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16 | A c a d e m i c  J o u r n a l  o f  S c i e n c e ,  E n g i n e e r i n g  a n d  T e c h n o l o g y  

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MEGAFOSSIL TREES REVEAL PAST LANDSCAPES OF THE SWEDISH SCANDES 

 

 

Maria Sofia Andersson and Olof Erik Johansson  
Department of Ecology and Environmental Science, Umeå University, SE 901 87 Umeå, Sweden 

 

Abstract: The structure and plant species composition of Late-glacial and Early-Holocene landscapes in the 

Swedish Scandes are poorly understood. Traditional pollen analytical inferences, glacier histories, and textbook 

narratives are beset with inaccuracies and uncertainties, particularly in high-mountain regions. More direct, 

robust, and reliable mega- and macrofossil records provide unambiguous evidence of the former early local 

presence of tree species at specific sites and elevations, far beyond modern treelines. 

Keywords: Megafossils, Treelines, Holocene, Scandes, Climate, Plant species 

  

Introduction   

The structure and plant species composition of Late-glacial andEarly-Holocenelandscapes in the Swedish Scandes 

are poorly comprehended. In this respect, traditional pollenanalytical inferences, glacier histories and textbook 

narratives are beset with inaccuracies and uncertainties, particularly in high-mountain regions (e.g.Lundqvist 

1969; Huntley & Birks 1983; Berglund et al. 1996; Karlén&Kuylenstierna 1996;Barnekow 1999; Johnsen 

2010).These methodological shortcomingsareevidenced by analyses of more direct, robust and reliable mega- and 

macrofossil records (cf. Helama et al. 2004; Paus 2013; Paus &Haugland 2017;Kullman 2017a). Unambiguously, 

these approaches are stating formerearly local presence of tree species at specific sites and elevations, far beyond 

modern treelines.  

Megafossil1 tree remains, representing former higher-than-present alpine treelines, mainly preserved in peat and 

lake mud, have for long been known and discussed in the Scandes. These records have contributed broad outlines 

of the Holocene history of high mountain landscape and climate evolution (Smith 1920; Lundqvist 1969;Karlén 

1976; Kullman 1995; Aas&Faarlund 2000; Kullman&Kjällgren 2000, 2006). However, studies of this kind are 

constrained by sparsity of peat as an efficient preservation medium at high elevations, which has urged for 

alternative megafossil archives when searching for the highest positions of tree growth during earlier epochs.  
1Megafossils are large pieces of wood, which are preserved near their growth places and which can be accurately 

determined to species and dated by the 14C-method. Ages are reported as calendar years BP (AD 1950), by 

intercept-values, and derive from sources, cited above.  

Megafossilrecords, originating from different elevations above the modern treeline, havedisplayed a discernible 

trend of treeline lowering throughout the Holocene, about 50 m per millennium(Kullman 1995; 

Kullman&Kjällgren 2000), in broadagreement with orbital forcing of insolation at the top of the atmosphere 

(Berger &Loutre 1991). Since thismechanism suggested a thermal maximum somewhat prior to the earliest and 

highest existingrecords, further search for megafossil wood remnants was extended to even higherelevations.  

Particular focus was on the fringes of currently melting glaciers and snow/ice patches alongthe entire Swedish 

Scandes (Fig.1). In addition, these efforts were inspired by positive results, basedon megafossils, from emerging 

proglacial sites worldwide, showing that forest trees hadprevailedat sites until recently covered by ice, during 

earlier parts of the Holocene,(Nicolussi&Patzelt 2000;Schlüchter&Jörin 2004; Benedict et al. 2008; IvyOchs et 

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al. 2009; Koch et al.2014). Prior to the studies reviewed in this study, no such discoveries had been reported from 

the Scandes.  

Results of these recent investigations, carried out in different parts of the Swedish Scandes, constitute the main 

core of this paper.The present review upsets the traditional comprehension of the late-glacial andearly postglacial 

climate as well asthe structure and composition of the high-elevation landscape (cf. Paus 2013; Kullman 2013; 

Luoto et al. 2014;Väliranta et al. 2015; Schenk et al. 2018). Possibly, the emerging views mayserve as proxy 

analogues of future subalpine landscapes in a potentially warmer world.In fact, the initial phase of such a course 

of change may be already underway in the Swedish Scandes (Kullman 2010, 2019). However, such projections 

should be treated cautiously, since future directions of the climate evolution remain uncertain.  

