Journal of Biogeography (J. Biogeogr.

) (2011) 38, 17921806

ORIGINAL
ARTICLE

Tree line identification from pollen data:

beyond the limit?
H. A. Binney1*, P. W. Gething2, J. M. Nield1, S. Sugita3 and M. E. Edwards1

Geography and Environment, University of

Southampton, Highfield, Southampton SO17
1BJ, UK, 2Spatial Ecology and Epidemiology
Group, Tinbergen Building, Department of
Zoology, University of Oxford, South Parks
Road, Oxford OX1 3PS, UK, 3Institute of
Ecology, Tallinn University, Uus-Sadama 5,
10120 Tallinn, Estonia

ABSTRACT

Aim The boreal tree line is a prominent biogeographic feature, the position of
which reflects climatic conditions. Pollen is the key sensor used to reconstruct
past tree line patterns. Our aims in this study were to investigate pollen
vegetation relationships at the boreal tree line and to assess the success of a
modified version of the biomization method that incorporates pollen
productivity and dispersal in distinguishing the tree line.
Location Northern Canada (307 sites) and Alaska (316 sites).
Methods The REVEALS method for estimating regional vegetation composition
from pollen data was simplified to provide correction factors to account for
differential production and dispersal of pollen among taxa. The REVEALS-based
correction factors were used to adapt the biomization method and applied as a set
of experiments to pollen data from lake sediments and moss polsters from the
boreal tree line. Proportions of forest and tundra predicted from modern pollen
samples along two longitudinal transects were compared with those derived from
a vegetation map by: (1) a tally of correct versus incorrect assignments using
vegetation in the relevant map pixels, and (2) a comparison of the shape and
position of northsouth forest-cover curves generated from all transect pixels and
from pollen data. Possible causes of bias in the misclassifications were assessed.
Results Correcting for pollen productivity alone gave fewest misclassifications
and the closest estimate of the modern mapped tree line position (Canada,
+ 300 km; Alaska, + 10 km). In Canada success rates were c. 4070% and all
experiments over-predicted forest cover. Most corrections improved results over
uncorrected biomization; using only lakes improved success rates to c. 80%. In
Alaska success rates were 7080% and classification errors were more evenly
distributed; there was little improvement over uncorrected biomization.
Main conclusions Corrected biomization should improve broad-scale
reconstructions of spatial patterns in forest/non-forest vegetation mosaics and
across climate-sensitive ecotones. The Canadian example shows this is
particularly the case in regions affected by taxa with extremely high pollen
productivity (such as Pinus). Improved representation of actual vegetation
distribution is most likely if pollen data from lake sediments are used because the
REVEALS algorithm is based on the pollen dynamics of lake-based systems.

*Correspondence: Heather Binney, Geography

and Environment, University of Southampton,
Highfield, Southampton SO17 1BJ, UK.
E-mail: h.a.binney@soton.ac.uk

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Keywords
Alaska, biomization, boreal forest, climatic change, northern Canada, palaeoecology, REVEALS, tree line, tundra, vegetation dynamics.

http://wileyonlinelibrary.com/journal/jbi
doi:10.1111/j.1365-2699.2011.02507.x

2011 Blackwell Publishing Ltd

Tree line identification from pollen data

INTRODUCTION
The boreal tree line is a prominent biogeographic feature on
both a regional and a continental scale; it reflects the
interaction of climate, physiography and species ecology and
physiology (Woodward, 1987; Prentice et al., 1992). The
position and character of this transition between forest and
tundra biomes, which are structurally distinctive, affects
terrestrial feedbacks to the global climate system via surface
properties such as albedo (Bonan et al., 1995; Foley et al.,
2003; Brovkin et al., 2009). Furthermore, with global warming,
some of the greatest changes that will affect future biodiversity
are likely to occur at such biome transitions (Callaghan et al.,
2005). The northern tree line is expected to respond strongly to
global warming, although the response may be locally variable
(Skre et al., 2002; ACIA, 2005), and empirical data suggest that
these changes are already under way (Sturm et al., 2001; Forbes
et al., 2009). Holocene records document dynamic shifts in the
position and composition of the tree line that largely reflect
climatic change (Payette et al., 2002), and an understanding of
past tree line dynamics and their critical drivers provides a
context for evaluating current and future change. Furthermore,
as a continental-scale vegetation feature that is strongly
climatically controlled, the tree line is a useful benchmark
for evaluating palaeoclimate model simulations and hence our
ability to project future changes in the tree line (Bigelow et al.,
2003; Kaplan et al., 2003; Wohlfahrt et al., 2004).
The Holocene boreal tree line is generally assumed to be
controlled primarily by growing degree-days (Woodward,
1987; Prentice et al., 1992), which in turn are governed by
hemispheric-scale climate controls. The modern boreal tree
line has been related to the median July position of the 10 C
isotherm (Koppen, 1931; Bryson, 1966), and the position of
the polar front (Pielke & Vidale, 1995), both of which are
correlated with the mean temperature of the whole growing
season. The boundary between forest and tundra is highly
variable, ranging from a broad ecotonal mosaic of forest and
tundra up to several hundred kilometres in extent (Payette
et al., 2001) to a sharp transition of a kilometre or less in
mountain areas. Trees at the limit of their climatic tolerances
may be stunted or prostrate (Elliott-Fisk, 1983). Any simple
concept of tree line is inadequate, and the phenomenon is
described via several spatially differentiated features: the forest
limit (or timber line), which defines the limit of closed forest;
the tree limit, which describes the farthest position achieved by
individuals of a given species; and the ecotone, which lies
between these two limits (Hustich, 1979; Callaghan et al.,
2002). Our usage, tree line, is shorthand for this complex
transition, which is described in more detail as necessary.
The interpretation of past tree line positions is normally
based upon the evidence of one or more fossil proxies, for
example pollen (Seppa, 1996), plant macrofossils (Binney
et al., 2009), fossil stomata (Leitner & Gajewski, 2004) and
fossil insects (Ponel et al., 1992). Pollen records are the most
abundant form of palaeoenvironmental data and are commonly used to infer past vegetation. However, the relationship
Journal of Biogeography 38, 17921806
2011 Blackwell Publishing Ltd

