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ORI GI NAL

ARTI CLE
Did tree-Betula, Pinus and Picea survive
the last glaciation along the west coast
of Norway? A review of the evidence,
in light of Kullman (2002)
Hilary H. Birks
1,2,3
*, E. Larsen
4
and H. J. B. Birks
1,2,3
I NTRODUCTI ON
Radiocarbon dates of late-glacial age obtained from megafos-
sil tree remains have recently been reported by Kullman (2000,
2001, 2002) from the central Swedish mountains. From this
evidence Kullman concludes that Betula pubescens Ehrh. ssp.
tortuosa (Ledeb.) Nyman, Picea abies (L.) Karst. and Pinus
sylvestris L. grew at 1360 m a.s.l. on Mt A

reskutan, Jamtland,
1
Department of Biology, University of Bergen,
Bergen, Norway,
2
Bjerknes Centre for Climate
Research, Bergen, Norway,
3
Environmental
Change Research Centre, University College
London, London, UK and
4
Geological Survey of
Norway, Trondheim, Norway
*Correspondence: Hilary H. Birks, Department
of Biology, University of Bergen, Allegaten 41,
N-5007 Bergen, Norway.
E-mail: hilary.birks@bio.uib.no
ABSTRACT
Aim We discuss the hypotheses proposed by Kullman [Geo-O