  

 
Figure 1. Subfossil pine, which inspired search for megafossils at exceptionally high elevations,particularly on 

the forefields of receding glaciers. The lower fringe of the glacier “Sylglaciären”, 1195 m a.s.l., about 400 m 

higher than the present treeline. Radiocarbon yielded 10 425 cal. yr BP. Photo: 1997-07-16. Source: 

Kullman&Kjällgren 2000.   

Results Late-Glacial and early-Holocenetrees as evidenced by megafossils-the new landscape perspective   

All main tree species of the current treeline ecotone were present on what has to be interpreted as ice-free 

nunataks(Fig. 2-5) already at the Late-Glacial/Early-Holocene transition, 17 000- 13 000years before the present 

day (BP) at unprecedented high elevationsalong the entire Swedish Scandes(Kullman&Kjällgren 2000; Kullman 

2002, 2004). These species are mountain birch (Betulapubescens ssp. czerepanovii), Norway spruce (Piceaabies) 

and Scots pine (Pinussylvestris). Henceforth,these will be cited as Betula, Picea and Pinus, respectively.   

The core site for discovery of the earliest tree megafossils is Mt. Åreskutan (1420 m a.s.l.) in the southern 

SwedishScandes (province of Jämtland), 1360 m a.s.l. and 510 m above the treeline of pine. This mountain has, 

close to its summit, until quite recently,harboured an ice patch or asmall glacier, which has gradually vanished in 

response to 20th century climate warming andconsequently megafossil trees have been exposed (Figs. 2 & 3).  

  

  

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Figure 2. Locus classicus, 1360 m a.s.l.,the site for the discovery of Late-glacial and Early-Holocene tree presence 

on early emergingnunataks in the Scandes (Kullman 2002). These findings have changed the conception of the 

earliestsubalpine/alpine landscape-history. a. A small glacier/ice patch prevailed here until quite recently. Its 

former position is indicated by the snow patch and the row of blocks in the foreground. The snow has completely 

melted by the end of late summer during most years of the past few decades. Photo: 2018-07-18.b. The former 

ice-distribution embraced the green moss-covered area in the center. Here, most of the megafossils have been 

recovered, reasonably dislocated downslope by snow avalanches from higher upslope positions. Photo: 2018-

0901. 

  
Figure 3. Overview of the earliest megafossils recovered at theÅreskutan-site, 1360 m a.s.l. a. Pinussylvestris, 13 

810 cal. yr BP. b. Piceaabies, 13 010 cal. yr BP. c. Betulapubescenss.lat., 16 810 cal. yr BP. These samples 

highlight earlier local deglaciation and tree instatement than previously inferred by more traditional approaches. 

Source: Kullman 2002.  

  

  

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Figure 4. Examples of Late-Glacial megafossil tree recoveries at sites along the entire Swedish Scandes. a. The 

glacier Helagsglaciären (province of Härjedalen), has receded areally by c. 40% during the past 100 years.  

Photo: 2015-08-13. b. Close to its former lower maximum range, 1150 m a.s.l., megafossils of Pinus were 

recovered and dated 13 145 cal. yr BP. Photo: 2008-07-04. Source: Kullman&Kjällgren 2000.c. The glacier 

Tärnaglaciären (province of Lapland). Currently, a large snow-field prevails in the mid-slope below the glacier, 

which extended down to the mid of the lake (1070 m a.s.l.) about 100 years ago(Lindgren &Strömgren 2001). 

Photo:2010-08-20. d. Preserved in a downwashed peat-cake 630 m a.s.l., a cone of Piceawas dated 11 200 cal. yr 

BP. Photo: 2017-09-01. e. The glacier Kårsaglaciären, 965 m a.s.l. (province of Lapland). Photo: 2009-08-21. f. 

A megafossil remnant of Pinus appeared in the outwash stream from beneath the glacierKårsaglaciären, 955 m 

a.s.l. It dated 11 760 cal. yr BP. Photo: 2008-09-17. Source: Öberg &Kullman 2011.g. Mt. Städjan (province of 

Dalarna), 1100 m a.s.l. h. MegafossilPinus protruding from a thin soil layer between boulders in the south-facing 

slope of Mt. Städjan. Dating yielded 12 425 cal. yr BP. Photo: 2007-07-14.  

The firm evidence, presented above, conflicts with traditional glacial geologic and paleobotanical opinions and 

have been questioned and opposed by proponents and defenders of these approaches, drawing on negative 

evidence (Birks et al. 2005) and refuted by Kullman (2006). 