between pollen and vegetation is not straightforward because

many factors can complicate the pollen signal, including
openness of the landscape, differential pollen production (in
boreal regions this is high for forest and relatively low for
tundra taxa), differential dispersal of pollen grains, and
prevailing wind directions (Sugita, 1994). Various studies have
demonstrated the utility of macrofossils in pinpointing the
position of actual trees (Spear, 1983; Kremenetski et al., 1998),
but the abundance of macrofossil data varies considerably
between regions. Pollen, therefore, is the key sensor used to
reconstruct forest/non-forest vegetation dynamics in most
regions, and here we assess whether we can improve the
sensing of continental-scale tree line patterns using pollen data.
Quantitative reconstruction of forest and non-forest
cover using pollen data
Palaeovegetation reconstructions from pollen data have been
attempted using several approaches. The ratio between arboreal and non-arboreal pollen taxa (AP/NAP) can be used to
indicate forest and non-forested areas (Faegri & Iversen, 1950),
although Sugita et al. (1999) concluded that NAP percentages
give only a first approximation of landscape openness. Modern
pollen datasets have also been compared with remotely sensed
estimates of canopy cover such as percentage of woodland
cover derived from advanced very high resolution radiometer
(AVHRR) data (Tarasov et al., 2007), and fossil pollen has
been used to reconstruct past variations in tree cover and leaf
area index (LAI), including those at the northern foresttundra
ecotone (Williams, 2003; Gonzales et al., 2008).
The biomization algorithm developed by Prentice et al.
(1996) is an objective method of assigning pollen spectra to
large-scale vegetation units (biomes) that are comparable with
classifications used in vegetation modelling. The method
relates pollen taxa to one or more plant functional types
(PFTs), and PFTs to biomes. The use of PFTs as an
intermediate step avoids dependence on regionally unique
pollen floras, so the method is, at least in theory, globally
applicable. A square-root transformation reduces the variance
in pollen abundance data and increases the sensitivity of the
calculation to less abundant taxa. The biomization routine
assigns affinity scores for all possible biomes to pollen spectra,
and the biome with the highest affinity score wins.
The success of the pollen biomization method can be
assessed by comparing the biome assignments of sites in
modern pollen datasets with modern or potential vegetation
maps. Williams et al. (2000) applied biomization to eastern
North America and found fairly high correspondence between
vegetation and pollen across a range of biomes, although forest
tended to be over-represented at the northern tree line and the
foresttundra ecotone, a pattern they attributed to the
occurrence of abundant Pinus pollen beyond the limit of pine.
Edwards et al. (2000) found that the distribution of tundra,
taiga and cool conifer forests in Alaska and north-western
Canada generally corresponded well to actual vegetation
patterns. However, in eastern Siberia, which is dominated by
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H. A. Binney et al.
Larix forest, there was an over-abundance of sites classified as
tundra, probably reflecting poor production and/or dispersal
of Larix pollen.
Bigelow et al. (2003) carried out a biomization for the
circum-polar north. They excluded pollen taxa that were
< 0.5% of the pollen sum (normal practice) and applied a
weighting to Larix ( 15) to compensate for the low pollen
productivity of this taxon. The major patterns of forest and
tundra matched observed vegetation well in virtually all
regions, as assessed by a visual comparison of biome and
vegetation maps. Site-by-site comparisons revealed classification errors, with misclassifications likely of deciduous
(Larix) forest as tundra in Siberia, and tundra as boreal
forest in central Canada. They noted that lacustrine samples
better reflected regional vegetation while moss polsters better
reflected local vegetation a function of the size of the
pollen catchment. Williams et al. (2000) experimented with
simple modifications to the procedure to overcome the
effects on biomization of over-representation (Pinus
excluded below 5%, Quercus below 2.5%). However, these
adjustments, and the Larix up-weighting by Bigelow et al.
(2003) described above, amounted to tuning of the
biomization to produce a better fit with the observed data
in a region, and were not based on empirical data or
palaeoecological theory.
Studies of pollenvegetation relationships and attempts to
calibrate these relationships through estimates of pollen
productivity, dispersal and pollen source area have significantly advanced our ability to reconstruct past plant cover on a
quantitative basis. Davis (1963) assigned correction factors
for over- and under-represented pollen taxa and Parsons &
Prentice (1981) produced theoretical models of pollenvegetation relationships; these were developed further by Sugita
(1993, 1994), who attempted to define the pollen source area
concept and proposed a simulation approach (POLLSCAPE).
Sugita (2007a,b) developed the landscape reconstruction
algorithm (LRA), which combines the modelling and simulation approaches for vegetation reconstruction at both local and
regional scales. The two models within the LRA reconstruct
vegetation at regional (REVEALS) and local (LOVE) scales.
The validation of these two models is currently under way in
Europe (Hellman et al., 2008; Nielsen & Odgaard, 2010) and
North America (Sugita et al., 2010). In this study we incorporate correction factors estimated from the REVEALS model
into biomization to account for differential pollen productivity
and dispersal using the North American northern tree line as
the test case.
Study regions
Two transects running approximately southnorth through the
boreal tree line were the focus of the study. Transect 1 is
located in northern Canada (Fig. 1) and runs from southern
Manitoba ()98.42 W, 52.17 N) to the southern shore of
Melville Island in the Northwest Territories ()110.63 W,
74.51 N); it is c. 2500 km long and 1000 km wide. The
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southern part of this transect is dominated by Picea mariana

