Vegetation Mediates Soil Temperature and Moisture in Arctic-Alpine

Environments
Author(s): Juha Aalto, Peter C. le Roux and Miska Luoto
Source: Arctic, Antarctic, and Alpine Research, 45(4):429-439.
Published By: Institute of Arctic and Alpine Research (INSTAAR), University of Colorado
https://doi.org/10.1657/1938-4246-45.4.429
URL: http://www.bioone.org/doi/full/10.1657/1938-4246-45.4.429

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Arctic, Antarctic, and Alpine Research, Vol. 45, No. 4, 2013, pp. 429–439

Vegetation Mediates Soil Temperature and Moisture in

Arctic-Alpine Environments

Juha Aalto*† Abstract

Peter C. le Roux* and Soil temperature and moisture are key determinants of abiotic and biotic processes in
arctic-alpine regions. They are important links to understanding complex ecosystem dy-
Miska Luoto* namics under changing climate. The aims of this study were to (1) quantify fine-scale soil
*Department of Geosciences and temperature and soil moisture variation, and (2) assess the influence of vegetation on
Geography, University of Helsinki, PO soil temperature and moisture patterns in a northern European arctic-alpine environment.
box 64, Gustaf Hällströmin katu 2a,
Inclusion of vegetation variables significantly improved models of soil temperature and
00014 Helsinki, Finland
†Corresponding author: moisture, despite abiotic variables (local topography and soil properties) being the most
juha.aalto@helsinki.fi influential predictors. Temperature varied by ⱖ5 ⬚C and moisture by ⱖ50% (volumetric
water content) over very short distances (ⱖ1 m), reflecting the extreme spatial heterogene-
ity of thermal and hydrological conditions in these systems. These results thus highlight
the biotic mediation of changes in abiotic conditions, showing how vegetation can strongly
affect local habitat conditions at fine spatial scales in arctic-alpine environments.

DOI: http://dx.doi.org/10.1657/1938-4246-45.4.429

Introduction and subsurface water flow, and the zonation of some plant commu-
nities (Brubaker and Entekhabi, 1996; Eugster et al., 2000; Ehren-
Soil temperature and soil moisture are key drivers of ecosys- feld et al., 2005; Legates et al., 2010). Vegetation regulates snow
tem functioning (Johnson and Billings, 1962; Bertoldi et al., 2010; distribution, radiation at ground level, and heat flux through evapo-
Cahoon et al., 2012), geomorphological activity (Broll et al., 1999; transpiration (Raich and Tufekcioglu, 2000; Körner, 2003). While
French, 2007; Malanson et al., 2012) and human activities (Post relationships among topography, soil characteristics, soil tempera-
et al., 2009) in arctic-alpine environments. Soil temperature and ture, and moisture are fairly well studied, the biotic impacts of
moisture have fundamental effects on the abiotic and biotic pro- vegetation on these properties are still being explored (see Wun-
cesses determining, for example, microbial activity, biochemical dram et al., 2010; Liancourt et al., 2012). In general, it is known
and carbon cycling, nutrient availability, plant growth and repro- that dense plant canopies may buffer abiotic conditions strongly,
duction, and earth surface processes (Chapin, 1983; Lloyd and Tay- reducing variability in soil temperature and moisture (Legates et
lor, 1994; Hodkinson et al., 1999; French, 2007; Starr et al., 2008; al., 2010; Gornall et al., 2011). By investigating the relationships
Pape et al., 2009; Saito et al., 2009; Legates et al., 2010; Olefeldt et among these three parameter groups, we expect to accurately pre-
al., 2012). While soil conditions strongly affect vegetation patterns, dict soil temperature and moisture across a range of scales.
plants may also have strong feedback effects on soil thermal and Recent studies have shown that soil surface temperatures can
hydrological properties (Ehrenfeld et al., 2005), with, for example, have remarkable fine-scale variation, with temperatures differing
shading from plant canopies reducing thermal extremes and evapo- by several degrees over distances of less than one meter (Scherrer
rative moisture losses (Salisbury and Spomer, 1964; Asbjornsen et and Körner, 2010; Graham et al., 2012; Lenoir et al., 2013). This
al., 2011). For this reason, studies of the impacts of vegetation on magnitude of thermal heterogeneity is ecologically significant as
soil temperature and moisture can provide important insights into it exceeds the amplitude of many climate warming projections,
the response of biotic communities and abiotic systems to changing suggesting that plants will have more potentially suitable habitats
climatic conditions. within regular dispersal distance (Christensen et al., 2007; Lenoir et
Models of soil temperature and moisture in arctic-alpine envi- al., 2008; Scherrer and Körner, 2011). Even though these variables’
ronments at fine spatial scales (resolution ⬃ 1m) need to incorpo- spatial and temporal variation is known to have great importance
rate the complex interplay between local topography, soil condi- to multiple abiotic and biotic systems (Billings and Mooney, 1968;
tions, and vegetation cover, as these three environmental Cahoon et al., 2012), surprisingly few studies have focused on
characteristics strongly affect local thermal and hydrological condi- explaining fine-scale spatial variation in these parameters in high-
tions directly and indirectly (Isard, 1986; Takahashi, 2005; Bertoldi latitude environments (e.g. Wundram et al., 2010; Graham et al.,
et al., 2010; Scherrer and Körner, 2011). For example, topography 2012). This fine-scale heterogeneity, probably driven by local topo-
has indirect effects on soil temperature and moisture by affecting graphical conditions, soil properties, and vegetation characteristics,
snow distribution, incident radiation, and wind exposure (Raupach may exceed coarse-scale (latitudinal and altitudinal gradients)
and Finnigan, 1997; Löffler, 2005; Beniston, 2006; Scherrer and variation over much greater distances (e.g. Billings, 1974).
Körner, 2011). Topographic conditions may also affect soil proper- The biota of arctic-alpine environments is particularly vulner-
ties, with, for example, fine sediments predominating in depres- able to climate change as species in these habitats are frequently
sions (French, 2007). In turn, soil conditions determine sensible highly specialized (Billings and Mooney, 1968; Chapin et al.,
and latent heat exchange between soil and the atmosphere, overland 2000), and the increases in temperatures are predicted to be highest