ko 21 (2000) 141;
Nordic Journal of Botany 21 (2001) 39; Journal of Biogeography 29 (2002) 1117] on
the basis of radiocarbon-dated megafossils of late-glacial age from the central
Swedish mountains that boreal trees survived the glaciation along the south-west
coast of Norway and subsequently migrated eastward early in the late-glacial to
early deglaciated parts of the central Swedish Scandes mountains.
Methods We assess these hypotheses on the basis of glacial geological evidence
and four lines of palaeoecological evidence, namely macrofossil records of the tree
species, vegetation and climate reconstructions from plant evidence, independent
climate reconstructions from other proxies for the late-glacial environment of
south-west Norway, and the patterns of post-glacial spread of the tree species.
Location South and west Norway, central Swedish Scandes mountains
(Jamtland).
Results and conclusions South-west Norway and the adjacent continental
shelf were under ice at the last-glacial maximum (LGM). The late-glacial
vegetation of south-west Norway was treeless and summer temperatures were
below the thermal limits for Betula pubescens Ehrh., Pinus sylvestris L. and Picea
abies (L.) Karst. Instead of spreading immediately after the onset of Holocene
warming, as might have been expected if local populations were surviving,
B. pubescens showed a lag of local arrival of 600 to > 1000 years, Pinus lagged by
1500 to > 2000 years, and Picea only reached southern Norway c. 1500 years ago
and has not colonized most of south-west Norway west of the watershed. Glacial
geological evidence shows the presence of an ice sheet in the Scandes at the LGM
and in the Younger Dryas, which was cold-based near or at the area where the
late-glacial-dated megafossils were recovered by Kullman. We conclude that the
samples dated by Kullman (2002) should be evaluated carefully for possible
sources of contamination. All the available evidence shows that the
biogeographical hypotheses, based on these radiocarbon dates taken at face
value, of late-glacial tree survival at the Norwegian coast and subsequent
eastwards spread to the mountains, are unsupportable.
Keywords
Betula pubescens, glacial survival of trees, late-glacial, macrofossils, megafossils,
Picea abies, Pinus sylvestris, pollen, south-west Norway.
Journal of Biogeography (J. Biogeogr.) (2005) 32, 14611471
2005 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi doi:10.1111/j.1365-2699.2005.01287.x 1461
central Sweden, during the late-glacial interstadial (Blling-
Allerd) and the following cold stadial (Younger Dryas)
between 14,000 and 10,200
14
C yr bp. Kullmans radiocarbon
dates are sensational and his conclusions have major implica-
tions for our current understanding of late-glacial plant
geography and vegetation history and of the deglaciation
history of the Scandinavian ice sheet.
Following from these results, Kullman (2000, 2001, 2002)
proposes that (1) the trees survived the glacial period on
exposed continental shelf areas west and south-west of
Norway, (2) the trees migrated into the Scandes mountains
from the west during the late-glacial period, and thus (3) the
Scandinavian ice sheet was thinner than previously suspected
and that late-glacial nunataks were available and suitable for
early tree colonization. These hypotheses are highly contro-
versial, going against the generally accepted pattern of glacial
and forest history of the area in the late-glacial and early
Holocene. Therefore, we consider that it is important to
examine the hypotheses critically in the light of the existing
palaeoecological fossil and glacial geological evidence.
TYPES OF PALAEOECOLOGI CAL FOSSI L
EVI DENCE
Previously Kullman (1995, 1996, p. 97, 1998a, pp. 425426,
1998b, p. 153, 2000, p. 165, 2001, p. 40) has criticized pollen
analysis as providing poor evidence for the local occurrence of
trees, asserting that small outlying tree populations are not
detected by pollen analysis because pollen analysts do not
generally consider that occasional pollen grains or tails in
pollen diagrams may originate from small locally present tree
populations. The only way to prove local presence in these
circumstances is through nds of macrofossils (including
conifer stomata) or megafossils. He suggests that by
combining mega-, macro-, and microfossil data, a more
realistic comprehension can be obtained concerning the critical
pollen percentage level, that must be exceeded before spatially
precise biogeographical range-limit reconstructions can be
inferred from pollen data (Kullman, 2000, p. 165). Kullman
(2002, p. 1117) states It is increasingly evident, however, that
fossil pollen data do not always accurately account for such
vital aspects of historical biogeography, such as, location of
glacial refugia geographical spread and elevational shifts.
We largely agree with these statements. Pollen analysis is a
difcult tool to use in treeless environments and in the
detection of tree-line movements. In these situations records of
macrofossils and megafossils provide additional complement-
ary evidence (Birks & Birks, 2000, 2003; Eide, 2003). Later,
however, Kullman (2002, p. 1121) returns to his previous
(2000) proposal of calibrating pollen data with macro- or
megafossil evidence of local presence, and, implying that the
calibration has already been made, states: pollen records
from south-west Norway (Kristiansen et al., 1988; Paus, 1989)
display pollen sums that, judging from comparisons of pollen
and megafossil evidence from other areas (Kullman, 2000),
may suggest presence of birch, pine and spruce at sheltered
localities since at least 12,000 bp. He maintains that the trees
spread from there eastwards during the late-glacial to the
central Swedish mountains, where, however, his earliest dated
remains (Betula) are very early in the late glacial, at
14,020 80 and 12,870 70
14
C yr bp.
Kullman (2000, 2001, 2002), based on this comparison of
pollen and megafossil evidence from other areas (above), uses
arguments based on pollen data to propose that tree-pollen
percentages in late-glacial sites along the Norwegian west coast
might derive from small locally surviving tree populations.
However, it could be argued as equally likely that they do not.
Winds were much stronger than today in full- and late-glacial
times (e.g. COHMAP members, 1988), thus facilitating long-
distance dispersal of pollen. This makes the presence of
macrofossils even more important as proof of local tree
presence (Birks & Birks, 2000, 2003; Birks, 2003). Small pollen
percentages of Pinus and Picea and small or sometimes quite
large percentages of Betula pollen (some at least of which
originated from B. nana L.; van Dinter & Birks, 1996) in the
west Norwegian late-glacial should not be taken as proof for
local tree occurrences without supporting macrofossil evi-
dence, especially when independent macrofossil evidence for
local vegetation and climate is available from several sites in
western Norway (Fig. 1).
EXAMI NATI ON OF THE EVI DENCE
Palaeoecological evidence
There are four main lines of palaeoecological evidence with
which we can test Kullmans proposals: (1) plant macrofossil
evidence, ignored by Kullman, (2) late-glacial vegetation
reconstructions in western Norway that show extensive
alpine-vegetation analogues, (3) independent climate recon-
structions from other proxies and (4) evidence from the routes
and timing of tree spread in the early Holocene in western
Norway.
Plant macrofossil and pollen evidence for late-glacial trees in
south-west Norway
In spite of numerous late-glacial macrofossil investigations in
south-west Norway (Fig. 1), no macrofossils of tree-Betula
have yet been found. Although it is unsatisfactory to argue
from negative evidence, it would appear that tree-Betula was
absent, or very localized and so far undetected in south-west
Norway (Birks et al., 1993; Birks, 1993; van Dinter & Birks,
1996; Birks & van Dinter, 1997; see Birks, 2003 for a review).
There is similarly no macrofossil or stomatal evidence for the
presence of Picea and Pinus to indicate that they survived the
glaciation in the south-west Norwegian region or that they
were present in the late glacial (e.g. Fgri, 1949; see Giesecke &
Bennett, 2004). If the highest late-glacial pollen percentages of
Betula (including tree-Betula) (e.g. Paus, 1989) were derived
from long-distance transport (see van Dinter & Birks, 1996), it
is probable that the percentages of Picea and Pinus pollen were
H. H. Birks, E. Larsen and H. J. B. Birks
1462 Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd
also long-distance transported. Pinus pollen is notoriously
long-distance transported. The percentage representation of
long-distance transported pollen depends on the local pollen
production. As macrofossil and other evidence suggests that
the west-Norwegian late-glacial vegetation was almost certainly
treeless, low amounts of long-distance transported pollen
would not have been masked by local pollen, and could have
reached high percentages in a total pollen sum (Birks & Birks,
2003). This effect is demonstrable today by the considerable
percentages of tree pollen recorded on Spitsbergen that derive
from areas far to the south (van der Knaap, 1987). A similar
long-distance component was reported in the Devon Island ice
core from arctic Canada (McAndrews, 1984). Pollen inux or
annual accumulation rates give more reliable information, for
example, in the late-glacial at Krakenes on the Norwegian west
coast (Birks et al., 2000) and in northern Scotland (Birks,
1984). It is curious that Kullman (2002) does not consider any
of the late-glacial macrofossil investigations in western Norway
and instead relies entirely on pollen analytical data, in contrast
to his own advice detailed above.
Evidence for the actual vegetation growing during the full-
and late-glacial
Plant macrofossil and pollen data from west Norwegian glacial
deposits (Barstadvik E. Larsen & H.H. Birks, unpubl. data;
Andya Alm & Birks, 1991) and late-glacial sites (on Fig. 1)
provide abundant evidence that the vegetation was treeless. In
the examples cited by Kullman (2002) no tree-Betula macro-
fossils were found and the local vegetation was dominated by
Betula nana or Empetrum L. [pollen diagram by Paus (1989);
macrofossils by van Dinter & Birks (1996); pollen diagram by
Kristiansen et al. (1988); unpublished macrofossil data of H.H.
Birks]. Other west-Norwegian sites (Fig. 1) have assemblages
dominated by Salix herbacea L. and taxa associated with snow-
beds and fjellelds, etc., all closely analogous to vegetation
above the tree-line in the oceanic west Norwegian mountains
today (Birks, 1993, 2003; Jonsgard & Birks, 1995; Birks & van
Dinter, 1997), or even polar desert (Alm & Birks, 1991).
Individual species recorded as macrofossils in the Norwegian
late-glacial include Salix polaris Wahlenb., Ranunculus glacialis
L., Saxifraga rivularis L., S. cespitosa L., Sagina intermedia
Fenzl, Koenigia islandica L., Papaver Sect. Scapiora (Reich-
enb.), Aulacomnium turgidum (Wahlenb.) Schwagr., Polytri-
chum sexangulare Brid., etc. Given the vegetation and the
environment suggested by such species it is improbable that
tree-Betula, Picea and Pinus could have grown in these
conditions.
The radiocarbon date of 10,360 170
14
C yr bp (T-5592)
on the layer containing squirrel (Sciurus vulgaris) bones, and
by inference the occurrence of conifer forest (Kullman, 2002),
at the top of the Skjonghelleren cave sequence on the west
Norwegian coast (Larsen et al., 1987) was measured on a large
bulk sample of bone material of marine mammals, birds and
sh. Thus the bone sample probably covers quite a large age
span and the Younger Dryas age obtained should not be taken
as a denitive age for squirrel (cf. Stewart & Lister, 2001). Also,
as most of the dated material is of marine origin, it suffers
from a marine reservoir age of at least 400 and up to 600
1000 years (Bondevik et al., 2001). Taking into account the
occurrence of lengthy radiocarbon plateaux in the early
Holocene (e.g. Bjorck & Wohlfarth, 2001) the squirrel remains
could easily have an early Holocene true age.
Independent climate reconstructions from late-glacial sites in
south-west Norway
Summer temperature reconstructions using chironomids [at
Krakenes (Brooks & Birks, 2001) and Bjerkreim (S.J. Brooks,
unpubl. data)], Cladocera [at Krakenes (Duigan & Birks,
0 10
70
65
60
20 E
G
K
Bl
U
Bj
E
L
A
N
Ba
400 km
N
Younger Dryas
Oslo
Kristiansand
Stavanger
Bergen
Trondheim
Troms
Figure 1 Map of Norway and Sweden showing Mt A

