A century of logging and forestry in a reindeer herding area in northern Sweden

Forest Ecology and Management 256 (2008) 1009–1020

Contents lists available at ScienceDirect

Forest Ecology and Management journal homepage: www.elsevier.com/locate/foreco

A century of logging and forestry in a reindeer herding area in northern Sweden ¨ stlund a, Jon Moen b, Johan Olofsson b Anna Berg a,*, Lars O a b

Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, SE-901 83 Umea˚, Sweden Department of Ecology and Environmental Science, Umea˚ University, SE-901 87 Umea˚, Sweden

A R T I C L E I N F O

A B S T R A C T

Article history: Received 17 December 2007 Received in revised form 30 May 2008 Accepted 3 June 2008

Boreal forest ecosystems are generally highly sensitive to logging and other forestry activities. Thus, commercial forestry has had major effects on the forests and landscape structure in northern Sweden since the middle of the 19th Century, when it rapidly extended across the region. Lichens (which constitute up to 80% of reindeer forage in winter and early spring) have often been amongst the most severely affected ecosystem components. The overall aim of the present study was to analyze how forestry has influenced the potential supply of ground-growing lichens as winter forage for the reindeer in this region over the past ca. 100 years. For this purpose, we analysed changes in forest and stand structure in Scots pine-dominated (Pinus sylvestris L.) reindeer wintering areas in the southern part of the county Norrbotten (covering ca. 58,000 ha) using detailed historical forest inventories and management plans. We found that the amount of the forest types considered potentially good pasture (mainly middleaged and old pine forests) decreased during the first part of the 20th Century. However, the quality of grazing grounds was improved by forestry during this time mainly because selective logging made the forests more open which benefits lichen growth. During the last part of the 20th century forestry impaired the quality of grazing grounds in several ways, e.g. by clear-cutting and intensified use of various silvicultural measures. We conclude that ca. 30–50% of the winter grazing grounds have been lost in the study area because of intensive forest management during the last century. The spatially precise historical information about the affects of forestry on lichen pasture provided in this study can be used to direct forest management which will facilitate and promote reindeer herding in the future. ß 2008 Elsevier B.V. All rights reserved.

Keywords: Cladina Reindeer pasture Forest history Forest structure Multiple use management

1. Introduction Forest ecosystems in northern Fennoscandia have supported human societies for millennia, notably the indigenous Sami people, who made their living as hunter-gatherers and reindeer herders in the vast boreal forests of Norway, Sweden, Finland and Russia. Reindeer husbandry has changed over time from a small-scale, family-based activity to a much larger scale enterprise during the 20th Century, in which reindeer herds are scattered over large areas in the mountains during summer and only handled at strategic moments, such as calf-marking and slaughtering. In the winter, the herds are moved or migrate to the boreal (Anon., 2006). In some areas, such as in this study, reindeer graze in the boreal forest all year round where they migrate between different grazing grounds within the forest. During the snow-free period of the year reindeer graze on a wide range of plant species (Baskin and Danell, 2003; Ma˚rell, 2006), but during winter the nutrient value of most

* Corresponding author. Tel.: +46 90 786 86 44; fax: +46 90 786 81 66. E-mail address: [email protected] (A. Berg). 0378-1127/$ – see front matter ß 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.foreco.2008.06.003

plant species decreases substantially. However, lichens have high carbohydrate contents even in winter. Furthermore, unlike most other animals, reindeer have a specialised microbial flora in their rumen that enables them to digest lichens (Storeheier et al., 2002; Sundset et al., 2004). Thus, in winter and early spring lichens may constitute up to 80% of reindeer forage (Heggeberget et al., 2002), and winter grazing grounds are generally the primary limitation on reindeer husbandry in Sweden (Sandstro¨m et al., 2006). During the last 150 years the forest ecosystems of boreal Sweden have been subjected to radical changes. Before the 1850s, exploitation of forestland in the northern parts of Scandinavia was largely restricted to spatially broad but low intensity uses by Sami ¨ stlund et al., 2003); only people and local agrarian activities (O ¨ stlund, 1998; small proportions of it were logged (Linder and O Axelsson, 2001). However, in the 1850s, the sawmill industry expanded and the search for large Scots pine timber progressed northwards in Scandinavia (Arpi, 1959). Since that time logging has had major effects on the forests and landscape structure in the region. In addition to logging, forest management methods have ¨ stlund et al., 1997). also affected the forests (Ebeling, 1955, 1959; O These developments have led to a complete transformation of the

1010

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

forest landscape since the 19th Century (Ericsson et al., 2000; ¨ stlund, 2001). Axelsson and O Since both the reindeer herding Sami and the forest owners use the same forest ecosystem, although for different purposes, the overlapping use has led to increasing conflicts between the two parties (Sandstro¨m et al., 2006). Furthermore, the impacts of the activities of the reindeer herders and especially the forest owners have increased during the last 100–150 years, and the management methods have also changed considerably. The resulting changes in the ecosystems, including the transition from predominantly fire-disturbance to forest management disturbance, have been complex and should be considered at several spatio-temporal scales. The ecological effects include, for example, fragmentation of older forests, loss of biodiversity (Linder and ¨ stlund, 1998) and overall increases in productivity. The groundO growing lichens in boreal forest ecosystems that reindeer feed upon are particularly sensitive to forest management for several reasons; reindeer lichens have poor dispersal and relatively low growth rates, so they are slow to regenerate after disturbance (Eriksson and Raunistola, 1990; Sunde´n, 2003; Bostro¨m, 2004); they are light-dependent, and when canopies close mosses tend to expand in the ground vegetation (Sulyma and Coxson, 2001); and they are easily out-competed by vascular plants at higher nutrient levels (Kellner and Ma˚rshagen, 1991). Forestry adversely affects the lichen resources required for reindeer husbandry in several ways. For instance, clear-cutting and soil scarification reduce the availability of grazing lands and the amount of lichens available (Eriksson and Raunistola, 1990). In addition, harvesting of old-growth forests strongly reduces the availability of arboreal lichens (Dettki and Esseen, 1998). However, although the generally negative effect of forestry on winter grazing grounds for reindeer is not disputed nowadays, very few data are available for quantifying this effect. Data from the Swedish National Forest Inventory suggest that as much as 50% of lichen-rich winter grazing grounds may have been lost since the 1950s (Anon., 2001; Sandstro¨m et al., 2006), but these are coarse regional estimates and are not related to the spatial scale of practical reindeer herding. The ongoing management of winter grazing grounds through improved consultation procedures between the two parties requires more detailed information on losses of grazing grounds and better understanding of the historical and ecological processes underlying these changes.

Thus, the overall aim of the study presented here was to analyze how forestry has influenced the potential supply of groundgrowing lichens as winter forage for reindeer over the past ca. 100 years. To do this, we analysed changes in forest and stand structure in Scots pine-dominated winter grazing areas in the southern part of the county Norrbotten (covering ca. 58,000 ha) using detailed historical forest inventories and management plans. We also evaluated possible losses in grazing grounds based on various assumptions or ancillary data on known or inferred relationships between forest structure and forest management, and the ecology of ground-growing lichens. The specific objectives were: (1) to analyse changes in forest structure (in terms of age, species composition, density and potential timber growth) in a reindeer herding area over the last 100 years, (2) to identify the silvicultural methods that have been used in the same area during this period, and (3) to estimate losses in potential reindeer grazing grounds in this area over time. 2. Materials and methods 2.1. Study site The study area lies at latitude 668000 in north-western Sweden, in the county of Norrbotten, in the middle boreal zone of Sweden (Sjo¨rs, 1965). It covers about 58,000 ha and consists of three former National Forests—Akkajaur, Abraur and Eggelats (Fig. 1). The borders of Akkajaur were extended to their current northern limits in 1936. Akkajaur and Abraur are adjacent to each other and will hereafter mostly be referred to jointly as the Akkajaur/Abraur area (the main study area), which today occupies 26,600 ha of forest land. For geographical comparisons, an additional area (Eggelats) is used, which is situated ca. 20 km further north-west and occupies 17,800 ha of forest land (Fig. 1). A major river, Pite a¨lven, runs through both areas from northwest to southeast. The central part of Akkajaur/Abraur consists of plains at approximately 400 m a.s.l., surrounded by hills that reach elevations of ca. 550 m a.s.l. In Eggelats the river is split into several minor channels. The landscape includes hundreds of small lakes at about 450 m a.s.l., surrounded by hills reaching 700– 800 m a.s.l. The temperature range over the year is large, with means of 14 8C in January and +13 8C in July. There are 200–225 days with

Fig. 1. Locations of the study areas. Gray areas are forest land, black areas are water and white areas are mires and unidentified lands.