䉷 2013 Regents of the University of Colorado J. AALTO ET AL. / 429

1523-0430/6 $7.00
in these regions (Anisimov et al., 2007). Thus, an improved under- STUDY SITES AND FIELD DATA
standing of fine-scale variation in soil temperature and moisture Two study sites are located approximately 100–200 m above
patterns, particularly in relation to topographical, soil, and vegeta- the tree limit on the Saana massif, both at an elevation of ca. 700
tion variables in arctic-alpine systems, is needed. This is particu- m a.s.l., but on different aspects (northwest- and southwest-facing
larly relevant as ongoing changes in vegetation cover in response slopes; Fig. 1). Six sampling grids were established at each site,
to environmental change are well documented in this region (Ep- with each grid comprising 160 1 m2 plots in a regular 8 ⳯ 20
stein et al., 2012). Specifically, Sturm et al. (2001), Tape et al. arrangement. Therefore, the fine-scale data set used in this study
(2006), and Kullman (2010) have reported increasing shrub cover
comprises 1920 cells surveyed systematically at one-meter inter-
in the circumpolar Arctic in response to warming. If vegetation is
vals.
a key driver of these systems, changes in land cover can trigger
Both response variables, i.e. soil temperature and moisture
feedback mechanisms, the impacts of which are unclear for the
(Fig. 2, parts A and B), were measured on two consecutive days
global climate system (Chapin et al., 2005; Tarnocai et al., 2009).
(NW site, 16 July 2012; SW site, 17 July 2012). Temperature mea-
Hence, the aim of this study is to (1) quantify fine-scale soil temper-
surements were made at a depth of 10 cm using a handheld digital
ature and soil moisture variation in arctic-alpine environment, and
temperature probe VWR–TD11 (VWR International, Radnor,
(2) assess the influence of vegetation on soil temperature and mois-
Pennsylvania, U.S.A.; accuracy of 0.8 ⬚C). Soil moisture was
ture patterns, after controlling for local topography and soil proper-
recorded through a 0- to 10-cm profile by using a time domain
ties. The study is based on a large field-quantified data set collected
reflectometry sensor (FieldScout TDR 100, Spectrum Technolo-
in alpine tundra in northwestern Finland.
gies, Inc., Plainfield, Illinois, U.S.A.; accuracy of 3.0% volumetric
water content [VWC]). Where possible, multiple soil moisture mea-
Data and Methods surements were taken within every cell, with mean values used in
STUDY AREA subsequent analyses. Where substrate was too rocky or shallow
The study area is located in northwestern Finnish Lapland (⬍10 cm deep) to conduct temperature measurements, linear inter-
(69⬚N, 21⬚E; Fig. 1). The climate of the area is strongly affected polation based on all cells within 2 m was used to estimate tempera-
by its high-latitude location and the proximity of the Arctic Ocean ture (required for 29 cells). Similarly, where soil moisture could
and the Scandes Mountains (Tikkanen, 2005; Aalto et al., 2012). not be measured to a depth of 10 cm (212 cells) or measured
The mean annual temperature at the nearby Kilpisjärvi meteorolog- at all (9 cells), moisture values were interpolated based on the
ical station (1981–2010) is ⳮ2.9 ⬚C (69⬚02′N, 20⬚47′E, 480 m surrounding cells, with observed values (i.e. based on soils shal-
a.s.l.). Mean annual precipitation was 487 mm over the same pe- lower than 10 cm) replaced by the interpolated values where the
riod, with seasonal snow cover persisting until late June and con- latter exceeded the former (ca. 70% of cases; average absolute
straining the length of the growing season (Pirinen et al., 2012). difference between interpolated and observed values 5%). Nonethe-
The treeline in this region is formed by mountain birch (Betula less, repeating analyses with a smaller data set excluding cells
pubescens ssp. czerepanovii), with vegetation above the treeline without measurements gave very similar results (results not shown).
characterized by shrubs, dwarf-shrubs, and graminoids almost ex- Re-measurement of soil temperature in 24 cells in the first study
clusively comprised of perennial species (Ahti et al., 1968). grid surveyed (ca. 12-h interval) showed an average increase of