Holocene megafossils in their settings  

Below, a representative sample of megafossils recovered in geomorphic glacial cirques, with currently receding 

ice cover,ispresented. This is a comprehensive and richly illustrated review of previously published data and 

updates, representing the Swedish Scandes, from south to north (Öberg & Kullman 2011, Kullman & Öberg 2013, 

2015; Kullman 2017a,b) (Figs. 5-20). A popular overview is given by Kullman & Öberg (2019). Surprisingly, 

little research on these issues has been carried out in Scandinavia by palaeoecologists, although archaeologists, 

particularly in Norway, are making rich findings of human artifacts on the forefields of melting glaciers and ice 

patches (e.g. Nesje et al. 2011).  

  

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Figure 5.The position of the glacier sites, particularly focused in this review. 1. Helags-Sylarna glaciers. 2.The 

glacier Tärnaglaciären with adjacent ice patches. 3. Glaciers and snow/ice patches in northern Lapland.  

1.Helags-Sylarna glaciers  

 
Figure 6. Downwashed stem of Betula, 1345 m a.s.l., which dated 8620 cal. yr BP. Mt. Helagsfjället. Photo: 

201008-11.  

  

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Figure. 7. Subfossil Betula, extracted from eroding moss-cover in a downstream proglacial delta below the glacier, 

1350 m a.s.l. Dating yielded 9520 cal. yr BP. Mt. Helagsfjället. Photo: 2006-10-15.  

  

  
  

Figure 8.a. Remnant of a fairly stout Betula-tree, uplifted from beneath the moss-cover in the delta below 

Storsylglaciären, 1275 m a.s.l. Presumably, the original growth position was higher upslope. Dating yielded 7170 

cal. yr BP. Photo: 2001-08-22.b. A downwashedPinus-remnant, 1210 m a.s.l., recovered well below its assumed 

original growth position, underneath the background Ekorrglaciären. Radiocarbon-dating gave9530 cal.yr BP. 

Photo: 2008-08-24. 

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2. The glacier Tärnaglaciären with adjacent ice patches  

  
Figure 9. “High-flying” Betula-megafossil at the margin of an ice-patch adjacent to Tärnaglaciären, 1425 m a.s.l. 

This is 635 m higher than the current local treeline. Dating yielded 9195 cal. yrBP.Murtsergure ice patch. Photo: 

2012-08-28.   

    
Figure 10. Piece of a Betula-stem, recovered 1410 m a.s.l., 700 m higher than the nearest present-day treeline. It 

is currently being washed downslope from a growth place close to an ice-patch, adjacent to the 

glacierTärnaglaciären. It dated 9365 cal. yr BP.Photo:2017-09-01.  

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Figure 11. Betula-megafossil, protruding from the snow rime at the glacier front, 1395 m a.s.l. This site is 685 m 

higher than the current local treeline. Dating yielded 9450 cal. yr BP. The glacier Tärnaglaciären. Photo: 2017-

0901.  

 
Figure 12. A virtually new source of past high-mountain vegetation composition is provided by outwashed 

”peatballs” of this kind. Here uplifted from behind a stone in the main melt-water stream. Their content of plant 

remains represents some of the former plant cover composition were ice prevailed until quite recently (see Fig.13). 

Ice-patch near the glacierTärna-glaciären, 1115 m a.s.l. Photo: 2012-09-22.  

  

  

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Figure 13. Tree remains of different species contained in “peat balls”, released from beneath glacier ice in the  

Tärna region (source: Kullman& Öberg 2013). Except for common forest bryophytes and dwarf-shrubs, cones of 

Larixsibirica and Piceaabies as well as leaves of deciduous boreal tree species, have been extracted and 

radiocarbondated; a. b. Larixsibirica, 7320 cal. yr BP.c. Piceaabies, 8450 cal. yr BP. d. Pinussylvestris, 7960 cal. 

yr BP. e. Sorbusaucuparia, 8460 cal. yr BP. f. Populustremula 8590 cal. yr BP.   

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3. Glacier and snow/ice patches in northern Lapland   

.  

Figure 14. Trunk of Pinus, dug out from glacier sediment, 940 m a.s.l. Obviously, it is worked by beaver (Castor 

fiber), indicative of a local forest environment at the dated time; 9280 cal. yr BP. The glacierKårsaglaciären. 

Photo: 2008-09-17.  

  
Figure 15. Megafossil remains of Betula, exposed just outside the lower glacer margin and much higher than the 

present local treeline, 990 m ö.h. They date 1950 cal. yr BP and support a general conception of a warmer-

thanpresent time, with a smaller glacier (Kullman 2013). The glacier Kårsaglaciären. Photo: 2013-09-12.  