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

snow every year, with an average snow depth of 60 cm in January (SMHI, Swedish Meteorological and Hydrological Institute, 2006). The dominant forest type is coniferous, and Scots pine (Pinus sylvestris L.) is the dominant tree species accounting for 80–90% of the standing volume. Norway spruce (Picea abies (L.) Karst.) is the second most common tree species. There is also a small component of deciduous trees (mainly Betula sp.). The ground vegetation is dominated by lichens, feather mosses and dwarf-shrubs.

1011

2.3. Sources of historical data This study is based on primary historical sources from the Swedish National Forest Service (Doma¨nverket) and data recently acquired by the present forest owners Sveaskog and Fastighetsverket. The primary sources used in the study were detailed forest inventories and management plans consisting of maps and stand descriptions. These records contain information on forest structure

2.2. General history of the study area The Sami people have lived in the area for thousands of years. They traditionally made their living from hunting reindeer and moose, fishing in streams and lakes and gathering vegetable food such as Angelica archangelica L., Rumex acetosa L. and Scots pine inner bark (Bergman et al., 2004; Quarnstro¨m, 2006). Numerous archaeological findings of Sami origin, primarily hearths, are found in the area (Karlsson, 2007). The Sami started to herd reindeer at least in the 17th Century (Lundmark, 1982). Originally this was an intensive form of herding in which the people lived close to the animals and moved with them between different areas in different seasons (Lundmark, 1982). The reindeer were used for various purposes, e.g. as transport animals and sources of meat and milk for food, fur for clothing and cots, and sinews for strings. This wide range of uses made the reindeer an essential part of Sami life and culture. The tradition of reindeer herding still exists in this area today, but since the late nineteenth century reindeer herding has become more extensive, involving larger herds and fewer people. The patterns of migration to find the best pastures are still the same, but today motor vehicles are used to keep track of the reindeer (Anon., 2006). As in the rest of boreal Sweden, the forests in the study area have been subjected to dramatic changes in land-use during the last 150 years. Before the 19th Century the timber in the forests had little real economic value, but during this century its value was increasingly realised. The forest land was delimitated and in northern Sweden the main parts became national forests owned by the Crown. In these areas forestry was practiced, but reindeer herders also had grazing rights. People from the rest of Sweden received special privileges from the crown if they settled in these northern inland parts of Sweden. Conflicts between the settlers and the reindeer herding Sami were common, mainly because the reindeer grazed crops the settlers used for their cattle. At the end of the 19th Century conflicts between Sami and forestry enterprises started to be more common, since foresters thought that reindeer grazing posed a threat to the regeneration of forests (Anon., 2006). In northern Sweden the sawmill industry expanded in the 1850s and the search for large Scots pine ¨ stlund, 1995). In 1870 a forest officer was timber progressed (O appointed to manage the forests in the study area (Holmgren, 1959) and in 1871 the Akkajaur area was set aside as national forest (at that time called Trollforsen). A general shift from highgrading of the largest Scots pines to more varied forest exploitation occurred in the early 20th Century, due to increasing demands for pulpwood, firewood, and smaller saw timber ¨ stlund et al., 1997). The forests close to the rivers were logged (O first. In the middle of the 20th Century forestry became more intensive, involving clear-cutting and forest management practices such as sowing, planting, prescribed burning, thinning, soil scarification, applications of herbicides, nitrogen fertilization ¨ stlund et al., 1997). Forest and introduction of exotic species (O management was based on the general concept of transforming the ‘‘wild forests’’ into modern, efficient, high-yielding domesticated stands. This conceptual shift made the forest and every tree in it economically valuable.

Table 1 Archival sources used in the study The provincial archives, Ha¨rno¨sand, Sweden (Landsarkivet) Kungliga doma¨nstyrelsen/Doma¨nverket arkiv (Archives of the Forest Service) Forest management plans with maps FIII.2a. Krp. Abraur A I-E Skogsindelningshandlingar (Stand descriptions) 1927/29, block 1–2 Skogsindelningskarta (Map) block 1 1910, block 3 1907 FIII.2a. Krp. Eggelats Skogsindelningshandlingar (Stand descriptions) 1925, block 1–2 FIII.2a. Krp. Trollforsen AI Skogsindelningshandlingar (Stand descriptions) 1894/1895, block 1–3 and 5 Skogsindelningskarta (Map) kronoparken trollforsen/JokkmokksArvidsjaurs kronopark 1893, block V Skogsindelningshandlingar (Stand descriptions) 1926, block 1–4 Malmesjaur revir Krp Akkajaur 1936, skifte 1–6 (Stand descriptions) Krp Abraur 1936, skifte 1–2 (Stand descriptions) Kartor 1936 (Map). A¨ldre skogskartor. Abraur sk I–II, Akkajaur sk I–VI. Kartor – a¨ldre skogskartor (1960) Abraur sk I–II, Akkajaur sk I–VI (Map) Kartsamlingen Kronoparken Akkajaur, sk 1–4, Nr: Nb 6–9, Plats XII:4 (Map) Arvidsjaurs revir (Volym utan signum). ‘‘A¨ldre kartor. Samtliga revir’’. Kronoparken Trollforsen, Jokkmokks-Arvidsjaurs kronopark 1894/1895, block I–III, V (Map) FXII. Skogsindelning, skoglig arealplanering 1976 Krp Akkajaur, skifte 1–4 (Stand descriptions) Arjeplogs revir HIV:158. Skogskartor-ortofotokartor 1973. Nr 1–21. Ajourha˚llna (Map) HIV:159. Skogskartor-ortofotokartor 1973. Nr 22–31, 34–42 Ajourha˚llna (Map) HII. Kronoparkskartor Kronoparken Eggelats sk I-II, 1936, 1923 (Map) H:340 Kartor (Map). Uppskattningshandlingar och avverkningsfo¨rslag fo¨r a˚r 1930, 1931, 1935 och 1936 kartlagda kronoegendomar. Besta˚ndsbeskrivning 1931, 1936. Krp Eggelats, skifte 1–2 H:340 (H:154). II. Kronoparkskartor 1960 a˚rs skogsindelning nr 1–10 (Map). A¨mnesordnade handlingar Skogsindelning 50.01 1973 a˚rs skogsindelning, Eggelats (Stand descriptions) Skogstaxeringsavdelningen FIVa. Indelningsplaner, uppskattningshandlingar, avverkningsplaner Arjeplogs revir 1960. Malmesjaur revir 1961. Krp Eggelats, skifte 1–2 (Stand descriptions) Kartsamlingen Kronoparken Eggelats, Block 1–2, Norrbottens la¨n – kartnummer: A 362, A363 (Map) Kronoparken Trollforsen, Block 1–4, Norrbottens la¨n – kartnummer: A 508–511 (Map) Documented silvicultural practices Malmesjaurs revir B:II. 1,2,3. Malmesjaur revir – A˚rsbera¨ttelser (Yearly reports) D:IV Liggare o¨ver a˚tga¨rder 1,4. Skogsodlingsliggare (site-specific information on silvicultural measures) EII:5 Sakordshandlingar Skogsva˚rdsa˚tga¨rder (site-specific information on silvicultural measures)