FIGURE 1. The location of the

study area in northern Fennoscan-
dia. The panel on the right shows the
location of the study sites (empty cir-
cles) on the slopes of Mount Saana,
with 100-m-interval contour lines in-
dicating elevation.

430 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH

FIGURE 2. Examples from the study grid number four (NW slope of Mount Saana; see Fig. 1 for details) showing the spatial variation
of the two response variables (at 1 m2 resolution), (A) soil temperature and (B) soil moisture; and some of the predictor variables (C)
meso-topography, (D) peat depth, and (E) vegetation volume. VWC ⴔ volumetric water content.

0.8 ⬚C by the end of the measurement period. Linear adjustment dictors (each comprising four variables) were measured and/or cal-
for soil temperatures against measuring time was conducted to take culated: Topography (T), Soil characteristics (S), and Vegetation
into account this warming of the soil during the measurement pe- (V). Topography is related to landforms and therefore, for example,
riod. Soil moisture values were not adjusted for measuring time as the radiation and hydrological conditions of the soil surface. The
the conditions in two consecutive days were similar (0.4 mm rain- four predictor variables related to topography were mesotopogra-
fall in previous 48 h), and the moisture content of the soil was not phy, slope angle, potential annual radiation, and elevation. Mesoto-
expected to change rapidly over the time span of measurements pography is a measure of local topography and reflects snow accu-
(ca. 12 h) (see e.g. Penna et al., 2009). mulation, solar radiation interception, and drainage patterns
In addition to the two response variables, three groups of pre- (Billings, 1973). It was scored from one (bottom of depression)