reskutan
(star), full-glacial sites (triangles) and late-glacial sites in south
Norway with macrofossil records (black dots). The limit of the
Younger Dryas re-advance is shown. A Andya (Alm & Birks,
1991), Ba Barstadvik (E. Larsen & H.H. Birks, unpubl. data),
L Lerstadvatn (J. Mangerud & H.H. Birks, unpubl. data),
G Gody (Birks et al., 1993), N Nordfjord transect (Birks &
van Dinter, 1997), K Krakenes (e.g. Birks et al., 2000),
Bl Blomy (Birks, 1993), U Utsira (Birks, 1993), E Eige-
bakken (van Dinter & Birks, 1996), Bj Bjerkreim (H.H. Birks,
Aa. Paus & S.J. Brooks, unpubl. data).
Glacial survival of boreal trees in south-west Norway?
Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd 1463
2000)], Coleoptera [at Gody (Birks et al., 1993) and Krakenes
(Lemdahl, 2000)], Oribatid mites [at Krakenes (Solhy &
Solhy, 2000)], and glacial geological evidence (Larsen et al.,
1984; Larsen & Stalsberg, 2004) are all consistent (e.g. Birks &
Ammann, 2000; Birks et al., 2000, 2005). Reconstructed July
temperatures in the Allerd and Younger Dryas are well below
the thermal limit for tree-Betula growth, which needs a mean
July temperature of c. 11 C at the coast (Odland, 1996) and
temperatures were therefore also below the thermal require-
ments of Picea abies and Pinus sylvestris. Although Kullman
(2000, 2002) records Picea krummholz growing at mean June
August temperatures of 67 C, krummholz is not a pioneer
growth form, but a persisting, poorly reproducing, relict of
former warmer conditions (e.g. Kullman, 1983, 2001; Payette,
1983; Holtmeier, 2000). Kullman (2002) reports a 50-year-old
Picea sapling growing at a mean JuneAugust temperature of
5 C (comparable with modern Svalbard temperatures) on
Mt A