1012

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

variables, such as tree species composition, tree age, timber volume, etc. Records documenting silvicultural measures, such as scarification, herbicide spraying, planting, etc., have also been used (Table 1). The historical sources all contain quantitative data and collectively cover a period of 111 years, from 1895 to 2006. Forest inventories are available concerning the Abraur area for the years 1895 (for the part north of lake Abraur), 1929, 1936, 1960 and 2006; from the Akkajaur area for 1895, 1926, 1936, 1960, 1976 and 2006; and from the Eggelats area for 1936, 1960, 1973 and 2005. The forest inventories contain stand-level data on tree species composition and age structure for all years. The recorded trees are classed as Scots pine (P. sylvestris), Norway spruce (P. abies) or deciduous trees (in all years except 1895, when only coniferous trees were counted). The most recent inventory also includes records on the introduced species lodgepole pine (Pinus contorta). Data on timber volume and potential timber growth are available for all years except 1895. For the inventory years between 1895 and 1936 there is also information about canopy openness. In some years sporadic information on field-layer vegetation is included. The documents from before the 21st Century are written documents, while the 2006 data have been extracted from digital forest stand databases maintained by Sveaskog and Fastighetsverket. Data on silvicultural measures applied at various spatial levels are available for different periods. Information regarding the whole district including Akkajaur and Abraur, covering 93,300 ha forest land in total, is available in yearly reports from 1924 to 1964, with incomplete information for the periods 1924–1936 and 1940– 1944. At stand level the nature of the data on silvicultural measures applied varies substantially (Table 1), but they all provide information on the types of measures applied, the dates they were applied and the areas concerned. The sources of this type cover the periods 1938–1952 and 1977–1984 for the Akkajaur area. The forest inventory maps of Akkajaur, Abraur and Eggelats also contain spatial and temporal information on silvicultural measures. 2.4. Data analysis A careful evaluation of the sources, their origin and their quality is essential when working with historical documents of any kind (Cipolla, 1991; Whitney, 1994). The historical records, forest inventories including maps and yearly management data, used in this study are generally of good quality and they provide detailed data on the forest structure relatively far back in time to a more detailed level than any other sources or methods can provide. These types of sources have been used for different purposes in many similar studies and have been found to be consistent and ¨ stlund reliable for temporal interpretations of forest change (O ¨ stlund, 2001; Gimmi and et al., 1997; Burgi, 1999; Axelsson and O Burgi, 2007; Gimmi et al., 2008). When using this kind of historical records it is important to consider that the forest management ideas have profoundly changed during the 20th Century and so have the forests. In turn this has influenced the methods of the ¨ stlund, 2001). Generally the forest inventories (Axelsson and O inventories before and after the shift from selective logging to clear-cutting in the middle of the 20th Century are less comparable ¨ stlund et al., 1997), and therefore the interpretation of primary (O data and subsequent reclassification is crucial. 2.4.1. Landscape level To compare different forest inventories, the data they contained ¨ stlund et al., 1997; Ska˚nes had to be generalized and reclassified (O ¨ stlund, 2001). Therefore, a and Bunce, 1997; Axelsson and O detailed classification system based on species composition and

Table 2 Age and species classes used to reclassify stands in forest inventories Forest stand type

Tree species

Tree age

1. 2. 3. 4. 5.

All >80% >80% >80% <80%

>90% >60% >60% >60% All

Clear-cuts Young Pine Middle-Aged pine Old Pine Others

pine pine pine pine

0 years 1–40 years 40–120 years 120 years

age was used to classify each forest stand in each of the years for which information was available in the surveys, according to five types of stands (Table 2). In the inventories from 1895 and from the 1920s the share of deciduous trees were not or inconsistently documented. It is therefore likely that the deciduous part is underestimated during this period. The age of forest stands have been documented in different ways at different inventory occasions. In the inventories before 1960 the stand age were presented in age classes, whereas after 1960 the age of a stand is given as a mean. This reflects the change from multi-aged stands as the norm in the beginning of the 20th Century to even-aged stands being the norm after the introduction of clear-cutting in the latter ¨ stlund et al., 1997). part of the century (O Between the years 1920 and 1973 potential growth was measured using the system of Tor Jonsson, giving the average timber-producing capacity of the forest in m3 timber ha1 yr1 in an ideal state (Jonsson, 1914). After 1973 another system for measuring potential growth was used; site index (Ha¨gglund and Lundmark, 2005). Thus, in order to compare data from all of the years, the data from before 1973 were converted into site indices, using a table in Ha¨gglund and Lundmark (2005). 2.4.2. Stand level We identified two sites with potentially good reindeer winter forage and analyzed the spatial and temporal changes that have occurred to the forests in them during the study period. Both sites are within the Akkajaur/Abraur area. Site Abmokheden was chosen because it was described as a poor heath with lichens in the forest inventories from 1926, and thus was a typical good area for reindeer foraging in winter. It is also situated along a traditionally important migratory route for reindeer along the river Pitea¨lven (Karlsson, 2007). Site Paulovuopme was chosen because it is known to have traditionally provided good winter forage for reindeer (Hedman, 2004). The Sami name of the area, Paulavuopme, indicates that it is a vast expanse of forest in which snow melts early in spring (Karlsson, 2007). It is a pine heath on glaciofluvial sediments – generally a good habitat for lichens – and was classified as lichen-rich in a recent vegetation survey of the county of Norrbotten commissioned by the County Administration Board. The same types of analyses as the landscape-level analyses described above were conducted in these areas to compare the changes that occurred in them with developments in the surrounding landscape. Table 3 Stand parameters in logged and non-logged stands in Akkajaur, 1895

Area (ha) Area (% of total area) Average tree age Pine (%) Spruce (%) Openness Growth index Average stand size (ha)

Not logged

Logged

23,509 96 204 93 7 0.63 0.63 58

869 4 55 100 0 0.52 0.6 109

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

3. Results 3.1. Changes in forest structure The forest in the studied area has undergone radical changes since the end of the 19th Century. The most obvious are the changes in tree age. In 1895 the mean age of the stands in Akkajaur/ Abraur was >200 years (Table 3), while in 2005 the mean age in this area was just 66 years. Furthermore, the proportion of Old Pine

1013

Forest in 2005 was only an eighth of the corresponding area in 1895. Reciprocally, the proportions of Young Pine Forest and Middle-Aged Pine Forest steadily increased. The proportion of clear-cuts peaked in 1960, when 40% of the pine forest was recorded as recently cut (Fig. 2). Yet another major change in forest structure that has occurred in Akkajaur/Abraur during the last ca. 100 years is that the density of the forest stands has increased. The standing volume in MiddleAged and Old Pine Forests has increased considerably since the

Fig. 2. Changes in age structure from 1895 to 2005 of stands in: (A) Akkajaur/Abraur (only Akkajaur is included in the data from 1976); and (B) Eggelats.

Fig. 3. Changes in standing volumes in four age-based classes of pine stands in Akkajaur/Abraur.