J. AALTO ET AL. / 431

to 10 (ridge top), following the methodology of Bruun et al. (2006) GAMs were fitted using the R statistics package mgcv with maxi-
(Fig. 2, part C). Slope angle, which affects moisture drainage and mum degrees of smoothing restricted to 5 (subsequently optimized
gravitational processes, was calculated from the height difference by the model fitting function; Wood, 2011) and assuming a Gaus-
between highest and lowest elevation points within each cell. Poten- sian error structure. To normalize the distribution of some explana-
tial annual direct radiation (MJ cmⳮ2 aⳮ1; assuming clear sky tory variables (rock cover, biomass, vegetation volume, lichen
conditions) was calculated from latitude, slope angle, and aspect cover, and moss cover) log-transformation was conducted. There
to describe the radiation conditions on the surface (McCune and was no strong multicollinearity among the predictor groups (maxi-
Keon, 2002), and reflects the maximum possible radiation input to mum Rspearman ⳱ ⳮ0.62).
a cell. Elevation (m a.s.l.) was extracted from a fine-scale digital Predictor groups were first tested separately against the re-
elevation model (spatial resolution of 1 m ⳯ 1 m) constructed sponse variables:
from field observations, in order to describe site-specific variation
in local conditions (e.g. meso-scale temperature and distance to GAMtopo ⳱ mesotopography Ⳮ elevation Ⳮ slope
snow accumulation sites). Ⳮ radiation (1)
Soil characteristics are related to the thermal and hydrological GAMsoil ⳱ soil moisture/temperature Ⳮ peat depth
properties of soil. The four predictors were soil temperature (when Ⳮ soil depth Ⳮ rock cover (2)
modeling soil moisture), soil moisture (when modeling soil temper- GAMvege ⳱ biomass Ⳮ vegetation volume
ature), peat depth, and the cover of rock. Peat depth (thickness of Ⳮ lichen cover Ⳮ moss cover (3)
the organic layer) and soil depth (cm; thickness of the mineral
layer; Fig. 2, part D) were determined by means of three measure- Thereafter, the GAMtopo and GAMsoil models were combined
ments inside each cell, using a thin metal rod to probe the soil to obtain the GAMabiotic model (i.e. only abiotic predictors), which
(following the methodology of Rose and Malanson, 2012). Due to was used as a baseline in subsequent analyses. Finally, GAMfull
high moisture-holding capacity, thick peat layers are expected to models, comprising predictors from all three groups (i.e. both abi-
buffer soil moisture and temperature more strongly than mineral otic and vegetation variables), were tested. These models were used
soils, thus describing some of the thermal and hydrological proper- to predict the variation in soil temperature and moisture patterns
ties of the soil. Cover of rock represents the percentage of bare across the study grids. Bootstrapping was used to test the signifi-
rock and coarse gravel within each cell with potentially positive cance of model improvement after the inclusion of additional pre-
effects on temperatures (thermal properties) and negative impacts dictor groups (i.e. explanatory power; 1000 repeats; R–package
on soil moistures (porosity). boot). Similarly, the models’ ability to predict soil temperature and
The impacts of vegetation on soil temperature and moisture moisture was assessed using cross validation with a semi-indepen-
are likely dominated by the effects of plants on incident radiation
dent data set (i.e. predictive power; 1000 repeats; 70% random
at ground level and hydrological conditions. The four predictors
sample).
in this group were vegetation volume, biomass, cover of moss, and
Spatial autocorrelation is a common property in fine-scale data
cover of lichen. Vegetation volume (Fig. 2, part E) was calculated
sets, which may cause uncertainty to model estimates’ significance
as vegetation cover (in mⳮ2 ) multiplied by median vegetation
testing and confidence levels (Legendre et al., 2002). Since our
height. Dry above-ground biomass from each plot was determined
analyses were not focused on the significance of individual param-
at the peak of growing season following the procedures described
eters, we did not account for spatial autocorrelation; nonetheless,
in Walker et al. (2003), using one randomly located 20 cm ⳯ 20
model residuals showed a marked reduction in spatial autocorrela-
cm clip harvest plot per cell. The two cover variables (moss and
tion relative to the raw data (Appendix Fig. A1).
lichen) describe some of thermal and hydrological properties of
Variation partitioning (VP) was used to determine the relative
the surface (Addison and Bliss, 1980; Gornall et al., 2011). Lichen
importance of different predictor groups (Borcard et al., 1992).
cover has effects on soil temperature, for example, by altering the
Variation in soil temperature and moisture was decomposed among
albedo of the soil surface (Stoy et al., 2012). Vegetation data were
the three groups of predictors, using a series of regression analyses
collected in July 2011 and July 2012 during the peak of the growing
implemented with generalized linear models (GLM), assuming a
season. Differences in canopy heights between vascular plants and
Gaussian error distribution (McCullagh and Nelder, 1989; Borcard
cryptogams may differentially affect abiotic conditions, as the mul-
tiple effects of the four vegetation variables on the soil temperature et al., 1992). Both linear and quadratic terms were included to
and moisture are thought to derive from the shading (lowering account for potential non-linear relationships (Heikkinen et al.,
temperatures), moisture capturing (increase in moisture, decrease 2004). The GLMs were fitted using the glm function in R with
in temperature), snow accumulation (increasing moisture), and automatic backward stepwise term selection procedure (based on
transpiration (lowering both temperatures and moisture) (Ehrenfeld Akaikes’ Information Criteria, using the stepAIC function for the
et al., 2005; Blok et al., 2010; Wundram et al., 2010). first three separate predictor models) (Akaike, 1974; Zimmermann
et al., 2007). When combining the best-fit models for the three
groups of predictors for further partitioning, no variable selection
DATA ANALYSIS technique was used.
We modeled the relationship between the two responses (soil VP partitioned variation into eight fractions (calculated fol-
temperature and moisture) and different predictor groups (Topog- lowing the procedures in Anderson and Gribble, 1998): (a) the
raphy, Soil characteristics, and Vegetation) using generalized addi- percentage of the total variation in soil temperature (or moisture)
tive models (GAM; Hastie and Tibshirani, 1990; Wood, 2006). that is explained by T but not by S or V; (b) the percentage of the

