as a response to fencing. Deschampsia ftexuosa (p

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 67

Tavavuoma MRB

90 Kr.lk Bla Ling Oris Cctr 80 Faclor Df Krus* * * ** 70 0 Area 4 60 50 Treatment *** * � *** � 40 Year u 30 Area ** ** 20!0 * YearYear* Treatment 4 *** **" ••• 0 1

MRB Son(jiillet

F Krus Krak Bla Lin Oris Ctllr aclor Df g *** Area 4 "** *** *** * ** Treatment I * ** * Year 1

Y ear* Area * "* Year* Treatment 4 * * * I

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1 45 Factor Df Bl!\ Ling Oris Krus KrAk Cetr Area *** *** * 4 *" "* Treatment 1 * ** *** *** Year 1

Year* Area 4 - Year* Treatment 1 * * �r)��-I�I:;�r'-t� . ����--��lf' l�i'-��:��' 0' L• ,� - , .. J . J ]' ,li,> _[-�, � .. .. i 95 98 j 95 98 i 95 98 ! 95 98 1 95 98 : 95 98 I 95 98 ..1 95 98 i 95 98 95 98 1 95 98 j 95 98 I ! f l ! .�ll i �J_JI t i � ! l r•l········--············ � .. ·· . l I i Krus I Grhg ! . Kn\k I Bl� i Ling i Lj un i Oris I Mos i Clad ;:�Cetr I �'t er... .. i ���Ovr. �.• I

M RB Fulufj!lle:i

25 -, Factor Df Krus BIA Lin� Oris Cetr *** Kclk*** *** * *"* Area 4 Treatment Year 1 * ** *** I *

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5. Cover % of species and groups of species in the moss-rich birch forest on the two inventory occasions at Tavvavuoma, Sonfjiillet, Fig. Langfjallet and Fulufjallet. Black columns show fenced plots, white columns show open plots, and the error columns show the standard error (SE). The tables show the result of the ANOVA calculated individually on each site. Significance is given with * for p<0.05, ** for p

Acta Phytogeogr. Suec. 87 68 0. Erikssson et al.

Tavavuoma RlS

40 - Factor Df Krak Krus Bia Ling 6ris Cetr Area 4 **• **• "'"'"' Treatment Year "'* ......

Year* Area * 4 jif��::tt:=r,�:r.�,��li Year* Treatment 1 ** j 95 99 j 95 99 : 95 99 i 95 99 i 95 99 i 95 99 I 95 99 i 9:5 99 i 9$ 99 ! 9S 99 i 95 99 i 95 99 ! i , : • : i IJun : : i : : ! ( Kn!S ! C.rhg KrAk ) RIH :! l.ing ! 1 Oris j Mos ! Clad [ Cetr [ Ster i Ovr i

SoafjlilletRIS

70 ' Factor Df Krus Knik Bla Ling 6ris Cetr · · - 60 +------······ ·-- ... --···· ··· - ·-·--·------Area 4 *** * *** *** *** *** Treatment Year *** ..... *• ***

* Year* Area 4 * * ** i��3� Year* Treatment 1 * *** 9s 98 i 9s 9& i 9s 98 i 9s 98 i j 95 98 j 95 98 i 95 98 ) 95 98 1 9s 98 ! 95 9& i 9s 98 i 95 98 i j ! ; : : : ! : l j l l i : Krus : Grhg i Krnk ! Bl

LAngfj�lletRIS

35 ! Factor Df Krus Krak Bh\ Ling 6ris Cetr Area 4 * ** *** *** *** *** *** -� .' -' �� �- -- ?r______Treatment I "'* ** * *** · · -- - Year - • ------U� �� I 1"J -----;�_2r�:_ 1'1,. - � 1 - - - -- I J-==-- - � r- _, : �-_: : � "' JO ; ------·:' ,. l ' -1J - - - - +------+ ��- ' ' ·f - ' ' I ' I I"" I I * * --- I - i Year Area 5 -�-- . : ----- . t Ill' liT 4 -- � ---�- - �- -::·:� --· r- - 1 ·J ' - Year* Treatment 0 � ��� · J J I�J ij1j : I - -- - Il. ,._lllj : 95 98 95 98 : 95 98 i 95 98 95 98 ! 95- 98 95 98 I 95 98 i 95 98 95 98 195 98 1 9598 I i i I -�i t�l-- j I i I i i I i I I ! 1lrr i i Clad i Ster i Ovr ! i Krus [ Grhg ! KrAk i Bla i Ling j Lj un i Oris [ Mos [ ) Cerr

FulufjlillerRIS

60; Df Krus Krak Bhi 6ris Cctr Factor Ling Area 4 *** * *** *** *** Treatment 1 *** ** Year

Year* Area 4 ** ** * Year* Treatment * * �[�-�:;���:.�]t;=� I 1 95 99 i 95 99 95 99 i 95 99 i 95 99 ' 95 99 195 99 ! 95 99 195 99 1 95 99 i 95 99 ! 95 99 i i Mos Celr i KrusI Grhg Krllk i Bla i Ling i Lj un I Oris I j Clad i [ Ster [ 6vr [

6. Cover % of species and groups of species in the dry heath at the two inventories at Tavvavuoma, Sanfjiillet, Langfjallet and Fig. Fulufjallet. Black columns show fenced plots, white columns show open plots, and the error columns show the standard error (SE). The tables show the result of the ANOVA calculated individually on each site. For explanations see Fig. 5.

treatment, Cetraria spp. and Va ccinium myrtillus increased, larger on the fenced plots. Between the two inventories, whereas Deschampsia jlexuosa decreased. Empetrum hermaphroditum increased its cover on the Most of the ground layer on Fulufjiillet is occupied open plots, and decreased on the fenced plots. No other by lichens belonging to the genera Cladonia/Cladina, species or species group shows similar significant differ­ followed by Empetrum hermaphroditum as the dominat­ ences between fenced and open plots. Cetraria spp. has ing dwarf shrub. Va ccinium vitis-idaea increased on both increased on both open and fenced plots. fenced and open plots, but the increase was significantly

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 69

Tavvavuoma MRB

100% r· Factor Clad 80% Df Area g 60% Treatment * Year "' ! 4(1% 20% · - Area * Year* ____ :1�11 1 : Year*Treatment 0% --'"""·c'-•-'-·r'"•·'-· -:: f : 1 ; ]1 -���,(i::J! l .. 95 99 j 95 99 ! 95 99 i 95 99 ! 95 99 1 95 99 95 99 195 99 i 95 99 i ::Jll .r.·.I t I : M I ; Krus ' Grhg : Ling ! Lj un j Oris 1 os 1 Clad I. Cetr I Ster i Ovr

Sonfjal!et. MRB

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80%

60% 40% J:( 20%

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Langtjit!let M RB

100% -. r- Factor Ljun Oris 80% ; I Df Area * i;' 60% fj I Treatment * 40% I Year �[ 0% 2 I !I Year* Area * * * ** 0"/o I t 11 - Year Treatment '95 9& 95 98 95 98 i 95 98 95 98 ! Krus Grhg Krak B!a Ling i

Fulutj!illetM RB

100%

-: 80% 1 n Factor Df Ljun Oris Area -1� -�� l Treatment 60% ·t - 1 r 40% I Year ! -t l 20% * j i = Year Area * i rf * ** * 0"/o J ,--- ' Year Treatment 99 1 9s 99 I 95 99 I i Cetr ! Ster 1 Ovr

7. The percentage of plots with occurrence of a species/species group in the moss-rich birch forest at the two inventories at Tav­ Fig. vavuoma, Sanfjal1et, Langfjallet and Fulufjallet. Black columns show fenced plots, white columns show open plots, and the error columns show the standard error (SE). The tables show the results of the logistic regression calculated individually on each site. For explanations see Fig. 5.

Frequency des/species groups that showed significantdiffere nces in frequency between years, as a response to fencing or both, Figures 7 and 8 show the number of plots with presence of are included in the tables. The results for each vegetation a species/species group in the moss-rich birch forest and types are presented separately below. the dry dwarf-shrub heath, respectively, at the two inven­ Birch forest-heath type with mosses: No large changes tories and on all four sites. The tables in the same figures occurred in the frequency of species/species groups in the illustrate the results of the logistic regression calculated for plots. At only two sites, Lfmgfjallet and Fulufjiillet, were each site and vegetation type individually. Only the spe- significant differences found as a response to fencing. At

Acta Phytogeogr. Suec. 87 70 0. Erikssson et al.

Tawavuoma RIS

100"/o

80%

[;· 60% g 40%

l 20%

0%

Sonfjallet RlS

100%

Oris 80% i 1 r Factor Df Krus Ling Art: a [ 60".10 Treatment * * * [ 40"/o Year ** ***

20o/o "" 1·.fl§j Year* Area ** * 0% :m Year*Treatment *

95 98 95 98 1 95 9sl_ 95 9s l1I 95 98 i 95 98 ! 95 98 ! 95 98 1 95 98 l Bla ! I ! I Krus Grhg , Kn\k i i Ling i Lj un I Oris i Mos I Clad Cetr I Ster

RlS IAngfjMIIet

Factor Df Bla Area I Treatment 1 r ·· · ** �- � ! Year I � ':40"/o �,·_• l ��r -l · �-··· ··· + ··· · ;r �-nr ···· rf·· J J--J'l-1- itf ·+r -fj --- -f I I •, ! � l i - i -·- -r_ -.- _,_ ' " • ,,·_ + : I 20% - . _· . l l \ i • \ ,__ -�- ,,,, \ Year* Area ** . I •I i ! I I . l I I I : I I i ! ! : I * **• . . . · " ' t ... Year Treatment Krus ! Grhg ! Kr.lk ! B!A l Ling j Ljun i Oris 1 M os ) Clad i Cetr I S!er i Ovr

fulufjallet RlS

Factor Krus Ljun Df Area 1 Treatment I I * Year

Year* Area Year*Trea tment **

The percentage ofplots with occurrence ofa species/species group in the dry dwarf-shrub heath on the two inventory occasions Fig. 8. at Tavvavuoma, Sonfjallet, Langfjallet and Fulufjiillet. Black columns show fenced plots, white columns show open plots, and the error columns show the standard error (SE). The tables show the result of the logistic regression calculated individually on each site. For explanations see Fig. 5.

Lfmgfjallet, Calluna vulgaris had become more frequent flexuosaand Va ccinium vitis-idaea increased on the fenced in general, but the increase was greater on the fenced plots but decreased on the open plots, whereas other dwarf plots. Other dwarf shrubs, on the other hand, decreased shrubs showed a general decrease in occurrence, regardless in frequency on both the open and the fe nced plots; the of treatment. Va ccinium myrtillus increased in frequency decrease was significantly greater on the latter. The same on the open plots, but decreased on the fe nced plots on applies to 'other dwarf shrubs' on Fulufjallet, where Cal­ Lfmgfjallet, and the same change was found for Calluna luna vulgaris decreased in frequency on the open plots, vulgaris on Fulufjallet. On Fulufjallet, Deschampsiaflexu­ but increased on the fenced plots. osa decreased significantly, regardless of treatment. Dry heath: On SamjaJ.let,the fre quency of Deschampsia

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 71

Tavvavuotm Mrb

Crown density Basal area Factor Df shrub Area 4 Treatment 1 Year I *** **"'

Year"'Area 4 Year*Treatment I 1995 1999 1995 1999 1995 1999 Crown Crown densityshrub densitytree ! Basalarea

Sonfj!lllet Mrb

� ��70 ��L�;i

1994 1998 1994 1998 1994 1998

Crown densityshrub Crown density tree Basalarea

UlngfjiUiet Mrb

70 �-······ -� ------r-- --: w ; 50 40

30 -·- - r ·· •-· no r � 20 ._ _ l=_ i 10 _ J•;· r'll ,,• 0 : 1rr 1994 1998 1994 1998 1994 1998

Crowndensity shrub · Crowndensity tree Basal area

Fulufiallet Mrh

60 50 -F�------Factor Df Basal area 40 +------J Area 4 Treatment 1 Year 1 ***

Year*Area 4 Year*Treatment 1 1995 1999 1995 1999 1995 1999

Crowndensity shrub Crown density tree Basal area

Crown cover,per cent, of shrubs and trees, and basal area of trees ha-1) at the two inventories at Tavvavuoma, Sanfjiillet, Fig. 9. (m2 Langfjiillet and Fulufjiillet. Black columns show fenced plots, white columns show open plots, and the error columns show the standard error (SE). The tables show the result of the ANOVAs calculated individually on each site. For explanations see Fig. 5.

Shrub- and tree-layer and Fulufjallet, there was also an increase in tree basal area. No significant changes, in either the crown cover of Figure 9 shows the crown cover of the shrub- and tree­ the shrub- and tree-layer or the basal area, were found at layer, together with tree basal area at the two inventories the other sites. in the birch forest-heath type with mosses at Tavvavuoma, Sfmfjallet, Lfmgfjallet and Fulufjallet. A significant de­ crease during the years in the crown cover of the shrub layer was found only at Tavvavuoma. At Tavvavouma

Acta Phytogeogr. Suec. 87 72 0. Erikssson et al.

Table 15. Number of reindeer droppings per hectare in fe nced and open plots in the birch forest-heath type with mosses and in the dry heath at Tavvavuoma, Sfmfjallet, Langfjiillet and Fulufjallet between 1994 and 1995. Values are means ± standard deviation.