  

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Figure 16.a. The glacierKåppasglaciären, c. 9 km west of Abisko in Swedish Lapland. Today, it should possibly 

be characterized as an ice-field. It released a megafossilPinus at its lower margin, 1030 m a.s.l.b. Dating yielded 

7860 cal. yr BP. Photo: 2010-08-28  

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Figure 17.a. An elongated snow/ice patch, located c. 15 km northwest of Abisko, extending 975-980 m a.s.l. At 

the lower margin, an extensive stone pavement indicates a prior more extensive size of this object. The front is 

currently disintegrating by “calving”, which exposes new mineral ground with some emerging megafossil tree 

remans. Photo: 2010-08-30.b. MegafossilPinus, dated 8900 cal. yr BP, up-raised from original position. Photo: 

2010-08-30.c. Basal part of a Betula-stem, possibly preserved in situ, 975 m a.s.l. Radiocarbon-dating gave 5800 

cal. yr BP.Låktatjåkka Ice Field. Photo: 2010-08-30.  

  

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Figure 18.a. The glacier Kitteldalsglaciären in the Kebnekaise-massif. The lower front is about 1190 m a.s.l. 

Megafossils of Pinus and Betula are recovered along the right-hand (east-facing) margin of the glacier. Photo: 

201308-11.b. Pinus-log melting out from the glacier, 1240 m a.s.l. It dated 9010 cal. yr BP and is located 690 m 

higher than the local present-day treeline.Photo: 2013-08-11.  

  

  
  

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Figure 19. The glacier Storglaciären in the Kebnekaise-massif is one of the most thoroughly investigated glaciers 

in the Swedish Scandes, although mainly with respect to size and mass balance changes. About 100 years ago the 

lower front was close to the lake, c. 1115 m a.s.l. (cf. Holmlund 2012). Photo: 2013-08-13  

  

  
Figure. 20. a.Megafossillog of Betula, which dated 8490 cal. yr BP, 1100 m a.s.l. Photo: 2013-08-

30.b.Downwashed peat cake, containing a Picea cone shell, 1105 m a.s.l. Photo: 2013-08-13.c. APiceaconeshell 

dated 8380 cal. yr BP. The glacier Storglaciären.  

Synthesis and discussion 

Demonstrably, alpine glaciers along the entire Scandes have been melting over much of the past century 

(Lundqvist 1969; Holmlund et al. 1996, Bakke et al. 2008). At their lower fronts, mega fossil tree remnants of 

different species are currently exposed. Radiocarbon-dated, these samples provide a new view of the LateGlacial 

and Early-Holocene high mountain landscape. It now stands out, that along the entire Swedish Scandes, all of our 

common tree species grew in small isolated populations, much earlier and at higher positions, than ever evidenced 

or contemplated. Main features are quantified and summarized in Table 1. The highest relative treeline positions 

and reasonably, the highest summer temperatures were attained 10 000-9500 cal. yr BP. This inference agrees 

with temperature reconstructions from other northern regions (e.g.Shenk et al. 2020;Mörner et al. 2020)  

 The discussion below draws on an “amalgam” of previously published original studies, based on megafossils 

retrieved from glacier forefields (Kullman 2004; Öberg & Kullman 2011, Kullman & Öberg 2013, 2015; Kullman 

2017a,b). These references provide additional detail and documentation to the images, which make up the core 

of this paper.  

Table 1. For each of the study sites (Fig. 5), age range of all megafossils, given as cal. yr BP, and corresponding 

relative elevation range of sample sites, displayed as altitudinal meters above the current treeline.  

  

Site Age-range Relative elevation 

range 

1 16 810-6100 115-585 

2 9530-4480 225-700 

3 11 760-1950 80-690 

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A particular noteworthy novelty is that boreal trees grew at sites of present-day glaciers already during the Late-

Glacial as early as about 17 000-12 000 years before the present. Analogous inferences are presented from the 

Norwegian Scandes (Paus et al. 2011). Taken together, these discoveries have a bearing both on glacier and 

vegetation history. The common wisdom, until present, has been that the high mountains were completely 

icecovered at this early stage (Lundqvist 1986, 1994), a viewquestioned (e.g. by Dahl et al. 1997; Follestad 2003; 

Hörnberg et al. 2006;Goeringa et al. 2008), and certainly not compatible with the presence of trees, if not assuming 

supraglacial tree growth. In fact, the last-mentioned option is discussed by different authors (Fickert et al. 2007; 

Zahle et al. 2018).  