1014

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

Fig. 4. Changes in Site Productivity Indices, derived by conversion factors in Ha¨gglund and Lundmark (2005) at Akkajaur/Abraur and Eggelats.

beginning of the 20th Century (Fig. 3). The site productivity index has also steadily increased (Fig. 4). In Eggelats the patterns of changes in the forests and forest use resemble those that occurred in Akkajaur/Abraur, but the forestry activities started later in Eggelats. The first data for Eggelats were from the 1930s, when the proportion of Old Pine Forest was 60%, compared to 40% in Akkajaur/Abraur at the same time, and this proportion is still higher in Eggelats now (Fig. 2). Similar increases in standing volume and site productivity index to those that occurred Akkajaur/Abraur were also detected in the Eggelats data (Fig. 4). 3.2. Changes in the use of silvicultural measures Forest management practices, primarily intended to improve regeneration, had already begun to be applied in the region in the early 1920s (Figs. 5 and 6). Measures applied at the time were mainly limited to sowing pine and spruce seeds, with some soil scarification (which became more common in the 1950s) (Fig. 6). To facilitate regeneration, prescribed burning on clear-cuts and seed tree stands was used in some cases, mainly during the period from 1949 to somewhere between 1963 and 1978 (Fig. 6). The forest has been thinned, and frequencies of unwanted tree species have been reduced since the 1920s, but use of these measures increased in the 1950s when the overall forest management regime became more intense. Chemical treatments of clear-cuts

with herbicides to suppress deciduous trees were introduced at the same time (Fig. 5). 3.3. Detailed stand analysis In Abmokheden the forest in 1895 was younger than the average for the area as a whole in the same period. Its age increased until the next inventory occasion in 1925, when the forests in Abmokheden were classified as purely Old Pine Forest. In 1973, 60% of the area was clear-cut and in 2005 only 5% of the area was classified as Old Pine Forest (Fig. 7). In Paulavuopme the forest was older than in Abmokheden in 1895; 75% of the forest there was classified as Old Pine Forest. In 1960, 40% of the area was clear-cut, but in 2005 as much as 33% was still classified as Old Pine Forest, which is a high proportion compared with the surrounding forests (Fig. 8). Neither Abmokheden nor Paulavuopme were affected by forestry measures during the years for which we have site-specific data (1938–1952 and 1977–1984). 4. Discussion 4.1. The development of forests Since the late 19th Century forestry has been the major factor influencing the forest landscape and forest structure in the study

Fig. 5. Changes in the use of pre-commercial thinning and chemical treatments with herbicides. Data cover the whole district including Akkajaur/Abraur available for the period 1936–1963.

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

1015

Fig. 6. Changes in the use of site preparation, prescribed burning and artificial regeneration in the whole district including Akkajaur/Abraur (shaded areas), and only Akkajaur (bars). Data for the whole district are available between 1924 and 1963, and data for Akkajaur are available for the periods 1938–1953 and 1977–1984.

area, as in other parts of boreal Sweden (Ericsson et al., 2000), and boreal Scandinavia (Kouki et al., 2001; Lofman and Kouki, 2003). Accordingly, trends in forest management and policy have had considerable effects on the forests. In this study three main periods of forest history could be distinguished during the last 150 years: the period before industrial forestry began (before ca. 1890), the period of selective logging (ca. 1890–1950); and the period of intensive forestry and clear-cutting (ca. 1950 to date). The changes that occurred in forest utilization are similar to those that occurred

in the rest of boreal Sweden, although the timing of the stages ¨ stlund, 1993; O ¨ stlund et al., 1997). The differed between areas (O general pattern is that industrial use of the forests started with large-scale cuttings that targeted the largest trees to provide the expanding sawmill industry with high quality saw timber (Arpi, 1959). The logging was mainly selective until the 1950s, when intensive forest management began, including clear-cutting as the main logging method (Holmgren, 1959). Silvicultural measures to ensure regeneration began to be used during the first part of the

Fig. 7. Changes in age structure in the forest stands in Abmokheden between 1895 and 2005.

1016

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

Fig. 8. Changes in age structure in the forest stands in Paulavuopme between 1895 and 2005.

20th Century (Holmgren, 1959), but their usage did not expand to a large-scale until the 1950s (Ebeling, 1959). 4.1.1. The period before industrial forestry (before ca. 1890) In the areas for which we have the earliest data, the Akkajaur/ Abraur area, the forests were predominantly Old Pine Forests at the end of the 19th Century (Fig. 2), with a mean age of more than 200 years (Table 3) and a generally heterogeneous age structure (Table 1, Stand description Trollforsen 1894/1895). Multi-aged pine forests, shaped primarily by recurrent low intensity fires, were common in boreal Fennoscandia before the beginning of ¨ stlund et al., 1997; Axelsson and O ¨ stlund, industrial forestry (O 2001; Kuuluvainen et al., 2002). Forest fires were the primary driving forces shaping and affecting the successional development of boreal forests in Sweden before industrial forestry began and efficient fire suppression measures were applied, beginning in the 19th Century (Zakrisson, 1977; Niklasson and Granstro¨m, 2000; Hellberg et al., 2004). In Akkajaur/Abraur many trees had scars from previous fires at the time of the first inventory in 1895 (Stand description, Trollforsen, 1895), confirming that fires were important ecological factors also in this area. The frequency of fires was dependent on factors such as biotope type, landscape topography, exposure (Zakrisson, 1977) and human activity (Niklasson and Granstro¨m, 2000). In a boreal forest landscape in northern Sweden the mean fire interval, before fire-suppressing policies were widely applied, was in general 80 years, with much shorter mean intervals, ca. 45 years, for dry pine forests of lichen-Calluna type on flat lands (Zakrisson, 1977). In the pine-dominated forests in Akkajaur/Abraur Calluna frequently occurred in the ground vegetation in 1895, sparsely mixed with other dwarf-shrubs and forest mosses, or (on heath lands) mixed with reindeer lichen (Stand description Trollforsen 1894/1895) indicating generally dry conditions. It is therefore likely that the fire interval was quite short at this site. Pine-dominated forests with fire intervals of 20–50 years develop into sparse forests with many very old pine trees (Kohh, 1975). Since fire kills most young trees it reduces tree density and hence benefits the surviving old pine trees (Wirth et al., 1999). Studies in Swedish boreal forests have shown that the standing timber volume was higher and concentrated in fewer, generally much larger, trees before the ¨ stlund and beginning of industrial forestry than it is today (O ¨ stlund, 1998). Linderson, 1995; Linder and O 4.1.2. The period of selective cutting (ca. 1890–1950) Like most forests in boreal Sweden, the forests examined in this study were affected by the timber frontier that swept across the country targeting the largest pine trees in the 19th Century

¨ stlund, 1993). Generally, the frontier moved from south to north, (O and from the coast up the major rivers. Accordingly, the timing of the first logging differed between the areas in this study. In the 1895 survey of Akkajaur/Abraur, 4% of the forest was reported as being previously logged. However, Eggelats was probably still not affected by logging at this time since the transport of timber was ¨ stlund, 2002), and dependent on floatable streams (To¨rnlund and O the river Pitea¨lven was only prepared for floating up to Akkajaur/ Abraur in 1895 (Table 1, Stand description Trollforsen 1894/1895). Eggelats was also less affected by logging in 1926, as indicated by its larger proportion of Old Pine Forest, than in Akkajaur/Abraur at the same time (Fig. 2). The first stands to be cut in Akkajaur, before 1895, were all pure pine stands (Table 3), and many of them were in the Paulavuopme heathland. The low proportion of Old Pine in Abmokheden in 1895 indicates that this area had also been logged before the 1895 inventory (Fig. 7), but since it is not recorded as having been previously logged in the 1895 inventory and it had a high proportion of Middle-Aged Pine and no Young Pine stands, it was probably logged long before 1895. The floatway was not ready at that time, thus the logging was probably done by local inhabitants, or alternatively the area may have been severely burnt. Selective cuttings reduced the standing timber volume to lower levels than those in with the forests primarily shaped by natural ¨ stlund, 1998; Myllyntaus and disturbance factors (Linder and O Mattila, 2002). In the Akkajaur/Abraur area the logging method was mainly selective, targeting the largest trees at least until 1936, as indicated by the decline in Old Forests and increase in MiddleAged Forests (Fig. 2), together with the reduction in standing volume in the older forest, between 1929 and 1936 (Fig. 3). During the first part of the 20th Century modern forestry practices, e.g. non-commercial thinning, soil scarification and sowing, were introduced at a small-scale (Figs. 5 and 6), following the general trends in forestry in northern Sweden (Holmgren, ¨ stlund et al., 1997). The methods for soil scarification used 1959; O during this period generally only moderately affected the ground vegetation (Holmgren, 1959). 4.1.3. The period of intensive forest management (ca. 1950 until today) Clear-cutting was the predominant logging practice in 1960, as indicated by the increases in clear-cuts at that time in both Akkajaur/Abraur and Eggelats (Fig. 2). Clear-cutting probably started during the 1950s, simultaneously with the increasing application of silvicultural measures (Figs. 5 and 6), following the ¨ stlund same trends as in the rest of boreal Sweden (Ebeling, 1959; O et al., 1997).