(Mill.) B.S.P., Picea glauca (Moench) Voss. and Pinus banksiana Lamb. The importance of Picea mariana increases
towards the northern tree line, where P. glauca becomes
restricted to well-drained situations and Larix laricina (Du
Roi) K. Koch to wetter locations (Elliott-Fisk, 2000). The
transect sampled the classic foresttundra ecotone of the
Canadian ShieldHudson Bay Lowland, where tree cover
declines gradually over several hundred kilometres. The
vegetation is a mosaic of patches of woodland and isolated
trees within tundra that decreases in stature northwards.
Successful tree growth depends upon mesoscale topographic
variation and soil qualities (Payette & Gagnon, 1985).
Transect 2, located in Alaska (Fig. 2), crosses a relatively
abrupt tree line, running from a location c. 300 km north of
Anchorage (150.17 W, 63.87 N) to 100 km south of the
northern Alaskan coast (153.43 W, 70.03 N). It is 700 km
long and 750 km wide. Much of transect 2 crosses the
interior boreal forest of Alaska, which lies between the
Alaska Range in the south and the Brooks Range in
the north. It is dominated by P. mariana and P. glauca
and stands of seral hardwoods, typically birch (e.g. Betula
papyrifera Marsh. var. humilis) and Populus spp. The
elevational tree line is at c. 800900 m, and alpine tundra
characterizes interior mountain ranges. The northern tree
line lies on the south slopes of the Brooks Range and is
abrupt, occurring in many areas as an elevationally controlled forest limit over a few hundred metres, although
gallery forest extends further to the north along north
south-trending river valleys (Viereck et al., 1992). Beyond the
elevational and latitudinal tree line, tall-shrub communities
are typical, dominated by alder [Alnus crispa (Ait.) Pursh]
and birch [Betula glandulosa (Michx.)] (Viereck et al., 1992).
North of the Brooks Range, vegetation is dominated by
shrubtussock tundra, dominated by birch, willow and
cottongrass (Eriophorum spp.).
Pollen representation in the study regions
At the broad scale in our study area forest tends to be overrepresented by pollen values. For example, for northern Alaska,
Anderson & Brubaker (1994) determined that spruce pollen
values of up to 10% typically indicated a location beyond the
tree line. Pollen productivity estimates (PPE, unit-less value)
describe the pollen production of a taxon relative to a standard
(Brostrom et al., 2008). Near the tree line, pollen productivity
of local stands may decline in response to less favourable
climatic conditions. Thus Ritchie (1980) estimates relative
Picea pollen productivity near the tree limit in north-west
Canada as < 1.0, while non-tree line values for Picea abies in
Europe are 1.8 (Brostrom et al., 2008). Hu et al. (1995) report
only 23% Picea pollen in western Alaskan gallery forests.
While pollen representation of boreal trees may be atypical
near the tree limit, the most common effect at sites recording a
regional pollen signal is for over-representation of boreal forest
pollen beyond the tree line.
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Tree line identification from pollen data

500

250

1000 km

70N

60N

50N
110W

100W

90W

Biomes

Pollen site
Cool mixed forest
Transect

Cold evergreen needleleaf forest

Cool evergreen needleleaf forest
Cold deciduous forest

Ecotone boundaries

Graminoid and forb tundra

Low- and high-shrub tundra
Erect dwarf-shrub tundra
Prostrate dwarf-shrub tundra
Cushion-forb, lichen, and moss tundra
Barren
Land ice

Figure 1 Location of transect 1 in northern Canada showing pollen sites with the underlying Kaplan & New (2006) vegetation map.

MATERIALS AND METHODS

Modern pollen and vegetation data
The modern pollen data are a subset of the Pan-Arctic
Initiative (PAIN) dataset (Bigelow et al., 2003). In total, 623
sites were selected for study, 307 in Canada and 316 in Alaska.
On the Canadian transect 31% of the samples are from
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lacustrine core tops and 66% from moss polsters. On transect 2

(Alaska), 50% of the samples are from lacustrine core tops and
49% from moss polsters. For both transects a small proportion
of the samples are from soil samples, peat core tops or air
pollen collectors, and for two samples the site information is
lacking.
Location maps of transects 1 and 2 are shown respectively in
Figs 1 & 2, at which scale some sites overlap. Full site details
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H. A. Binney et al.

250

1000 km

500

70N

60N

Brooks Range divide

160W

150W

140W

Figure 2 Location of transect 2 in Alaska showing pollen sites with the underlying Kaplan & New (2006) vegetation map. See Fig. 1 for
legend. The location of the Brooks Range divide is indicated by a dashed line.

are provided in Appendix S1 in the Supporting Information.

The vegetation map used and shown in Figs 1 & 2 was
produced by Kaplan & New (2006) and is based upon a
compilation of remotely sensed data. For this compilation the
Circumpolar Arctic Vegetation Map (CAVM Team, 2003;
Walker et al., 2005) is used to map the distribution of tundra
(fivefold classification). Other vegetation types and the location of the forest limit are based upon the Global Land Cover
2000 (GLC2000) map (JRC, 2003). The map was interpreted
and classified by regional experts and subsequently checked
extensively by ground-truthing (Kaplan & New, 2006). The
projection is Lambert azimuthal equal area and is based upon a
10-km grid.
Biomization approach
We adopted the taxon/PFT and PFT/Biome tables used by
Bigelow et al. (2003) and described in detail in Kaplan et al.
(2003). Bigelow et al. used a two-step approach to biomization. The first step differentiated forest biomes from a single
tundra biome, and a second step assigned tundra to one of five
1796

types. This two-step approach reduced misclassifications

between forest and tundra over a single-step approach (see
Table S1 in Appendix S2 for biomes and characteristic taxa).
We used the two-step approach in our biomization. Unlike
Bigelow et al. (2003), we did not use any multipliers for taxa
that are particularly poorly represented in the pollen spectra,
but we used the > 0.5% inclusion threshold for all taxa in the
pollen sum as recommended by Prentice et al. (1996). The
Bigelow et al. (2003) pollen biome classification we used
corresponds to the classifications used in the Kaplan & New
(2006) vegetation map. Tundra biomes are mapped in the
vegetation map and the biomized pollen data, but we do not
address patterns in tundra biomes further in this paper.
The biomization affinity scores for the pollen dataset were
calculated using an adapted biomization algorithm written in
matlab v. 7.3.0.267 (MATLAB, 2006; Mathworks Inc., Natick,
MA, USA), which is available from the authors on request. The
algorithm provides the option of conventional square-root
transformation and applies specified correction factors after
taxon percentage calculations. In cases where a pollen sample
has multiple highest affinity scores, the biome assignment is
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Tree line identification from pollen data

made using a prescribed order, whereby the biome with the
least diverse subset of taxa is assigned (Prentice et al., 1996;
Bigelow et al., 2003).
Landscape reconstruction algorithm (LRA)
The LRA is a framework for pollen-based quantitative
reconstruction of vegetation and land cover based on palynological theory (Sugita, 2007a,b). It corrects for biases in
pollenvegetation relationships: basin size, inter-taxonomic
differences in pollen dispersal and production, and spatial
heterogeneity of source plants (Prentice, 1985; Sugita, 1994)
and generates quantitative estimates of vegetation cover. The
REVEALS model is the first step and generates estimates of
regional vegetation composition, which are necessary in the
second step (LOVE) to calculate background pollen derived
from beyond the relevant source area for small-sized sites
(< 1050 ha; Sugita, 2007b). The basic premise behind
REVEALS, which is supported by both numerical simulations
and empirical studies, is that within a given region, pollen
records from large sites (> 100 ha) do not differ significantly
(Sugita, 2007a; Hellman et al., 2008; Sugita et al., 2008;
Soepboer et al., 2010). We apply a simplified version of the
REVEALS model as a correction method to reduce representation bias in pollen spectra with the expectation that this
should improve accuracy of the biomization procedure. The
original model and our modifications are described below.