Birch forest-heath type with mosses Dry heath

Site Plots Reindeer Moose Hare Reindeer Moose Hare

Tavvavuoma Fenced 1995 129 ±47 Fenced 1999 Open 1995 92 ±45 3 299 ± 989 Open 1999 2167 ± 241 Sanfjallet Fenced 1994 200 ± 101 Fenced 1998 Open 1994 614 ± 174 Open 1998 276 ± 159 322 ± 257 202 ±92 Langfjallet Fenced 1994 69 ± 42 62 ± 5 426 ± 39 Fenced 1998 Open 1994 14 ± 14 7±3 256 ± 38 Open 1998 312 ± 175 Fulufjallet Fenced 1996 147 ± 97 * * Fenced 1999 Open 1995 * Open 1999 18 ± 18

* sporadic occurrences.

Droppings the Mittadalen Saami village (Figure 3). On Langfjallet, the reindeer droppings found in 1994 on the open areas that Table 15 shows the number of piles of droppings per were later fenced, had disappeared by 1998. An increase hectare of reindeer, elk and hare for the two types of in the number of piles of reindeer droppings was noted vegetation and on the study plots at all sites. In the birch on the open plots, but it is not possible to link this with forest-heath type with mosses at Tavvavuoma, reindeer the number of reindeer, which remained fairly constant in droppings were found only in 1995, when all plots were ldre New Saami village during the 1990s (Figure 3). The open. Droppings found on areas that were subsequently inventories on the Fulufjalletdry heath in 1995/96 recorded fenced had disappeared by 1999. A similar pattern was only sporadic occurrences of reindeer and hare droppings. also found for Langfjilllet, where reindeer and elk drop­ At the re-inventory in 1999, no droppings were found. pings were found in 1994, but none were found at the re-inventory in 1998. On Sanfjallet, reindeer droppings were found only in 1998 on the open plots. At Fulufjallet, 4.2 Birch forest-heath type with lichens only elk droppings were found on the fenced plots in 1996, and on the open plots in 1999. No droppings of hare were Species present found in the birch forest-heath type with mosses at any of the four study sites. Two sites with the vegetation type birch forest-heath type No droppings of elk or hare were found on the study with lichens are included in the project, Tavvavuoma and plots on the dry heath at Tavvavuoma, and the amount of Fulufjallet. Table 16 shows the number of taxa (species, reindeer droppings decreased on the open plots between species groups, genus, etc.) found at the two inventories 1995 and 1999. The reduction by half of the reindeer on the two sites, and the appendix lists all taxa found. (The droppings on the open plots on Sanfjallet's dry heath cor­ fact that species which are difficult to determine, e.g. ju­ responds with the reduction in the number of reindeer in venile lichens belonging to the genus Cladonia, have been

Table 16. Number of taxa, for fenced and open plots at Tav­ Table 17. Shannon's diversity index for fenced and open plots vavuomna in 1995 and 1999, and at Fulufjallet in 1995, 1996 at Tavvavuoma in 1995 and 1999, and on Fulufjallet in 1995, and 1999. 1996 and 1999.

Plots Tavvavuoma Fulufjallet Plots Tavvavuoma Fulufjallet

Fenced 1995 44 29 * Fenced 1995 2.487 2.145 * Fenced 1999 46 29 Fenced 1999 2.736 2.238 Open 1995 50 30 Open 1995 2.504 2.390 Open 1999 48 29 Open 1999 2.642 2.345

* Inventory, 1996. Inventory, 1996. *

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 73

grouped together when it was impossible to determine the species to which they belonged, makes it difficult to see from the species list whether a certain species was actually present, or if it disappeared over the years.) At Tavvavuoma, Arctostaphylos alpina and Phleum alpinum disappeared from the open plots, whereas An­ tennaria dioica and Phleum alpinum disappeared from 0:2 the fenced plots. Calamagrostis lapponica and Calluna ("\ \__/ vulgaris were two species that had arrived on the open 0:3 • plots, whereas Hieracium spp., Anthoxanthum odoratum, • \_j("\ 1:2

]uncus trifidus and Peltigera aphthosa were new species 0:1 on the fenced plots. On Fulufjallet, there was no particular change in spe­ cies composition between the two inventories. Polytrichum Droppings spp., however, had arrived on both the fenced and the open plots.

-1

Shannon's diversity index Treeheight

Table 17 shows the species diversity for fenced and open 10. Ordination diagram (PCA) for study plots in the birch Fig. plots, calculated according to formula 1. At Tavvavuoma, forest-heath type with lichens at Tavvavuoma and on Fulufjallet. the diversity on both the open and fenced plots increased The first figure in the identification code refers to the inventory slightly over the years (p

Fulufjlillet LRB

Factor Df Krus Ling qun Moss 6vr Bla Cetr Area * ** *** ** *** *** 4 Treatment 1 Year "'* I

Year* Area 4 Y ear*Treatment * I 0.06

Tavavuorna LRB Factor Df Krus Grh Ling Ovr s Bla Moss Area *** *** 4 Treatment I Year ** *** *** I

Year* Area ** 4 Year* Treatment l

11. Cover, per cent, of species and species groups at the two inventories in the birch forest-heath type with lichens at Tavvavuoma Fig. and on Fulufjlillet. Black columns show fenced plots, white columns show open plots and the error columns give the standard error (SE). The tables show the results of ANOVAs calculated individually for each site. Abbreviations, see Figure 5.

Acta Phytogeogr. Suec. 87 74 0. Erikssson et al.

where both sites were tested, have been included. Table 18. Number of reindeer droppings per hectare on fe nced Mosses dominate strongly at Tavvavuoma, whereas the and open plots in Tavvavuoma's Birch forest-heath type with most common lichens are represented by species belong­ lichens in 1995 and 1999.

ing to Cladonia/Cladina. The result of the analyses of Plots Dropping piles variance show that no species/species group had changed significantly in degree of cover as a response to fencing. Fenced 1995 331 ± 125 Other grasses/sedges, Va ccinium myrtillus and vitis­ Fenced 1999 0 V. Open 1995 275 ±90 idaea increased in general at Tavvavuoma, whereas Des­ Open 1999 643 ± 210 champsiaflexuosaand mosses decreased. Mosses represent the largest decrease in cover degree; they decreased from 1995 to 1999 from about 50% to slightly less than 30%. The reduction is of approximately the same magnitude for The tables in the same figures show the results of the both fenced and open plots. logistic regression calculated individually for each site. The largest proportion of the ground layer in the Fu­ Only the species group (other dwarf shrubs) that showed lufjallet birchfo rest-heath type with lichens consists of li­ significant differences in frequency between years, as a chens belonging to the genera Cladonia/Cladina, followed response to fencing or both, are included in the tables. No by Empetrum hermaphroditum, which is the dominating species or species group showed any significantchange in dwarf shrub. Between the two inventories, Cetraria spp. frequencyin response to fencing, neither on Fulufjallet nor increased its degree of cover on fenced plots, and decreased at Tavvavuoma, but 'other dwarf shrubs' were less frequent on open plots. No other species or species group shows on Fulufjallet, regardless of treatment. similar significant differences between fenced and open plots. Va ccinium myrtillus, V. vitis-idaea and 'other' in­ Shrub- and tree-layer creased on both the open and the fenced plots, regardless of treatment, whereas Calluna vulgaris decreased. Figure 13 shows the crown cover of the shrub- and tree­ layer and tree basal area (m2 ha-1) at the two inventories in the Birch forest-heath type with lichens at Tavvavuoma Frequency and Fulufjallet. At both places, the crown cover in the Figure 12 shows the number of plots with occurrence of a shrub layer increased on the open plots, but decreased on species/species group at both sites at the two inventories. the fenced plots, and the difference is significant.

Tavvavuoma LRB

100"/o Factor Df Oris 80% Area 4 ** Treatment ** 60% 1 i Year 1 * �� 40% 20% Year*Area * 4 1 0% Year* Treatment

Fulufjllllet LRB

Factor Oris Df Area 4 Treatment * * 1 Year 1

Year* Area * 4 Year* Treatment 1 0.06

12. Number of plots with occurrence of a species or species group at the two inventories in the birch forest-heath type with lichens. Fig. Black columns show fenced plots, white columns show open plots and the error columns give the standard error (SE). The tables show the results of logistic regressions calculated individually on each site. Abbreviations, see Figure 5.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 75

Droppings Table 19. Number of taxa for fenced and open plots at Ritsem (dry heath and low herb meadow) and on the Langfjallet grass No droppings of reindeer, elk or hare were noted in the heath. study plots on Fulufjallet. At Tavvavuoma, only reindeer droppings were found (Table 1 8). At the inventory in 1995, Plots Langfja.Ilet Ritsem Ritsem grass heath low herb meadow dry heath all plots were open and the reindeer droppings found on the plot subsequently fenced had disappeared by 1999. Fenced 1995 31 110 57 Fenced 1997 35 124 I) 64 Lfmgfjallet's dry grass heath and Ritsem's dry dwarf­ 2> Open 1995 34 1 3 66 shrub heath and low herb meadow: 1 Open 1997 39 2> 118 72 The vegetation types low herb meadow and grass heath I) Inventory, 1999; are each found only at one place, Ritsem and Langfjallet, 1l Inventory, 1998. respectively, and have been treated separately. As men­ 2) tioned above, the dry dwarf-shrub heath at Ritsem has also been treated separately.

Stereocaulon paschale and Festuca ovina on the fenced 4.3 Ritsem meadow with low herbs and plots. Loiseleuria procumbens and Sphagnum were not dry heath found at all in 1995, but in 1998 Sphagnum was found on both fenced and open plots, whereas Loiseleuria procum­ Species present bens was found only on fenced plots. The meadow with low herbs at Ritsem is the vegeta­ Table 19 shows the number of taxa (species, species tion type with by far the largest number, and turnover, groups, genus, etc.) found at the two inventories on the of taxa. Saxijraga cernua, Lychnis viscaria, Tr ientalis different sites and vegetation types, and all taxa are listed europaea, Botrychium lunaria, Nephroma arcticum and in the Appendix. Salix glauca are examples of species found on the open Pleurozium schreberi and Festuca ovina were found plots in 1995, but which had disappeared in 1999. For on the open plots on La:ngfjallet's grass heath in 1995, fenced plots, Phyllodoce caerulea, Cassiope hypnoides but not at the inventory in 1998. The same applies to and Tr ientalis europaea had disappeared over the years.

Lrb Tnvvavuoma

Factor Df Crown dens. shrub Area 4 *** Treatment 1 Year ***

Year* Area 4 *** ����0lt�:���� Year* Treatment 1 * Crown density shrub ! Crown density tree I Basal area

Fulufj!illet Lrb

Df Factor Crown dens. shrub Area 4 ** Treatment I 1 Year 4 Year*Area Year*Treatment 1 ** 1995 1999 ! 1995 1999 1995 1999 Crown shrub density ! Crown density tree Basalarea

13. Crown cover, per cent, of shrubs and trees, together with basal area (m2 ha·1) of trees at the two inventories in the Birch forest­ Fig. heath type with lichens at Tavvavuoma and on Fulufjallet. Black columns show fenced plots, white columns show open plots and the error columns give the standard error (SE). The tables show the results of ANOVAs conducted on each site individually. Abbreviations, see Figure 5.

Acta Phytogeogr. Suec. 87 76 0. Erikssson et al.

Examples of species that have arrived are Rhodiola rosea, Eriophorum scheuchzeri, Saxifragafoliolosa and Cladina a .. 0: 3 rangiferina on the open plots, and Tr iglochin palustris, and on Eriophorum scheuchzeri Saxifraga oppositifolia .1:3 the fe nced plots. 0:2 0 Having been absent from the dry heath at Ritsem in 1995, Tr isetum spicatum, Va ccinium uliginosum, Hy loco- 0: 1 . 1: mium splendens, Sphagnum spp. and Peltigera aphthosa 20 0:5 0 were found on both fenced and open plots at the inventory Droppings �- . in 1997. Rumex acetosa ssp. lapponicus and Gentiana • 0;6 on the other hand, were found only in the • nivalis, 1995, 1:1 latter only on open plots. 1:5°

• <> Shannon's diversity index 1:6 0:4

Table 20 shows the species diversity for fenced and open plots, calculated according to formula 1. On Langfjallet's grass heath, diversity increased slightly on fenced plots, �:4 -1.0 +1 . 0 but decreased on open plots; the difference was significant (p<0.001, Mixed-model ANOVA). At Ritsem, there were no significant differences in species richness between b 0: 1 1 fenced and open plots, but diversity had decreased in gen- 1:3 eral on the dry heath (p<0.001, Mixed-model ANOVA).

5 1: 1 Ordination

The result of the ordination shows that all plots on Langfj al- 1: 1 1 1 :4 0 ,o:s let's grass heath (Fig. 14a) have moved downwards in the ·-.--·· - �- ... . Droppings ordination diagram. Additionally, at both inventories, plots 0 0: D 11 4 1, 5 and 6 grouped themselves together, which suggests 0:6 1:6 some ftoristicsimilarity betweenthe plots, and that changes in the vegetation have been fairly similar.Plots that diverge most from the others are plots 3 and 4. D 2 0:2 01 : is not possible to identify clearly from the diagram It 10: 3 in Fig. 14b any definitepattern for the study plots on the meadow with low herbs at Ritsem. On Ritsem's dry heath

(Fig. 14c), the open plot 2 and the fenced plot 5 have - 1.0 +1 .0 approached each other, and the same applies to the two fenced plots 1 and 3. The movements have not gone in c 0: 6 the same direction, which indicates that there have been 11 different changes in the composition of the vegetation on 1:6 D these plots.