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

Various reforestation measures, such as prescribed burning and soil scarification, were used to improve the conditions for tree seeds and seedlings during the second half of the 20th Century (Fig. 6). Prescribed burning was used from 1947 until at least 1963 (Fig. 6), and according to the forest owner Doma¨nverket, prescribed burning should have been mostly used on moist ground with a thick layer of raw humus (Anon., 1976). We have local information concerning silvicultural measures applied in the Akkajaur area during two periods: 1938–1953 and 1977–1984 (Fig. 6). During the first period soil scarification was applied to on average 0.04% of the forested land per year, but in the second period this figure had increased to 0.39% (Fig. 6). It was not possible to identify the types of methods used for soil scarification in the area during this time from our data sources. However, in the instructions from the forest owner Doma¨nverket three types of soil scarification methods were recommended in 1976: mounding and disc trenching in dry to moist sites and ploughing on soils with thick, inactive humus layers (Anon., 1976). Thinning started in the beginning of the 20th Century, but its use expanded in the 1950s when also chemical treatments to remove unwanted tree species started to be used (Fig. 5). Thinning made the forests more open, but usually only for a short while, since large trees with high growth potential were left to expand after thinning (Anon., 1976). No data on nitrogen fertilization were ¨ stlund et al. (1997) have provided available for Akkajaur, but O evidence showing that nitrogen fertilization was used from the 1970s to 1990 in their study area in Lycksele further south. Forestry in this period of intensive forest management has fragmented the forest landscape, turning it from a landscape of old multi-storied forest into a patchwork of even-aged forests (Lofman and Kouki, 2003), as reflected by the decrease in Old Pine Forests (Fig. 2) and the switch from selective logging to clear-cutting. Forestry practices also included building roads and ditches, which added to the fragmentation. 4.2. Effect of forest changes on lichen abundance and availability Forestry affects the quantities of ground-growing lichens present in forest stands both directly (by removing lichens) and indirectly (by changing the abiotic conditions in forests and thus their growth conditions). Furthermore forestry can affect the accessibility of lichens to reindeer in winter. Fire and soil scarification have direct effects on lichen abundance by removing ground-growing lichens to various extents. While severe fires consume almost all the lichens (Webb, 1998), soil scarification affects 35–90% of the vegetation cover, depending on the method used (Eriksson and Raunistola, 1990). In this study area fire had substantial effects on the forest during two periods: the period before industrial forestry began and the period of prescribed burning during the 1950s and 1960s (Fig. 6). Between 1949 and 1953 on average 0.33% of the forested area in Akkajaur was burnt yearly (Fig. 6). No comparable figures on fire intervals are available for the area in the period before industrial forestry began, but they are presumed to have been similar to intervals inferred by Zakrisson (1977) in the valley of the river Vindela¨lven south of our study area, where ca. 1.2% of the forested land appears to have been burnt annually. If so, fires were never as frequent during the era of prescribed burning in Akkajaur as they were before the introduction of industrial forestry and active fire protection. Since lichens often are totally consumed by fire and lichens mainly disperse by fragmentation, their recolonization after fire is slow. In Canada reindeer lichen was found to establish ca. 10 years after fire (Webb, 1998), and then increase until they peaked 40– 120 years later (Ahti, 1959; Kershaw, 1976; Morneau and Payette,

1017

1988; Thomas et al., 1996). Webb (1998) found that the cover of reindeer lichens was on average 5.3% in burnt sites compared to 25.2% in undisturbed sites 15–16 years after fire. Despite lichen’s slow regeneration, many Pinus-lichen dominated stands developed on sites with moist soils in northern Sweden after severe forest fires in the past that reduced the nitrogen level of the soil. In these types of stands succession will eventually lead to more productive vegetation and thus reductions in lichen abundance (Ebeling, 1978). In our study area such development towards more productive lands following fire suppression is reflected in the increases in site productivity indices that have occurred since the beginning of the 20th Century (Fig. 4). Ho¨rnberg et al. (1999) also found that fire had been an important factor causing the depletion of soil nutrients, thus creating Picea-lichen dominated stands, and that these types of stands were probably more common before the start of commercial timber cutting. The lichen cover has also been found to decrease with stand age in old pine forests on dry sites (Bra˚kenhielm and Persson, 1980; La¨hde and Nieppola, 1987). These observations indicate that lichen abundance was probably positively affected by the active fire suppression policy initially, since it gave lichen more time to regenerate. However, as time passed succession in moist lichen-rich stands led to increased nutrient levels and lichen were out-competed by dwarf-shrubs (Ebeling, 1978). This process appears to have been further promoted by the introduction of silvicultural measures (Fig. 6). The main effects of forestry on forest fires were definitely mediated by the support of the fire suppression movement, and compared to that the effects of fire during the short period of prescribed burning appear to have been minor in this area. In contrast to fire, soil scarification always leaves some proportion of the ground undisturbed (Eriksson and Raunistola, 1990; Roturier and Bergsten, 2006), which promotes recolonization by lichens. Various methods of soil scarification are used at different types of sites (Anon., 1976), with varying effects on the vegetation cover (Eriksson, 1976a). Disc trenching, which is applied in poor to medium fertility sites, is the most common method used in Sweden. The forest owner Doma¨nverket recommended this method on dry to moist grounds (Anon., 1976), which are the dominating ground moisture classes in the study area and which are grounds where lichens commonly grow in the ground vegetation (Ebeling, 1978). Disc trenching has been found to affect 45–55% of the plant cover (Eriksson and Raunistola, 1990), and full recovery of lichens in a lichen-rich site is predicted to require about 50 years (Sunde´n, 2003). Open forests with abundant old trees generally have greater lichen cover than denser forests (Pharo and Vitt, 2000), and logging affects abiotic variables, notably light availability and moisture levels, that strongly influence the growth of lichens (Arnstro¨m, 1975). During the period of selective logging, cutting of the largest trees made the forests in our study area more open, which should have favoured lichen growth. However, the effects are likely to have been more severe following clear-cutting, which results in greater changes in the ground flora than selective cuttings since no tree layer is left (Nieppola, 1992; Hannerz and Ha˚nell, 1997; Bergstedt and Milberg, 2001), although the effects of clear-cutting depend on the site type. At dry sites lichens expand after clearcutting since light conditions improve (Bra˚kenhielm and Persson, 1980; Kardell and Eriksson, 1992; Nieppola, 1992; Olsson and Staaf, 1995; Bra˚kenhielm and Liu, 1998), but at moister sites pioneer species like Deschampsia flexuosa increase substantially because nutrients are released (Hannerz and Ha˚nell, 1997; Palvainen et al., 2005; Uotila and Kouki, 2005). Furthermore, when logging residues are left on the ground at dry sites D. flexuosa is favoured and reindeer lichens are disfavoured (Bra˚kenhielm and Persson, 1980; Olsson and Staaf, 1995; Bra˚kenhielm and Liu, 1998).