Original model
The REVEALS model (Sugita, 2007a) is a modified and more
generalized version of the original R-value model (Davis, 1963)
and is expressed as:
 R
1
Z max
ni;k ^ai R
gi zdz
^ i;k 
V
1
 R
1 :
Pt
Z max
gj zdz
aj R
j1 nj;k ^
^ i;k is the
The symbols in equation 1 are defined as follows: V
estimate of the regional vegetation composition of species i in
proportion, based on pollen data from site k (dimensionless),
ni,k is the pollen count of species i at site k (grains), ^ai is the
estimate of pollen productivity of species i relative to a
reference taxon (dimensionless; pollen productivity is a

measure of the pollen produced by a given species relative to

a set of pollen taxa within a given region), R is the radius of site
k (in metres), z is the distance from the centre of site k (in
metres), t is the total number of species used to calculate
vegetation composition, Zmax is the maximum spatial extent of
the regional vegetation in a radius from the centre of site k (in
metres), and gi(z) is a species-specific pollen dispersal-deposition function for species i (a function of distance z).

Modified REVEALS
To make the REVEALS model applicable to the biomization
process we simplified it and assumed the following.
1. Pollen dispersal and deposition followed Prentices model
(Prentice, 1985), which was developed for describing the
pollenvegetation relationship on bogs and mires. This is
simpler than other models and thus appropriate for a first
approximation method such as the approach we are describing
here.
2. All pollen came from within a 500-km radius from each site,
which corresponded to the 9095% characteristic source
radius of very light pollen types (Table 1) (Prentice, 1985;
Sugita, 1993).
3. Basin radius, R, was set to 500 m at all sites to portray all
sampling basins as large sites (see above) and thus appropriate for the REVEALS applications (Sugita, 2007a). In reality
many sites were smaller, particularly as some samples were
from moss polsters. Nevertheless, we used all samples because
modern and fossil pollen datasets tend to have mixed sources
(from moss polsters and lakes, and from small to large basins),
and we wished to assess whether our approach could detect a
signal regardless of basin size and type.
Then equation 1 becomes,
ai Ci
^ i;k  Ptni;k =^
V
aj Cj
j1 nj;k =^

where
Ci  e2:174bi  e5:157bi
and
4:0vg;i
bi  p
n pCz u
(Prentice, 1985).

Table 1 The assignment of five pollen dispersal factors (C; value shown in column 5 and named in column 1) based upon the fall speed (vg,
derived from pollen size, mass and morphology, column 2); the resulting mean values for types assigned into that category (column 3); and a
standardized value relative to the intermediate value (column 4, rounded in column 5).

Pollen type

Fall speed of pollen,

vg (m s)1)

Averaged
C value

Standardized relative
to intermediate value

C value assigned to each

class (i.e. pollen type)

Very heavy
Heavy
Intermediate
Light
Very light

vg 0.065
0.065 > vg 0.050
0.050 > vg 0.035
0.035 > vg 0.020
0.020 > vg

0.01416
0.04400
0.09652
0.19627
0.28874

0.147
0.455
1.000
2.003
2.991

0.15
0.45
1.00
2.00
3.00

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H. A. Binney et al.
The mean wind speed, u, was set to 3.0 m s)1, the empirical
constant to express the atmospheric condition, n, was set to
0.250 (i.e. neutral conditions) and the vertical diffusion
constant, Cz, was set to 0.12 (Tauber, 1965; Prentice, 1985).
Fall speed of pollen, vg,i, is species dependent (i.e. for species i).
Therefore, Ci (hereafter we term this the dispersal factor of
species i) is a function of fall speed of pollen of this species.
Given pollen counts, dispersal factors and productivity values
of pollen grains of constituent taxa, pollen proportions were
converted to the vegetation proportions that were less biased
than uncorrected estimates. We call equation 2 the REVEALSbased correction for the biomization method.
To increase the ease of application of the REVEALS
correction, we made further simplifications. The fall speed of
pollen grains, vg in (m s)1), was classified into five pollen type
categories based on pollen size, mass and morphology: (1) very
heavy, vg > 0.065; (2) heavy, 0.065 vg > 0.050; (3) intermediate, 0.050 vg > 0.035; (4) light, 0.035 vg > 0.020; and (5)
very light, 0.020 vg. When measured and observed values of
fall speed were not available for a given taxon, values were
estimated by Stokes law (Gregory, 1973), based mostly on
pollen size and morphology (i.e. long and short axes, and
shape factor). For each category, we calculated the mean value
of the dispersal factor, Ci, within the range of each of the fall
speed categories given above. We then standardized the
dispersal factors relative to that of the intermediate value, as
shown in Table 1.
Accurate pollen productivity estimates are available only in a
limited number of regions in the world (Brostrom et al., 2008),
although boreal and tree line taxa are among the most studied
to date. Here, the pollen productivity, a, was set to one of only
three classes: in our initial experiments these were 2.0, 1.0 and
0.5 for high, intermediate and low pollen producers, respectively. This simplified system allowed us to portray semiquantitatively relative differences in pollen productivity of
constituent plant taxa in a region where knowledge of their
ecology, phenology and reproductive biology is, though
incomplete, relatively well understood. To explore the importance of the relative values of the three classes of a we also
experimented with additional classes that expand the range of
the corrections for high and low pollen producers: (1) a
3:1:0.5; (2) a 3:1:0.333; and (3) a 4:1:0.25, the latter was the
most extreme correction for high and low pollen producers.
The original biomization algorithm includes a square-root
transformation of pollen percentage calculations, which stabilizes variance and increases the methods sensitivity to less
abundant taxa (Prentice et al., 1996). It is also potentially a
way of reducing the relative importance of high pollen
producers (i.e. a correction). We therefore also included
experiments where square-root transformation was not used in
the biomization calculations.
Finally we investigated how Pinus affects the location of the
tree line in northern Canada. Empirical evaluations of pollen
productivity in Europe showed that Pinus sylvestris has a
higher pollen productivity estimate than Picea abies (Brostrom
et al., 2008). Whilst the species in North America are different,
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Table 2 The combinations of the dispersal (C) and the pollen

productivity (a) factors for the key taxa used in the initial
REVEALS-based biomization experiments. See Appendix S3 for
the full list of taxa.