432 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH

total variation in responses that is explained by S but not by T or variables alone explaining 21.1%) (Fig. 5, part A). The bootstrap-
V; (c) the percentage of the total variation in responses that is ping result clearly demonstrates how the addition of vegetation
explained by V but not by T or S; (d) the percentage of the total predictors to baseline models (i.e. with abiotic variables only) sig-
variation in responses that is explained by T and/or S, but which nificantly improved the performance (Fig. 5, part A) and predictive
cannot be allocated between the two predictor groups (also referred power (Fig. 5, part B; Appendix Fig. A3) of the models.
to as joint effects); (e) the percentage of the total variation in For both soil temperature and moisture, the large majority
responses that is explained by T and/or V; ( f ) the percentage of of explained variation was accounted for by soil characteristics,
the total variation in responses that is explained by S and/or V; (g) topography, and their joint contribution (Fig. 6). In both sets of
the percentage of the total variation in responses that is explained analyses, vegetation properties had the smallest unique contribution
by any of the three groups of explanatory variables (i.e. cannot be (soil temperature: 4.2%; soil moisture: 6.3%).
partitioned among the three predictor groups); and (h) unexplained
variation.
Discussion
Incorporating vegetation characteristics into soil temperature
Results and moisture models significantly improved their fine-scale predic-
Both soil temperature and moisture varied notably over short tions. This suggests that vegetation has a clear role mediating soil
distances. Soil temperatures in the NW study site (Fig. 1) varied temperature and moisture patterns in arctic-alpine systems. For both
from 2.4 to 13.7 ⬚C and in the SW site from 5.3 to 13.1 ⬚C (Appen- variables it is evident that soil characteristics have the dominant
dix Table A1), respectively. Similarly, the soil moisture values effect, with topography clearly being the second most important
varied from 4.6 to 90.3% VWC and from 8.1 to 65.2% VWC, group of predictors. Additionally we demonstrate notable spatial
respectively. The largest within-grid variation for soil temperature variation in soil temperatures and moistures at the spatial scale of
was 8.1 ⬚C and for soil moisture 71.8% VWC (Fig. 3). Soil tem- 1 m2, in agreement with Wundram et al. (2010) and Scherrer and
perature and moisture were significantly negatively correlated Körner (2011).
(RSpearman ⳱ ⳮ0.6, p ⱕ 0.001) (Appendix Fig. A2). The improvement in model performance after the inclusion
All the predictor groups were significantly related to soil tem- of vegetation variables is greater for soil moisture than for tempera-
perature and moisture (examples of bivariate relationships illus- ture, suggesting that vegetation has a stronger direct impact on
trated in Fig. 4). The baseline model (i.e. topography Ⳮ soil charac- moisture patterns than temperature. Vegetation may affect soil tem-
teristics) explained 67.8% of the variation in soil temperature. perature and moisture through a variety of mechanisms (Cahoon
Adding vegetation predictor variables to the abiotic model signifi- et al., 2012; Graham et al., 2012). For soil temperatures, vegetation
cantly improved the proportion of variance explained to 71.8% cover operates as an insulator, leveling out temperature variations.
(with the vegetation variables alone explaining 30.5% of the varia- Vegetation also acts through shading, decreasing the amount of
tion in soil temperature) (Fig. 5, part A). For soil moisture the direct solar radiation reaching the ground surface (Pielke, 2001;
inclusion of vegetation variables also significantly improved the Blok et al., 2010) and subsequently lowering and buffering soil
models’ performance; the baseline model explained 55.5% and full temperatures (Salisbury and Spomer, 1964; Graham et al., 2012).
model 64.2% of the variation in soil moisture (with the vegetation For soil moisture, vegetation generally increases the soils’ moisture

FIGURE 3. The maximum differences in soil temper-

ature and soil moisture as a function of distance be-
tween cells.