reskutan, thus illustrating the unique local microclimates


apparently present there today. However, in hollows such as
that where the wood remains were found, temperatures may be
several degrees lower. In the A

reskutan area, hollows in the


conifer forest are occupied by birch trees, or in the subalpine
birch forest, the hollows are treeless and contain tundra
polygons (J. Lundqvist, pers. comm.).
Routes and timing of tree spread in the early Holocene
Kullman (2002) proposes that trees survived in scattered
cryptic (Stewart & Lister, 2001) refugia with locally favour-
able climatic conditions. If so, these populations would have
been expected to expand immediately after the climate became
generally favourable after the start of the Holocene (11,530 cal
yr bp; Gulliksen et al., 1998). However, they did not, as
discussed below.
The pattern of Holocene tree-Betula spread in southern
Norway is summarized in Fig. 2b. Tree-Betula was the rst tree
to arrive and was locally present (as shown by macrofossils
and/or pollen inux values) in the Oslo area and in
southernmost Norway by 10,800 cal yr bp (Eide et al., 2005;
H.J.B. Birks & S.M. Peglar, unpubl. data) and at the west coast
near Bergen at 10,600 cal yr bp (J. Larsen, S.M. Pegiar, H.J.B.
Birks., unpubl. data). The high-resolution radiocarbon chro-
nology and macrofossil stratigraphy at Krakenes revealed that
tree-Betula took c. 650 years to expand locally (10,900 cal
yr bp) after the start of the Holocene (Birks et al., 2000).
Betula arrived near Trondheim at c. 10,6009800 cal yr bp
(H.J.B. Birks & S.M. Peglar, unpubl. data). Macrofossils of
Betula indicate its presence in the Dovre mountains by
10,000 cal yr bp (Eide, 2003) and on the southern Hard-
angervidda plateau by 10,80010,000 cal yr bp (Eide et al.,
2005). The oldest megafossils (large wood remains) of Betula
in the S. Norwegian mountains are dated to c. 9250 in Dovre
(Barth et al., 1980) and c. 7950 cal yr bp on the Hardangervi-
dda (Aas & Faarlund, 1988). Tree-Betula appears to have
migrated from its nearest proven (by macrofossils) Younger
Dryas refugia in Denmark and south Sweden (e.g. Jensen,
1985; Liedberg Jonsson, 1988). More than 600 years were
needed for Betula to spread round the Norwegian coast and up
the valleys into the mountains.
Pinus spread rapidly across southern Norway some 800
1000 years after Betula (Fig. 2c). Radiocarbon dating of its rst
macrofossil occurrences and pollen inux increases shows that
Pinus reached the Norwegian south coast and also the southern
Hardangervidda (Setesdal) at 10,0009600 cal yr bp (Eide
et al., 2005) and the Oslo area by 96009200 cal yr bp (H.J.B.
Birks & S.M. Peglar, unpubl. data). Pinus expanded near Bergen
on the west coast at 9160 cal yr bp (J. Larsen, S.M. Pegiar,
H.J.B. Birks, unpubl. data). The earliest Pinus megafossils in the
south-central Norwegian mountains near Trondheim were
dated at c. 9250 cal yr bp and on the Hardangervidda at
c. 9650 cal yr bp (Aas & Faarlund, 1988). At Lake Spaime
c. 45 km SW of A