Pollen taxon

Pollen
Fall speed productivity,
Dispersal
of pollen, a (initial
classes)
factor, C
vg

aC
(combination)

Larix
Picea
Pinus
Juniperus
Betula
Alnus
Populus
Salix
Poaceae
Cyperaceae
Ericales
Asteraceae
All other NAP

0.100
0.056
0.031
0.016
0.024
0.021
0.025
0.022
0.035
0.035
0.038
0.051

0.15
0.45
4.00
3.00
4.00
4.00
1.00
1.00
0.50
0.50
0.50
0.225

1.0
1.0
2.0
1.0
2.0
2.0
0.5
0.5
0.5
0.5
0.5
0.5
0.5

0.15
0.45
2.00
3.00
2.00
2.00
2.00
2.00
1.00
1.00
1.00
0.45
Depending
on vg

NAP, non-arboreal pollen.

many observations confirm the particularly high productivity

of North American boreal species of Pinus (e.g. Jackson &
Smith, 1994). Pinus proved problematic for Williams et al.
(2000) in their biomization of eastern North America, and it
was present at the southern part of the Canadian transect (but
not in the region covered by the Alaska transect). We
employed the simple modification of omitting Pinus from
the pollen sum in one experiment (no other corrections except
square-root).
Table 2 shows the combinations of the dispersal (C) and the
initial productivity factors (a) for key taxa we used for the
REVEALS-based biomization experiments. As described above,
different a values were assigned in subsequent experiments. (A
list for all taxa contributing to the pollen sum is available in
Appendix S3.)
Vegetation and pollen biome comparisons
We made a series of comparisons between the vegetation- and
pollen-derived biome classes, using just forest and tundra
biome classes for selected combinations: the PAIN (uncorrected) biomization (Bigelow et al., 2003), plus a range of
REVEALS-based values of a and C, singly and in combination,
with and without a square-root transformation ().
Analysis was carried out using a geographical information
system (GIS). The vegetation map was imported into ArcGIS
v. 9.3 (ESRI Inc., 2008, Redlands, CA, USA) as a raster layer
with a spatial resolution (cell size) of 10 10 km, and the set
of 623 pollen sites was imported as a point vector file with
each site located according to the latitude and longitude
coordinates recorded in the field. The two datasets were
Journal of Biogeography 38, 17921806
2011 Blackwell Publishing Ltd

Tree line identification from pollen data

aligned spatially using a common Lambert azimuthal equal
area projection.
While visual comparison of the two datasets in the GIS
facilitated qualitative assessment of major spatial patterns of
misclassification, an aim of this study was to develop a new
methodology that would allow the more formal representation
and assessment of these spatial patterns across the latitudinal
tree line and the foresttundra ecotone. The method we
devised is designed to quantify this correspondence along any
transect of interest.
A bespoke algorithm was written in the R programming
language (R Development Core Team, 2008) that generated,
for any specified transect across the northern tree line, a
graphical summary of the southnorth transition between
different biomes as derived from both the vegetation map and
the pollen data. Rather than capturing this information along a
one-dimensional line, we defined transects with a specified
width as well as a length in order to capture a sufficient sample
of pollen sites. The resulting transect region was divided along
the southnorth axis into (n) narrow segments of band-width
(b) such that n b = l. Within each segment, the proportion
of pollen sites assigned to each biome was computed.
Correspondingly, the proportion of raster grid cells of the
vegetation-derived map assigned to different biomes was
calculated within each segment. Estimated biome proportions
could then be plotted against transect position for any given
biome or set of biomes, and the plots for vegetation- and
pollen-derived biomes compared. Both transects were divided
into 15 segments, giving an approximate bandwidth of 167 and
47 km for Canada and Alaska, respectively. The data were
smoothed using a moving average in R.
The basic pixelbiome comparisons and transect-segment
data were then used in further analyses (described below with
their results for efficiency) to assess misclassification bias,
spatial variation in classification error and the role of basin size
(essentially a comparison of lake and moss-polster samples).
RESULTS
Latitudinal transects: graphical representation of the
tree line
The decline from predominantly forest cover (dark grey area)
to predominantly tundra cover (light grey area) for both
transects is shown in Fig. 3a(i) and Fig. 3b(i). The most steeply
sloping portion of the curve approximates the latitudinal
extent of the ecotonal zone in the regions covered by the
transect. A point where the forest and tundra proportions are
each 50% is used by Larsen (1974) and Elliott-Fisk (1983) to
define the boundary between arboreal/non-arboreal vegetation; here we also use the 50% point as the tree line for the
purposes of comparison.
On the Canadian transect (Fig. 3a(i)) the mapped vegetation shows a gradual decline in the proportion of boreal
forest, and the 50% point (marked by an arrow on the
figure) falls c. 700 km north from the southern origin. The
Journal of Biogeography 38, 17921806
2011 Blackwell Publishing Ltd

pollen values show a comparatively sharp drop in contrast

to the gradual decline in the vegetation, and the decline is
displaced northwards in all REVEALS experiments and the
original PAIN biomization. The best performing correction
factor is shown by a (productivity) correction 4:1:0.25 plus
square root (Fig. 3a(iii)), in the sense that the northward
displacement of the mapped 50% point is most reduced
(300 km too far north). The correction that includes both a
2:1:0.5 and C (dispersal) (with square root) is slightly poorer
(550 km, not illustrated), and the correction for C alone
(with square root) performs the worst (700 km too far
north).
The Alaskan transect (Fig. 3b(i)) crosses the tree line along
the southern Brooks Range at c. 340 km from the southern
origin; this area is shown by a relatively steep decline in boreal
forest cover. The biomized pollen data matched this pattern
well but showed a small tail of forest northwards, a feature
that is not affected by any REVEALS experiments. The a
correction (3:1:0.33 with no square root) performed the best
(Fig. 3b(iii)), with a northward displacement of the 50% point
of only 10 km.
Error rates and spatial bias: Canada
We assessed overall error rates, whether there was a bias in
misclassification and whether that bias was spatially dependent. Table 3(a) shows the four best overall percentage correct
matches of forest and tundra biomes for all Canadian sites,
plus percentage contribution to the error of the mismatch in
the forest zone (i.e. forest vegetation shown as tundra by the
pollen). The a 4:1:0.25 (with ) was best at 67.1% correct (this
also gave the best result on the transect see above). There was
little difference in the overall performance in many of the
REVEALS-based correction factors, but a few factors performed particularly poorly, notably C-only and the original
PAIN results. Errors were predominantly due to the assignment of forest biomes to pollen samples within the tundra. The
removal of Pinus from the Canadian pollen samples (not
shown) showed a marked improvement of forest/non-forest
biome assignment (45.8% for original PAIN; 64.7% with the
removal of Pinus). Only correction factors with a = 4 (applied
to Pinus) gave a higher number of correct biome assignments
than the no-Pinus experiment. For a full list of results see
Table S2 in Appendix S2.
The spatial distribution of the error was examined for the
best performing correction factors. Pollen samples were
assigned to one of three sectors (see Fig. 1) that correspond
approximately to the forest, ecotone and tundra portions of
the transect and combined to give a sector error using the best
performing correction factor (Table 4). The error rates were
relatively low in the forest zone and the ecotone (c. 20%); all
errors within the forest zone were due to forest sites being
misclassified as tundra, and the error was evenly distributed in
the ecotone. In the tundra zone (40% error rate) virtually all
errors were due to the tundra being misclassified as forest by
pollen.
1799