J. AALTO ET AL. / 433

FIGURE 4. Relationships between the two response variables (soil temperature and moisture) and explanatory variables from every
predictor group (Topography, Soil characteristics, and Vegetation), as modeled using generalized additive models (see text for details). Meso-
topography ranged from 1 (bottom of depression) to 10 (ridge top).

content as it mediates the soil water fluctuations by reducing the sheltered valley bottoms determines the wind and radiation condi-
evaporative losses through shading and holding both horizontal tions subsequently controlling the presence, thickness, and duration
and vertical water flow in the soil (Daly and Porporato, 2005; of the snow cover with multiple effects on two parameters studied
Asbjornsen et al., 2011). On the contrary, abundant vegetation can (Löffler, 2005; Litaor et al., 2008; Penna et al., 2009). The tempera-
promote soil drying through intensified transpiration (Horton and ture relationships with soil characteristics are mainly related to
Hart, 1998). different thermal properties of the soils (Graham et al., 2012). Soil
Our results show that soil temperature and moisture are mainly moisture is predominantly a function of water-holding capacity
controlled by local topography and soil characteristics (in agree- (related to soil texture and porosity; Legates et al., 2010) and there-
ment with e.g. Isard, 1986; Wundram et al., 2010; Scherrer and fore highest soil moisture contents are found in soils with high
Körner, 2011). The toposequence from wind-blown ridge tops to organic content (e.g. peat lands). In agreement with Wundram et

FIGURE 5. Bootstrapped estimates of (A) model performance (adjusted R2 ) and (B) predictive power (adjusted R2 ) for the soil temperature
and soil moisture generalized additive models based on 1000 samples. All models differed significantly ( p ⱕ 0.001).

434 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH

 FIGURE 6. The results of the varia-
tion partitioning for (A) soil temper-
ature and (B) soil moisture in terms
of the proportion/fraction of varia-
tion explained. The variation of the
response variables is explained by
three groups of predictor variables:
Topography (T), Soil characteristics
(S), and Vegetation (V); a, b, and c
are unique effects of T, S, and V, re-
spectively; d, e, f, and g represent
their joint effects.