reskutan, the earliest Holocene ora was of an


alpine, pioneer type and B. pubescens macrofossils are rst
recorded at 9500 cal yr bp (Hammarlund et al., 2005).
Although Pinus pollen reached c. 70% between 9500 and
c. 8000 cal yr bp, probably indicating its local presence,
macrofossils were not found until c. 6000 cal yr bp. The pollen
situation is similar in sites in the Handol valley 40 km SE of Mt
A

reskutan, but specic macrofossil records are lacking


(Segestrom & von Stedingk, 2003). Given the lower elevations
of Lake Spaime and the Handol valley, the lack of a rapid
expansion of both Betula and Pinus at these sites argues against
nearby late-glacial refugia at Mt A

reskutan (1360 m a.s.l.).


This evidence, based on macrofossils, megafossils and pollen
inux values, does not support a spread of Betula and Pinus
from coastal refugia or from the Swedish Scandes in the Allerd
and Younger Dryas but illustrates a post-glacial spread from the
south. The documented establishment delays of 600 to
> 2000 years would not be expected if these pioneer trees were
already locally present in small populations in the mountains
during the late-glacial or earliest Holocene or on exposed
continental shelf and coastal areas of western and south-
western Norway during the late-glacial. The nearest Younger
Dryas occurrences of Pinus documented by macrofossils, apart
from Kullmans records, are from southern England (Lambert,
1964), and south Sweden [Kullaberg (Liedberg Jonsson, 1988);
Skane (Gertz (1926); Blekinge (Berglund, 1966)].
The case of Picea abies is different. Picea can be a pioneer
tree on open soils (Giesecke & Bennett, 2004) but its seedlings
are shade tolerant and well adapted to regeneration within
forest, where it grows best on acid soils with adequate nutrients
and competes strongly with other forest trees (Nikolov &
Helmisaari, 1992). However, Mt A

reskutan has virtually no


soil, just bare bedrock (Lundqvist, 1969). Picea has relatively
low pollen production. Picea pollen records show that it spread
from the east late in the Holocene towards Jamtland and
western Norway and it never naturally reached the coast except
in the lowlands in the Trondheim area (e.g. Huntley & Birks,
1983; Hafsten, 1992; Kullman, 2001; Giesecke & Bennett,
2004). This pattern suggests that temperatures in the
mountains had already fallen below the thermal limit of Picea
before it could cross the watersheds and its southward and
H. H. Birks, E. Larsen and H. J. B. Birks
1464 Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd
westward spread in south Norway was blocked, so it has never
been native in south-west Norway (e.g. Fgri, 1949; Huntley &
Birks, 1983; Hafsten, 1992). In contrast, Kullman (2000, 2001)
interprets early Holocene dates of Picea megafossils to indicate
that small populations of Picea grew at high elevations in
moist, snow-rich habitats in the Swedish Scandes. He proposed
that Picea spread downwards and eastwards. As climate
became cooler and moister around 3000
14
C yr bp, the
scattered Picea populations increased in the eastern lowlands,
thus accounting for its apparent late spread from the east as
deduced from pollen records (see Giesecke & Bennett, 2004).
Pollen analyses of Holocene peat sections 4045 km SE of Mt
A

reskutan (Segestrom & von Stedingk, 2003) record single


Picea pollen grains back to c. 9000 cal yr bp, but not earlier.
Their records of coniferous wood macrofossils from one site
could be ascribed either to Picea (one pollen grain), Pinus
(whose pollen was abundant since c. 10,000 cal yr bp), or
Juniperus, which has a continuous pollen record at 23%.
Without specic macrofossil evidence, the record of conifer-
ous wood should not be used to conrm local presence of
Picea from small numbers of pollen grains.
The nearest last-glacial Picea macrofossil record to the
Scandinavian mountains is from Byelorussia (Punning et al.,
1983), which supports the spreading pattern based on pollen
records (Giesecke & Bennett, 2004). There are no other
documented records of late-glacial Picea in the Swedish
Scandes from mountains that might be more suitable
ecologically than Mt A