0:4 11

1:1 1:4 1:3 0 I 1 0:5 I • Droppings I 0: 1

1 0: 3 14. Ordination diagram (PCA) for study plots on: a) Fig. 1 I Ul:ngfjiillet's dry grass heath, b) Ritsem's low herb meadow, :5 D and c) Ritsem's dwarf-shrub heath. The first figure in the plot 1:2 0 identification code refers to the inventory (O=first inventory, 0:2 1 =second inventory), whereas the second figurerefers to the plot

- . number. Black symbols represent fenced plots. 1 0 +1 .0

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 77

Table 20. Shannon's diversity index for fenced and open plots Only the species/species groups that showed significant at Ritsem (dry heath and meadow with low herbs) and on differences are shown in the tables. Uingfjallet's grass heath On Lfmgfjallet's grass heath, Deschampsia jiexuosa and show significant signs that they Plots U.ngfjallet Ritsem low Ritsem Cladonia!Cladina grass heath herb meadow dry heath thrive better on the fenced plots. Although D. flexuosa decreased on both open and fenced plots, the decrease is Fenced 1995 2.671 3.346 2.775 less pronounced on the fenced plots. Cladonia/Cladina Fenced 1997 2.682 3.484 2.467 2) l) decreased on open plots, but increased on fenced plots. Open 1995 2.855 3.457 2.824 Open 1997 2.485 2) 3.524 1) 2.674 Further significant differences are that the group 'other' (litter, stones, bare soil, and water) decreased on the fenced 1) Inventory, 1999; plots, but remained fairly constant on the open plots. Inventory, 1998. 2l Empetrum hermaphroditum, Va ccinium vitis-idaea and Cetraria all increased their degree of cover on the grass heath, regardless of fencing. On the meadow with low herbs at Ritsem, grass and Cover degree sedges together make up about 40% of the ground layer, herbs about 20% and mosses a few per cent less. Lichens Figure 15 shows the ground layer's percentage composition occur only sporadically, whereas dwarf shrubs such as of the three types of vegetation at the two inventories. The Va cciniummyrtillus and Calluna vulgaris are completely tables in the same figure show the results of the ANOVAs absent. Other herbs appear to have been favoured by calculated for each individual site and vegetation type. grazing, as they increased on the open plots but decreased

Ritsem RIS

60 l ...... 50 ·1·------·-----·--·--- Factor Df Clad 6vr

.. *"'* L-�------...... Area 4 .. 40

q: I - � 30 +-, ------YearTreatment 1 ______1 8 20 +-1 - - Year* Area 4 *** ** _____ ...... ___ Year* Treatment I *** i ..... �� • : 0.05 i , , I I I 1 i i I 1 - i i ' 9s 97 95 97 95 97 9s 97 , 95 97 •1 9s 97 95 97 'I 95 97 r 95 97 . 95 97 , i I ! I ---� �i�i i�==:� I -=�� jTitl Krus Grhg j Knlk ! BIA i Ling I Ljun I Oris 1 Mos i Clad I Cetr &' ter '! Ovr i I 95 97 95 97 I ! Ritsem ANG

35 ·-···· ··-········-�------ID�r--�------·------·-·····-···-···-··.. • Factor Df Ort Oris Moss --�;- 25 Area Treatment 20 Year 0:: ** *** � 15 ' Year* Area 4 w ' Year*Treatment I 5 11 * I �� ' ·---H 0 : 14! � � I 99 4 � 9 5 95 99 99 ..· 5 99 I I 9Ort i 1 9 I 95 9 � I �����95 ���95� 99 11 95��� 99 95 99 19 �1�15 95 99 1 Ungtjdllet ORH

Clad Cetr Ov Factor Df Krus Krllk BIA Area 4 Grilg*** *** Treatment I ��g Year I 0.06

Year* Area 4 * * *** *** *** I 0.05 0.05 . ..j Year*Treatm..:nt ..

15. Cover, per cent, of species and species groups at the two inventories on Langfjiillet'sgrass heath and at Ritsem (dry heath and Fig. meadow with low herbs). Black columns show fenced plots, white columns show open plots and the error columns give the standard error (SE). The tables show the results of ANOVAs conducted at each site individually. Abbreviations, see Fig. 5.

Acta Phytogeogr. Suec. 87 78 0. Erikssson et al.

RitsemRIS lOO% . ' . Factor Of Krus Krflk Ling . ·----··--- -H.,·------1 Area 4 .... "'"'* 60% . � Treatment I *** ** ** [>..[ 40% Year I

20% Year*Area 4 *** * I •-ca.,.L._L..,.L.CLA,' * 0% 11 �! - I Year*Treatment l ** ** ** 1 1 ·1, 95 97 95 97 95 97195 97 95 97 5 97,95 97 95 97 95 97 I1 ,9597 95 97' 1 ! ;; ;; ; 1 I ' I I ! Krus I Grhg 1 Kr6k .I Dla Ling 1 Ljun 1 Oris I Mos I Clad I Cetr i Ster I Ovr

Ritsem ANG

Factor Df Ling Area 4 Treatment Year

oo1o 11 1 n i I Year*Area 4 .1 Year* Treatment l ** 95 99 95 11 :1 99 l1 9599 1 qn ! Krus : 6ws

Ungfjilllet GRH · - 1

m � 01 � 95 98 ! I os 119 Krus5 98 19Orhg598 1 KrAk 95DIA98 J 95.Lmg 98 19Ljun� 98 956r1s �8 I19 MS9 8 9H8Clad 119 CctrS 98 95Stcr98 956vr98 1 16. Proportion of study plots with occurrence of a species or species group at the two inventories on Langfjiillet's grass heath. Fig. Black columns show the fenced plots, white columns show the open plots and the error columns give the standard error (SE). The tables show the results of logistic regression conducted on each site individually. Abbreviations, see Figure 5.

on the fenced plots. The cover of mosses, on the other between years, as a response to fencing or both, are in­ hand, increased on fenced plots but remained unchanged cluded in the tables. on open plots. The differences between open and fenced On the meadow with low herbs in Ritsem, Va ccinium plots in changes in the degree of cover are significant in vitis-idaea shows a significant change in occurrence as both cases. Regardless of fencing, the other dwarf shrubs a response to fencing. The frequency increased on both decreased their degree of cover, whereas the group 'other' fenced plots and open plots, but the increase was somewhat (litter, stones, bare soil and water) increased. larger on open plots. On the dry heath, the frequency of Mosses, and other dwarf shrubs, make up most of the Empetrum hermaphroditum increased on fenced plots but ground layer of the dry heath at Ritsem. Only Cladonia/ decreased on open plots, whereas Va ccinium vitis-idaea Cladina show significant changes in cover as a result of and Deschampsia jlexuosa increased on open plots but fencing. They increased on open plots but decreased on decreased on fenced plots. fenced plots. No species or species group showed any significant change in frequency on Langfjallet's grass heath. Frequency Droppings Figure 16 shows the proportion of study plots containing a species/species group of the three vegetation types at the No droppings of elk or hare were found on the study two inventories. The tables in the same figures show the plots on the three types of vegetation (Table 21). At the results of the logistic regressions individually calculated inventory in 1995, all plots on Langfjallet's grass heath for each site and vegetation type. Only the species/species were open, and reindeer droppings found on the plots groups that showed significant differences in frequency subsequently fenced disappeared by 1998. On the open

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 79

Table 21. Number of piles of reindeer droppings per hectare on Table 22. Number of taxa for fenced and open plots at Tj uoma­ fenced and open plots at the two inventories on Langfjallet's jaure and Poullanvare. grass heath and at Ritsem (meadow with low herbs as well as dry heath). Plots Tj uolmajaure Poullanvare

Plots Ungfjallet Ritsem low Ritsem Fenced 1997 42 37 Grass heath herb meadow dry heath Fenced 1999 39 39 Open 1997 43 41 Fenced 1995 399 ± 7 909 ± 201 Open 1999 38 35

Fenced 1997 - 2) - I) Open 1995 728 ± 42 1150 ± 194 5167 2017 ± Open 1997 294 ± 242 2l 643 2076 ± 1415 On the open plots at Puollanvare, ]uncus trifidus was present in 1997, but not in 1999, and the same applies to Va ccinium myrtillus on the fenced plots; this was also a plots, the frequency of reindeer droppings had more than new species on the open plots at the inventory in 1999. halved between the years, and the same applied to the meadow with low herbs at Ritsem. Also on the dry heath, there was a marked reduction in piles of droppings; here Shannon-'s diversity index there appears to be a relationship between the number of Table 23 shows species diversity for fenced and open plots, reindeer in Sorkaitum's Saami village (Figure 3) and the calculated in accordance with formula 1. At Tjuolmajaure number of piles of droppings. and Puollanvare, diversity had increased significantly, on both open and fenced plots (p<0.01 and p<0.05, respec­ tively, Mixed-model ANOVA). 4.4 Tj uolmajaure and Puollanvare Ordination

Species present In the ordination diagram (Fig. 17), the fe nced plots are separated from the open plots. This applies to both sites and The study plots at Tjuolmajaure and Puollanvare are both inventories. The fenced plots are also those that have all in the vegetation type named dry dwarf-shrub heath. moved least between years, which suggests that the changes Table 22 shows the number of taxa (species, species groups, genus, etc.) found in the two inventories at both sites, and all taxa are listed in the Appendix. The factthat Poullanvare 0 some 'difficult' species (e.g. lichens belonging the genus Cladonia) sometimes have been grouped together, causes 1]uohmjaure D some difficulty in deciding from the species list whether one of these species has actually occurred, or whether it has disappearedover the years.

Festuca ovina, Pedicularis lapponica,. Empetrum her­ 0 maphroditum and Va ccinium uliginosum are examples of 0

I species found on the open plots at Tjuolmajaure in 1997, I ------�------but not in 1999. Within the fenced plots, Solorina crocea 1 • 0 • 1 shows the same pattern.]uncus trifiduswas present during 0 1 the initial phase only on the fenced plots, but in 1999 was also present on the open plots.

• • 1 : 0 Table 23. Shannon's diversity index for fenced and open plots at Tjuolmajaure and Puollanvare.

Plots Tj uolmajaure Poullanvare -1.0 +1 .0

Fenced 1997 2.163 2.394 17. Ordination diagram (PCA) for study plots at Puollanvare Fig. Fenced 1999 2.915 2.719 and Tj uolmajaure. Filled symbols represent fenced plots, and the Open 1997 2.407 2.763 figurein the identificationcode of the plots refers to the inventory Open 1999 2.759 2.875 (O=first inventory, l=second inventory).

Acta Phytogeogr. Suec. 87 80 0. Erikssson et al.

Tjuolmajaure

35 T ------30 .:..

Factor Df Krak Oris Clad Year ** * Treatment ***

* * l��=�j�iffj�]JJtf��� Year Treatment 0.06 : . : ' ' ' ' : : 1 , I : f97 99[97 99!97 99 ! 97 99 !97 99:97 99 ! 97 99 !97 99 97 99 i 97 99 [97 99 t 1��� 1�! �1� ful�:���l�l�l

Poullanvare

Factor Df Ling Clad Ovr *** ** *** Year 1 Treatment * * ***

Year* Treatment *

Cover, per cent, of species and species groups at the two inventories at Tj uolmajaure and Puollanvare. Black columns refer to Fig. 18. fenced plots, white columns to open plots, and the error columns show the standard error (SE). The tables show the results of ANOVAs calculated individually for each site. Significance is shown by * for p<0.05, ** for p

in the vegetation have been small. It is also possible to see response to fencing, as it disappeared from the open plots, that, at the time of the firstinventory, the open plots at both but increased slightly on the fenced plots. 'Other dwarf sites were fairly similar, but at the second inventory they shrubs' increased on fenced and open plots, regardless of had moved further from each other in the diagram. treatment. The largest difference between Puollanvare and Tj uol­ Cover majaure is that the formerhas considerably more Empetrum hermaphroditum, on both fenced and open plots. Other­ Figure 18 shows the percentage composition of the ground wise, the distribution of species/species groups at these cover at the two study sites at the two inventories. The two sites is fairly similar. Cladonia/Cladina increased tables in the figure show the results of AN OVA, calculated on the open plots, but remained at a uniform level on the individually for each site. Only the species/species groups fenced plots, whereas Va ccinium vitis-idaea and 'other' that showed significant differences in the analysis of vari­ (bare soil, litter, stones, etc.) in general decreased at Puol­ ance are shown in the table. lanvare over the years. At Tjuolmavare, the ground layer on the open plots is dominated by 'other dwarf shrubs', mosses and 'others ' (bare soil, litter, stones, etc.), whereas the occurrence of lichens is relatively low. Within the fenced plots, on the other hand, lichens belonging to the genera Cladonia! Cladina, Cetraria and Stereocaulon make up more than half of the ground layer. Empetrum hermaphroditumis the only species that shows a significantdiff erence in cover in

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 81

Tj uo!majaure

100"/o

80% >. u 60% :::1� er� 40"/o u: 20"/o

0"/o 91 99 91 99 97 99r9799

Clad Cetr Ster i Ovr

Poullanvare

Factor Df Grhg Year Treatment

Year*Treatment *

The proportion of study plots with occurrence of a species/species group at the two inventories at Tj uolmajaure and Puollanvare. Fig. 19. Black columns refer to fe nced plots and white columns to open plots. The tables show the results of the logistic regression calculated individually for each site. Significance is shown by * for p<0.05. Where nothing is shown, the result is not significant. Abbreviations, see Figure 5.