H. A. Binney et al.
(a)
Transect 1 (Canada)

(b)

Transect 2 (Alaska)

(i) 1.0

1.0

50:50 = c. 700km

0.4

0.2

0.2

500

1000

1500

1.0

2000
(PAIN)
PAINPollen
(with
)

0.8
0.6

1300km
1350km

100

(ii)

400

500

600

700

PAIN (with )
360km

+40km

0.4
0.2
0

500

(iii) 1.0

1000

1500

2000

correction (4:1:0.25) (with )

0.8
0.6

1000km

(iii)

100

200

1.0

300

400

500

600

700

correction (3:1:0.33) (with no )

0.8
0.6

+300km

0.4

330km

+10km

0.4

0.2

0.2
500

1000

(iv) 1.0

1500

initial

0.8

2000

correction (2:1:0.5)

1200km

+500km

0.4

100

200

(iv) 1.0

(with )

0.6

300

400

500

600

700

0.8

initial correction (2:1:0.5)

(with )

0.6

305km

-15km

0.4
0.2

0.2
0

300

0.6

0.2

200

1.0
0.8

+650km

0.4

50:50 = c. 320km

0.6

0.4

Kaplan & New vegetation

0.8

0.6

(ii)

(i)

Kaplan & New vegetation

0.8

0
500

1000

(v) 1.0

1500

C correction (with )

0.8

100

2000

(v)

200

300

400

1400km

+700km

0.4

600

700

1.0

C correction (with )

0.8

0.6

500

0.6

425km

+105km

0.4

0.2

0.2

0
500

(vi) 1.0

1000

1500

C & initial correction (with no )

0.8
0.6

2000

1275km

100

300

400

500

600

700

C & initial correction (with no )

0.8

+575km

0.4

200

(vi) 1.0
0.6

365km

+45km

0.4

0.2

0.2

500

1000

1500

2000

100

200

300

400

500

600

700

Transect Distance (km)

Transect Distance (km)

Forest

Tundra

Figure 3 The proportions of forest and tundra recorded for (a) transect 1, Canada, and (b) transect 2, Alaska, by (i) the vegetation map
(sampled by grid squares) and (ii) the Pan-Arctic Initiative (PAIN) pollen biome assignments, compared with the results obtained using
four correction factors [(iii) to (vi)]. Panel (iii) for both transects represents the best performing correction factor for that transect. The
arrow shows the 50% transition point for each transect with the offset from the vegetation forest/tundra transition shown to the right.
a, pollen productivity. C, pollen dispersal.

1800

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Tree line identification from pollen data

Table 3 Comparison of the overall percentage correct match for
pollen biomes versus vegetation for (a) all Canadian sites and (b)
the lake-only sites. The four best performing REVEALS-based
biomization methods (a, pollen productivity categories) are
shown [plus original Pan-Arctic Initiative (PAIN) and pollen
dispersal (C)-only results for all site types]. Column 3 shows error
due to tundra vegetation shown as forest by the pollen. The
opposite error (forest shown as tundra by pollen) is 100 minus the
value in column 3.

Biomization method
(a) All Canadian sites
a 4:1:0.25 
a 4:1:0.25 no 
No Pinus or factors with 
a 3:1:0.33 
Original PAIN
C only with 
C only with no 
(b) Canadian lake sites
a 4:1:0.25 
a 4:1:0.25 no 
No Pinus or factors with 
a 3:1:0.33 

Overall
success
rate (%)

Tundra vegetation
shown as forest
by the pollen (%)

67.1
66.8
64.7
64.1
45.8
43.1
39.7

85.6
93.9
94.2
91.5
100.0
100.0
100.0

79.1
84.6
83.5
82.4

47.4
85.7
73.3
68.8

The REVEALS model is underlain by the assumption that a

large site (R = 500 m) was recording regional vegetation; this
assumption was partly violated by the inclusion in the dataset
of moss polsters enclosed by forest. We therefore examined the
results for a lakes-only subset of the data that included 90 sites
and meets the assumption better (Table 3(b)). There was a
marked improvement in the overall predictive ability of the
data, with the majority of REVEALS-based correction factors
giving a success rate of > 80% (the four best results are again
shown). The spatial error for lakes using the a 4:1:0.25 (with )
correction factor showed that the improvement was within the
tundra zone (84.9% correct) (Table 4). The error rate for the
lake sites was highest within the ecotone (30% error) where
tundra sites tended to be misclassified as forest. Within the
forest zone all the errors were due to forest sites being
misclassified as tundra by the pollen.
Error rates and spatial bias: Alaska
The best-performing a 3:1:033 (no ) combination, which also
gave the best transect result, had a success rate of 79.4%
(Table 5). When considering both lakes and polsters, the
Alaskan transect showed a higher proportion of correct pollen
biome assignments than the Canadian transect for all experiments (c. 7080% and c. 4070%, respectively). The types of
mismatches were more varied and errors less biased. As with
the Canadian transect, correction for dispersal alone did not
perform well. Notably, the original PAIN biomization (78.8%
correct) was barely less effective than any REVEALS-based
Journal of Biogeography 38, 17921806
2011 Blackwell Publishing Ltd

Table 4 Analysis of the spatial distribution of the misclassification errors of tundra vegetation shown as forest by the pollen data
for the three sections of the Canadian transect (forest, ecotone and
tundra) using the best performing REVEALS-based correction
factor (pollen productivity, a 4:1:0.25 with square root) for (a) all
sites on the transect and (b) a subset of lake-only sites. Column 3
shows error due to tundra vegetation shown as forest by the pollen
(the reverse error is 100 minus the value in column 3).