reskutan for its growth. Picea pollen


peaks associated with Picea macrofossils (needles) occurred
transiently in eastern and southern Finland in the early
Holocene before 9000
14
C yr bp (e.g. Vasari, 1962; Tolonen,
1967; Giesecke & Bennett, 2004), showing that this area could
be a source for early Holocene spread and establishment of
scattered small Picea populations to the west, even though
Picea subsequently became extinct in Finland and recolonized
later in the Holocene. Spruce can spread extremely quickly, as
~1.5 (0.75 mac)
0.9
0.7
2.8
Picea
(d)
Trondheim
reskutan
Oslo
Kristiansand
Bergen
S
e
t
e
s
d
a
l
YD
N
400 km
10 E
60 N
Dovre
v
H
(a)
Pinus
9.65
10
9.6
9.9
~10.0
9.6
9.16
9.0
9.25
(c)
>10.0
10.8
10.6
10.8
10.6
10.6
10.9
10.0
9.25
9.5
7.95
Betula
(b)
H
Spime
Figure 2 Maps showing the earliest
14
C-dated macrofossil/pollen-inux increase
records (black dots) and megafossil records
(stump symbol) in the Holocene for 2b
Betula L., 2c Pinus L., and 2d Picea A.Dietr.
Ages are given in calendar years bp
(1950) 10
)3
. Place names mentioned in the
text are shown in 2a (Hv Hardangervidda,
H Haukeligrend). Mt A

reskutan is shown
by a star. The Younger Dryas (YD) ice
re-advance limit is shown.
Glacial survival of boreal trees in south-west Norway?
Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd 1465
the seeds are shed in winter and travel long distances across a
smooth snow-covered landscape (Ritchie & MacDonald, 1996;
Giesecke & Bennett, 2004). However, its late-Holocene
expansion through already forested northern Europe was slow.
An example of actual spreading routes can be traced in detail
by combined Picea macrofossil and pollen inux records in
Setesdal, southern Norway (Fig. 2d). It entered Setesdal from
the south near Kristiansand by 1500 cal yr bp (Eide et al.,
2005) and spread some 60 km north up the valley by 980 cal
yr bp. In contrast, Picea reached northern Setesdal from the
eastern lowlands, arriving in the Haukeligrend area by 2800 cal
yr bp (H.J.B. Birks & S.M. Peglar, unpubl. data), but it never
reached the southern Hardangervidda plateau (Eide et al.,
2005). Spruce is sparse in the upper pine forest today and its
current altitudinal species limit is similar to that of Pinus at
around 900 m. The lowest pass across the mountains is at
1150 m. Picea penetrated northern Setesdal as far south as the
Hovden area by 700 cal yr bp. The central part of Setesdal
lacks native Picea where the two invading populations have not
yet met and coalesced (Fig. 2d). If Picea had survived during
the late-glacial on the south-west coast of Norway, it is
surprising that it went extinct at the start of the Holocene, and
was, in fact, absent from all of north-west Europe except at
high elevations in the Swedish Scandes.
Genetic studies also suggest a westward Holocene spread of
Picea abies from the east and south. Vendramin et al. (2000)
identied two main genetic types, a central European-alpine
and an east European-Scandinavian, that relate to the
locations of the glacial refugia proposed from the pollen
record by Huntley & Birks (1983) and Giesecke & Bennett
(2004). Further analyses by Bucci & Vendramin (2000)
delimited a genetically homogeneous area in central Sweden
and the Trondheim area, which was closely related to an
adjacent area in south-east Norway and south Sweden. They
propose that genetic variability increases with geographical
isolation (distance). Thus one might expect as indeed was
predicted as a generalization by Kullman (2002) that
populations of spruce derived from proposed refugia in
Figure 3 The maximum extent of the
Scandinavian ice sheet, the extent of cold-
based ice at the last-glacial maximum, the
limit of the Younger Dryas re-advance, and
the areas of cold-based ice at the start of
deglaciation. Mt A

reskutan is shown by a
star.
H. H. Birks, E. Larsen and H. J. B. Birks
1466 Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd
western Norway and central Sweden would be genetically
distinct from the populations that spread from refugia in
south and east Europe. However, this hypothesis is not
supported by the genetic data available so far.
Glacial geological evidence: the extent and character
of the Scandinavian ice sheet
How do the late-glacial megafossils in central Sweden relate to
the extent of the Scandinavian ice sheet? Kullman (2002)
quotes evidence to support a proposal that deglaciation was
more complex than previously supposed, with early emerging
nunataks. Trees could occupy these, perhaps in krummholz
form, during the Allerd and Younger Dryas, having migrated
from glacial refugia in western Norway.
The most recent reconstruction of the maximum extent
and conguration of the Scandinavian ice sheet (Svendsen
et al., 2004) parallels earlier reconstructions (e.g. Kullman,
2002, Fig. 1) (Fig. 3). The ice sheet was thickest (> 2500 m)
over southern Scandinavia, including the A

reskutan area.
The central part was cold-based, namely frozen to the
substrate. Little erosion occurred, and by mapping uneroded
landscapes and ribbed moraines, Kleman et al. (1997) and
Kleman & Hattestrand (1999) have delimited the cold-based
area, both for the last-glacial maximum (LGM) and for the
start of deglaciation (see Fig. 3). Mt A