Frequency differences in frequency between years, as a response to fencing or both, are included in the tables. Figure 19 shows the proportion of plots with occurrence At Puollanvare, 'other grass/sedges' has become less of a species /species group on the study sites at the two frequent on fe nced plots, but more frequenton open plots, inventories. The tables in the figureshow the results of the which is the only significant difference between the two logistic regression calculated individually for each site. sites. Only the species/species groups that showed significant

Acta Phytogeogr. Suec. 87 5. The WWF-project: Discussion

Shannon diversity indexes, ordinations and analyses of the reference areas, when studying their effect on vegetation species cover showed that exclosure of reindeer and elk composition. At ungrazed sites, the lichen cover was 42%, for three to four years had a marginal or no effect on the whereas it was only 4% at grazed sites; for mosses the investigated vegetation types. This was expected, because figures were the opposite, 4% and 22%, respectively. In the dynamics of grazed vegetation in general, and grazed another descriptive study, Helle&A spi (1983) also showed vegetation in subarctic and alpine areas in particular, is that the relative abundance of Cladina spp. increased slow (e.g. Hill et al. 1992, Manseau et al. 1996, Aus­ with decreasing grazing pressure in Finnish Lapland. trheim & Eriksson 2001), and the duration of the present These examples accord with several other studies of the experiment was short. In plant communities such as those effects of reindeer on vegetation, reviewed by Suominen studied here, where reindeer have grazed since the latest & Olofsson (2000). ice age, the species compositions and local species pools In theWWF-study, the three-way interactions between are determined by their grazers (Zimov et al. 1995, Mulder yearxvegetationxtreatment were not significant,indicating 1999, Olofsson et al. 2001, see also Jefferies et al. 1994 ). that the effectgrazers have on the vegetation was uniform, Succession within these communities is slow. Examples of or more correctly, that the absence of effects was uniform. reindeer-grazed systems with low resilience are given in a Once again, what would have been the result had the present study from northern Finland by den Herder et al. (2003). study been running for 30 years instead of three to four They investigated vegetation changes in exclosures in a years? On the dry heath at Puollanvare (Tavvavuoma in Scots pine (Pinus sylvestris) forest and a subarctic heath­ the WWF-study), in the 30-year study by Eriksson et al. land; after 13 years, their conclusion was that the vegeta­ (1998), Cetraria nivalis, Cladina arbuscula andStereocau­ tion in both communities was still in succession. Another lon paschale increased owing to the exclusion of reindeer example of a grazed system with low resilience is from and elk, but in the Birch forest heath type with lichens at North Wales (Hill et al. 1992). In a sheep exclosure experi­ Jarfuna, 10 kmwest ofPuollanvare,F estuca ovina, Cladina ment, Hill with eo-workers observed that the community stellaris, C. arbuscula and Stereocaulon paschale were structure was still not stable after 30 years. Moreover, it more abundant inside exclosures than outside. At Puol­ was shown that the succession strongly depended on the lanvare, C. nivalis, which is an important fodder lichen on initial vegetation (Hill et al. 1992). wind-exposed sites, had increased inside the exclosure. At What will the results be if the present study continues Jarfuna, C. stell aris, also an importantfo dder lichen on sites for 25 more years? We do not know: but probably they will not exposed to wind (Eriksson et al. 1981, Warenberg et al. resemble those from the 30-year exclosure experiment by 1997), showed increased abundance inside the exclosure as Eriksson et al. ( 1998), that revealed conspicuous effects compared with the control plot. There are more examples on the vegetation composition. The results of Eriksson of differences between responses of vegetation types to et al. (1998) study demonstrated, among other things, terminated grazing. In a 7- and in an 11-year-old exclosure an increase in the lichen carpet cover at the expense of experimentin Finnmark,northern Norway, changes in plant bryophytes, which accords with what other studies of the community structure were studied (Moen & Oksanen 1998, effect of grazers on subarctic and alpine vegetation have Olofsson et al. 2002). Two arctic-alpine plant communities shown. For example, a 50-year exclosure experiment in were used, an unproductive snowbed and a considerably the Finnish boreal forest gave similar results (Vare et al. more productive tall-herb meadow. In the latter, the effect 1995). In the Finnish forest, the cover of Cladina stellaris of exclosure was not as extensive as in the low-productive was 63% inside the fence, and only 4% outside; the total snow bed. The effect of summer reindeer grazing on tundra cover of lichens was 92% inside and 75% outside. The total heath vegetation at four sites with different climate and cover of mosses was 1% and 12% at ungrazed and grazed differentvegetation composition, was studied by Olofsson sites, respectively. Similarly to the results of Eriksson et et al. (200 1 ); they used a fence erected 30 years earlier. At al. (1998), Dicranum spp. benefitted most fromgrazing two sites, graminoids dominated the grazed areas, while (Vare et a!. 1995). In the 13-year exclosure experiment by ericoid dwarf shrubs had almost disappeared. At the other den Herder (2003), C. stellaris, C. rangiferina, C. mitis, sites, neither graminoids nor ericoids increased, owing to C. uncialis, Cladonia spp. and Cetraria nivalis decreased grazing. In the study by Manseau et al. (1996), the lichen with grazing, whereas they saw no significant changes in carpet on the shrub tundra was reduced, whereas it was bryophyte cover. At Ungava Bay in Canada, Manseau et not affected in stands of Betula glandulosa. al. (1996) used sites naturally inaccessible to reindeer as

Acta Phytogeogr. Suec. 87 Use and abuse ofreindeer range 83

Species diversity forests (Finnish Lapland), where several studies of di­ versity after reindeer grazing have been performed, the The most commonly used hypothesis on the effectof her­ results suggest that grazing increases diversity and that bivory on species diversity is the intermediate disturbance the increase is due to reduced competition of the dominant hypothesis (Grime 1973, Fox 1979). According to this, species, Cladina spp. (e.g. Helle &Aspi 1983, Viire et al. diversity is low in the absence of disturbance, because 1995). However, in the 30-year exclosure experiment by of competitive exclusion, and high at intermediate levels Eriksson et al. ( 1998), in the same area as the present study, when competitive exclusion is hindered. Diversity is also no significant differences in species number or diversity expected to be low at high levels of disturbance, when between open and fenced plots were detected, although only the most tolerant species persist. One problem with other conspicuous changes in the vegetation within fe nced the intermediate disturbance hypothesis is that it is diffi­ plots had occurred. cult to determine a priori which disturbance level is low, The dryheath at Ritsem has more taxa (57 -72) than the intermediate, or high, and thereby test the hypothesis. Mil­ same vegetation type at other sites (18-43), which is prob­ chunas et al. (1988) modifiedthe intermediate disturbance ably due to the calcareous bedrock and the humidity of the hypothesis, and suggested that a disturbance is determined site. It is difficultto compare, accurately and meaningfully, by the grazing history of the vegetation and the mode of the number of species or species groups between sites, but plant competition within it, i.e. for a community adapted it may be noted that the number of species found in a Pinus to a certain grazing pressure, every deviation from that sylvestris forest in northwest Finland was between 11 and pressure is a disturbance. Furthermore, Olff & Ritchie 22 (Helle &Aspi 1983), whereas in the study by Eriksson (1998) theorise that large herbivores always affect plant et al. (1998) the number of taxa found was between 28 diversity positively through several mechanisms. However, and 39. There were conspicuously large increases in the at high densities, such as in intensive livestock grazing, number of species between inventories on the dry heath in large herbivores can graze non-selectively, leaving only a Tavvavuoma, Sfmfjallet and Langfjallet. In the open plots few tolerant plant species, hence reducing plant diversity. on Sanfjiillet, the number of recorded taxa increased by Thus, according to the intermediate disturbance hypothesis, more than 60%. The reason for this is unknown to us, but we would expect a smaller number of species within our might involve the reduced number of reindeer that periodi­ fenced plots compared with open control plots, provided cally appear in the area, or that the team that carried out that the grazing pressure had been on an intermediate level. the inventory in 1998 was better able to separate lichen However, if the grazing pressure has caused a high level species, than the team that made the initial inventory in of disturbance to the vegetation, which is implied by many 1996. Indications of the latter can be seen on the taxa­ observers e.g. Lundh (1998), Oksanen (1992), and Ihse lists in the Appendix. The weather conditions before, or & Allard (1995), the number of species should increase between, inventories might also have had some effect, but within the fenced plots, as a result of endurable conditions the data available are insufficientlydetailed to allow this to for more species. An increased number of species within be determined. Whatever the reason for the increase, the fenced plots is also to be expected according to the theory result shows the importance of including different types of Olff & Ritchie ( 1998). of vegetation in surveys, to avoid generalizations. The However, as has already been pointed out, there were Shannon diversity indices for the dry heath on Sanfjiillet no particular effects on the number of taxa present or on increased by 34% between inventories of open plots, and Shannon diversity indices, in consequence of terminated 36% between inventories of fenced plots, which is more reindeer and elk grazing; this is an expected result, likely than twice as much as if the proportions between taxa were to be caused by the short duration of the experiment. Large the same; this indicates that the degree of evenness between numbers of species had not time to become dispersed and taxa has increased. This may also be an effect of splitting established on the plots during the two to four years for a large group of taxa into several smaller ones. which the fences have existed, and no large changes in the proportions between species had time to develop. A Ordinations species that has disappeared from the local species pool will have considerable difficulty in re-establishing via Within plant ecology, ordination is often used to arrange seed and other long-distance dispersal options, such as habitats in a two-dimensional space. Here we use the fragmentation, in arctic and subarctic areas (Amen 1966, method to discover changes in species presence and cover Kudo 1993, St6cklin & Biiumler 1996, Pysek 1997). Thus, due to changes in grazing. No conspicuous effects of the although our results imply that diversity in the investigated cessation of grazing by large herbivores were seen in the area is not affectedby the grazing of large herbivores, this PCA -ordination. The different sites with vegetation type is probably not the case. In general, grazing appears to 'Birch-forest heath type with mosses', 'Birch-forest heath increase diversity on heathlands (Suominen & Olofsson type with lichens' and 'Dry heath' differed distinctly in 2000, Austrheim & Eriksson 200 1). In Pinus sylvestris vegetation composition. This dispersal in the ordination

Acta Phytogeogr. Suec. 87 84 0. Erikssson et al.

space was unknown, but is very significant for the study, mer grazing by reindeer in northern Norwaychanged the as it facilitates some generalisation of future results, since vegetation, from moss-rich heath tundra into graminoid­ the vegetation types used represent a larger part of the dominated steppe-like tundra vegetation (Olofsson et al. vegetation of the mountain range (Table 7). 200 1). The basal meristems of grasses, their large resource In an interim-report on the present study (Eriksson et reserves belowground, together with their high photosyn­ al. 1999), 'Fencing' was used as an environmental factor thetic rate, quick leaf turn-over and nutrient absorption, in an ordination of all vegetation types on Langfjallet and and low production of secondary compounds, all increase Sanfjallet, and this factor was, unsurprisingly, not statisti­ their capacity for regrowth after grazing (Archer & Tieszen cal! y significant. However, the environmental factor 'Time' 1980, Chapin 1980). Grasses may respond to grazing indicated significant changes in the plant community, ir­ through several mechanisms, e.g. by increasing axillary respective of grazing treatment. Thus, the open and fenced shoot production, increased number of leaves per shoot or plots moved similarly in the ordination space. This can by delayed senescence. Furthermore, reduced self-shading be explained by the reduced grazing in the areas, and is and an increased proportion of young leaves may also be also implied by the reduction in reindeer droppings at all advantageous for the growth rate of grasses. However, sites, except the vegetation type 'Birch-forest heath type low nutrient availability limits plant growth in subarctic with lichens' at Langfjallet. At this site, exclosur es were and alpine areas, especially of grasses (Chapin et al. 1986, suggested to affectthe vegetation. McKendrick et al. 1980). Consequently, grasses are usually ofless importance in plant communities in these areas. But, according to Zimov et al. (1995), a high grazing pressure Cover of the species on tundra may change the vegetation towards a grass­ Grasses are generally expected to be resistant to trampling dominated steppe owing, for instance, to increased nutrient and grazing (Emanuelsson 1984, Hik et al. 1992, Wegener turnover and thaw-depth of the ground. This hypothesis & Odasz 1997, Mulder 1999); grasses might even be fa­ was supported by a recent study of the vegetation along a voured by grazing (e.g. Moen et al. 1993, Olofsson et al. 40-year-old fence separating different grazing regimes in 2001). Crawley (1997) states that the ability of grasses to northern Fennoscandia (Olofsson et al. 2004). withstand grazing is their most famous trait. Intense sum-

Fig. XXIV. The mountain vegetation is highly sensitive to trampling.Here, a path has been created in the space of two or three years by tourists who have diverged from the Linnaeus trail on L�mgfjallet, to read the sign beside a fenced sample plot. The vegetation type at the site is Dry heath with lichens.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 85