  

The ice-free glacier cirques displayed the character of outlying forest enclaves, high above the closed and 

continuous forest below. Prior to 10 000 BP, the dated samples are too few to form definite opinions about the 

relative abundance of Betula,Pinus and Piceain the concerned habitats, although all three species were present at 

high elevations during that period, as evidenced by this review. Possibly, the contemporary sparsity of recoveries 

related to still incomplete deglaciation and relatively small areas available for tree growth on the nunataks.   

Since about 10 000 years BP, Betula appears to have formed the upper treeline in these habitats, although Pinus 

is found to have joined Betula at the highest elevations, 600-700 m above current treelines, as these appeared 

during the past 10 years or so (Kullman & Öberg 2009; Kullman 2013). Except for the dominating tree species 

particularly focused and depicted in this study, the early tree vegetation contained an array of sub-ordinate boreal 

tree species, which today prevail sparsely in the mountain forest below. These species are Sorbusaucuparia, 

AlnusincanaandPopulustremula, all documented by megafossils (Kullman& Öberg 2013, 2015).  

Presumably, the Late-glacial and Early-Holocene nunatak tree groves may have served as dispersal nodes for 

trees and other plants, enabling their rapid subsequent downslope spread and establishment over the ice-free 

landscape as it gradually emerged (cf. Väliranta et al. 2011, 2015).  

It is of particular interest to find that,Piceaabies occurred on a regular basis and at unprecedented high elevations 

above its current treeline, and so even during the Late-Glacial. This contrasts with the orthodox view (based on 

pollen analysis) of spruce as a particularly late postglacial immigrant to western and high-elevation Sweden (Moe 

1970; Giesecke& Bennett 2004; Seppä et al. 2009). Encouraged by the megafossil evidence presented above, 

some researchers, drawing on microfossils and DNA-technique, support the option of early Holocene presence of 

Piceain the high-mountains(Segerström& von Stedingk 2003;Hörnberg et al. 2006;Paus 2010; Paus et al. 2011; 

Parducci et al. 2012; Carcaillet et al. 2012). In addition, during the early Holocene, the concerned tree groves 

harboured a tree species not growing spontaneously in Sweden today, namelyLarixsibirica(Kullman 2018), which 

also occurred outside the present kind of habitats along the entire Scandes, both in Sweden and Norway (Kullman 

1998; Bergman et al. 2004; Paus 2010;Carcaillet et al. 2012).Possibly this light-demanding species was 

outcompeted by advancing denser populations ofBetula and Picea by the mid-Holocene, as these species were 

favored by the evolving Neoglacial climate, which then turned to a more oceanic and snow-richcharacter (cf. 

Kullman 2018). By analogy with the rich tree flora, it is reasonable to assume that plant species richness in general 

was high in these, obviously sparse high-elevationtree stands. This gains support from analyses of plant remains 

in peat-cakes released from beneath the glacier ice (cf. Kullman & Öberg 2013, 2015). The presence of tree 

assemblages is suggested also from the fact that beaver (Castor fiber), an obligate forest dweller, utilized trees 

growing in these sites (Fig. 14).  

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31 | A c a d e m i c  J o u r n a l  o f  S c i e n c e ,  E n g i n e e r i n g  a n d  T e c h n o l o g y  

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Tentatively, the maximum difference by 700 m between the early-Holocene treeline position and the present 

treeline may be translated into summer temperature change decline over this period of time. Based on a general 

temperature lapse rate of 0.6 °C per 100 m altitude (Laaksonen 1976), it may be inferred that the temperature has 

lowered by 4.2 °C since 9500 cal. yr ago. However, this figure, has to be adjusted due to the effect of subsequent 

glacio-isostatic land uplift, which here may be in the order of 200 m (Påsse&Andersson 2005). This reduces the 

figure on which temperature change may be calculated to 500 m and consequently a temperature 3.0 °C higher 

than at the present day.  

A warmer climate in the future, as commonly alleged, may turn the high mountain landscape into a state 

envisioned by the findings for the early Holocene, as depicted in this study. Tentatively, this implies a 

highmountain landscape, virtually without glaciers and large late-lying snow/ice patches. The former sites of these 

elements are likely to stand out as isolated treed oases high above the continuous forest. The surrounding more 

wind-exposed and snow-poor terrain remains virtually untreed, by analogy with the reluctance of trees and forests 

to colonize this type of habitats in response to the warming of the past 100 years (Kullman & Öberg 2009).   

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