reskutan lies close to


the boundary between cold- and warm-based ice during the
glaciation and deglaciation and may have experienced
both states, as there are glacial striations and scouring on
the abundant bare bedrock up to the summit (Lundqvist,
1969).
Seasonally sea-ice free conditions in the Norwegian-Green-
land Sea occurred during the full-glacial period (e.g. Sarnthein
et al., 1995) corresponding to warmer periods of the
Dansgaard-Oeschger cycles (Dokken & Hald, 1996). Dokken
& Hald (1996) and Siegert & Marsiat (2001) proposed that
moisture collected by mid-Atlantic storms passing over these
open seas and guided up the Norwegian coast built up the
Scandinavian ice sheet at modelled rates of accumulation of
c. 500 mm yr
)1
water-equivalent at the west coast. Although
conditions were relatively warm, the modelled annual temper-
atures along the west coast are between )5 and )12 C (Siegert
& Marsiat, 2001). The ice margin oscillated during the
Weichselian, allowing biota to spread periodically (e.g. during
the A

lesund interstadial 40 ka; e.g. Forsstrom & Punkari, 1997)


but it is universally agreed that the ice sheet extended beyond
the Norwegian coast to the continental shelf edge at the LGM
(c. 22,000
14
C yr bp) (Fig. 3), both from geological evidence
and modelling evidence (e.g. Siegert & Marsiat, 2001; Svendsen
et al., 2004), leaving nowhere for trees or other plants to grow
(Brochmann et al., 2003).
One of the earliest deglaciated areas close to the ice-sheet
margin (Svendsen et al., 2004) was Andya (north-west
Norway), but macrofossil evidence from soon after the LGM
(c. 19,000
14
C yr bp) shows that the vegetation there was polar
desert (Alm & Birks, 1991). Although the summer sea-ice
margin retreated north during the early deglaciation, allowing
open water to penetrate along the Norwegian coast, the reverse
happened during the Younger Dryas, with persistent sea-ice
south to the Lofoten-Vring Plateau area (Koc et al., 1993;
Birks & Koc, 2002; Birks et al., 2005). On land, the ice withdrew
from the coast during the early deglaciation (c. 1513 ka) but
re-advanced during the Younger Dryas to the limits mapped by
Andersen et al. (1995) and in coastal cirques (Fig. 3). A narrow
coastal strip of land was ice-free during the Younger Dryas
where climatic conditions were severe (Birks et al., 2005). Mean
July temperatures were c. 7 C lower than today at Krakenes
(i.e. mean July c. 56 C) (Larsen et al., 1984; Birks et al., 2000;
Larsen & Stalsberg, 2004) and at nearby Gody (Birks et al.,
1993). These temperatures typify the high- or mid-alpine
Norwegian vegetation zones and central Spitsbergen today.
DI SCUSSI ON ALTERNATI VE HYPOTHESES
The hypotheses of tree survival and migration can now be
considered in the light of all the available evidence. At the
LGM, the Scandinavian ice sheet covered western Norway
and the continental shelf. Where could the trees survive?
Plant-macrofossil evidence, inferred past vegetation, and
climate reconstructions, all falsify the hypothesis of tree
survival in south-west Norway during the late-glacial. The
occurrence of ice-free nunataks is still controversial (Nesje
et al., 1987; Larsen et al., 1995), but if present, these were
certainly mountain peaks with little or no soil and a harsh
climate. The existence of locally favourable environments for
tree growth in the mountains during the early late-glacial is
not supported by geological evidence or by the geomor-
phology of Mt A

reskutan, with its predominance of bare


bedrock and sparse soil development (Lundqvist, 1969): an
unlikely habitat for Picea. Therefore, from all these lines of
evidence, Kullmans (2000, 2001, 2002) hypotheses of glacial
survival of boreal trees at the south-west Norwegian coast
and their subsequent eastward migration to mountain
nunataks during the Allerd and Younger Dryas cannot be
supported.
The main problem in rejecting the tree-survival hypotheses
is the radiocarbon-dated wood remains implying late-glacial
tree growth on Mt A

reskutan. We can ask if the dates are


correct ages? Examination or treatment of the material for
possible sources of contamination was not reported, so we
have to take the dates at face value. If the dates are not
correct ages, they must be either too old or too young.
Although it seems to be an obvious idea that the trees are of
Holocene age and the wood has been contaminated with old
carbon, it is difcult to propose a mechanism for this. The
main ice-movement from the east may have transported
limestone to the non-calcareous Mt A