Results from the few studies of the re growth capacity of elk, may reduce the abundance of bryophytes and increase forbs aftergrazing are not conclusive; for example, Mulder the abundance of their competitors, such as lichens of the Ruess (1998) and Olofsson et al. (2002) showed quick genus Cladina, probably as a consequence of changes in & re growth of forbs after ex closure of grazers, whereas Moen the microclimate close to the ground towards drier condi­ et al. (1993) conclude that forbs generally suffer from tions (Suominen et al. 1999). grazing. When the vegetation along fences was studied, In our study, there were no significantthree- or four-way herbs were the only growth forms that did not respond interactions in the models. This means, among other things, significantly to summer grazing by reindeer (Olofsson et that no taxa responded significantlydifferently to exclosure al. 200 1). In the same study, dwarf shrubs were replaced by treatment owing to the surrounding and competing species. graminoids in consequence of grazing, which is consistent Moreover, no significant differencesin responses of taxa to with previous findings (Clarke et al. 1995, McKendrick et fencing between different sites were detected, i.e. latitude al. 1980). Compared with grasses and forbs, dwarf shrubs did not affect the outcome. However, as the duration of have a poor ability to compensate for defoliation and graz­ the present study is so short, it was highly improbable that ing (Mulder 1999). However, in a recent study, Va ccinium such differences would be detected. Seven taxa/groups: myrtillus and vitis-idaea increased most after herbivore Deschampsia fiexuosa, Herbs, Va ccinium myrtillus, V. V. exclosure (Olofsson et al. 2004b ). vitis-idaea, Cetraria spp., Cladina spp./Cladonia spp., and Lichens have a low growth rate (e.g. Karenlampi et al. 1975). A calculated maximum dry mass production of reindeer lichens in northern Finland is 175 kg ha-1 yr-1, or 0.06 g per gram lichen and year (Kumpula et al. 2000). These figures are in accordance with those presented in another study in the same area (den Herder et al. 2003), and predictions in the North Swedish mountain region (Eriksson 1979, Eriksson 1984a). However, the growth of Cladina stellaris may occasionally be as much as 0. 17 g g-1 yr-1 (den Herder et al. 2003), which is almost the same as the growth of the dwarf shrub Empetrum hermaphrodi­ tum (Weih Karlsson 1999). With these, on average, low & growth rates, it is understandable that dense mats of lichens are considered as one of the most sensitive vegetation types in Fennoscandia. Moreover, results indicate that heavy grazing reduces the growth rate of lichens to even lower values (den Herder et al. 2003 ). The abundance of lichens is not reduced just by grazing, but by trampling as well, since trampling easily destroys dry lichens (Manseau et al. 1996). Not only does lichen abundance decrease in areas utilised by reindeer (Arseneault et al. 1997, Allard et al. 1998, Eriksson et al. 1998, Suominen Olofsson 2000), & but there may also be a change in the relative abundance of different lichen species in such areas, since reindeer graze selectively (Danell et al. 1994), and resistance to grazing and trampling varies between lichen species (Klein 1987). Response of the bryophytes to reindeer grazing is inconsistent. In some studies, their performance has been favoured by grazing, whereas in others it is hampered. Since they are mostly weak competitors, their abundance in ground vegetation may increase, owing to selective grazing of their competitors (Helle & Aspi 1983, Oksanen Virtanen 1995, Vare et al. 1995). However, as they are & sensitive to trampling (Chernov 1980 in Zimov et al. 1 995), Fig. XXV. Some sites have less snow than others, which consider­ intense grazing/trampling may reduce the abundance of ably affectsthe vegetation. These patches/sites are more exposed bryophytes and even promote a transition of moss-rich to cold, because they lack an insulating snow cover in winter. heath tundra into graminoid-dominated steppe-like tundra The cold prevents the growth, e.g. of crowberry (Empetrum). The (Olofsson et al. 200 1). Furthermore, paradoxically, brows­ contrasting phenomenon, the snowbed, is better known. ing by the other large herbivore in subarctic Sweden, the

Acta Phytogeogr. Suec. 87 86 0. Erikssson et al.

Fig. XXVI. After years without grazing and trampling by reindeer and elk, the lichen vegetation has changed markedly. 30 The fe nced sample plots are situated (a) at Jaramii and (b) at Poullanvare; the fences were erected in 1968. Photo: Eriksson, 1998. 0.

Mosses, responded to fencing (i.e. timexfencing), but no the Birch forest heath type with lichens, there were no major changes of vegetation communities were detected. significant changes between open and fenced plots. In the vitis-idaea was the most responsive taxon in our study, present study, the response of Cladina sp. to fencing was V i.e. it had the highest number of significant changes due not consistent; the abundance increased in the Grass heath to fencing; however, the response was ambiguous. but decreased in the Dry heath ofRitsem. Cetraria sp. also V vitis-idaea increased within ex closures in the Birch forests responded inconsistently to fencing, increasing at two sites heath type with mosses at Tavvavuoma and Fulufjallet, and decreasing at a third. Our inconclusive result regard­ but decreased on Sanfjallet. At the same site, vitis-idaea ing the response of lichens to fencing was expected, since V increased on the Dry heath. In a recently performed four­ the treatment period is so short compared with the long year exclosure study in northern Fennoscandia, Olofsson lifespan of these 'plants' (J efferies et al. 1994 ). However, et al. (2004b) found increased abundance of vitis-idaea Olofsson et al. (2004b) recorded increased abundance of V when reindeer, Norwegian lemmings (Lemmus lemmus), Cladina mitis (a species close to C. arbuscula) on fenced and Grey-sided voles (Clethrionomys rurocanus) were plots after only four years without reindeer grazing. . prevented from grazing, but not when only the reindeer were excluded. In the 30-year study (Eriksson Raunistola & Shrub and tree canopy cover 1996), vitis-idaea decreased due to fencing in a Birch V forest heath type with lichens (at the Kalatonjarvi site), Although browsing of shrubs and trees is important in the whereas in the same study, another evergreen dwarf shrub, reindeer diet (Warenberg et al. 1997, Warenberg 1977, Empetrum hermaphroditum, tended to be more abundant Eriksson et al. 1981) especially during springtime, ef­ inside the fence than on the open plot, adjacent to the plots fects of herbivores on them are poorly studied (Mulder on the Dry heath of Tavvavuoma in the present study. 1999). The results of the studies known to the authors In all four plots of the 30-year study (Eriksson & are inconsistent. In a Canadian study, the ground cover Raunistola 1996), Cladina spp. increased in abundance, of Betula glandulosa was significantly lower on grazed whereas mosses decreased, e.g. Dicranum scoparium. The areas than on ungrazed areas (Manseau et al. 1996). lichen Cetraria spp. had higher abundance on fenced plots Warenberg (1977) described negative effects of reindeer than on open ones at two Dry heath sites in this study. At grazing on Salix glauca and Sorbus aucuparia at Ottfjall

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 87

Fig. XXVII. Moss-rich mountain birch forest below Sonfjallet, in which Va ccinium myrtillus dominates the fi eld layer. On some birch trunks, the snow lichen (Parmelia olivacea) indicates the man-high winter snow depth.

in Jamtland, Sweden. It has also been shown that reindeer had been able to move between countries with their herds, grazing may prevent development of new forest after out­ as was most favourable for pasturage. Subsequently, the breaks of autumnal moth (Epirrita autumnata) (Lehtonen borders of the national states were closed, in consequence of which competition for grazing increased markedly. Im­ & Heikkinen 1995). However, in north Scandinavia, the cover of B. puberscens ssp. czerepanovii, Salix glauca and migration to northern Lapland from the Grand Duchy of Sorbus aucuparia was not changed by a 7 -year exclo sure Finland took place at the same time as the ability of the Swedish Saamis to move at will to the summer grazings of folivorous mammals (Moen & Oksanen 1998). Rein­ deer grazing and trampling may also favour birches via in Norway was restricted. Overcrowding in the northern increased germination of seeds, owing to gap formation in parts of the Swedish reindeer-husbandry area gradually the vegetation (Helle et al. 1998 in Olofsson 2001). became so great that the authorities began to organise There were unexpected reductions inthe shrub canopy­ extensive, obligatory 'dislocations'. cover in the vegetation type 'Birch forest heath type with Those Saamis affected by the obligatory moves took lichens' in Tavvavuoma and Fulufjallet, subsequent to with them an extensive form of reindeer husbandry, based fencing. The cause is unknown, but a probable reason is principally on meat production, and developed in the that larger shrubs have grown into trees and only small northern part of the Scandes where, during the snowfree shrubs are left in the class. period, the terrain allows rather free movement of the herds, determined mainly by the availability of pastur­ age, by wind direction, by topographic features such as 5 .1. Concluding thoughts precipices, valleys and glaciers, and to some extent by the need to watch over the reindeer herds. As appears from the accounts of early travellers in Lapland, In the intermediate and southern Saami communities, in former historical circumstances reindeer-grazing was a the form of reindeer husbandry was traditionally intensive, considerable asset. and implied controlled herding of the reindeer during most During the 19th century, and at the beginning of the of the year. This form of husbandry was based on all of 20th century, the grazing situation changed radically. the benefits that could be obtained from the reindeer, but Previously, the Saamis, irrespective of their citizenship, included hunting and trapping. The collision between the

Acta Phytogeogr. Suec. 87 88 0. Erikssson et al.

various forms of husbandry was intense, and led to the ent from today. Probably, we would have much larger and permanent overexploitation of the grazing resources, in thicker lichen mats and fewer mosses. It is also probable particular the lichen grazings. that the abundance of graminoids would be lower, as these The measures so far applied to bring about a palpable are favoured by grazing and trampling. However, it is far and permanent reduction of the reindeer stock- whether from certain that diversity would decrease. undertaken on the initiative of the reindeer owners (the The annual air-temperature means have fallen in Saami communities) or by the State-have had little posi­ Scandinavia during the two latest 30-yearstandard periods tive effect in the longer term. Neither the biomass nor the (1931-60 and 1961-90), and the glaciers on the west side species composition of the grazed plants appears to have of the Scandes are growing rapidly, whereas the size of been favourably affected. glaciers on the east side is unchanged (Bjornet al. 2000). A short period of prevented reindeer and elk grazing Nevertheless, the global mean air-temperature is predicted does not change the diversity or species composition of to rise by l-3.5°C by the year 2100 (Houghton etal. l996) the field layer in the Swedish mountain range. The results and may push isotherms 150-550 km farther north; con­ from the present and earlier studies imply that, in order sequently, the distribution, abundance and accessibility to understand the effect of large herbivores on vegetation of most plant species will be affected. Further, increased in the Fennoscandic mountain range, there is a need for temperatures lead to increased nutrient availability for the long-term experiments in differenttypes of subarctic and vegetation. Grasses will gain from this, whereas slow­ alpine vegetation. Thus, the succession in the investigated growing species will decrease; the abundance of dwarf areas was slow. shrubs will probably increase, which may be negative We do not know what the vegetation of the Swedish for the reindeer, as lichens may be out-competed by the mountain chain would look like without managed large dwarf shrubs (Bjornet al. 2000). Thus, the future climate grazers. Nevertheless, according to theory and results from in Swedish mountain areas will determine the vegetation earlier studies discussed above, it would look quite differ- there at least as much as the large herbivores.

Acta Phytogeogr. Suec. 87 References

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Resa genom Sveriges och Norriges Lapp­ berggrunden i Jamtlands lan. Ser Ca no 52. Sheet 1. - Liber marker, forrattad ar 1821. - Lund. Kartor, Stockholm. Zetterstedt, J.W. 1833. Resa genom Umea lappmarker i Ve sterbot­ tens Ian, forrattad ar 1832. - 6rebro. Zetterstedt, J.W. 1822. Resa genom Sveriges och Norriges Lapp­ marker, forrattad ar 1821. - Lund. Zetterstedt, J.W. 1833. Resa genom Umea lappmarker i Ve sterbot­ tens Ian, fOrrattad 1832. - 6rebro. ar Zimov, S.A., Chuprynin, V.I., Oreshko, A.P., Chapin, F.S. Ill, Reynolds,J.F. &Chapin,M.C. 1995. Steppe-tundra transition: A herbivore-driven biome shift at the end of the pleistocene. - Am. Nat. 146: 765-794.

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 95

d Appen ix I. Recorded species.

Birch forest type with lichens Swedish common Tavvavuoma Fulufjallet Species/taxa and author name C95 C99 095 099 C96 C99 096 099

Va scular plants Antennaria spp. kattfot Anthoxanthum odoratum L. varbrodd Arctostaphylos alpinus (L.) Sprengel ripbar Betula nana L. dvargbjork Betula pubescens ssp. czerepanovii N. I. Orlova fj iillbjork Calamagrostis lapponica (Wahlenb.) Hartm. lappror Calluna vulgaris (L.) Hull ljung Deschampsiaflexuosa (L.) Trin. krustatel Empetrum hermaphroditum Hagerup knikbar Festuca ovina L. farsvingel Hieracium spp. fibbla ]uncus trifidus L. klynnetag Juniperus communis L. en Linnaea borealis L. linnea Lycopodium alpinum L. fj iillummer L. annotinum L. revlummer L. clavatum L. mattlummer L. complanatum L. plattlummer Phleum alpinum L. fj iilltimotej Phyllodoce caerulea (L.) Bab. lappljung Salix glauca L. rip vide Solidago virgaurea L. gullris Tr ientalis europaea L. skogsstjarna Va ccinium myrtillus L. blabar uliginosum L. odon V. vitis-idaea L. lingon V. Other grasses Ovrigt gras

Lichens Cetraria cucullata (Bell.) Ach. strutlav C. ericetorum Opiz smal islandslav C. islandica (L.) Ach. islandslav C. nivalis (L.) Ach. snolav Cladina arbuscula (Wallr.) Hale & Culb. gulvit renlav C. rangiferina (L.) Nyl. gra renlav C. stellaris (Opiz) Brodo fOnsterlav Cladonia sulphurina (Michaux) Fr. poslav C. bellidiflora (Ach.) Schaerer blombagarlav C. coccifera (L.) Willd. kochenillav C. cornuta (L.) Hoffm. syllav C. crispata (Ach.) Flotow taggig bagarlav C. .fimbriata (L.) Fr. naggig bagarlav C. furcata (Huds.) Schrader rislav C. gracilis (L.) WiHd. stangellav C.squamosa (Scop.) Hoffm. fnaslav C. uncialis (L.) Wigg. pigglav Cladonia spp. bagarlavar Icmadophila ericetorum (L.) Zahlbr. vitmosslav Nephroma arcticum (L.) Torss. norrlandslav Ochrolechiafrigida (Swartz) Lynge nordlig ornlav Peltigera aphthosa (L.) Willd. torsklav P. canina (L.) Willd. filtlav Pertusaria dactylina (Ach.) Nyl. fingerlikporlav Solorina crocea (L.) Ach. saffranslav Stereocaulon paschale (L.) Hoffm. P.iiskrislav Other lichens Ovriga lavar

Mosses Dicranum scoparium Hedw. kvastmossa Hepaticae Levermossa Hylocomium splendens (Hedw.) Schimp. husmossa Paludella squarrosa (Hedw.) Brid. piprensarmossa Pleuroziumschreberi (Brid.) Mitt. vaggmossa Polytrichum juniperinum Hedw. enbjornmossa piliferum Hedw. hiirbjornmossa P. Polytrichum spp. bj ornmossor Ptilidium ciliare (L.) Hampe franslevermossa Other mosses Ovrig mossa

Acta Phytogeogr. Suec. 87 96 0. Erikssson et al.