Sector
(a) All sites
Forest
Ecotone
Tundra
(b) Lakes only
Forest
Ecotone
Tundra

Overall
success
rate (%)

Tundra vegetation
shown as forest by
the pollen (%)

80.0
82.0
61.0

0.0
44.4
98.8

78.4
70.0
84.9

0.0
66.7
100.0

Table 5 Comparison of the overall percentage correct match

for pollen biomes versus vegetation for each of the four best
REVEALS-based biomization methods for sites on the Alaskan
transect. Column 3 shows error due to tundra vegetation shown as
forest by the pollen (the reverse error is 100 minus the value in
column 3).

Biomization method

Overall
success
rate (%)

Tundra vegetation
shown as forest by
the pollen (%)

a 3:1:0.33 no 
a 2:1:0.5 
a 3:1:0.5 
Original PAIN 

79.4
79.1
79.1
78.8

55.4
40.9
40.9
74.6

correction. For a full list of results see Table S3 in Appendix S2.

In the three-sector comparison for Alaska, the northern
portion of the interior boreal forest and the southern Brooks
Range are defined as the ecotone for the purposes of
comparison. Vegetation in this area formed a large-scale
mosaic of forest and tundra, primarily controlled by topography (Fig. 2). The best-performing factor gives 74.2%
correct in the forest of the southern interior, with the majority
of the errors due to forest being misclassified as tundra by the
pollen (Table 6). The success rates in the ecotone and the
tundra zone (north of the Brooks Range tree line) were 54.0%
and 94.1%, respectively, with slightly more than half of the
errors being due to tundra misclassified as forest by the pollen.
The use of lakes sites alone for the Alaskan data (not shown)
had little effect on the results. In the forest zone the success
rate in using only lakes was unchanged as there were no mosspolster sites within the forest. In the ecotone 57.1% of biome
calls were correct, while 92.3% were correct north of the tree
line.
1801

H. A. Binney et al.
Table 6 Analysis of the spatial distribution of the misclassification errors of tundra vegetation shown as forest by the pollen data
for the three sections of the Alaskan transect (forest, ecotone and
tundra) using the best performing REVEALS-based correction
factor (pollen productivity, a 3:1:0.33 with no square root, ).
Column 3 shows error due to tundra vegetation shown as forest by
the pollen (the reverse error is 100 minus the value in column 3).

Sector

Overall
success
rate (%)

Tundra vegetation
shown as forest by
the pollen (%)

Forest
Ecotone
Tundra

74.2
54.0
94.1

25.0
60.9
54.5

We determined that small-scale variability in pixel ID on the

vegetation map was not affecting the comparison of vegetation
and pollen data by comparing the vegetation of a single pixel
that represents a pollen site with that aggregated from 9and 25-pixel areas (the vegetation cover being defined by a
simple majority of forest or tundra pixels). For both the 9- and
25-pixel aggregated vegetation there was only an 8% change in
vegetation assignment.
DISCUSSION
The REVEALS model accounts for both pollen dispersal and
variation in pollen productivity. In this simplified REVEALSbased correction approach we have experimented with
correction factors that appear reasonable, given incomplete
information on pollen production in the North American
boreal forest. The effectiveness of the REVEALS model also
depends upon assumptions about vegetation homogeneity and
basin size, and thus their impact on this study also requires
examination. Below we discuss the differences in success of the
correction factors we tested and features of spatial scaling that
may affect the results. Then we assess the overall effectiveness
of correction factors in the two regions and the potential for
REVEALS-based correction biomizing and/or tree line transect
analysis to locate the tree line effectively, including their
application to fossil pollen data.
Relative performance of different correction factors
(and no correction)
The Alaskan experiments had a fairly small range of classification success and exceed 50% in all cases (66.579.4%).
However, for Canada the range was wider (39.767.1%) and
the worst-performing factors had success rates below 50%,
including PAIN. The square-root transformation had no
consistent effect, possibly as it affected pollen values of minor
taxa more strongly than it suppressed those of high producers.
Correction for dispersal alone was not effective in either
region. This may appear slightly counter-intuitive, as explanations of the tree line pollen problem often invoke dispersal
of tree pollen into tundra.
1802

Values calculated for Picea in Alaska show why this might

be. When dispersal is strictly defined on the basis of the
aerodynamic properties of pollen, the rapid fall speed of Picea
generates a low C value and thus actually augments Picea
values, even though it is clear from observation that Picea was
over-represented in the pollen spectra (see above). In contrast,
assigned productivity (a) values always work to reduce the
impact of high pollen production.
In Canada the highest ranges of a values gave the best
success, probably because they best mitigated the high
production of Pinus pollen. This was supported by the fact
that merely removing Pinus from the pollen sum and not
applying correction factors was the third best result. In Alaska
the higher success rates were also mostly shown by a-based
corrections, but they varied as to whether there was a bias
towards over-prediction of forest or over-prediction of tundra;
in general the over-prediction of tundra was more prominent
than in Canada. In Alaska the original PAIN biomization had a
success rate only slightly less than the best a-corrected
biomization. In a Picea-dominated region such as northern
Alaska, where pollen production and dispersal values were not
extreme (compared with Pinus in Canada, for example), the
square-root transformation functioned quite effectively to
reduce the impact of high producers and boost the values of
low producers (which, typically, were abundant and rare taxa,
respectively). In such a situation REVEALS-based biomization
has little extra effect on biomizing outcomes.
The transect-based calibration of past tree lines
The heterogeneity inherent in any vegetation mosaic is likely to
reduce the effectiveness of any 1/0 classification system such as
biomizing. Thus the use of transects, which average vegetation
over an area that approaches the scale appropriate to
REVEALS spatial assumptions, is a productive approach to
testing the REVEALS methodology and to the reconstruction
of past tree lines using spatial arrays of pollen data.
The pollen-based curves for Canada (Fig. 3a) are generally
steeper than the vegetation curve, suggesting that the bias
leading to over-interpretation of forest in tundra is maintained
for some distance north and is then rapidly reduced. Our
analyses point to the strong influence of Pinus pollen, which is
derived from the more southerly boreal forest. The sharp
decline in pollen-defined forest may reflect the transport limits
of Pinus pollen, perhaps mediated by meteorological control
such as the summer location of the Arctic Front. In Alaska, a
less productive taxon, Picea, dominates the forest signal and,
unlike Canada, the tree line is relatively abrupt. Thus, perhaps
not surprisingly, the best-corrected pollen transect closely
reflects the overall position of the tree line in Alaska.
Data quality
We investigated two possible data-related causes of bias: the
quality of the map, and the scale of the map as it relates to
choice of study sites used in the dataset. The dominant error,
Journal of Biogeography 38, 17921806
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Tree line identification from pollen data