al. (2010), we argue that the soil temperature (and soil moisture) potential habitat shifts or changes in frost-related processes under
is even more strongly driven by soil properties than local topo- warming climate, it is important to appreciate the notable fine-
graphical variables. They made an important finding, however, that scale variation and complex interplay of these variables caused by
high soil moisture content evens out the extreme temperature varia- numerous environmental factors and not to rely only on coarse
tions in soil due to wet soils’ thermal conductivity and the high resolution climate models based on mean air temperature data.
heat capacity of water. Soil moisture correlates negatively with soil
temperatures as the increase in soil temperatures increases evapora-
tion and in turn lowers the moisture content of the soils. On the Conclusions
contrary, increased moisture intensifies evapotranspiration, which Our results indicate that vegetation has an important role in
in turn lowers soil temperature (Legates et al., 2010). mediating soil temperatures and moisture variation at fine spatial
Climate warming will affect multiple feedback loops related scale in the arctic-alpine system. Thus, when modeling soil temper-
to soil temperatures and moisture. For example, the prolonged ature and moisture, vegetation properties need to be explicitly con-
snow-free period in the Arctic has already caused shifts in vegeta- sidered. Extreme variation in temperature and moisture was ob-
tion cover and composition (Stow et al., 2004; Chapin et al., 2005; served over short distances, reflecting the strong spatial
Tape et al., 2006; Kullman, 2010; Epstein et al., 2012). The expan- heterogeneity of thermal and hydrological conditions in these sys-
sion of shrub-dominated vegetation into areas previously domi- tems. As the ongoing changes in pan-Arctic tundra vegetation are
nated by dwarf-shrubs and graminoids may have particularly pro- highly dependent on both soil temperature and soil moisture, the
nounced consequences for ecosystem dynamics due to the potential understanding of these patterns is crucial for fine-scale climate
for species of this growth form to affect soil moisture and tempera- change impact studies.
ture conditions (Sturm et al., 2001; Bonfils et al., 2012). The green-
ing of the arctic-alpine regions has notable effects on surface energy
budget as the reduced albedo in both winter and summer causes Acknowledgments
more radiation to be absorbed and turned into sensible heat flux, We thank A. Niskanen, S. Suvanto, H. Mod, S. Jääskeläinen,
subsequently promoting local warming (Chapin et al., 2005; Sturm and A. Kulonen for helping collect the data. We also thank Dr.
et al., 2005). Arctic areas are major sources of organic C in the Christophe Randin and one anonymous reviewer for their construc-
soil, and the potential increase in soil temperatures will accelerate tive comments and suggestions. Aalto was funded by the Geog-
the rate of C cycling in ecosystem as it intensifies, for example, the raphy Graduate School of the Academy of Finland, a University
soil respiration rate with possibility for major global implications of Helsinki grant, and a Societas pro Fauna et Flora Fennica grant.
(Lloyd and Taylor, 1994; Raich and Tufekcioglu, 2000; Hiltbrunner Le Roux and Luoto were funded by the Academy of Finland
et al., 2012; Olefeldt et al., 2012). (Project Number 1140873).
Therefore, using field-quantified biotic and abiotic data fine-
scale variability in soil temperatures and moistures could be accu-
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 APPENDIX

FIGURE A1. Correlograms for raw data and

residuals of two generalized additive model
(GAM) specifications (GAMabiotic and GAMfull )
indicating presence or absence of spatial auto-
correlation in the terms of Moran’s I. (A) Soil
temperature, (B) soil moisture. Statistical
significance is presented with filled boxes ( p
ⱕ 0.05) and empty circles (not significant).

TABLE A1
The means and standard deviations for all of the measured variables. The significance of the differences between the study sites (see Fig.
1 for details) was tested using Mann-Whitney U-tests and indicated as: *** p ⱕ 0.001; ** p ⱕ 0.01; n.s. ⴔ not significant.

Category Variable Unit NW site SW site

Topography Mesotopography Index 5.0 Ⳳ 2.1 ** 5.2 Ⳳ 1.8
Elevation m a.s.l. 701 Ⳳ 7.3 *** 729 Ⳳ 15.5
Slope Radians 0.39 Ⳳ 0.2 *** 0.60 Ⳳ 0.2
Radiation MJ/cm2/a 0.26 Ⳳ 0.1 *** 0.72 Ⳳ 0.1
Soil characteristics Soil temperature ⬚C 8.5 Ⳳ 1.6 *** 8.9 Ⳳ 1.7
Soil moisture % VWC 31.3 Ⳳ 14.8 n.s. 28.7 Ⳳ 9.1
Peat depth cm 4.4 Ⳳ 3.4 *** 7.1 Ⳳ 5.2
Rock cover %/m2 9.2 Ⳳ 13.2 *** 20.6 Ⳳ 24.2
Vegetation Biomass g 9.9 Ⳳ 10.5 *** 15.6 Ⳳ 16.1
Vegetation volume m3 0.013 Ⳳ 0.015 *** 0.031 Ⳳ 0.029
Lichen cover %/m2 5.5 Ⳳ 9.2 *** 1.2 Ⳳ 1.9
Moss cover %/m2 23.0 Ⳳ 17.5 *** 6.7 Ⳳ 7.3

FIGURE A2. The relationship between soil temperature and soil

moisture based on bivariate GAM modeling. VWC ⴔ volumetric
water content.

438 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH

FIGURE A3. Observed and predicted patterns of (A) soil temperature (ⴗC) and (B) soil moisture (% VWC) inside the study grid number
four (NW slope of Mount Saana; see Fig. 1 for details). The predictions are based on two GAM specifications (GAMabiotic and GAMfull )
with semi-independent calibration set (i.e. no observations from grid four was used for predictions). Spearman’s correlation coefficients
(Rs ) between observed and predicted values are also presented.

J. AALTO ET AL. / 439