Birch forest type with mosses Swedish common Tav avuoma Sonfjallet Ungfjallet Fulufjallet Species/taxa and author name C95 C98 095 098 C95 C98 095 098 C95C98 095 098 C95C99 095 099

Andromeda polifo lia L. rosling Arctostaphylos alpinus (L.) Sprengel rip bar Bartsia alpina L. svartho Betula nana L. dvlirgbj6rk Betula pubescens ssp. czerepanovii N. I. Orlova fj hllbjork Calamagrostis spp. ror Calluna vulgaris (L.) Hull lj ung Carex spp. starr Deschampsiaflexuosa (L.) Trin. krustatel Empetrum hermaphroditum Hagerup krfrbar Equisetum sylvaticum L. skogsfrliken Eriophorum vaginatum tuvull L estuca ovina L. fa rsvingel F Hieracium spp. fibbla Huperzia selago (L.) Bemh. ex Schrank & Mart. lopplummer Juniperus communis L. en Linnaea borealis linnea L. Luzula pilosa (L.) Willd. vanryle Lycopodium alpinum L. fj hllummer L. annotinum L. revlummer L. clavatum L. mattlummer L. complanatum L. plattlummer Melampyrum pratense L. angskovall Mo linia caerulea (L.) Moench blatatel Nardus stricta L. stagg Pedicularis lapponica L. lappspira Phyllodoce caerulea (L.) Bab. lappljung Pinus sylvestris L. tall Rubus chamaemorus L. hjortron Salix herbacea L. dvargvide Solidago virgaurea L. gullris Sorbus aucuparia L. ronn Tr ientalis europaea L. skogsstjlirna Va ccinium myrtillus L. blil.blir V. uliginosum L. odon V. vitis-idaea lingon L. Lichens Cetraria cucullata (Bell.) Ach. strutlav C. ericetorum Opiz smal islandslav C. islandica (L.) Ach. islandslav C. nivalis (L.) Ach. snolav Cladina arbuscula (Wallr.) Hale & Culb. gulvit renlav C. rangiferina (L.) Nyl. gra renlav C. stellaris (Opiz) Brodo fonsterlav Cladonia sulphurina (Michaux) Fr. p6slav C. bellidiflora (Ach.) Schaerer blombagarlav C. coccifera (L.) Willd. kochenillav cornuta (L.) Hoffm. syllav C. crispata (Ach.) Flotow taggig bagarlav C. C.fimbriata Fr. naggig bagarlav (L.) C. gracilis (L.) Willd. stangellav C. pyxidata (L.) Hoffm. trattlav C. squamosa (Scop.) Hoffm. fn aslav C. uncialis (L.) Wigg. pigglav Cladonia spp. bagarlavar lcmadophila ericetorum (L.) Zahlbr. vitmosslav Nephroma arcticum Tors s. norrlandslav (L.) Peltigera aphthosa (L.) Willd. torsklav Stereocaulon paschale (L.) Hoffm. paskrislav Other lichens bvriga lavar

Mosses Dicranum scoparium Hedw. kvastmossa Hepaticae Levermossor Hy locomium splendens (Hedw.) Schimp. husmossa Paludella squarrosa(Hedw.) Brid. piprensarmossa Pleurozium schreberi (Brid.) Mitt. vaggmossa Polytrichum juniperinum Hedw. enbjomrnossa piliferum Hedw. harbjomrnossa P. Polytrichum spp. bj 6mrnossor Ptilidium ciliare (L.) Hampe franslevermossa Sphagnum spp. vitmossa Other mosses bvrig mossa

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 97

Dry heath Tjuolmajaure Poullanvare Swedish C97 C99 097 099 C97 C99 097 099 Species/taxa and author common name

Betula nana L. dvlirgbjork Betula pubescens ssp. czerepanovii N. I. Orlova fj allbjork Calamagrostis lapponica (Wahlenb.) Hartm. lappri:ir Carex bigelowii Torr. Ex Schwein styvstarr Cassiope hypnoides (L.) D.Don mossljung Empetrum hermaphroditum Hagerup krakbar Festuca ovina L. farsvingel ]uncus trifidus L. klynnetag Luzula multiflora ssp.jrigida (Buchenau) V. I Krecz. sliterfryle L. spicata (L.) DC. axfryle Pedicularis lapponica L. lappspira Phyllodoce caerulea (L.) Bab. lappljung Va ccinium myrtillus L. biabar uliginosum L. odon V. vitis-idaea L. lingon V.

Lichens Alectoria nigricans (Ach.) Nyl. upprlitt tagellav A. ochroleuca (Hoffm.) Massal. fj lilltagellav Cetraria cucullata (Bell.) Ach. strutlav C. ericetorum smal islandslav Opiz C. islandica (L.) Ach. islandslav C. nivalis (L.) Ach. snolav Cladina arbuscula (Wallr.) Hale & Culb. gulvit renlav C. rangiferina (L.) Nyl. gra renlav C. stellaris (Opiz) Brodo fonsterlav Cladonia amaurocraea stor pigglav C. bellidiflora (Ach.) Schaerer blombligarlav C. coccifera (L.) Willd. kochenillav C. cornuta (L.) Hoffm. syllav C. crispata (Ach.) Flotow taggig bligarlav C. .fimbriata (L.) Fr. naggig bligarlav C. gracilis (L.) Willd. stlingellav Cladonia subfurcata (Nyl.) mossbligarlav Am. C. uncialis (L.) Wigg. pigglav Cladonia spp. bligarlavar Icmadophila ericetorum (L.) Zahlbr. vitmosslav Nephroma arcticum (L.) Torss. norrlandslav Ochrolechiafrigida (Swartz) Lynge nordlig ornlav Peltigera canina (L.) Willd. filtlav scabrosa Fr. strlivfiltlav P. Th. Pertusaria dactylina (Ach.) Nyl. fingerlik porlav Solorina crocea (L.) Ach. saffranslav Sphaerophorus globosus (huds.) Vainio korallav Stereocaulon paschale (L.) Hoffm. paskrislav Thamnolia vermicularis (Swartz) Schaerer masklav Other lichens Ovriga lavar

Mosses Dicranum scoparium Hedw. kvastmossa Hepaticae Levermossor Polytrichum juniperinum Hedw. enbjornmossa piliferum Hedw. harbjornmossa P. Polytrichum spp. bjornmossor Ptilidium ciliare (L.) Hampe franslevermossa Other mosses Ovrig mossa

Acta Phytogeogr. Suec. 87 98 0. Erikssson et al.

Swedish Grass heath common Ungfjallet Species/taxa and author name C95 C98 095 098

Betula nana L. dvargbjork Callluna vulgaris (L.) Hull lj ung C. bigelowii Torr. Ex Schwein styvstarr Cassiope hypnoides (L.) D.Don mossljung Deschampsiafiexuosa (L.) Trin. krustatel Empetrum hermaphroditum Hagerup nordkrdkbiir Festuca ovina L. farsvingel Hieracium L. sect. alpina (Griseb.) Gremli fj allfibbla Hieracium spp. ovriga fibblor Juncus tri.fidus klynnetag L. Juniperus communis L. en Loiseleuria procumbens (L.) Desv. krypljung Lycopodium procumbens (L.) fj iillummer Phyllodoce caerulea (L.) Bab. lappljung Solidago virgaurea L. gullris Tr ientalis europaea L. skogsstjiirna Va ccinium myrtillus L. blabar vitis-idaea L. lingon V.

Lichens Bryocaulon divergens (Ach.) Kiirnef. spiirrlav Cetraria cucullata (Bell.) Ach. strutlav C. ericetorum Opiz smal islandslav C. islandica (L.) Ach. islandslav C. nivalis (L.) Ach. snolav Cladina arbuscula (Wallr.) Hale & Culb. gulvit renlav C. rangiferina (L.) Nyl. gra renlav C. stellaris (Opiz) Brodo fonsterlav Cladonia sulphurina (Michaux) Fr. poslav C. bellidijfora (Ach.) Schaerer blombagarlav C. coccifera (L.) Willd. kochenillav C.cornuta (L.) Hoffm. syllav C . .fimbriata (L.) Fr. naggig bagarlav C. gracilis (L.) Willd. stiingellav C. unicialis (L.) Wigg. pigglav Cladonia spp. bagarlavar lcmadophila ericetorum (L.) Zahlbr. vitmosslav Lepraria membranacea (Dickson) Vainio mjollav Ochrolechia.frigida (Swartz) Lynge nordlig ornlav Stereocaulon paschale (L.) Hoffm. paskrislav Other lichens bvriga lavar

Mosses Dicranum scoparium Hedw. kvastmossa Hepaticae Levermossa Pleurozium schreberi (Brid.) Mitt. vaggmossa Polytrichum piliferum Hedw. harbjommossa Polytrichum spp. bj ommossor Sphagnum spp. vitmossa Other mosses Ovrig mossa

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 99

Meadow with low herbs Swedish common Ritsem Species/taxa and author name C95 C99 095 099

Agrostis mertensii Trin. fj allven Antennaria spp. kattfot Anthoxanthum odoratum varbrodd L. Betula nana L. dvargbjork Bistorta vivipara (L.) Gray ormrot Calamagrostis lapponica (Wahlenb.) Hartm. lappror Carex atrata svartstarr L. C. bigelowii ex Schwein styvstarr Torr. C. canescens grastarr L. C. lachenalii ripstarr L. Carex spp. ovrig starr Cassiope hypnoides (L.) D.Don mossljung Cerastiumfontanum ssp.jontanum Gartner riparv Deschampsiaflexuosa Trin. krustatel (L.) Empetrum hermaphroditum Hagerup nordkr3.kbar Equisetum arvense L. 3.kerfriiken Festuca ovina farsvingel L. Gentiana nivalis L. fj allgentiana Gnaphalium supinum L. fj allnoppa Hieracium spp. ovriga fibblor Huperzia selago (L.) Bemh. ex Schrank & Mart. lopplummer ]uncus trijidus L. klynnetag Loiseleuria procumbens(L.) Desv. krypljung

Luzula multiflora(Ehrh.) ej . angsfryle L L. spicata (L.) DC axfryle Ly copodium alpinum L. fj allummer L. annotinum L. revlummer L. clavatum L. mattlummer Pedicularis lapponica L. lappspira Petasites frigidus fj allskrap L. Phleum alpinum fj alltimotej L. Phyllodoce caerulea Bab. lappljung (L.) Poa pratensis ssp. alpigena (Fr.) Hiitonen nordgroe Ranunculus nivalis L. fj rulsmorblomma Ranunculus spp. ovriga smi:irblommor Rhodiola rosea L. rosenrot Rubus chamaemorus hjortron L. Rumex acetosa L. angssyra Rumex acetosa ssp. lapponicus (Hiitonen) Jalas lappsyra Salix glauca L. ripvide S. herbacea L. dvargvide Sibbaldia procumbens L. dvargfingeri:irt Solidago virgaurea L. gullris Ta raxacum spp. F.H. Wigg maskros Tr ientalis europaea skogsstjarna L. Tr isetum sp icatum (L.) K. Richter fj allhavre Va ccinium myrtillus L. blabar uliginosum L. odon V. vitis-idaea L. lingon V. Va hlodea atropurpurea (Wahlenb.) Fr. ex Hartm. lapptatel Ve ronica alpina L. fj allveronika Violabiflora L. fjallviol

Lichens Alectoria nigricans (Ach.) Nyl. uppdi.tt tagellav A. ochroleuca (Hoffm.) Massal. fj alltagellav Cetraria ericetorum Opiz smal islandslav C. islandica (L.) Ach. islandslav C. nivalis Ach. snolav (L.) Cladina arbuscula (Wallr.) Hale & Culb. gulvit renlav C. rangiferina (L.) Nyl. gra renlav C. stellaris (Opiz) Brodo fo nsterlav C. bellidiflora (Ach.) Schaerer blombagarlav C. coccifera Willd. kochenillav (L.) C. cornuta (L.) Hoffm. syllav C.jimbriata (L.) Fr. naggig bagarlav C. uncialis (L.) Wigg. pigglav Cladonia spp. bagarlavar

Acta Phytogeogr. Suec. 87 100 0. Erikssson et al.

Meadow with low herbs, cont.