reskutan from strata


situated 46 km east and south-east and 800900 m lower
than the summit (Sveriges Geologiske Undersokning, 1984).
Calcareous contamination may then have entered the
groundwater from unweathered till (Lundqvist, 1969, pers.
comm.).
Glacial survival of boreal trees in south-west Norway?
Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd 1467
Alternatively, we ask if the tree remains are ancient, perhaps
older than the limit of radiocarbon dating? Remains of all
three species are recorded near the end of the last interglacial,
both near sea-level in western Norway and south Sweden, and
in the Hardangervidda mountains at 900 m (summarized by
Donner, 1995). Remains of the trees that grew at high altitudes
in the central Swedish mountains could have been preserved
under cold-based ice, where erosion was minimal, during the
full-glacial and the Younger Dryas (Fig. 3). However, Picea did
not grow in Jamtland during MIS 5a or 5c when the area was
free of ice, and it seems improbable that tree remains from
MIS 5e (interglacial) have been preserved through three glacial
episodes and two interstadials (J. Lundqvist, pers. comm.).
Kullman (2002) proposed that a small cold-based ice body
existed for some part of the later Holocene (Neoglaciation) on
Mt A

reskutan, contributing to the preservation of the ancient


plant remains. However, such snow-beds are rare on
Mt A

reskutan today and would probably have disappeared


during the Holocene thermal maximum (Lundqvist, 1969).
Whatever their true ages, the fact is that the wood remains
were preserved by cold temperatures and waterlogging in a
snow-bed area and a small pond at A

reskutan through the


Holocene.
Relatively small amounts of modern contamination by
organic material in very old samples could result in late-glacial
and early Holocene radiocarbon dates. We can only speculate
about the amount of undetectable contamination that may
have taken place on wood buried for the length of the Holocene
or longer in snow-beds or under shallow water and overgrown
by moss. Betula and Picea wood especially could have been
penetrated by rootlets and microbes. If such contamination is
suspected, individually teased out wood bres should be dated
(M. Friedrich, pers. comm.). However, if the wood remains
were all of interglacial age, it seems unlikely that they would
have received the same 2025% modern contamination neces-
sary to result in late-glacial and early Holocene radiocarbon
ages. A wider range of ages from 10 to 30 ka bp might have been
expected (S. Bjorck, pers. comm.).
Independently, if the pine remains are well enough
preserved, their ring sequences could be analysed and any
match with the Scandinavian Holocene tree-ring record
(K. Briffa & M. Friedrich, pers. comm.; e.g. Briffa & Osborn,
2002) would determine if they are of Holocene age. A non-
match could mean that the trees were not Holocene, or that
they did not match the Holocene tree-ring pattern.
At present, the radiocarbon dates reported by Kullman
(2002) remain uninterpretable except at face value as
documenting local presence of the trees in the central Swedish
mountains during the late-glacial and early Holocene. In light
of the evidence discussed here, we suggest that these dates
should not necessarily be taken at face value. The wood
samples should be carefully assessed for possible sources of
contamination and submitted (if suitable) to independent age
assessment by tree-ring analysis before basing unsupported
hypotheses on them about glacial and late-glacial survival of
the tree species at the south-west coast of Norway.
ACKNOWLEDGEMENTS
We are grateful to many people for helpful discussion and
input of ideas to this paper, in particular Keith Bennett, Anne
Bjune, Svante Bjorck, Keith Briffa, Wenche Eide, Michael
Friedrich, Jan Lundqvist, Heikki Seppa, John Inge Svendsen
and Ruth Terhuerne-Berson. We thank Jorunn Larsen and
Sylvia Peglar for the use of unpublished fossil data and Cathy
Jenks and Bjrg Svendgard for help with the gures. This is
publication Nr A91 from the Bjerknes Centre for Climate
Research, Bergen.
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BI OSKETCHES
Hilary H. Birks is a Quaternary palaeoecologist at the
University of Bergen. She specializes in plant macrofossil
analysis, and has used this technique to investigate the changes
in past vegetation and climate in several parts of the world, but
particularly during the late-glacial and early Holocene in
Norway.
Eiliv Larsen is a Quaternary geologist at the Geological
Survey of Norway, Trondheim, Norway. His main research is
on the reconstruction of glaciations during the last-glacial
cycle in Scandinavia and Russia and the progress and processes
of deglaciation.
John Birks is a plant ecologist and Quaternary palaeoecol-
ogist at the University of Bergen, Norway, and University
College London, UK. His research interests are centred on late-
Quaternary environmental and biotic change and on quanti-
tative palaeoecology, ecology and plant geography, particularly
in arctic and alpine areas.
Editor: Mark Bush
Glacial survival of boreal trees in south-west Norway?
Journal of Biogeography 32, 14611471, 2005 Blackwell Publishing Ltd 1471