1018

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

Logging residues left on the ground after logging also constitutes a physical obstruction for reindeer foraging (Eriksson, 1976a), and for this reason about a third of the logged area will be inaccessible for reindeer forage for at least 5 years after logging (Arnstro¨m, 1975; Eriksson, 1976a). Logging residues were left on the ground during the period of selective logging, as well as after clear-cutting. However, the type of residues at that time were somewhat different, since whole tree crowns were left that were noticeable above the snow surface, making it easier for reindeer to avoid them. Residues are also left on the ground when pre-commercial thinning is applied to young stands (Eriksson et al., 1987). Precommercial thinning has been applied to ca. 5% per year of the forest land in our study area since the 1950s (Fig. 5). Another type of physical obstruction to reindeer digging for lichen is the increased snow density on clear-cuts than in uncut forests (Eriksson, 1976a,b; Pruitt, 1979), and the adverse effects of the denser snow exceed the positive effects of increased lichen growth after clear-cutting at some sites. It has also been shown that reindeer avoid grazing on clear-cuts and in young forests, preferring to graze in forests >100 years old (Helle et al., 1990; Thomas et al., 1996), and that reindeer grazing is negatively correlated with stand density (Helle et al., 1990). Hence, clear-cuts and young stands are not good for reindeer winter pasture for several reasons. The proportion of potentially suitable areas for reindeer grazing (i.e. Middle-Aged and Old Pine stands and stands that becomes Mixed forests after 1895) decreased slightly between 1895 and

1936 as shown in Fig. 9A. However, the picture is more complicated than an analysis of only the stand age shows. At the beginning of the 20th Century logging was selective and generally opened up already sparse forests and therefore improved the conditions for lichen growth. Fire was also prevented allowing lichens to regenerate (Fig. 9B). After 1960 the proportion of potentially suitable grazing grounds has increased again (Fig. 9A). However, the Middle-Aged and Old Pine stands of today are not comparable to such stands of the past since the standing volume has almost doubled since the beginning of the 20th Century (Fig. 3). Further, they have been subjected to various silvicultural measures (Figs. 5 and 6), their site productivity index has increased (Fig. 4), they are single-storied rather than multi-storied, and they are no longer repeatedly burnt (Fig. 9B). The Old and Middle-Aged Pine stands of today are thus denser and darker, compared to such stands 100 years ago, favouring mosses over lichens in the ground layer (Pharo and Vitt, 2000; Sulyma and Coxson, 2001; Sedia and Ehrenfeld, 2003) and the absence of fire allows succession towards more nutrient-rich conditions at moist sites which further reduce the lichens. We estimate that the extent of suitable grazing lands in the study area has been reduced by ca. 30–50% during the last century, and the quality of the remaining grazing lands has been severely impaired (Fig. 9). This supports data from the National Swedish Forest Survey which shows that the amounts of lichenrich grounds have declined by ca. 50% during the last 50 years (Sandstro¨m et al., 2006). In the two separately studied areas with good lichen pasture in the late 19th Century, the history of logging impact differ from to the general trends in the area. In Abmokheden the logging has been very intensive especially in the 1970s, and today only fragments of Old and Middle-Aged Pine Forest remains (Fig. 7), while in Paulavuopme less intense logging has resulted in more Old and Middle-Aged Pine forest than the average for the whole area (Fig. 8). The loss of grazing grounds in sites like Abmokheden has accordingly led to changes in the migratory routes for reindeer in the area. Traditionally it was common to move along the rivers because it was easy and lichen pastures along rivers were generally good. The general decrease in overall winter grazing grounds for reindeer in the studied landscape is likely to lead to increased grazing pressure in remaining lichen-rich sites, such as Paulavuopme. A very high grazing pressure may lead to a long-term degradation of the lichen pasture if other plants become dominant as has been suggested for Arctic tundra vegetation (van der Wal, 2006). 4.3. Implications for forest management

Fig. 9. Principal figure showing impact of forestry on reindeer winter pasture in the study area during the 20th Century, as discussed in the article. The first graph (A) shows the proportion of forest types potentially good for reindeer winter pasture at the different inventory occasions, i.e. areas classified as Old or Middle-aged Pine forest and areas that becomes Mixed forests after 1895 (mostly because of selective logging of large pine trees in the first part of the century). The second graph (B) shows the trend in how forestry has affected the quality of the grazing grounds.

Forestry has been the major driving force in changing the forests in this region since the late 19th Century, and it will continue to do so. Climate change will most likely also have a substantial impact on the forest ecosystem, partly by changing the abiotic conditions and partly by changing the direction of forestry. Under these circumstances information on baseline conditions and trajectories of forest development is needed. The spatially precise historical information about the effects of forestry on lichen pasture provided in this study can be used to direct forest management to facilitate and promote reindeer herding in the future. Forestry improved reindeer winter pastures in the study area during the first part of the 20th Century, while the pastures were impaired during the second part of the century. To reverse this trend the forestry methods used today would have to be changed. A problem is that there is an inherent conflict between successful forestry and good lichen pasture since forestry aims at increasing forest productivity which highly disfavour lichens. A way of

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

improving growth conditions for lichens would be to adapt forestry on areas with poor coarse soils that are naturally good habitats for lichens and of relatively low economical importance to forestry companies. Selective logging and whole tree harvesting on these poor soils would improve growth conditions for lichens. These can be further improved if fertilization and soil scarification is avoided. On sites with moist soils re-introduction of fire would be beneficial for lichens, since it would stop the succession towards more nutrient-rich conditions and reduce competition from dwarfshrubs and herbs. The areas Abmokheden and Paulavuopme that had good reindeer winter pasture at the end of the 19th Century (i.e. sparse old pine forests on poor soils) could fairly easily be returned to that state if adapted forestry was applied. A positive trend in forestry for reindeer herders is the renewed interest in continuous cover forestry (CCF) to meet requirements for certifications for sustainable forestry. CCF includes silvicultural systems which involve continuous maintenance of forest cover and avoids clear-cutting (Pommerening and Murphy, 2004). Such methods, especially if standing volume and tree density is kept fairly low, resembles the silvicultural techniques used at the beginning of the 20th Century when pine forests were selectively logged and would hence improve growth conditions for ground lichens. Many of the measures to improve winter pasture for reindeer, e.g. re-introduction of fire, avoidance of fertilization and the creation of old open stands, are also consistent with general nature conservation goals in boreal forests and these two interests could therefore be functionally combined. 5. Conclusions The studied forest ecosystem, which is used jointly for forestry and grazing reindeer, has changed dramatically during the last ca. 100 years. Before industrial forestry began to affect the study area (-1895) grazing of ground-growing lichen by reindeer in the winter was largely controlled by recurrent forest fires that not only regularly consumed the lichen mat, but also created good growing conditions for lichens by reducing the amounts of available nutrients at some sites. The effect of forestry on lichen availability and abundance has not always been adverse. During the period of selective logging (ca. 1895–1950) cutting of the larger trees created more open forest conditions, which improved conditions for lichens. During this time forest fires were also prevented, allowing lichens to regenerate for longer times. Logging residues presented minor problems and silvicultural measures were only applied at a smallscale. Thus, forestry created favourable conditions for reindeer winter grazing at this time. During the ongoing period of intensive forestry (from ca. 1950 to date) clear-cutting and intensified forest management has radically changed the forest landscape and ecosystem. Clear-cutting has led to increases in lichen biomass at some sites, but overall it has also had much greater adverse effects, mediated by associated changes in snow quality and logging residues, increases in soil scarification and dense regeneration of forests. Natural succession in the absence of fire, together with efficient silvicultural measures, has resulted in increases in productivity and standing volume, and thus further exacerbated growing conditions for lichen. We argue that ca. 30–50% of the potentially good winter grazing grounds for reindeer has been lost because of intensive forest management during the 20th Century and furthermore the quality of the grazing grounds are considerably impaired. The study also shows that historical information can provide baseline data that could facilitate forest management regimes that promote the interests of reindeer herding in northern Scandinavian forests.