Swedish common Ritsem Species/taxa and author name C95 C99 095 099

Diploschistes scruposus (Schrever) Norman groplav /cmadophila ericetorum (L.) Zahlbr. vitmosslav Nephroma arcticum (L.) Torss. norrlandslav Ochrolechia.frigida (Swartz) Lynge nordlig ornlav Peltigera aphthosa (L.) Willd. torsklav Solorina crocea (L.) Ach. saffranslav Stereocaulon paschale (L.) Hoffm. paskrislav Other lichens Ovriga lavar

Mosses Dicranum scoparium Hedw. kvastmossa Hepaticae Levermossor Hy locomium splendens (Hedw.) Schimp. husmossa Pleurozium schreberi (Brid.) Mitt. vliggmossa Polytrichum piliferum Hedw. harbjornmossa Sphagnum spp. vitmossor Other mosses Ovriga mossor

Acta Phytogeogr. Suec. 87 Use and abuse of reindeer range 101

Appendix 11. Saami names for utility goods, botanical names, villages and study sites, and list of persons who provided information

Utility goods

English Saami Shovel staff or ski stick Goaivosoabbi Lapland sleigh Gieres Cradle Gietka Smudge fire Suovva

Species and groups

Latin or English Saami Alectoria sp. Uigejeagil Angelica archangelica boska Astragalus alpinus duottarsaphal Betula nana skierre Betula pubescens ssp. czerepanovii lages Bryophyte seamul Bryoria fuscescens guossalahppu Calluna vulgaris livdnju Cetraria nivalis fiskesjeagil Cladina arbuscula roancejeagil C. rangiferina ninesjeagil C. stellaris oaivejeagil Cladonia coccifera bohccejeagil Empetrum hermaphroditum cahppesmuorji Epidendric lichen lahppu Ep ilobium angustifolium horbma Equisetum sp. gorddet Fungus guoppar Lactarius terminosus riesaraski Lichen jeagil Menyanthes trifoliata Muoska Nephroma sp. Jorgqujeagil Parmelia olivacea beassegatna Rhododendron lapponicum duottarcila]a Rumex acetosa ssp. lapponicus joupmu Salix phylicifolia ruksessieoga Solidago virgaurea beatnatjuovccarassi Stereocaulon paschale smarvejeagil Usnea filipendula lrugejeagil

Saami villages

English Sa ami Talma Tal m a Lainiovuoma Lainiovuoma Saarivuoma Saarivuoma Mellanbyn Baste Cearru Stirkaitum Unnatjarro Mittadalen Mihte ldre nya sameby ldre nya sameby Fulufjallet Fulufjallet

Study sites English Saami Tj uolmajaure Cuolmmajavri Poullanvare Boullanvarri Ta vvavuoma Davvavuopmi Ritsem Rijtjem Sanfjallet Langfjallet Fulufjallet

Acta Phytogeogr. Suec. 87 102 0. Erikssson et al.

Persons who provided information

Baer, Lars. herder, chairman. Saarivuoma Berggren, Lars. assistant bailiff. Vittangi Blind, Olof. herder, chairman. Talma Idivuoma, Per. herder, chairman. Lainiovuoma Mellkvist, Hannes. senior officer, county administration. Dalarna Mikaelsson, Stefan. herder. ROdingtrask Ronnback, Jan-Ivar. senior officer, county administration. Norrbotten Ruong, Israel. professor. Uppsala Skum, Nickolaus. herder, chairman Norrkaitum

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1. E. Almquist. 1929. Upplands vegetation och flora. (Vege­ Takem. ISBN 91-7210-012-5. Price: SEK 160. tation and flora of Uppland.) Out of print. 13. Vaxtgeografiska studier tillagnade Cart Skottsberg pa 2. S. Thunmark. 1931. Der See Fiolen und seine Ve getation. sextioarsdagen 1112 1940. (Geobotanical studies dedicated ISBN 91-7210-002-8. Price: SEK 240. to C. Skottsberg.) 1940. ISBN 91-7210-013-3. Price: SEK 3. E. Du Rietz. 1931. Life-forms of terrestrial flowering 290. G. plants. I. ISBN 91-7210-003-6. Price: SEK 160. 14. N. Hy lander. 1941. De svenska formema av Mentha gen­ 4. B. Lindquist. 1932. Om den vildviixandeskogsalmens raser tilis L. coil. (Zusammenf. : Die schwedischen Form en der och deras utbredning i Nordvasteuropa. (Summ.: The races Mentha gentilis L. sensu coil.) ISBN 91-7210-014-1. Price: of spontaneous Ulmus glabra Huds. and their distribution SEK 160. in NW Europe.) Out of print. 15. E. Hasselrot. 1941. Till kannedom om nagra nordiska T. 5. H. Osvald. 1933. Ve getation of the Pacific coast bogs of umbilicariaceers utbredning. (Zusammenf. : Zur Kenntnis North America. ISBN 91-7210-005-2. Price: SEK 160. der Ve rbreitung einiger Umbilicariaceen in Fennoscandia.) 6. Samuelsson. 1934. Die Ve rbreitung der hoheren Wasser­ ISBN 91-7210-015-X. Price: SEK 240. G. pflanzen in Nordeuropa. 1934. Out of print. 16. Samuelsson. 1943. Die Verbreitung derAlchemilla-Arten G. 7. G. Degelius. 1935. Das ozeanische Element der Strauch­ aus der Vulgaris-Gruppe in Nordeuropa. ISBN 91-7210- und Laubflechtenflora von Skandinavien. Out of print. 016-8. Price: SEK 160. 8. R. Sernander. 1936. Granskiir och Fiby urskog. En studie 17. Th .Arwidsson. 1943. Studien iiberdie Gefasspflanzen in den over stormluckornas och marbuskarnas betydelse i den Hochgebirgen der Pite Lappmark. ISBN 91-7210-017-6. svenska granskogens regeneration. ( Summ.: The primitive Price: SEK 240. forests of Granskiir and Fiby. A study of the part played 18. Dahlbeck. 1945. Strandwiesenam siidostlichen Oresund. N. by storm-gaps and dwarf trees in the regeneration of the (Summ.: Salt marshes on the S. E. coast of Oresund.) ISBN Swedish spruce forest.) ISBN 91-7210-008-7. Price: SEK 91-7210-01 8-4. Price: SEK 160. 240. 19. E. von Krusenstjerna. 1945. Bladmossvegetation och 9. R. Sterner. 1938. Flora der Insel Oland. Die Areale der bladmossflora Uppsalatrakten. (Summ.: Moss flora and i GefasspflanzenOlands nebst Bemerkungen zu ihrer Oeko­ moss vegetation in the neighbourhood of Uppsala.) ISBN logie und Soziologie. Out of print. 91-7210-019-2. Price: SEK 290. 10. B. Lindquist. 1938. Dalby Soderskog. En skansk lovskog 20. N. Albertson. 1946. Osterplana bed. Ett alvaromracte pa i forntid och nutid. (Zusammenf.: Ein Laubwald in Kinnekulle. (Zusammenf. : Osterp1anabed. Ein AI vargebiet Schonen in der Vergangenheit und Gegenwart.) ISBN auf dem Kinnekulle.) ISBN 91-7210-020-6. Price: SEK 91-7210-0 10-9. Price: SEK 240. 240. 11. Stalberg. 193 9. Lake Vattern. Outlines of its natural 21. H. Sj ors. 1948. MyrvegetationiBergslagen. (Summ.: Mire N. history, especially its vegetation. ISBN 91-7210-011-7. vegetation in Bergslagen, Sweden.) ISBN 91-7210-021-4. Price: SEK 160. Price: SEK 290. 12. E. Du Rietz, A. G. Hannerz, Lohammar, R. Santesson 22. S. Ahlner. 1948. Utbredningstyper bland nordiska barrtrads­ G. G. & M. W 1939. Zur Kenntnis der Ve getation des Sees lavar. (Zusammenf.: Ve rbreitungstypen unter fennoskan- rern.

Acta Phytogeogr. Suec. 87 104 Svenska ViixtgeografiskaSiillska pet

dischen Nadelbaumfiechten.) ISBN 91-7210-022-2. Price: of the Baltic basin and in lakes from Preboreal time to the SEK 240. present day.) ISBN 91-7210-038-9. Price: SEK 160. 23. E. Julin. 1948. Ve ssers udde, mark och vegetation i en 39. M. Fries. 1958. Ve getationsutveckling och odlingshistoria igenvaxande lovang vid Bjarka -Saby. (Zusammenf. : Ve ss­ i Varnhemstrakten. En pollenanalytisk undersokning i ers udde. Boden und Ve getation in einer verwachsenden Vastergotland. (Zusammenf. : Ve getationsentwicklung Laubwiese bei Bjarka-Saby in bstergotland, Stidschweden.) und Siedlungsgeschichte im Gebiet von Vamhem. Eine ISBN 91-7210-023-0. Price: SEK 240. pollenanalytische Untersuchung aus Vastergotland (Si.id­ 24. M. Fries. 1949. Den nordiska utbredningen av Lactucaalpi­ schweden).) ISBN 91-7210-039-7. Price: SEK 160. na, Aconitum septentrionale, Ranunculus platanifolius och 40. B. Pettersson. 1958. Dynamik och konstans i Gotlands flora Polygonatum verticillatum. (Zusammenf. : Die nordische och vegetation. (Zusammenf. : Dynamik und Konstanz in Verbreitung von Lactuca alpina, Aconitum septentrionale, der Flora und Ve getation von Gotland, Schweden.) ISBN Ranunculus platanifolius und Polygonatum verticillatum.) 91-7210-040-0. Price: SEK 400. ISBN 91-7210-24-9. Price: SEK 160. 41. E. Uggla. 1958. Skogsbrandfalt i Muddus nationalpark. 25. Gjarevoll. 1949. SnlZ)leievegetasjonen i Oviksfjellene. (Surnrn.:Forest fireareas in Muddus National Park, northern 0. (Summ.: The snow-bed vegetation of Mts Oviksfjallen, Sweden.) ISBN 91-7210-04 1-9. Price: SEK 160. Jamtland, Sweden.) ISBN 91-7210-025-7. Price: SEK 42. Thomasson. 1959. Nahue1 Huapi. Plankton ofsome lakes K. 160. in an Argentina National Park, with notes on terrestrial 26. H. Osvald. 1949. Notes on the vegetation of British and vegetation. ISBN 91-721 0-042-7. Price: SEK 160. Irish mosses. ISBN 91-7210-026-5. Price: SEK 160. 43. Gillner. 1960. Ve getations- und Standortsuntersuchungen V. 27. S. Selander. 1950. Floristic phytogeography of southwestern in den Strandwiesen der schwedischen Westktiste. ISBN Lule Lappmark (Swedish Lap land). I. 1950. ISBN 91-7210- 91-7210-043-5. Price: SEK 240. 027-3. Price: SEK 240. 44. E. Sj ogren. 1961. Epiphytische Moosvegetation in Laub­ 28. S. Selander. 1950. Floristic phytogeography of southwestern waldern der Insel bland, Schweden. (Surnrn. : Epiphytic Lule Lappmark (Swedish Lapland). II. Karlvaxtfloran i moss communities in deciduous woods on the island of sydvastra Lule Lappmark. (Summ.: Vascular flora.) ISBN bland, Sweden.) ISBN 917210-044-3 (ISBN 91-7210- 91-7210-028- Price: SEK 160. 444-9). Price: SEK 160. I. 29. M. Fries. 1951. Pollenanalytiska vittnesbord om senkvartar 45. G. Wistrand. 1962. Studier i Pite Lappmarks karlvaxtflora, vegetationsutveckling, sarskilt skogshistoria, i nordvastra med sarskild hansyn till skogslandet och de isolerade fjallen. Gotaland. (Zusammenf. : Pollenanalytische Zeugnisse (Zusarnrnenf. : Studien iiber die Gefasspflanzenflorader Pite der spatquartaren Ve getationsentwicklung, hauptsachlich Lappmark mit besonderer Beri.icksichtigungdes Wa ldlandes der Waldgeschichte, im nordwestlichen Gotaland, Stid­ und der isolierten niederen Fje1de.) ISBN 91-7210-045-1 schweden.) ISBN 91-7210-029-X. Price: SEK 240. (ISBN 91-7210-445-7). Price: SEK 240. 30. M. Warn. 1952. Rocky-shore algae in the bregrundArchi­ 46. R. lvarsson. 1962. Lovvegetation i Mollosunds socken. pelago. ISBN 91-7210-030-3. Price: SEK 290. (Zusarnrnenf. : Die Laubvegetation im Kirchspiel Mol­ 31. Rune. 1953. Plant life on serpentines and related rocks losund, Bohuslan, Schweden.) ISBN 91-7210-046-X (ISBN 0. in the North ofSweden. 1953. ISBN 91-7210-03 1-1. Price: 91-7210-446-5). Price: SEK 160. SEK 240. 47. Thomasson. 1963. Araucanian Lakes. Plankton studies K. 32. P. Kaaret. 1953. Wasservegetation der Seen OrHi.ngen und in North Patagonia, with notes on terrestrial vegetation. Trehomingen. ISBN 91-7210-032-X. Price: SEK 160. ISBN 91-7210-047-8. Price: SEK 240. 33. E. Hasselrot. 1953. Nordliga lavar i Syd- och Mellans­ 48. E. Sj ogren. 1964. Epilitische und epigaische Moosvege­ T. verige. (Nordliche Flechten in Slid- und Mittelschweden.) tation in Laubwaldernder Insel bland, Schweden. (Surnrn. : ISBN 91-7210-033-8. Price: SEK 240. Epilithic and epigeic moss vegetation in deciduous woods 34. H. Sjtirs. 1954. Slatterangar i Grangarde Finnrnark. (Surnrn.: on the island of bland, Sweden.) ISBN 91-7210-048-6 Meadows in Grangarde Finnrnark, SW Dalarna, Sweden.) (ISBN 91-7210-448-1). Price: SEK 240. ISBN 91-7210-034-6. Price: SEK 160. 49. Hedberg. 1964. Features of afroalpine plant ecology. 0. 35. S. Kilander. 1955. Karlvaxternas ovre granser pa fj all i (Resume fran�ais.) ISBN 91-7210-049-4 (ISBN 91-7210- sydvastra Jamtland samt angransande delar av Harjedalen 449-X). Price: SEK 240. och Norge. (Summ.: Upper limits of vascular plants on 50. The plant cover of Sweden. A study dedicated to G. Einar mountains in southwestern Jamtland and adjacent parts of Du Rietz on his 70th birthday by his pupils. 1965. ISBN Harjedalen (Sweden) and Norway.) ISBN 91-7210-035-4. 91-7210-050-8. Price: SEK 460. Price: SEK 240. 51. Flensburg. 1967. Desrnids and other benthic algae of T. 36. Quennerstedt. 1955. Diatomeerna i Langans sj o­ Lake Kavsjon and Store Mosse. SW Sweden. ISBN 91- N. vegetation. (Surnrn.: Diatomsin the lake vegetation of the 7210-05 1-6 (ISBN 91-7210-45 1-1). Price: SEK 240. Langan drainage area, J amtland, Sweden.) ISBN 91-7210- 52. E. Skye. 1968. Lichens and air pollution. A study of crypto­ 036-2. Price: SEK 240. gamic epiphytes and environment the Stockholm region. in 37. M. -B. Florin. 1957. Plankton of fresh and brackish waters ISBN 91-7210-052-4 (ISBN 91-7210-452-X). Price: SEK in the Sodertalje area. ISBN 91-7210-037-0. Price: SEK 240. 160. 53. J. Lundqvist. 1968. Plant cover and environment of steep 38. M. -B. Florin. 1957. Insjostudier i Mellansverige. Mikro­ hillsides in Pite Lappmark. (Resume: La couverture vege­ vegetation och pollenregn i vikar av Ostersjobackenet och tale et l'habitat des flancs escarpes des collines de Pite insjoar fr:'inpreboreal tid till nutid. (Summ.: Lake studies in Lappmark.) ISBN 91-7210-053-2 (ISBN 91-721 0-453-8). central Sweden. Microvegetation and pollen rain in inlets Price: SEK 240.