particularly in Canada, was where a forest pollen biome
occurred in mapped tundra. Errors would arise if map grid
cells commonly recorded tundra where ground observation
confirmed that forest was dominant. We might also expect a
forest biome assignment when the map was correct (tundra
dominant) but the sampling site itself was surrounded by trees,
as may be the case for a lake in an ecotonal region, or where
trees are sparse but nevertheless producing pollen. Author
descriptions of some sites are available as metadata; a small
proportion of sites have local vegetation descriptions and the
majority have regional vegetation descriptions. We used these
to test the possibilities outlined above.
In Alaska, out of 41 author-described forest-tundra or tree
line sites, 39 are mapped as tundra in the Kaplan & New
(2006) vegetation map, but only eight of these (20%) are
defined as a forest biome by pollen. Here, then, we might
conclude that neither the nature of the ecotonal vegetation nor
any author preference for tree-surrounded lakes appears to
have unduly biased biome calls towards forest.
In Canada, of 18 sites described as forest-tundra by
authors, 14 are mapped as tundra and approximately half of
these 14 are assigned to a forest biome by the pollen. Of 113
sites described as boreal forest and barren by authors, 83 are
mapped as tundra, but of these only six are assigned to a
tundra biome by the pollen. In isolation, this result might be
taken to indicate erroneous mapping, but as this pattern was
also observed with the set of samples that lie north of the
ecotone, a region which is described and mapped as tundra
with a high degree of certainty but where sites are still in many
cases assigned to forest by biomization, we conclude that the
observed bias is more likely to be an issue of regional pollen
representation than due to mapping errors or site choice.
Spatial scaling issues
That the REVEALS model is underlain by the large lake
assumption implies that the use of moss polsters may violate
this assumption and reduce the effectiveness of the correction.
Reducing the modern dataset to only lakes markedly increased
the success rate in Canada (from 67.1% using all sites to 79.1%
using lakes only). In Alaska there was little effect. Bigelow et al.
(2003) examined the success of pollen spectra from moss
polsters versus lakes in predicting the vegetation of the pixel in
which they were located. At this scale of spatial resolution, lake
spectra did better because polster spectra sense the local finescale (subpixel) vegetation mosaic, which may or may not
reflect vegetation as represented by the pixel. Bigelow et al.
(2003) also showed that where local vegetation at the sampling
site was recorded, polster spectra predicted this more successfully than the regional vegetation description. Thus in
homogeneous vegetation (particularly tundra), when local
and regional signals are similar, polster spectra may represent
regional vegetation quite well, and the Alaska result may be
explained by the fact that the majority of the polster samples
derive from tundra regions north of the Brooks Range while
lake samples predominate in other areas.
Journal of Biogeography 38, 17921806
2011 Blackwell Publishing Ltd

CONCLUSIONS
While pollen corrections have been used since the 1960s, they
have typically been used for small-scale vegetation reconstructions. The combination of REVEALS-based correction
and biomization increases the accuracy of broad-scale vegetation reconstructions, which may have relevance for use in
assessing long-term land-cover change and vegetationclimate
interactions. In this study, the most notable improvements in
pollen representation of vegetation are seen across the
relatively flat, broad mosaic landscape of the Canadian
foresttundra ecotone. High pollen productivity appears to
be the critical factor to correct, rather than dispersal. As
expected from theory, pollen data from lakes respond best to
REVEALS-based correction. The method will probably be
most applicable to regions and time periods for which there
is relatively good site coverage, such as the Holocene of
Europe and North America.
ACKNOWLEDGEMENTS
This work was funded by the Natural Environment Research
Councils (NERC) Quantifying and Understanding the Earth
(QUEST) programme (Theme 2), PalaeoQUMP (to M.E.E.
and H.A.B.) and the Estonian Science Foundation Mobilitas
Programme (MTT3) (to S.S.). The work was partially inspired
by the participation of H.A.B., S.S. and M.E.E. in the
POLLANDCAL network (Nordforsk) and LANDCLIM project
and network sponsored by the Swedish Science (VR) and
Nordic (NordForsk) Research Councils, respectively. We thank
Jed Kaplan (Ecole Polytechnique Federale de Lausanne,
Switzerland) for permission to use the vegetation data (shown
in Figs 1 & 2) and to Jack Williams and three anonymous
referees for their comments which improved the manuscript.
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H. A. Binney et al.
SUPPORTING INFORMATION
Additional Supporting Information may be found in the
online version of this article:
Appendix S1 Site names, grid reference and original PAIN
biome assignment for the modern pollen samples from Alaska
and Canada.
Appendix S2 Description of characteristic taxa of biomes
used in this study (Table S1) and comparison of the overall
percentage correct match for pollen biomes versus vegetation
for Canada (Table S2) and Alaska (Table S3).
Appendix S3 Dispersal (C) and initial productivity factors
(a) for taxa contributing to the pollen sum.
As a service to our authors and readers, this journal provides
supporting information supplied by the authors. Such materials are peer-reviewed and may be re-organized for online
delivery, but are not copy-edited or typeset. Technical support
issues arising from supporting information (other than
missing files) should be addressed to the authors.

1806

BIOSKETCH
Heather Binney is a post-doctoral research assistant at the
University of Southampton. She is a palynologist with special
interests in late Quaternary pollenvegetation relationships
and vegetation dynamics since the Last Glacial Maximum.
Author contributions: H.A.B. undertook the analyses and took
primary responsibility for writing the paper; P.W.G. and
J.M.N. provided theoretical input and undertook the data
computations; S.S. provided the theoretical input for the
adaptation of the biomization procedure; M.E.E. had primary
responsibility for the project; she conceived the study and
contributed significantly to the writing.

Editor: Jack Williams

Journal of Biogeography 38, 17921806

2011 Blackwell Publishing Ltd