1019

Acknowledgements We wish to thank Per Linder at Fastighetsverket and Ronny Lo¨fstrand and Lars-Erik Bergstro¨m at Sveaskog for generously providing us with recent forest inventory data, Kristoffer Sivertsson for his sterling work collecting and compiling the historical documents used in this study, John-Erik Hansson and Niklas Wiberg at the Provincial Archive in Ha¨rno¨sand and the staff at Forskningsarkivet at Umea˚ University for professional help and guidance in the archive, John Blackwell at Sees-editing, UK, for improving the language, Ingela Bergman at Silvermuseet, Arjeplog for valuable information about the history of the studied area and one anonymous reviewer for constructive comments on the manuscript. CMF (Umea˚, Sweden) and FORMAS (Sweden) financed this study. References Ahti, T., 1959. Studies on the caribou lichen stand of Newfoundland. Ann. Bot. Soc. Vanamo 30, 1–44. Anon., 1976. Doma¨nverket skogsva˚rdshandbok. Doma¨nverket. Anon., 2001. En ny renna¨ringspolitik. SOU 2001:101, Stockholm, p. 658. Anon., 2006. Samernas sedvanemarker. SOU 2006:14, Stockholm, p. 550. Arnstro¨m, 1975. Effekter av kalavverkning pa˚ renbetesmark - na˚gra fa¨ltfo¨rso¨k. Va¨xtbiologiska Inst, Uppsala, p. 40. Arpi, G., 1959. Sa˚gverkens virkesfo¨rbrukning. In: Arpi, G. (Ed.), Sveriges skogar under 100 a˚r. Ivar Haeggstro¨ms Boktryckeri, Stockholm, pp. 129–159. Axelsson, A.-L., 2001. Forest Landscape Change in Boreal Sweden 1850–2000—a multiscale approach. Department of Forest Vegetation Ecology, Swedish University of Agricultural Sciences, Umea˚, p. 39. ¨ stlund, L., 2001. Retrospective gap analysis in a Swedish boreal Axelsson, A.-L., O forest landscape using historical data. For. Ecol. Manage. 147, 109–122. Baskin, L.M., Danell, K., 2003. Ecology of ungulates. In: A Handbook of Species in Eastern Europe and Northern and Central Asia, Springer-Verlag, Berlin, p. 434. ¨ stlund, L., Zackrisson, O., 2004. The use of plants as regular food in Bergman, I., O ancient Subarctic economies: a case study based on Sami use of Scots pine innerbark. Arctic Anthropol. 41, 1–13. Bergstedt, J., Milberg, P., 2001. The impact of logging intensity on field-layer vegetation in Swedish boreal forests. For. Ecol. Manage. 154, 105–115. Bostro¨m, M., 2004. Renbetestillga˚ng pa˚ ba¨rris-, ljung/kra˚kba¨r- och lavmarker ca 50 a˚r efter brand/bra¨nning: en studie utfo¨rd ovanfo¨r odlingsgra¨nsen i Norrbotten pa˚ uppdrag av Statens Fastighetsverk och Tuorpons sameby. In, Examensarbeten - SLU, Sveriges Lantbruksuniversitet. Institutionen fo¨r skogssko¨tsel, p. 23. Bra˚kenhielm, S., Liu, Q., 1998. Long-term effects of clear-felling on vegetation dynamics and species diversity in a boreal pine forest. Biodiver. Conserv. 7, 207–220. Bra˚kenhielm, S., Persson, H., 1980. Vegetation dynamics in developing Scots pine stands in central Sweden. Ecol. Bull. 32, 139–152. Burgi, M., 1999. A case study of forest change in the Swiss lowlands. Landscape Ecol. 14, 567–575. Cipolla, C.M., 1991. Between Two Cultures—An Introduction to Economic History. w. w. Norton & Company, New York, p. 198. Dettki, H., Esseen, P.A., 1998. Epiphytic macrolichens in managed and natural forest landscapes: a comparison at two spatial scales. Ecography 21, 613–624. Ebeling, F., 1955. Den moderna skogssko¨tseln i Norrbotten. In Rensko¨tseln och den nutida norrla¨ndska skogshanteringen. Lappva¨sendet, renforskningen, pp. 44– 48. Ebeling, F., 1959. Skogarna och deras va˚rd i o¨vre Norrland fra˚n och med 1930-talet. In: Arpi, G. (Ed.), Sveriges skogar under 100 a˚r. Ivar Haeggstro¨ms Tryckeri, Stockholm, pp. 413–443. Ebeling, F., 1978. Nordsvenska skogstyper. Sveriges Skogsva˚rdsfo¨rbunds Tidskrift 76, 340–381. ¨ stlund, L., Axelsson, A.L., 2000. A forest of grazing and logging: Ericsson, S., O deforestation and reforestation history of a boreal landscape in central Sweden. New For. 19, 227–240. Eriksson, O., 1976a. Silvicultural practices and reindeer grazing in northern Sweden. Ecol. Bull. 21, 107–120. Eriksson, O., 1976b. Sno¨fo¨rha˚llandenas inverkan pa˚ renbetning. Va¨xtbiologiska Inst, p. 22. Eriksson, O., Raunistola, T., 1990. Impact of soil scarification on reindeer pastures. Rangifer 3 (Special Issue), 99–106. Eriksson, O., Sandewall, M., Wilhelmsson, E., 1987. Virkesproduktionens inverkan pa˚ rensko¨tselns lavbete—en metodstudie. Rangifer 7, 15–32. Gimmi, U., Burgi, M., 2007. Using oral history and forest management plans to reconstruct traditional non-timber forest uses in the Swiss Rhone valley (Valais) since the late nineteenth century. Environ. History 13, 211–246. Gimmi, U., Burgi, M., Stuber, M., 2008. Reconstructing anthropogenic disturbance regimes in forest ecosystems: a case study from the Swiss Rhone Valley. Ecosystems 11, 113–124.

1020

A. Berg et al. / Forest Ecology and Management 256 (2008) 1009–1020

Ha¨gglund, B., Lundmark, J.-E., 2005. Bonitering, Definitioner och Anvisningar. Skogsstyrelsen, Jo¨nko¨ping, p. 53. Hannerz, M., Ha˚nell, B., 1997. Effects on the flora in Norway spruce forests following clearcutting and shelterwood cutting. For. Ecol. Manage. 90, 29–49. Hedman, S.-D., 2004. Skog och Historia i Norrbottens la¨n 2004. Norrbottens museum, p. 39. Heggeberget, T., Gaare, E.M., Ball, J.P., 2002. Reindeer (Rangifer tarandus) and climate change: importance of winter forage. Rangifer 22, 13–32. Hellberg, E., Niklasson, M., Granstro¨m, A., 2004. Influence of landscape structure on patterns of forest fires in boreal forest landscapes in Sweden. Can. J. For. Res.— Revue Canadienne De Recherche Forestiere 34, 332–338. Helle, T., Aspi, J., Kilpela, S.S., 1990. The effects of stand characteristics on reindeer lichens and range use by semi-domesticated reindeer. Rangifer 107–114. Holmgren, A., 1959. Skogarna och deras va˚rd i o¨vre Norrland intill a˚r 1930. In: Arpi, G. (Ed.), Sveriges skogar under 100 a˚r. Ivar Haeggstro¨ms Tryckeri, Stockholm, pp. 375–412. ¨ stlund, L., Zackrisson, O., Bergman, I., 1999. The genesis of two PiceaHo¨rnberg, G., O Cladina forests in northern Sweden. J. Ecol. (Oxford) 87, 800–814. Jonsson, T., 1914. Om bonitering av skogsmark. Sveriges skogsva˚rdsfo¨rbunds tidstift 12, 369–392. Kardell, L., Eriksson, L., 1992. Contortatall och renbete - studeir inom Mala˚ skogssamebys marker. Dep. of Environmental Forestry, Swedish University of Agricultural Sciences, p. 63. Karlsson, N., 2007. Bosa¨ttning och resursutnyttjande. Umea˚ University, Umea˚, p. 172. Kellner, O., Ma˚rshagen, M., 1991. Effects of irregation and fertilization on the ground vegetation in a 130-year-old stand of Scots pine. Can. J. For. Res. 21, 733–738. Kershaw, K.A., 1976. Studies on lichen dominated systems. XX. An examination of some aspects of the northern boreal lichen woodlands in Canada. Can. J. Bot. 55, 393–410. Kohh, E., 1975. A study of fires and hard pan in the forests of A¨lvdalen. Sveriges Skogsfo¨rbunds Tidskrift 73, 300–336. Kouki, J., Lofman, S., Martikainen, P., Rouvinen, S., Uotila, A., 2001. Forest fragmentation in Fennoscandia: linking habitat requirements of wood-associated threatened species to landscape and habitat changes. Scand. J. For. Res. 27–37. Kuuluvainen, T., Maki, J., Karjalainen, L., Lehtonen, H., 2002. Tree age distributions in old-growth forest sites in Vienansalo wilderness, eastern Fennoscandia. Silva Fenn. 36, 169–184. La¨hde, E., Nieppola, J., 1987. Vegetation changes in old stands of Pinus sylvestris L. In southern Finland. Scand. J. For. Res. 2, 369–377. ¨ stlund, L., 1998. Structural changes in three mid-boreal Swedish forest Linder, P., O landscapes, 1885–1996. Biol. Conserv. 85, 9–19. Lofman, S., Kouki, J., 2003. Scale and dynamics of a transforming forest landscape. For. Ecol. Manage. 175, 247–252. Lundmark, L., 1982. Uppbo¨rd, utarmning, utveckling. Det samiska fa˚ngstsamha¨llets o¨verga˚ng till rennomadism i Lule lappmark. Arkiv fo¨r studier i arbetarro¨relsens historia, Lund, p. 230. Ma˚rell, A., 2006. Summer Feeding Behaviour of Reindeer: A Hierarchical Approach. Department of Animal Ecology, Swedish University of Agricultural Sciences, Umea˚, p. 31. Morneau, C., Payette, S., 1988. Postfire lichen-spruce woodland recovery at the limit of the boreal forest in northern Qubec. Can. J. Bot. 67, 2770–2782. Myllyntaus, T., Mattila, T., 2002. Decline or increase? The standing timber stock in Finland, 1800–1997. Ecol. Econ. 41, 271–288. Nieppola, J., 1992. Long-term vegetation changes in stands of Pinus sylvestris in southern Finland. J. Veg. Sci. 3, 475–484. Niklasson, M., Granstro¨m, A., 2000. Numbers and sizes of fires: long-term spatially explicit fire history in a Swedish boreal landscape. Ecology 81, 1484–1499. Olsson, B., Staaf, H., 1995. Influence of harvesting intensity of logging residues on ground vegetation in coniferous forests. J. Appl. Ecol. 32, 640–654. ¨ stlund, L., 1993. Exploitation and structural changes in the north Swedish boreal O forest 1800–1992. In Dissertations in Forest Vegetation Ecology, p. 30 + 102 pp. ¨ stlund, L., 1995. Logging the virgin forest: northern Sweden in the early-nineO teenth century. For. Conserv. History 39, 160–171.