Phytogeogr. Suec. Acta 87 Svenska Viixtgeogra.fiska Sallskapet 105

54. Conservation of vegetation in Africa south of the Sahara. to climatic conditions. ISBN 91-7210-070-2 (ISBN 91- Proceedings of a symposium held at the 6th Plenary meeting 7210-470-8). Price: SEK 240. oftheAETFAT, UppsalaSept. 12-16, 1966. 1. &0. Hedberg 71. C. Johansson. 1982. Attached algal vegetation in running (eds.) 1968. ISBN 91-7210-054-0 (ISBN 91-7210-454-6). waters of Jamtland, Sweden. ISBN 917210-071-0 (ISBN Price: SEK 290. 91-7210-471-6). Price: SEK 240. 55. L.-K. Kbnigsson. 1968. The Holocene history ofthe Great 72. E. Rosen. 1982. Ve getation development and sheep grazing Alvar of Oland. ISBN 91-7210-055-9 (ISBN 91-72 10- in limestone grasslands of South Oland, Sweden. ISBN 455-4). Price: SEK 290. 91-7210-072-9 (ISBN 91-7210-472-4). Price: SEK 290. 56. Hallberg. 1971. Vegetation auf den Schalenablager­ 73. Zhang Liquan. 1983. Ve getation ecology and population H. P. ungen in Bohuslan, Schweden. (Summ.: Vegetation on biology ofFritillaria meleagris L. at the Kungsangen Nature shell deposits in BohusHi.n,Sweden.) ISBN 91-7210-056-7 Reserve, eastern Sweden. ISBN 91-7210-073-7 (ISBN 91- (ISBN 91-7210-456-2). Price: SEK 240. 7210-473-2). Price: SEK 240 . 57. S. Fransson. 1972. Myrvegetation i sydvastra Viirmland. 74. Backeus. 1985. Aboveground production and growth I. (Summ.: Mire vegetation in southwesternViirrn land, Swe­ dynamics of vascular bog plants in central Sweden. ISBN den.) ISBN91-7210-057-5 (ISBN91-7210-457-0). Price: 91-7210-074-5 (ISBN 91-7210-474-0). Price: SEK 240. SEK 240. 75. E. Gunnlaugsd6ttir. 1985. Composition and dynamical 58. G. Wa llin. 1973. Lovskogsvegetation i Sjuharadsbygden. status of heathland communities in Iceland in relation to (Summ.: Deciduous woodlands in Sjuharadsbygden, recovery measures. ISBN 91-7210-07 5-3 (ISBN 91-7210- Vastergotland, southwestern Sweden.) ISBN 91-7210- 475-9). Price: SEK 240. 058-3 (ISBN 91-7210-458-9). Price: SEK 240. 76. Plant cover on the limestone Alvar on Oland. Ecology­ 59. D. Johansson. 1974. Ecology of vascular epiphytes in West sociology-taxonomy. E. Sjogren ( ed.) 1988. ISBN 91-7210- African rain forest. (Resume: Ecologie des epiphytes vas­ 076-1 (ISBN 91-7210-476-7). Price: SEK 320. H. Bjarnason. culaires dans la foret dense humide d 'Afrique occidentale.) 77. A. 1991. Vegetation on lava fields in the ISBN 91-7210059-1 (ISBN 91-7210-459-7). Price: SEK Hekla area, Iceland. ISBN 91-7210-077-X (ISBN 91- 290. 7210-477-6). Price: SEK 290. 60. H. Olsson. 1974. Studies on South Swedish sand vegetation. 78. Algological studies of nordic coastal waters -Afestschrift ISBN 91-7210-060-5 (ISBN 91-7210-460-0). Price: SEK to Prof. Mats Wrern on his 80th birthday-. I. Wallentinus 240. & P. Snoeijs (eds.). 1992. ISBN 91-7210-078-8 (ISBN 61. Hy tteborn. 1975. Deciduous woodland at Andersby, 91-7210-478-3 ). Price: SEK 290. H. eastern Sweden. Above-ground tree and shrub production. 79. Ta mrat Bekele. 1994. Ve getation ecology ofremnantAfro­ ISBN 91-7210-061-3 (ISBN 91-7210-461-9). Price: SEK montane forests on the Central Plateau of Shewa, Ethiopia. 240. ISBN9 1-7210-079-6 (ISBN 91-7210-479-1). Price: SEK 62. H. Persson. 1975. Deciduous woodland atAndersby, eastern 290. Sweden. Field-layer and below-ground production. ISBN 80. M. Diekmann. 1994. Deciduous forest vegetation in 91-7210-062-1 (ISBN 91-7210-462-7). Price: SEK 160. Boreo-nemoral Scandinavia. ISBN 91-7210-080-X (ISBN 63. S. Brakenhielm. 1977. Ve getation dynamics of afforested 91-7210-480-5). Price: SEK 290. farmland in a district of south-eastern Sweden. ISBN 91- 81. Plant root systems and natural vegetation. H. Persson & 7210-063-X (ISBN 91-7210-463-5). Price: SEK 240. I.O. Baitulin (eds.) 1996. ISBN 91-7210-081-8 (ISBN 91- 64. M. Ammar. 1978. Vegetation and local environment 7210-081-3). Price: SEK 290. Y. on shore ridges at Vickleby, Oland, Sweden. An analysis. 82. R. Virtanen & S. Eurola 1997. Middle oroarctic vegeta­ ISBN 91-7210-064-8 (ISBN 91-7210-464-3). Price: SEK tion in Finland and middle-northern arctic vegetation on 240. Svalbard. ISBN 91-7210-082-6. (91-7210-482-5). Price: 65. L. Kullman. 1979. Change and stability in the altitude of SEK 290. the birch tree-limit the southernSwedish Scandes 1915- 83. E. Kaimierczak. 1997. The vegetation of kettle-holes in in 197 5. ISBN 91-7210-065-6 (ISBN 91-7210-465-1 ). Price: central Poland. ISBN 91-7210-083-4. (91-7210-483-X). SEK 240. Price: SEK 290. 66. E. Wa ldemarson ens en. 1979. Successions in relationship 84. Swedish plant geography - dedicated to Eddy van der Maarel J to lagoon development in the Laitaure delta, North Sweden. on his 65th birthday-. H. Rydin, P. Snoeijs & M. Diekmann ISBN 91-7210-066-4 (ISBN 91-7210-466-X). Price: SEK (eds.) 1999. ISBN 91-7210-084-2 (ISBN 91-7210-484-8). 240. Price: SEK 400. 67. Tu hkanen. 1980. Climatic parameters and indices in plant 85. Succession and zonation on mountains, particularly on S. geography. ISBN 91-7210-067-2 (ISBN 91-7210-467-8). volcanoes - Dedicated to Erik Sjogren on his 65th birth­ Price: SEK 240. day-. E. van der Maarel (ed.) 2002. ISBN 91-7210-085-0 68. Studies in plant ecology dedicated to Hugo Sjors. E. Sjogren (ISBN 91-7210-485-6). Price: SEK 290. (ed.) 1980. ISBN 91-7210-068-0 (ISBN 91-7210-468-6). 86. S. Fransson. 2003. Bryophyte vegetation on cliffs and screes Price: SEK 290. in We sternV armland, Sweden. ISBN 91-7210-086-9 (ISBN 69. C. Nilsson. 1981. Dynamics of the shore vegetation of a 91-7210-486-4). Price: SEK 290. North Swedish hydro-electric reservoir during a 5-year 87. Eriksson, M. Niva & A. Caruso. 2007. Use and abuse 0. period. ISBN 91-7210-069-9 (ISBN 91-7210-469-4). Price: of reindeer range. ISBN 978-91-7210-087-9 (ISBN 978- SEK 240. 91-7210-487-7). Price: SEK400. 70. K. Wa renberg. 1982. Reindeer forage plants in the early grazing season. Growth and nutritional content in relation

A limited number of cloth-bound copies of Acta 44, 45, 46, 48, 49, 51, 52, 53, 56, 57, 61, 63, 66-87 is available at an additional cost of75 SEK per volume. (Use ISBN n°S. within brackets to order.) Acta Phytogeogr. Suec. 87 106 Svenska Viixtgeografiska Siillskapet

STUDIES IN PLANT ECOLOGY (VOL. 1-20)

1. S. Bnlkenhielm & lnge!Og. 1972. Ve getationen i Kungs­ 10. A. Miiller-Haeckel. 1976. Migrationsperiodik einzelliger T. hamn-Morga naturreservat med f6rslag till skotselplan. Algen in Fliessgewassern. ISBN 91-7210-810-X. Price: (Summ. : Vegetation and proposed management in the SEK 112. 11. Sjodin. 1980. Kungshamn-Morga Nature Reserve south of Uppsala.) A. Index to distribution maps ofbryophytes ISBN 91-7210-801-0. Price: SEK 112. 1887-1975. I. Musci. (hard-bound). ISBN 91-7210-811-8. 2. Inge!Og & M. Risling. 1973. Kronparken vid Upp­ Price: SEK 160. T. 12. Sj odin. 1980. sala, historik och bestandsanalys av en 300-arig tallskog. A. Index to distribution maps of bryophytes (Surnrn.:Kronparken, history and analysis of a 300-year old 1887-1975. II. Hepaticae. (hard-bound). ISBN 91-7210- pinewood near Uppsala, Sweden.) ISBN 91-7210-802-9. 812-6. Price: SEK 112. Price: SEK 112. 13. Eriksson, Palo & L. Soderstrom. 1981. Renbetning 0. T. 3. H. Sj ors et al. 1973. Skyddsvardamyrar i Kopparbergs vintertid. U ndersokningar reran de svensk tamrens niirings­ Hin. [Surnrn.: Mires considered for protection in Kopparberg ekologi under snoperioden. ISBN 91-7210-81 3-4. Price: County (Prov. Dalarna, central Sweden.)] ISBN 91-7210- SEK 112. 803-7. Price: SEK 112. 14. G. Wistrand. 1981. Bidrag till Pite lappmarks viixtgeografi. 4. L. Karlsson. 1973. Autecology of cliffand scree plants in ISBN 91-7210-814-2. Price: SEK 112. Sarek National Park, northern Sweden 1973. ISBN 91- 15. Karlsson. 1982. Euphrasia rostkoviana i Sverige. (English T. 7210-804-5. Price: SEK 160. surnrn.). ISBN 91-7210-815-0. Price: SEK 160. 5. B. Klasvik. 1974. Computerized analysis of stream algae. 16. Theory and models in vegetation science: Abstracts. R. ISBN 91-7210-805-3. Price: SEK 112. Leemans, I.C. Prentice & E. van der Maarel (eds.) 1985. 6. Dahlstrom-Ekbohm. 1975. Svensk miljovards- och ISBN 91-7210-816-9. Price: SEK 160. Y. omgivningshygienlitteratur 1952- 1972. Bibliografi och 17. Backeus. 1988. Mires in the Thaba-Putsoa Range of the /. analys. ISBN 91-7210-806-1. Price: SEK 112. Maloti, Lesotho. ISBN 91-7210-817-7. Price: SEK 160. 7. L. Rodenborg. 1976. Bodennutzung, Pftanzenweltund ihre 18. Forests of the world - diversity and dynamics (Abstracts) Ve riinderungen in einem alten Weidegebiet auf Mittel­ E. Sjogren (ed.) 1989. ISBN 91-7210-81 8-5. Price: SEK Oland, Schweden. ISBN 91-7210-807-X. Price: SEK 290. 112. 19. E. Sj ogren. 1994. Changes in the epilithic and epiphytic 8. H. Sj ors & Ch. Nilsson. 1976. Vattenutbyggnadens effekter moss cover in two deciduous forest areas on the island of pa levande natur. En faktaredovisning overvagande fran bland (Sweden). - A comparison between 1958-1962 and Umealven. (Surnrn.: Bioeffects of hydroelectric develop­ 1988-1990. ISBN 91-7210-819-3. Price: SEK 200. ment. A case study based mainly on observations along 20. Vegetation science in retrospect and perspective (Abstracts) the Ume River, northern Sweden.) ISBN 91-7210-808-8. 1998. E. Sjogren, E. van der Maarel & G. Pokarzhevskaya Price: SEK 160. (eds.) ISBN 91-7210-820-7. Price: 160 SEK. 9. Lundqvist & G. Wistrand. 1976. Strandftora inom ovre 1. och mellersta Skellefteiilvens vattensystem. Med en sam­ manfattning betraffandebotaniska skyddsviirden. (Surnrn.: Riverside vascular florain the upper and middle catchment area of the River Skellefteiilven, northern Sweden.) ISBN 91-7210-809-6. Price: SEK 112.

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ISBN 978-9 1-721 0-087-9 (ISBN 978-91-7210-487-7) ISSN 0084-5914