¨ stlund, L., Ericsson, T.S., Zackrisson, O., Andersson, R., 2003. Traces of past Sami O forest use: an ecological study of culturally modified trees and earlier land use within a boreal forest reserve. Scand. J. For. Res. 18, 78–89. ¨ stlund, L., Linderson, H., 1995. A dendrochronological study of the exploitation and O transformation of a boreal forest stand. Scand. J. For. Res. 10, 56–64. ¨ stlund, L., Zackrisson, O., Axelsson, A.L., 1997. The history and transformation of a O Scandinavian boreal forest landscape since the 19th century. Can. J. For. Res. 27, 1198–1206. Palvainen, M., Fine´r, L., Mannerkoski, H., Piirainen, S., Starr, M., 2005. Responses of ground vegetation species to clear-cutting in boreal forest: aboveground biomass and nutrient contents during the first 7 years. Ecol. Res. 652–660. Pharo, E., Vitt, D., 2000. Local variation in bryophyte and macro-lichen cover and diversity in montanae forests of western Canada. The Bryologist 103, 455–466. Pommerening, A., Murphy, S.T., 2004. A review of the history, definitions and methods of continuous cover forestry with special attention to afforestation and restocking. Forestry 77, 27–44. Pruitt, W.O., 1979. A numerical ‘‘snow index’’ for reindeer (Rangifer tarandus) winter ecology (Mamalia, Cervidae). Annales Zoologici Fennici 16, 271–280. Quarnstro¨m, E., 2006. De tycka emellertid av gammal vana att det smakar gott, och tro dessutom att det a¨r bra fo¨r ha¨lsan’’ - Samiskt va¨xtutnyttjande fra˚n 1600talet fram till ca 1950. In: Examensarbeten, Department of Forest Vegetation Ecology, Umea˚, p. 77. Roturier, S., Bergsten, U., 2006. Influence of soil scarification on reindeer foraging and damage to planted Pinus sylvestris seedlings. Scand. J. For. Res. 21, 209–220. ¨ ., 2006. Progressing toward coSandstro¨m, C., Moen, J., Widmark, C., Danell, O management through collaborative learning: forestry and reindeer husbandry in dialogue. Int. J. Biodiver. Sci. Manage. 2, 326–333. Sedia, E.G., Ehrenfeld, J.G., 2003. Lichens and mosses promote alternate stable plant communities in the New Jersey Pinelands. Oikos 100, 447–458. Sjo¨rs, H., 1965. Forest regions. Acta Phytogeografica Suecica 50, 48–63. Ska˚nes, H.M.H., Bunce, R.G.H., 1997. Directions of landscape change (1741–1993) in Virestad, Sweden—characterised by multivariate analysis. Landscape Urban Plann. 38, 61–75. Storeheier, P.V., Mathiesen, S.D., Tyler, N.J.C., Olsen, M.A., 2002. Nutritive value of terricolous lichens for reindeer in winter. Lichenologist (London) 34, 247–257. Sulyma, R., Coxson, D.S., 2001. Microsite displacement of terrestrial lichens by feather moss mats in late seral pine-lichen woodlands of north-central British Columbia. Bryologist 104, 505–516. Sunde´n, M., 2003. Re-establishment Rate of Reindeer Lichen (Cladina spp.) After Soil Scarification in Scots Pine-Lichen Forest Types in Boreal Sweden. Dept. of Forest Management, SLU, Swedish University of Agricultural Sciences, Umea˚, p. 16. Sundset, M.A., Cann, I.K.O., Mathiesen, S.D., Mackie, R.I., 2004. Rumen microbial ecology in reindeer—adaptations to a unique diet. J. Anim. Feed Sci. 13, 717–720. Thomas, D.C., Barry, S.J., Alaie, G., 1996. Fire–caribou–winter range relationships in northern Canada. Rangifer 16, 57–67. ¨ stlund, L., 2002. Floating timber in Northern Sweden: the construcTo¨rnlund, E., O tion of floatways and transformation of rivers. Environ. History 8, 85–106. Uotila, A., Kouki, J., 2005. Understory vegetation in spruce-dominated forests in eastern Finland and Russian Karelis: Successional patterns after antropogenic and natural disturbances. For. Ecol. Manage. 215, 113–137. van der Wal, R., 2006. Do herbivores cause habitat degradation or vegetation state transition? Evidence from the tundra. Oikos 114, 177–186. Webb, E.T., 1998. Survival, persistence, and regeneration of the reindeer lichens, Cladina stellaris, C. rangiferina, and C. mitis following clearcut logging and forest fire in northwestern Ontario. Rangifer 41–47. Whitney, G.G., 1994. From Coastal Wilderness to Fruited Plain—A History of Environmental Change in Temperate North America From 1500 to the Present. Cambridge University Press, Cambridge, p. 451. Wirth, C., Schulze, E.D., Schulze, W., von Stunzner-Karbe, D., Ziegler, W., Miljukova, I.M., Sogatchev, A., Varlagin, A.B., Panvyorov, M., Grigoriev, S., Kusnetzova, W., Siry, M., Hardes, G., Zimmermann, R., Vygodskaya, N.N., 1999. Above-ground biomass and structure of pristine Siberian Scots pine forests as controlled by competition and fire. Oecologia 121, 66–80. Zakrisson, O., 1977. Influence of forest fires on the North Swedish boreal forest. Oikos 29, 22–32.